Brigham Young University BYU ScholarsArchive

Theses and Dissertations

1951-06-01

Galls and gall insects of Artemisia tridentata Nutt

Solomon Lander Brigham Young University - Provo

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BYU ScholarsArchive Citation Lander, Solomon, "Galls and gall insects of Artemisia tridentata Nutt" (1951). Theses and Dissertations. 7805. https://scholarsarchive.byu.edu/etd/7805

This Thesis is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Galls anj Gall Insects

of

A.rtEtisia tridentata l'utt.

A Thesis

�;ubmited to the

Depart;;ient of Zoology and Entomology

of

lrigham Young University in Partial Fulfill.'.ent of the Hequire,lients

for the deo::ee

of

Laster of Art,s

by

Jolo;:;::on La:rider

,;,me 1951 ':'he assistance and encourare:,,~nt, fro:i, others has eased the writer's task in prr;,par.-inp, thls -;:,aper, a.n.d he ther-efore ·wishes to ackno;.;ledite the heln of the following:

Dr. "'.rasco • rar:ner, Depart1;;ent of' Zoolor,y and ';~ntomoloi:ry, under whoae direction this work was done, and whose .,:uidance and inspi:rat.ion lightened tho burden; Dr. ~iertrantl I<. Harrison, Department of ..iotany, who sui 0 ,;ested referfmces for the 0otanicnl aspncts, and er.coura,~ed the writer at all tir;,es; ;;r. t,. Lyr~n Har>'lard, Departu,ent of 1.:ooloty

nr. 1\o 1.:arl '·'ritchard, Goller::e or Ar,:riculture, University of California.

(>,er.'teley), for identificati.on of tne Uonididae; r. G. F. ~:. 1.ueaebeck and: r. A. :3. ;:;ahan of th<~ u. ,1. rati:.i11al Luseu.1t, for idenLi.fication of the t;halc:l.Joidea and ::-:cGlionidae; Ir. Lewis i:. ·,,eld, for supplying na:1,es of ;:,eople 1n the .field and ::-eferences, and for sending six types of galls of Artemisia tridentata, tli.!'!.t Dr. ::. P. Felt had collect,ed; Dr.

ey, Institute For ')e:x. itesca.rch, llniversity of Jr:dirma, for me

he:r collection o.f m,:iterials, data and aid in d.issection of the ;:alls; friends an:J fellow stuchmts for tJrirn1::i.ng in 'alls at . .:very opport1mity.

iii p, age TJTLE • • • • • . . . . • • • • ... •· . • • • • • • • • i • • • • • • • • • • • • • • • • ii • • • • • • • • • • • • • • • • • • • • iii • • • • • • • • • • • • • • • • • • • • iv I: TH.ODUCTION• • • • • • • • • • • • • • ...... , . l • • • • • • • 3 Nf,;THOJ)S . .. . • • • • • • • • • • • • • • • • • • • • 6 • • • . . . • • • • 0 • • • • • 8 • • • • • • • • • • • 10

• • • • • • • • • • • • • • 13

K'JX 1'0 GALLS • • • • • • • • • • • • • • • • • • • • 0 • 16

ful'lliform, Jla';,;rous, ,,r-own, SteL'. .itll • • •• • • • • lS !Lt:~elrmt-sb,pe:l, !~all • • • • • • • • • • ••• • 20 Oval, R.ufous, Leaf Gall • • • • • • • • • • • • • • • 22 Globose, Jlad.Jery., Puffy Leaf (}all • • • • • • • • • 24 GloLiulsr, 1Jlabrous, Urown, .Stem Gall •••••• • • 26 1'0,:->-shnped, '/Joolly, Whitish-brown, ;;t,13n, Gall • • • • 29 'top-shaped, :,colly, ·_,rhite, Stem Gall •••••• • • 32 Jlobular, Fibrous, Yellowish, Spotty, Stem :}all • • • 34 Globular, F'ibrous, ¥'1bite, Spotty, '.1tem :}all • • • • • 36 Globular, 0ark Purple, Sten Gall ••• • •• • • • • 38 Diarthronomyia sp. F'rom Unknown Gall • • • • • • • • 40

• • • • • • • • • • • • • • 42

SUL;AHY • • • • • • • • • • • • • • • • • • • • • • • • • 46

Llr~~HATIJ'i)~ CIT En • • • • • • • • • • • • • • • • • • • • 47

iv t,re.

l.

•

l.-

•

l 2

'"al.ls. WhAt the ~n--iter nas done is to reflect on the de,,:rees of cor:1- olexity And specia.li:u1.ti.on an,1 points of of the f;a.lls collected in light, of th~ conclusions of (!~i.., l s (1()''1.')r:. and i\insey (1923). /\lthow~h ,,·:alls and gall innects have !Jeer: associated with IJlar:ts

since time :i1:ii'r,er;;orial1 productive scientific obl'H~rvation first jeveloped in the ln.t,e lE~oo•s. '~ven in these ::.odern L.imes one finds few and greatly s~at t.ered refer(mces 0n t,hj,, subject in compa.rison with other plr.rnt-in- sect rolatiorn:;hips.

At the outset ·,dtchcra.ft was ur-oposed to 1:1.f!COunt i'or t,he origin of ;·:alls (':o:1ena 1 1913), cut later Cecidolo::y (the study of 1:~alls Md gull in.sects) developed into a true science. the early workers responstblA !.'or this :l.evelopment tt,e narnP.s of ;;i.ley (l:>13), Osten ~'.ac.K- en (t. s. :.;eol. Sur·. cf !'err., lt~F'?), :jeut,e:·1:,mll•:;r (l'tOh), a1;d Cook (190t,;.) st.a,:d out; Lhe first Uiree for dtrnc:d Jtions of ralls and i;~all insects I and tbfJ la:-,ter for his ~xcelhmt work on discussei proof of the fact Lhat the ovi:oosition of tne 1~yr1inid .fern.ale was not t,·'.",

beyerniek (lE??) \'l!.'l.S ,'.imong thf~ first to sug,r,est di vi.ding galls into indefinite A.nd :!efinite groups. {,uster (1911) .~onsidered the in- def1nit.e ,ieveloptilent to be a pri,,d tive comiit :ton as o;)poseJ. t.o the def- inite type which he vinweri as advarn!ed. 1\e used the terfos 11kataplasrnas" and 1prosQnl~maas' 1 respectively to desc.ribe tl1e r;;.anr,t:ir of gall for·ma.tion ar:d in Jefininr and delim1 tine; t•;e use of these terms furthered tlie under-

3 standing of i1ow r,alls develoo and of the reLi.tionship of the t,3.11-n:.akers.

He defined the t,~r:ns ,"ls follows:

Kataolasn;as - those ::l.miefinite galls whose stl'.'Ueture is dev~looed throut;h hy;:'lerpla.sia of emoryonic tissue,, the end '.,iroduct not i:.H?.CO,:l- :ir:p; in its diI'fnrcnt.Lat.ion, orient,ation an:i for:ri of tissues, fml'ia- mentally :iif'ferect frca .. the nor;.Ml-1 ,il"int p.:1.rt; they represent inhi- b 1tlon cf the normal dif!'erenti.ation, thf, ::,ore ,,,Jvancf:d ones thus arnroacr:ing nomo,c.;er;ity. i":·osonlQ.s:::.tin - definlte r::alls whose struct,,1re fers fun:.la,ller;tally· froni the norr:;.":tl pl;'.l.nt tX,rt, the tissue itt their form :m,i orientation chc1racters conatitutin,-: an a.::-':reeat,ion of now qualities.

lationshi ps nffiCnF ,'.,alls.

The work of Houar:i (1922) on i~alls or Africa, 1\sia, the l'•,edi- ter:ranean area, and South and ~ent..ral America was an important contri- bution to '.Jecidoloi,r,y-.

r:osena (1913) contributed some important concepts when hf? pre- sented the view that ar.tong the Cynipid ,,alls the reason for- Lyoert roohy of the plwt part attack£,id lies in the fact that extra nourinh:nent is pro- vided f'or the plant tin sue in :.he forn; of sur;~r that.; was ch:u1rsd. from starch by lArval enzymatic action.

In more recant tis;;cs such r:reat workers as Felt ( , 1918, 1925,

1940), 1:eld (1926), and Kinsey (1925) h:iv~ 1_~-reatly increased our knowledge on t,he 1,;echanisi:,s of t11e :1la.nt and the activit:r of i:.he insect i:1 t:all ,:iro- duct::.on. l':insey (193,S) developed tho study of the 'Jynipidae to its high-

?r,t ;:;:,int. Felt (191 , 19'.?5) dealt with Ced.·lology frorr; a broader- vinw

The studies of ':;alls of t be 5

CockE>rell (1909) described a ga11-e:nat on f•!"ter:iisia. Felt (191+0) in his book (oJ,. cit.) sum carized 1::.ost, of the 1"!?.rlier work., in adiition to his coll,,.ctimH3 ·,:hich i-;ere :.:B ie in the interi:,ountain ;,.;est. To the wri-

field. ,;,earirir t.lle insects from the talls proved t.o be a difficult task.

()f the thrf:!e ;nethods em:::loyed t.he vial method described below, though the si,;1:)lest, was most productive. ':ihcre tu,e i~ :i factor the br'.:?eding box method. should not be followed. The bell-type jar :r.ethod is ideal, since it clom,,ly ;:;i.mulates the natural environment.

'i'hrec nethods for rearing the insects from the galls were employ- ed, as foll0ws:

1. Hre":ldin,;:s: r;1ethod

A portion of the ster,1 and lP.e ·es, with ,:;all or ,,alls att.l¼ched, was ,)laced i.'1 a v5r,.i l1~,lf fjll~xi with ,vatar. Great care i,,;;;u3exercised to ::i:-1:td,ain a r;ioist condition for the i,ase ,.,ft,he stefa. Gotton was stuffed around tl,r:l sterH fillinq the ,,,outh of the vial, tne ga.Ll or ealls situated above the cotton, so that Lhe insects wm:-e uncible to enter the vial. The vial was then pll'l.ced in a cigar c.ox, from which a test tube protruded, its n.outh opening inside the oox. iubber bands were then snugly fitted around the box. The insects when ei::,er;dng from t.he galls \<1ere attractPd by the light shi.ling t hrou~h the test tube, and flew into it. The insects were then placed in cyanide jars. llalf of those insects from a single type gall w:1s prEH3fH•ved in iseventy uer cent alcohol, while~ the other half was pl

6 ? 2. Y.:!.e!:net.hod

1'hc stem and ln:'.'l.VAs with gall or ralls attached were placed in a vitl. l\ wad of 0noist cotton W'lS inserted ir; the vial and w-as

't'his conrlisted of a ±'in,:er bowl three-quarters filled with !':,oist

S!ind. '!'he st!~ms of the plant with ::all:, attached were ol.1:1cedin the sand and tnen a bell-typA ,jar c:ovor W,3.s :.1lacPd over the f'iPger bowl and ;;:;alls.

The top of t,he jar was flat with the ;lans cut away, and covered wl th a vtl'rv f.ine gau-ze. mnor{l:i.nr; insects eould not escape through this

1ra.uze. 'l'hey were killed and tr~a.ted as ;,,entioned above.

Slides of Lbe ltonid:i..uae 'tiere prepared by Dr. Pritchard., wh"ile slides o.f the parasites on the <1all-makers were p1 ,pared Ly U1e writer.

The wings of the lat.ter grouµ were rer.ioved from tiried soecimens and

directly in Ganada balsarr,, while the antennae were boiled in hot caustic potash., cleared in 3.lcobol, and ii,ounted iri t,alsau,.

i' :ication of

s and :,,A.le ,,.slia ·,cere

on of lOOX. T'he

) procedu r-e. 'l'he r::aterial that fo.rms tht> basis of this ;)aper was collfcteJ in fJt;:i.h County. Gert::dH a:re!l!'i were coll·~cted .,,o:re extensively than others

collection ar~ao arc descrited below.

'.:mryon - two ,:1Hes n0rtheast of Pr·ovo; ;'l.ltitu:Je 5 feet;

by oa.k brush.

Foothills of ••yn ,,onntain - one ,d.le easL of ?rovo; altitude 1/!DO ft~et; thick stl'l!1d or saH::l:rush asso:!iated with :~ror:i'.ls tectorun~ L.

~ak~, ~.Ue - twelve ::d.les west cf '.'rovo; altitude 430G feet; large stand of bi.;~ s:u;ebrush presEmt with T's.marix f/J.llic:ila growing close to the lake.

l\spen (}rove - T,,ie:ity miles northeast of Provo; altitude -- ' sLand of sagebrush bordering road, edeed with 01-lk brush.

Canyon :aeadow - .";ituat"d in e

IJ:.~ - five niileA northeast of ?rovo; iltitu:ie I.. feet; 9 i:obGle Greek - nine ·,iiles southeast of Provo; altitude 5000 f'e(';t,; big saf;ebrush present atrnociated with A;(ro:i;)!ron sp.

Strawberry Jlesorvoir - twenty-five n:dles southeast of Provo; 111.ti- tu:ie 75'.)0 feet; sa/ebrush growinf in associa,tion with ;:iromus sp. m!.i.tely one to six feet. :Xce:.ptiona.l stands as higH as ter, feet are oc- casiona.lly found. lt is ,-1 deep:l.y-roote:i, widE~l:r-:;,r.ancning plant, 1tdth

+· ..+- ·, I ,.r:i.cnorairn,,, . ' ~!l'l;flf,f=.l.Y' .., c ,... o.., t h. in:~ t .ne 1 eaves ( .1e:,,,E)L,,') . ' ..1..)•:i37 .,, \ g'.l.Ve' ....co1e plarJ, 1 ts s::i.lver·y- ·rey appea:·ance. ''i.'he lcavt':s are ty:..1icall.y Lhree-toothed

yellow p.!l.ni- culate flownrs ( ::rd.berg, 1917), proiuced in ,,i,t,und.ance, t:loo,n ir1 the late

for :.ev,mty-f:l.ve to ono hurnlred yea.rs.

Distribution ---and ~?.,uH,e rrter:d.idn .:!:_ddantata is or;e of the moflt widely distributed shruos in the T.lnitad Sta.tt1s. According to :Jiettert (1937) the range exbmds as far north as .::iritish .;olu::ibia, south to Arizona and ;·'ew i,exico, with the

10 11 wc:-.:Lnr-n "Jakot.'1.s and the Pacific CoaRt for.111in? i.ts east-west uoundaries.

The plrmt, is ,::oirnuonover ;":uch of t:tti.h, es('.)edally Jn the low foothills of t,he 1<1estern half, which lies in thE1 ,·; r.:,at :,a sin. ln fact this plant is one oft

hi'oitP.d by any fori1ation west of the prairies and plains.

well-:irained. f res. of appre-

•

~, the µreference

uvrcnlf:i _,_.,___ spic,."rnr· (, 1urnh.1'· 'icr:\.,Jn, an(i :::~ru.lth, and COlHfoonly with ,auch other 12 dart and Srr,ith, l'JJiJ).

i"a.xono,uy

'\rtc.:;iisia tridentFi.t.a is part, of a com9..i.~1xof closf1ly related For the purpose of clarity all ,~all descriptions follow the same pattern. CharMters described include exterior construction, color, ef- fects of age, position of gall and r,,anner of at tachwent on pl~t, kata.- plasrr.ic or pro£1oplasmic type of development, sinrle or clustered growth, measurements, interior construction, and !)osit:i.ot1 of hole from which the insect emer{':es•

Of the ten galls collected, seven were produced by members of the

Itonidida.e (Cecidomyiidae) or gall midges. This group of small, delicate flies, which rarely exceeds one quarter of an inch in length, and are usually much s:r:1aller (F~lt, 1740), 'ire highly specialized in gall-ruaki:ng.,

They are 1r.a.tched., if not exceeded, in numbers and in diversity of c'.all- production only by the Gynipi,:if,.e. T:1ei r wings are usnif\lly :1airy, and have retained only three or four of the Principal veins. I'he writer has in some cases (Pltite I\/, figs. 2, 4; plate V., fi 6• 2; pl;:,,te Vl, fig. 2; plate VIII, fig. :.?) ccmp'lrod t.hc venation and structure of the wings. 'rhe antenr.n.e of this group art1 :remarkable for t:::.tlir- divers:Uy of structure.

Thread-like., hoa,ogeneous, chitinous structures nn.med circuu:bilia are present on ~a.ch ant~nnal ae2~~:€!nt. Tt1ese circm1;filia. are distinc-'.:-i ve in their arrltn;:re,nont f'or e,3,ch species. No analysis of these ;;t,ructures has been m.ade, rut some drawfo;::s (Plate IV, fig. 3; plate V, fig. 4; plate IX,

!'iit:. 2, plate x, fig. 2) of the ant~nnn.1 segments have be,:m included. 13 14

I\ weevil, Apion sordidum Smith (Fall, H~9P.) was the gall-rnak\"r of the oval, rufous, le~.f r:all ( Plate III, fig. lA). Of t.he Curculionidae a. number of members of the ,c:l'?nus l'qion have ueen associated with fills.

Two meJ::'vers of the Tr-y;,.,etidae were included witr, the gall-iliakers.

However, one was not reared f'rom the ralls, but other evidence indicates

or ·trypetid,':l,e.

\';here further :iete:rrr,ination was it1possiblcJ only the ,'.enus of the insect is .bren. ln one C,ise, only thf'.! pa:r;i.sites on the eail-mclker were reared, The writer is aw-R-:reof the iesirability of' obtaining the adult gall-rr.~KfH' beforo descricim'. the gall., llut the presence of a record of the parasites on the gall-maker, plus the ruling 1 of the committee on nomenclature in the 1909 1::iecting of the :,:ntomologica.l Society of America

( Committee, 1909), induces him. t.o incltrie this description.

Cecidology is n study which ma.kes use of a ,Hstinctive vocabulary.

':'he writer therefore, includes a glossary for the sa..

Glossary

Cecidomyia. or ltonid - feneral tHrms applied to any species r~.ferable to the ,gall midvea or I tonid Liae.

t::ynipid - a e~~ner,,.l terni ~ppliert to any species referable t.o the

lJn.. tlrJ.e.1.,. . ,,, +,:.ne ruling states that if the 6a.ll-maker has not ueen reared the i!a.11 :ll'l.Y be nar'l

Inseparable - a_ tern: apolied to galls th!lt are so closely united with

t ls sues, dif-

•

md orientation

;17'1_ • la. :1te;:~, gall

2a. :~e.;:>arable

3a. Posit:i.on on stem terminal or lateral; fibrous and/or leafy ( 1.mdi vid ed r;,arzins) , woolly, or g;labrous.

4a. Fibrous, and/or leafy, or woolly; variously colored. 5a.. Fibrous and/or leafy; variously colored; occurring singly or in loose clusters.

6a. Gonsists :mainly of fibers; a few leaves 11.i.a.y be interapersed among the fibers; color yellowish or whitish.

7a. Whitish in color; diameter 7-11 mm; length 7-11 mu,••• (36) Globular, fibrous, !hfu, spott¥, s;all Diarthroncu,yia s p.

?b. Yellowish in color; dia ..,eter 10-15 ima.; length 10-14 mw.. • • (34 ) Globular, fibrous, yellowish, snott1, gall

6b. Consists mainly of' leaves; may have a, few fibers :i.nterspersed ~wn;;t, the laaves; color purple • • • • • • • • • • • (38) Globular, dark pur;ele., i~all Diarthronomyia sp.

5b. :Joolly, whitish or brownish; usually in clusters of three.

11-.1\.11colors :iven a:::ply at he.ilht of e~rowth of t,he itall. Golor ct,rmges with age are given in the descriptions. 1'.he number preceding the nacie of the rc)J.l irn.lic::ites the page in th:is paper on which a description of the ga.11 a:.:,pears.

16 17

6a. ,;;xtrer:iely white; rat.her rough-woolly Hppearance • • • • • • • • • (32) l'.2£-sh::rned, woolly, gall Rhopalonwia sp.

6b. ,;hitish-brown; rather Vf.llvety in a;:pearance •• • • • • • • • • (29) ~-shaDed, woolly, whitish- orown r;all Diarthrunom~ria, n. sp.

1.. b. Glabrous, brown • • • • (26) G1ooular, §lai..,rous, brown, ~f".':J,11 ~l;_arthronomyla, r.. sp.

3b. Posit.ion on tAr'.·iinal, replacing a bud • • • • • • • • • • • . . . . • • • (20) Viazelnut-shaoed, r;all . . Zutreta diana. u. S.

2b. lnseµarable • • • • • • • • • • • (18) Fusiform, gla.brous 1 ~, stem gall TrJpetid lb. Leaf gall

2a.. 'tufous I grow singly or in twos on OJJposite sides of leaf •••• • • • • • • • • • • • • • • • • • (22 ) ~, rufous, !£!!. i3all Aeion sordi:iurn Smith

2b. Greenish-white to rose, often in clusters of 2-t'; generally larger tha.n oval, rufous, leaf gall • • • • • • • • • • • • • . . . . . • • • • • ••• (24) .;.lobose, ~laddary 1 2':!f~;y:, . . . . le~f r;a.11 Dia rth ron?11~ia. arte1i.1s1ae Felt ~-Fusiform, :}labrous, ~, 9.!!!,

This gall (Fla.ta I, fig. l) is a lighter L-"'ownth.an the surround- ing stem, and ls C;,.'i:,remely hard. i::ith age it assumen a. deeper brown color, the extedor rei:iaintnr:; hard, while the interior tissue becou,es brittle. Occasionn.lly a. braEch grows ouc fror:1 the side of the gall. It is a swelling of the stem proper, and therefore inseparable. Development of tlle gall is of a kat.aplasmic t.ype. ':'he gall ra11e::es from. 9 to 11 r,;.m. in diameter, and from 15 to 18 mr,,. in length. Internally it cons3 ..sts of a pulpy layer surrounding the s.inp:le l::irval cavity, the cavity itself lying -;-,arallel to t,he si::Ies of this lHO.nothalamic ty:::,e of c~,:i.11. The hcle from which the irrnect en,er,:.:es is variaLly distriL,ut ~1d over U,tc surface or

'rryDetid. !.'o gall-m;;ker was reared from this type .~:all, l..>ur,Felt (1:,40)

:reports th11t a member of the Trypet,i,:la.e was reared f'rorr: the galls he collected. ·rt1e writer had a~cess to these talls, and 5t,udied Felt •s

(1940) descriDtion. .~ comparison of Felt •s galls and those collected by the wrj.te:r 1:mgfested that they were both the same type.

'rorvmu.s k:inse~i Huber. Ghalcidoidea. {: ''l a.,et .." , figs. 2, 3).

18 19

h.emarks

This type gtll begins t,o form in februar,v. The galls '

F'ig. l. Pusiform., glabrous., brown., stera 1Y,all.

Fig. ~Iin~ of Tory711uskinse;yi lluber.

Fig. J. i\;1tenna of '?orvr.mls kinseyi Huber. Fie. 2

Fie.. 3 20

Gall - !iazelnut-shapc'td, 1.md ·Jall ------__...__ - - bud i:mll (?late fig. 1), wit,h t:rowniatl lines Rxter 1din::: across thf' fall dividinr it ir,to ::,nree

t~r:i.or is riilose., and co,1sists of overlappin:.:: lay~rs oi' ., reen:i.sh-rone

and ?1.3 mm. in lenrrth. The irLt.e:rio:r consists o:r a lr,yer of tendor f;reen- ish tissue, with:tn whlch lier, a layer of white 0ven Jno:re ttm:iP.r tissue surrounding the larval cavity. This cavity lies ;)arallel t.o thf' Rides of t,h1.s monothalamic type of tal.l... •~'he hol,~ fco1i, which the lnsect eir,erges is U$Ually at the miripoint of the i:'.:,,i.ll.

Ga.11-I,.aker

'Sutr.eta diana 0. s. "I'rypetidae.

'1'he larva is y~llowish-brown, and in its late staises averages

4 mm. in length. ln the early stages the larva. t.1pers t.owa:rd tllf' hea.d rer1,ion, civini:I: it a :.wri.forn:, shape; it then rra,hrn.lly asBur;ies an almost cylind!'.'ic;il shape.

the <:H·.irtna.l dP.scriotion of Ud.s insect (P. :>. Geol. '?ur. Of 21

Ter., 1877) the f'er;m,lc is ccr:siderei to be 5 mr::. in ler:gth (including oviposito:r), nnd the 1:.F1lt1 i.:. 'foscri~i(';d r:J.S 6 to 7 :r,m. in length. from tile wrlter•s oi.rnervations the ;,:ale is note,'.l to be 4 to 4.5 i.lii,1. iri lEmgth,

this species exist. /', se?.rch through the literat,ure :inriicates that this is the firat rei,)ort c,.f :.hillips, 19~'3).

,._ ')3) Chalcidoidea. (Pl,tte 1 1'ltS• ,,,,, , •

Tor;vrm1s kinzeyi lhn:,~r. Ch;;lciioidea.

'I'l1is tyi::ie fall bed ..ns to ro~11 in l\pril, the insc-:cts (3li,er 1d,,g any- time from the ;-.nd of i-a.y unt:il early July. Felt (19/,1,0) describPd the color of the gall as yellowish-brown. The examination t;y Felt may h,qve been made after thf-1 ,:alls had oeen removed from t.hl:'; plant, for only during the late sta)!es when the insects h;1ve enierged, and the tissue is dying 1oes the gall aasu.;ne this color. Pig. l. Hazelnut-shaped, bud gall.

Flg. 2. 1::ing of llabrocystus sp. F'ig. 3. Antenna of Habroczstus sp. Fie. f

Fie, 2

Ftc. 3 22

---Ci'l-:..1- --')val, r;111 f'ous, -----Leaf ia.11

b:rmm color; th11 su.::-i'ace w:rinJ-des, while the int~riot· becomes driec1. The

wMch are parallel to tho sides of th1o ;_all, and s:i.tuatl:'!d :Lr, the lower half of the rr.all. Two to four pupa ca.st~G an, present Jn this :.1olytha.larrdc type o.f gall. F'ro:;l the. ba.stl of the pupa cases there t,xtrmd let1ef~tr1y fibers tow;q,rd the 'trpp!'!r cxt,erior of the ga:l.l. The hole::, 1'rom wtLict1 t,bti inr;ects

found nnar the base. -(}all-!'.aker Anion sordictum ~;r:LLth. Gurculionidae. (}all, lW/$).

,fo lsrv?.1 or pupal forms wore found. ilowev•.n·, the exuviac of one Anion ,,·as discovered in the :1upa c.?.iJe.

__'i'orv;::us.....,_ k:tni:;evi l:ui.Jnr. r;h".tlcLl.oidea.

Formation of the i;all begins in arch, the insects emergimI any- time from : ay unt:i.l the en6. of June.

Tills 1,:uf ;;n.11 can be distinguished from the glooose, bladdery 23 puffy, leaf gall by its smaller si1.,e, more redd1.sh appe"lrance, and also by the fact thst thouzh two may grow on opposite eidP-s of a leaf, tney do not. ?row in clusters. 24

:.xa.1.., , 1 - 1j·'l ct,oac,. ··1~, ac,t~r:y,' . •Ju_ ff .. v, .1..ee..1.' ,,. ,.,u·a ll ------___, - - ,,,,...; ,r1s ga. 11 (rl.· cl~i,t Lu.,·•·r·· f' 1.g. l''',.,) is' rr.,ern.sri-w.. t·tiJ. ·e w:i'th ..• a roee tit n

throughout. so.rt hair:: ::re sc~,.ttere:i over the surface. lt 5.s cormected

to thr leaf' ~s n sep-<1rable outrrowt.h, end ~t the r;,oint of attachment the

di.a.met~r, and from 9 to 13 nm. in

Two tn

often located ntmr the !;.op of th-: ;;rnll, 0ut a.re inconi:ii:-it,:mtly arrar1red.

DiR·r-throtton:,vi9: artemisiae ff!lt. (Felt, 1925), (r>late IJI, fig.2).

·:~he la.rva is yellowish-orange, and averages 1. 5 mm. in length in its late stages.

:'be i,ta.le genitalia (::>late Ill, .f'ig. 2) of this s 1.Jecies average

in irddth, or approximat,Hly h,tlf the wiJ1,h cf the new species a.ssocia.ted with the top.-slia.pod, woolly, whitisb-b:::-own stem g:all. The aedeagus, in dorsal view 1 can barely t:iH seen in the crotch of the cercd.

A.:ninverted 11Y" typifies the appear;, ..nce of t.he cerei, with the arms of the 11Y11 very wide and separated Uy a deep crotch. Ledium-sized hairs extend .from the edges of the ~lightly co.nv~x loter'll borders of the gonostipes. 1t:hilo th-c1 cl tisty.H d!'" fo s.t. tlte"i.r _;;:id,~io1nts they taper

5, 6).

~- (Burks, 1938). 4'' 1'et.rastlchua sp. (Plate , .I. l.gS • 1l I t:)) •

1Ul these forms b?lonf;:; to thP, auJierfamily Ghalcidoidea.

•r,1-• 1 ·") , r,1.s ga 1..,... was ucscr1. .. t>(:",.l' oy '-'• e... "it (·•9... ;:iv , 't.;ou ...n a.c.b -rev1.a . t ea'~ 1 om.

"1'ha writer, thnrei'ore, corndd.f'.lred a e,:ora ::ietailed descrif.~tion a.pprop-r-iate.

time from Apri.1. until late in ,rune. It was fcnmd in al.moat all ti,F: aretts coll,,.,dnd, but w:ts the onl;r tyu~ t,a.l"L found on the west side of tJta.h L.ake.

list of p~. .rasit('ls it can be readily seen how diffi- cult H, wn.1>to oht~in the g<.all-nake.r. U::iost t"!:Very parasite that attaeked th!"! other species of ga.ll- 1,1akers, also 2.ttacked Diart.hrono::9::.ia arLemisiae

?elt. PLATEIII

Fig. 1, A. Oval, rufous, leaf gall. B. Globose, bladdery, puffy, leaf gall.

Fig. 2. Genitalia, male. Jiarthronomyia artemisiae Felt. Fig. 3. Wing of Lepta.cis sp. Fig. 4. Antem:a of Leptacis sp. Fig. 5. Wint; of Pla.tygaster sp.

I Fig. o. Antenna of Plat:z::.!'.aster sp. A B

FIG, I

FIG . 3 FIG . 4

FIG, 5 FIG. 6 ?6

( 1'·1~ .. H,i.,t, ., c:if'i.:,en with l~~ve~ rrojectinr

easily, 1,hil,, th" int,,.:rior tissue rf1i:.1tlns tendrr for a lonr; period.

Dtnrelopment o.r ga.lJ. :u is conr:nctcd to

fro:r: the st.er,, but is ;;ot a swelbnr of

, are present.

Lost often a~-e J~ro1wc;; towu:rds tne lower centl'al J_'.,ortion of this

variably distributed over th,~ surface of the gall.

--~;all-Laker Db.rthr-onon:yia, new species. (='late IV, fii,rs. 2, 3, 4, 5).

the early stages, twfore pupa.ting. it tapers slightly ard is r~mnded at tne ;,ost.erior abdornina..l. section. '/:he head re.:,;ion tapers more sharply.

rhe pupa is brownisr., and averages 2.5 mm. in length in its late

P-Xoskeleton. This 11µ~11:1.,,sto all 1iu'.,a. of the Ito,lididae collected. ( l!':~l.1!i'11.t'1f:.. ' '' ?~Yl it a.1 11'.?)· \ •

5) Of t}-,j S DPW

crotch.

,..,\ . ,,_) . dict:ist,1li

chitini7,ed tir:s ,::.re 1~,resent on the iist.ir;tyi.:.; t of

;?.rter.,isiae , clt.

point wher-e vdn ? :~,,,,,et5 1.t. Vein E1 fuses with the costal vr.:'in rm.st the :ddnoint of thB costnl margin. Vein E curves upwar::i sli1:,1.htly and then 5 slo::Jes down to r.if!et the costal vein. Vein Gu1 slopes towal:"d the anal mar- gin of tho wit:''. m,d t lien curves even morr-; chtc.rply towards that corder, but fades out before rci1ches the :,:J-::rcin.

T'he fer:iale avera:::;cs 5 ,,ii:,.in ler; 2;t.h ( includinf; ovipositor). The •

a:re n0 nairs. Tl,e, bys.line sectio!',s of the ·,dr.if, hf;,ve r,,:d.n:,, i:ivt, n0t

h.6 m~et. vein and then curves downward as it :aeet-s the costal v0ir1. Vein Cui is also thicker, but extends little past. the ,dd:,1oint of tr1e wing.

193.9).

Tetr:1st,,icrms sp. is present :ir: three othor tyi·)es of /.:alls, whi.l~

t~ present in all tn,t two of t~he t~nlle.

latter ;x:.rt cf :~eccrnbor until Lhe middle o,,. ay.

'l'he starid of uii"'. sa;1:eb.rush on the foothills o.f ,ryu u:.mmtain was the source of the 5:ro:d,t.1st rn,i.;r,'ber of theso calls. However they were found in alraost every area ex.ce::,t the west side of Utah Lake. PLA'l'E IV

Fig. 1. 3lobular, glahrous, brown, stem gall.

Fig. 2. itJin1~, fenale. Diarthronomyia, n. sp.

Fig. 3. SegrnP.nts l~ ,3,nd 5 of ant~rma.

Fig. h. ldng., male. Oiarthronomyia., n. sp.

Fig. 5. Genitalia, male.

... FIG. I

C

,.- /

R,

I r-- I / --- Rs - -- I -- F,c. 2

Cu,

Rt C

Fie. 3 i:-,c. 4 Cu1

Fie. 5 29

i.'he exterior of tbc gall ( ?late V, fig. l} cor~sists of whitish- brown velvet,y fib~rs, which are very mucb entangled. It is connected to the lateral portion of trie ste1ii, as a separable outgrowth from the stern, hut is not a swelline of the stem proper. Development of th~ gall is of a prosoplasmic type. ;;it,h age the g13.ll a.ssurnes a darker brown color; th~ velv~ty surface is retained, while the interior tissue becomes brown and brittle. 'rhey grow in clusters of two to twelve, the smallest clus- ters ranginr: from 7 to 13 mm. in diameter, an:t from 7 to lJ mm. in length, while the l,qrger clusters rant~e from 9 to 16 m.11.in diameter and from

9 to 17 mm. in len~th. \Nhon growing in clusters of two approximately ha,lf the stem is exposed at the point of at tachr:,ont; in larger clusters, the whole stem is surrounded. Internally the gall consists of a greenish, succulent tissuP.. One to five larval cavities, irregular in outline, are interspersed. in the tissue. They lie close to each other, parallel, and in the lowP.r half of this polythalamic type of .,;,all, rather close to the

;)oint of attach.rnent of the p;all to the stem. The holes from which the insects e,,,erge are variably jistributed over the surface of' the gall.

Diarthronoi::t;ria, new speeies. (Plate V, figs. 2, 3, l.).

·rhe larva is orar::.p;e, n.nd averages l.5 rr.ru. in its late stages. lt. ta,pers slightly at the head rerion. The pupa is brown, and avcirages 2 mril. in lene;th in its late sta.ces.

1'he oale a,v~rar,es 4 mm. in length; the antennae (Plate V, fig. 4) 30 avera~e l. 5 rc;11.in length.

'l'he ,::ale genitalia (?la.t.e V, fig. 3) of this new species n,easured

.5 mii1. in width, considerably larger than the three other species examined.

Only one specimen of n iuale was obta.ir.ed. Extending slifhtly past the distal portion of the cerci is the aedeagus. ll•i-sh:ipe:.:l cerci a.re present, with a wide rounded border in the crotch. Lon,: hairs extend from the very convex lateral bor:iers of the e,onostipes. S:1orter nairs extend from the dististyli, cerci, and t:1n border between the posterior edge of the dis- tistyli rmd the cerci. The di.stistyli are alfaost, tubular in appear.a.nee and are widely separated in comparison with the other species. Along the dist~a.l margin of the dististyli are heavily chitinized areas.

'!.'he fHrnale ~\Vera,t,ea 3 .5 nu:i. in length; the sntern,ae average 1 ,nm. in length.

'l'he wing of the fer:,nle (;>late V, f'tg. 2) averages 3 mm. in length.

Long hairs extend i'ron; the lHise of the wing along the costal border up to a point before the costal vein starts curvine downward. A group of very long hairs extend from slightly past the bas~ of the wing up to a ;)oint where vein Cui begins tapering, along the ,anal margin. Shorter hairs are inte:rsp~rsed in the hyaline sections of t.he wing. The costal vein is fairly thin and is met by veins H1 tci.n:i :\.1 at the outer margin of' the wing.

Vein r:t1 extends from the base of the wing, curving upward toward t,he costal

1:1ar!".in, and then curves downward to meet the coetal vein and vein H5 • Vei.n H extends from the base of the wing in a straight, line, and tapers 5 as it ,:1eet~ the outer ,:,art:in. Vein Cu1 ~xtarids from the base of t,he wing t,owl:l:rds t,he :.uial itia.r[:tn., tapers, and fades out quite a dist;:,,.,nca from .31 this :-:,arg:in.

?arasites 2!! !d!!! Gall-~ ·retrastichus sp. - approaches hibua.

T'orymus kinsexi Huber.

Eemarks

•rhe gall ber,ine to form in J anua.ry, the insects emerging anytime from the latter part of .February until the latter pa.rt of April.

ln general appearance this type gall resembles tho top-shaped, woolly, white, stem gall, especially when growing in clusters of three.

The outstanding distinguishing characteristics are the velve::.y appearance, and +:.he ti.nee of brown in t.he top-3haped 1 woolly 1 whitish-brown stem gall~

;'\not.her intP.!resting f'eat,ure is the fact tba.t the lat,ter gall was never Fig. 1. TOP-shaped, woolly, whitish-brown, stem gall.

Fig. 2. Wing, female. Dia rt hronomyia, n. sp.

Fig. 3. Genitalia, male. Diarthronomzia, n. sp. Fig. 4. Segments 4, 5, 6, 7 of antenna. FIG. I

'' I' i Cu,

---G

FIG . 3 FIG. 4 'l'he exterior of this gall (Plate VI, fig. 1) consists of extre;;,e- ly whit.e fibrous t.l,rea..is. It is connected to t,he btcral f)Ortion of the stem, as o. !'1.op;irable outii:ro,-rth from the stem., but ia not a swelling of the stei.;,1 ;::iroper. Uith age the eall a.ssu:1tes a .,;ra.yi.sh color; the exterior r~:icn.ins woolly, ,vhilr3 the int e:-:i.or tissue beco:,~es o:rown. i)evelopment of the gall is of a prosoplaewd.e typi;J. They grow singly or in clusters of two to nine. The> sinr,le galls r/'it.nge from P to 16 c:;:n. ir1 diameter, and from. to 13 mm. in l

C:Tierge ar~ most often located n.t the top of the t,;all. -;all-taker !1.hopalo:,,yin s:;J. - a_·J.-:,roaches Ii• alticola G~ll. (Plate \iI, fig. 2). 33 late '3tages. In the early stages the abdomen is three t.imes a.a thick as t.he anterior portions, later ta.;)€cing shl'.lrply from. mid-abdomen to the genitalia. lfoen irritati~d in the larv".\l cavity it beco:i1es very active.

The pupa is lrrown in the early stages, and then assumes a black color. lt aver.'lges 2.5 ::a,1. in lengtn in Us late st~ges.

The wing of Htf~ few.u.e (Plate VI, fig. 2) avera,~es l.5 mm. in length. Hairs extend from a.11 the tiargins of the wing, and .are inter- spersed in the hyalinR s~ctiona of 4',he wlng. The CO!i!tal vein is very

th~ costaJ. vein slightly ne!'oc•e it be,0:ins tapering. VP-in extends

out before it reaches t,he cuter m~.rgin. Vein cu1 e:xt>?nds from the base of t. he wing in a st raip;ht line towards the anal margin, tapers, and fades out before it reaches this r::argin.

( 1''1 ar,e. fir,s. 3, 4.).

!'ost Hnit~d ~tates species of this gienu.s are undescribed. A pecl1lir1r hook-like i;tructure on the metathora.x is d.istinct for this group. 'l'his insect is specific for this type gall.

:teu,arks

Tho distribution of this type gall is seer:1ingly lirr.ited to the roadside area of fr.eek G,.myon. 'J'h>?.gall begins to form in the latter part of Decnmber, the iasects r,r:;ergint anytime from the latter part of FebMiary until the l~tt!'>,t' PLATE VI

F':ig. 1. Top-shaped, woolly, white, stem g,J.l.

Fig. 2. Wine;., female. Rhopalomyia sp. Pig. 3. Wing of Inostemma, n. sp.

Fig. 4. Antenna of lnostemma., n. sp. FIG.

F1G. 3

F1G. 4 34

'.•·,1· 1 - i,"l oou' l ar, Pf'. orous, v., e 11 owis.· h , !.P~,,, ·t "t .=!_·"' 11

The fibers are grP-i:mjsh-white to y~llow in appearance. Other- wise, th:i.s gall ( 0 late VII, fig. 1) is quite sim.ilar to the dark ;mrple gall. lt is connMted to the lateral ?Ori.ion of the ste."n as a separable outgrowth from the ater:i., but is not a swelling cf the stein proper. t'lith

,'l;'!e th~ ra.11 assui,:es a ,iull violet color; th0 f'i.bers dry rapidly and crumble to thi,1 touch. Jevelopr1ent of the gall i3 of a y)roso_;)lasmic type.

'l'hey grow singl:,r, or /Cl.regrouped in twos anJ ttlr.ees in 11 clooe .linee.r

rru1r:e from 10 to 18 :nt:. in ti:iHmetc:r, and fro,,, 10 to 19 rm;,.

qu,1,rtl'lr cf t.11e stE>.n1i~ cxposc1 at t.he point oi' attache1cnt., or occasionally the stP-r:i is COc.!l;Jl~tcl7; encircled; i:1 close 2~rou;)s th-, stom is conroletely f3Urrounded. Intm:·n~lly it consist~ of the :.mpa case, fron. which extend

113aves wH,h entire m,,'lrgins to the 1;.n::t!"!rior, ~nd some carmine-re1 fibers.

The pupa caM lie~ clos0ly apposed and parallel to the eten at the pcint of attachment of thi~ ;iol:.,thrtlam:ic type of' gall. 1'he hole from which the insect ernp,rges is usually near the top of t,he gall.

viarthror,omyia sp.

This is a different. species than an;v of the others .:~(!ntioned.

'i'he larva is yellowi eh-orange, a.rd averages 1 • 5 n:rrn. in lene,th at about 1:.ho midpoint of it,R drwelo;;ment. t'o pu.:,:;awera found. '1 t: .~-.,,.!

Tcrvrnus kinseyi Huber.

Tetrastichus sp.

'Phese three bAlong to the r,:uperfamlly Ghalcidoidea.

'l'his tall bogi.ns to form in A;:n•il, the insects encrr:inz; anytime from the latter part of ray until the early part of July. PLATEVII

Fig. 1. Globular, fibrous, yellowish, spotty, stem gall.

Fig. 2. i~ing of Tetrastichus sp. Fie. 3. Antenna of 'fetrastichus sp. Fig. 4. Wing of Amblymerus sp.

Fig. 5. Antenna of Ambll!,!!;erus sp. F°IG. I

Fie. 2

Fie. 3

Fie. 4

Fie. 5 36

1,.... 11.;J.,. 1) is the

t'ror,' +::.h":c•orte:::- of the _e;.~ll, ,.nd tirotrnd.ing from the surface. ~ihite

to the lat-?.r°'1.l nortion of the stem, as !l. "!!~parable outt~rowth f:rom the stem,

r;rnyish color and cru1:1bles to the touch. Develo!)ra~nt is of a prosoplasmic type. Thesl". ralls grow sin_i;;ly or in clusters of two to seven, the lar,-:est clusters r-1:1.nr;ingfro:\, 11 to 15 m.rn. in :li?..meter, arid from 12 to 24 m.n,. in le~1gt,h, whereas th~ others r;..nir.e from o to 12 mm. in diameter, and from

the point of attnchment; :i.r: clusters the entire ~tem is surrounded. Tan pup~ cases, usually two to five in number, lie elosnly apposed lengthwise, and parallel to the st.err; ri.t the r,oint of att~chment, of this polytha.lamic

trH;.uted, },11t r.,ost often t:l..relc,ce:ted ne4r the U:)per parts of tho f;all.

Jiarthronomyia sp. fig·. 2).

is s. .Ji.:'ferent species than the others mentioned. The larva

is brown :ln color_. arHt A.Vr>:rat,res 2 mm.• in l~ngth in its late stares.

';'bB winr of the fentle U'l1te VIJI, fig. 2) averae:es 2 ,nm. in lenr:;th. Il:J.ir~ erl.erv:l f'-:-o:i,all marf".ins of the wing, 1:.nd '.lre interspersed 37 a;;,one the hya,llne sections of the wing. ,Hl the veins are rather thick.

'i'hf'l costal vein tapers 'ls it iueets vein Vein ::L1meets the costal vein ~t a point s.lir·htly l:;efore th,i costal vein curves :fowr,w,ird. Vein

R8 ext<-mds in from the base of the wing in a straight line, curves down- w~1.rdand taoers as it re1chee the outer margin. Vein Cu1 extends from the base cf the winB; toward the anal mar,~in, tapers, and fades out quite

9. distance .froD this mHrgin.

TP-tra.s+.ichus sp. - reseJ;ibles !• ~. Chalcidoidea. This apecies is different from the parasite in the globular, g-labrous, stem g.all.

Remarks This type gall does not occur in abundance in comparison with other galls. They eta.rt developing in January, the insects e;:1erging anytime :from early harch until the latter part of June. vnc ga.11 1

0 collected £,arch 30, and dissected on June 51 revealed a very active

Itonid larva. Fig. 1. Globular, fibrous, white, spotty, stem gall.

Fig. 2. Wing, fer,'lale. Diarthronomyia sp. C

I cu,' F,c.. 2 )a.11 - ,:aotul:CJ.r, Dark hu·ole, --t:tfrt,1 Gall 1'he gall (Pl::i.te IX, fi,;:. 1)------is very fibrous in appearance, the fibers interspersf'!d ,'3.inonr:the leaves ,and aridng from within the gall but near the surface. lt is connected to the later:,.l portion of the stem as a separable outgrowth from the stern, but is not a swelling of the stem proper. With age the gall loses little of its color, and crt1Jribles to the touch. lJevelop:cr,ent ia of a prosoplasmic type. 'rhey e:row singly or occasionally in clusters of four to five. Singly the talls range fro,;1 7 to 10 m:r•• in dia!11eter, and from 8 to 11 m.m. in length; in clusters., t!1ey range from 10 to 13 mm. in diameter, and fro1n lO to 12 :1mt. in length. When r;rowine singly approxima.tely one third of the stem is exposed at the point of' at ta.chment., but occasionally tbe entire stei":, is encircled; in clusters., tht~ entire stem is endrcL,d.. Internally it consists of a. centrally loc,1ted pupa case, covered with hairs; len.ves with entire i:,areins grow out from this monothala.'Uic tyc,e of gall. T'he hole from which thl':l insect nmerges is vnriably dist :ribut,ed over th!3 surface of tho gall.

Ga.ll-tla..'l

Diarthronomyia. sp. (Plate lX., fig. 2).

This is n lif'ferent species than t,he others mentioned. The larva is yellowish-orange., and averages 1.5 r:m:n.in length in its h,te stages.

The thorax is ,'! darker orown than the abdomen. In later stages the thorax assum~s an exceedingly dark brown color. 39

!.'his type gall beg.ins to fc.rm in t.ho part of J rtt!uary, i,he insects emcrj_et:1ng:1n;ftimo from early l'•,arch ur:t.il the latter part of Luy. PLA'l'E IX

Fig. 1. Globular, dark !)urple., steir, tsall.

I<'ig. 2. Segments 4 and 5 of antenna. ...

Fie.

Fie. 2 40

The fi,:ures ( Ph.te X, fi.1:,s. l, 2) a:re of Diarthronomyia sp.

(a,_1pro,q,;::hGs Q. occi:Jental:ls felt). i'ositlve association between it

and a gall ·was r,ot, est,:1ulislled. Seer,tingly it was reared from a thin-

walled, ovate, white-woolly, leaf gall. 1t was collected wltb another

type {fall,

'l'he male ,genitalia (flate X, fig. l) of this species measure

.3 :nui. in width. 1)1'.:tending noticeably past the distal edges of the

cerci is the aedeagus; the cerci appear as an inverted heart. Laterally the edi;tes of the gonosti?es are smooth, as is the border between the

posterior edge of t,he dististyli and the cerci. Distally the 1:dges of' the dististyli are very convex, and slightly concave along the o;:poiriite borders. l'.oticeable hooks are formed by the chitinized tips of the

dististyli, and trtese almoAt, ;::,eet at a point close to the aedeagus. Fig. 1. Genitalia, Kale. DiaJ:•thron{i:nyi~ sp.

Fig. 2. 0P-l:Jnents 4 and 5 of I'Jiarthronomna sp. •- --G

FIG .

Frc . 2 -~-----~17:xplan:ltory --;, ote for --Plate Xl- 7wo ?roups of 0alls are 9ictured on Plate Xl. 'l'hose in outline for.:n are oxam~::ilesof ltonid galls from Wells I paper (1921); tbos1-1 de- picted in detail are the ton r,alls collected by the writer. 'dells (1921)

"'Xplains the reh.tionshi?s of those in outline form in the following

:::rrnnP-r: ( 1\11 numbers in the followi.ng pn::ra.rra.ph are figures on Plate XI).

'J.'he most prfatit.ivu gall in this grou!) is a sir,,ple leaf fold, a ;;rinci ryn.l vein constit,iting the gall a.xis (1). From the initial leaf fold was evolvmi th~ va.riAble ill-defined pocket type (2) 1 whiel"1 i.n t~1rn I:ave rise to the si,n71le prosopla.s:ta (3) • This advances either throuP,"h th!'! differentiation within it of a sclerf'nchy,na layer (4) • or through the p~rtial transfer of the gall toward the upper side of the leaf (5). F'rom this type we have a striking advance through the sclcrenchymatized cecidium (6), to the type characterized by a de- hiscing larval cell formed by the lignified tissue (7). The majority of up-walled ealls show the advanced state, hm,1ever, through their containing a scleride zone ( 9). From the UP-walled type ( e) there has also been derived the interesting double-chambered form (11), with the type shown in (10) as an intermediate condition. Originatintz eith~r :i.ndepondently or from the ltonid insect which produces the shplest leaf type., we have the burrowing form which causes the •tbliat,::-r" gall (ll.,). A form of this kind attacking the embryonic fruit gives the situation shown in (15), or beco.r;iing more specialized on thB leaf' gives u.s the prosoplasma (16}, or att.ackin1; the stem initiat,es a series of ster11 galls (17-20), going over into the proso- plasma ro,'.ion with the att,:i.inrnent of sclerenchyma layers around the ln.rval chambers. If the apical merbtem is used as a basis of o~era.tfon we get such a primitive F,;all as shown in (21) 1 and either through a burying ~Jrocess on the part of the larva or throu;:;h the up..walling r.1ode of overgrowth, we pass throut:h such .s. sta:'e as (22), ~nding in such a hiehly [email protected] type a.s (2.4).

1'he writer ha.s insrirted the drawings of the gn.lls of Artemisia t eidentata to de:.,onst rate t~1(c.ir posi.tion in the :iegree of specializa- tion, ani ~,hnir possible ori;:;:in. '.{ea.sons for thl':3ir position on t,he plate are p:i."OSf'nted in the (msuint; discussion. .PU,T.E XI

1'~igs. 1-lh, 16 • • Leaf galls.

Figs. 17-21} • . • Stem galls. 1'"ig. 15 • • • . • F'ruit gall. Pig. A • • • • •• Fusiform, glabrous, brown stem gall.

Fig. B • • • • • • Hazelnut-shaped, bud gall. Fig. C • • • • • • Oval, rufous leaf gall. Fig. D • • • • • • Globose, bladdery, puffy leaf gall. Fig. E • • • • • • Globular, glabrous, brown, stem gall. Fie. F • • • • • • TOP-shaped, woolly, whitish-brown, stem gall.

Fig. G. • • • •• Top-shaped, woolly, white, stem gall.

Pig. H. • • • • • Globular, fibrous, yellowish, spotty, stem gall. Fig. I • • . . • • Globular, fibrous, white, spotty, st.-,.'llgall. Fig. t,T • • • • • • Crlotmlar, dark purple, steii:1 gall. Key Labels .2£Galls £f. Artemisia. tridentata ~• F - r'ib1·ous tissue

r'I Fibers

GS - Green succulent tissue L - Larval cavity

LV - Leaves (with undivided margins)

01 - Overlapping layers of gall tissue

P - Pupa. case

S Succulent tissue

WT - White tender tissue PROSOPLASMAS KATAPLASMAS 0. I I .. :z::/ , li:

• > I ~·~-~-/Q. - ..J 0. 0. llo

~ I I I ' I I l'.) CD -----

I I I I ----- I I I I ..J "> ... _, C') w o u~ ~C\I ' Q.

1 ' , <{ ..J

'&.. I, II) l'he ten £1.:alls collPcted will b~ discussed starting from the type showing the least, specializ:'3tion, ,1nd progr-essi.ng to the more specializ,ed.

Differences and sfoiilar:tties are exhibited by these r:<1lls, anci they will also he pointed out.

the ten ,•alls, th<~ only k3.ta;:ilasr::ic t~ne gall collected is

C!lvity. Accord:i.ng to · ells (1921) a stet:. r:all ,himonstra.ting a polythala- r:d.c condition is mo,,.e sDcci.::ilized than the r10nothalrudc st('.n gall. V,in- sey (1925), in referrinr: to the :::yninidae, considers the reverse t,o be

cause it is in,ieparable, a recor;nized less-spl:'lcialized condition, and because it does not exhibit a true cell condition, lmt is rat,h'!'!r a mere cavity in the stem. Additional supportive arfum~nt is the fact tha.t membP-rof the 1'rypetidae is the gall-maker, and in i'L;," 1 lt is not as specialized as members of tho ltonididae in gall-making. The point of orip;in of this type gall, accord.inc; to Wells, is from a highly s;H-Jciali- zed leaf gall, which either attacks the en;.bryonic fruit ( Plate XI, fig.

15) or irritates the stern (PLs.te XI, fig. 17) •

l•ihether the hazelnut-shr1ped tmd p;all ( r'late XI I fig. d) or the leaf r~a.lls (Plate Xl, figs. G, U) are :next in degree of specialization is in doubt. ':'he writer ifl inclined to consider the bud gall less spP,cialized, since it tc

V9.llCEdt1 since they 9,risc :in a specialh:ed raar:ner fro,t th~ leaf gall. ;\ polyt.halo.uic condition present a:r,ong the leaf :;;,tlls D.lso favors their

(1921) st,:i.tef'l ths.t all bud galls ~re proso:;.,lo.smas,

whic~: is rm ex:u:,ple of the apical meristc:r. of the st.em scent t.ype

., fig. 24), moro layers develop an up-w.rtl.led condition a..'1d thereby f?nclose the larva. 'l'he l~af galls would, in the 'll."riter• s esti- mation, a.dvarn.:e frot;, tho condition of t)late X.11 fig. 9 by merely having the lateral wallR meet and fuse.

rhe glohular, gla.brous, stem gall (Plate Xl, fig. 1") and the top- shn.;)t:!d, woolly, whitish-brown stem gall {Plate Xl, fig. B:) are considered next. 'rhough they differ exterrn ..'¼.lly., one being rJ.abrous and the other woolly, the interlor is very s:i1dlar., botn boing polythalamous, possessing 4h distinct larval cases, and both having an interior consisting of green succulent tissue. rrhe wr.iter considers tbese two less specialized tnan the tor,-ahapAd, woolly, white, stem g,111 (Pl.1t,1 lntter has :m additional layin• of tissue, and the larval case:e are more r~t~ularly ri.rran.r:,e.d. 1'he arrangement in the first two mentioned is more

J;,·1,:1h:,l:z;;.rd. 'i'hcsc two falls, in the writer's opinion, would also arise

J'1~omthe t,ype shown in ?late :KI, fig. 23, where th.-; apical :neristem of t;,he stem is the pa.rt attacked. This ..muli involve a fusion of the np-

't'his condition, acco to 11.insey l•1q~~)., , ~- ,, J ,:tv,;:;,.~ft•a . ,_ - ""Pf.f~ .. . ..·,~t , coMddcrat ion of thE'! /all as more specialized •

Two new species of Di1rthrono;nyia were collected (Plates 1V, V), :md both were reared fro11c these two galls.

The top-sL.:iped, woolly, white stert. gall (:?late Xl, fig. i}) ex- bibits more speciali?.:P.tion than those preceriing it. tis in the two galls coni-,id~red above, a thick 11:iyer of ereen succulent ':issue surrounds the larval cases, but i'1 addition there is a ,rore h'3.rdern•d layer surrounding this t,issue. Also the arrange:1,ent of the cases, which can be two to five

'rhis type t:nll would possioly be dc.rivr-d fr-011, the two mentioned above,

fibrous, yellowfah, ::wotty, strm 1~:i.ll ( i'late n, fi1.:'.• H), tne ,r;lobular, l,5 fibrous, white, apotty, stem gall (:'late XI, fig. I), !'lnd the r;lobultir leafy, dark purp.'. ~, stc>Jii fit~• tl')' • Their commor.feat\lre

!,ccG,~iJ,." .. to ''7ells (1921) it is the essential newne!;S of p.rosoplasHktS

this be thB C'lse, tlwn +:,hcs,e three gallo ( Plate Xl, .ti;:s. B, I, J) must bo connidered zaore s;)i<:1cialized fen:· here is no reshaping or addition of

consist ma.ir:l;v of

ir1:i.epen-

fig. 21). t.hrou

thro 1:i13hthis 15near series. 1. Galls of ~rtemiaia tridentata riutt. of Utah County were collected and 1A.scriL>sd in detail. 2. ur the ten r;alls collf'Ctl'.'d, seven were caused by members or the

Itonididae, two by m,~nbers of the ·I'rypatidae, and one by a member of the Cnrculionidae. 3. or the seven Itonid galls, six were caused by species of Diarthro- nomvia, and one by a species of ;iliopalomyia.

4. Two of the galls were caused by new species of Diarthronom.yia.

5. 'Three genera of the Scelioniclae (hyi;;enoptera) and four 1•.encra of the Chalcidoidea (l:ymenoptera) were also reared from the galls.

6. A kay to the ten galls was constructed, but does not necessarily

include all the galls of big sagebrush of Utah County.

7. 'ro the writer's knowledce this is a first report of Eutreta

d1.a.na.. u. ,, ,).<'." ('T, • ,J." ,:100" l • C,ur. o f .,,,ierr., l'""'?}~·r in. Ut a,.h 8. The position of the gcl.lls in t:.heir relationships as to complexity

1u1:~spt"cializat.ion .3,nd possible ori,,;in has been pointed out.

4.6 Adat:1s, A. :~, and Shantz, ll. L. 1921._ Tyµos of \/eretation in the '.1ewiarid Portion of the u. s., and i'heir rc;conomic ;iirnii'icanee. Jour. Agric. {es., Vol. :2p, p;). 99-129.

The Insect Galls of tho Vicinity of New York City. Jour. A.mer. :,,us. S:at. Hist. Vol. I+, no. 4, p. 33, fi.g. 79.

jeyernick,

18?'7. Ober Pfl~u1zengallan. Uot. Zeit. Vol. :35, po. 17-23. Burks, B. D.

193$. A Study of Cha.lcidoid ~:ings. Ann. of 1•~nt.Soc. of Amer. Vol. 31, No. 2 1 pp. 157-161, pl. 1, fig. 7 •

.-,, r::lemonts, F. B.

1920. Plant lndicators. 'l'he 1.ielation of Plant Communities to Process and Pract;ice. Carn. Inst. of ~~anh. ~;o. 2)0, P?• 152-154, 158, 257, 301.

Gockr!rell, ·r. D. A.

1909. A New Gall-gnat on Artemisia. '.;an. t;:nt. Vol. 41, No. 5, pp. 150-151.

192B.; !1 '.1tudy of tne 'l'ertl.inal r,bdomin:tl :1tructures of Lale Diptern. Proc. of the Calif. '\cad. of Sci., 4th Series, Vol. 16, to. 14, p. L;l 7, pl. l, fi}~s. 4k-l+5•

Co.. ;r.dttee.

1909. lleport of the Cormr.ittlc\e on r omenclature of Gall Insects. i'\n.n. of tr,e F;nt. f1oc. of t\lnor., Vol. 2 1 pp. $-9. Cook, 1:. T.

1901~. Galls n.nd Insects Pro

:~osens, A.

1912. A Contribution to the Horphology and ciiolor:Y of Insect '.xalls. 'i.'rans. Gan. Inst. Vol. 9, pp. 297-378, pls. 11, 12, 13. 1913. Inaect Galls. Can. Ent. Vol. 45, No. 11, pp. 380-383. Dice, L. R.

1943. The isiotic Provinces of' North America. Univ. of' !0 ,ich., VIII, 78 p., Map.

Diettert, H. A.

1937. 'I'he Forpholo,sY of /1.rtew.isia tr:i.dentata Nutt. Lloydia, Vol. l, pp. 4-9, 15, 16, lS-32, 35-43. !?all, H. c.

lF:'98. A H.evision of the Species of Apion. Trans. Amer. f::nt. ~;oc. Vol. :25, p;). 127-1 '?.

Felt, r:::.P. 1913. Adaptation frt the Gall Hidges. Can. Ent., Vol. 45, No. 11, PP• 376-378, pl. 121 lJ, figs. 1-11.

1918. l'Cey to American Insect Galls. N. Y. t3tate hus. iJull. No. 200, pp. :WR-212, figs. 239-242, 24l+-2451 247-248.

1925. t1ey to Gall Eidgee. N. Y. State l'.us. Bull., No. 257, 239 p., ft pls. 1940. Plant Galls and Gall Kakers. Comstock Publ. Company, Inc., ii. Y ., pp. 3-35, 333-335, figs. 332-339.

Hall, H. H., P..nd Clements, I<'• 1~.

1923. 'fhe North .American Species of krtemisia., Ch!"lsotha.rnnus, and Atriplex. Carn. Inst. of ~iash., No. 326, PP• 42, 44, 135-150. 49 i''.J.nsey, A. .,.r 1923. ?he Gall ',:asp Genus f~euroterus (i,ymonoptera). lnd. Univ. ',tudies., 7,o. 5f:, pp. 11-13.

19J6. ?he Origin of High("r Categories in Cyrdps. Ind. Univ. Publ., s~ience Series i\lo. 4, pp. 1-16.

1911. Oie '.}allm1 iJer !'flam;;en. Leipzig.

Lar:1bert, f': ?1rlyle B.

1940. '1'he Distribution of Plant Types in the T::scalante Desert of Utah with ::tela.t.ion to '.~oil Conditions. lJnpualished L,~stcr's 'l"heaus. L:righa.m Young Pniversity, pp. 40, 42, 51, 52, 55.

?hi.Hips, V. T.

1923. 1\ H.aviaion of the Trypetidae of Northeastern Arnerica. t.lOllr. of N. Y. r;nt. Soc., Vol. 31, p. 141, pl. 19, fig. 41.

,~iley, c. V •

lPE.n. On Oviposition in the Gynipida.e. Proo. 1~nt. Boe. of ash., Vol. 3, pp. 254-263.

Yiydb~rg, P. A•

1917. f'lora. of the itocky ;,ountains., Publ. uy the Author, p. 973. Stoddart, L. A., and ,3;.:iith, A. n.

1943. ;1anr:;e ha.na€:er..ent. Lc1.;raw-l!ill Uook Co:i,pany, Inc., l';. Y. and London, pp. 8l-e2, 87, .:3il, 92, 94, 95, 179., l~O, 25$.

'J'idestrom, Ivar.

1925. Flora of Utah and fwvada. Bull. of T1. S. Nat'l. Lua., vol. 25, ?• 601,.

U. S. :ieoloqJ.cal c,urvey of 'l'erri tories.

l[-l;??. ~)escription of' t:rype~-~(Futreta) diana o. s., '1.'ash. ~,ov•t. :"rinting Office, Vol. 3, P• Jl,.l. 50

es.ver, J. , and Clemnnts, F. l~.

1938. Plant "•"'--~''"-'. i,c ~raw-Hill Hook Inc., r:.. Y. and London, pp. 4t>6, 476.

1926. l;ield on Ga.11-lnh.i:i.biting Cynlpid with Descrip- tiona of f"2Aw Species. Proc. o:t: the u. ~. t,at•l. , 7ol. Art. lo, PP• 131, r:.a. •

Wells, H. • 1921. :nvolution of Zoocecidia. Jot. Gaz., Vol. ?l, l·to. 5, pp. , pls. 21-22.