STA lt FOH.D . 11 E R TTY P BL I · 1'1 0 NIVERSIT Y ER I E

BI OLOG I AL VOLU..a VIU

eIE!\" ES N U WJHR 1

A Contribution to The Biology of North American

Vespine Wasps

B)'

CARL D. DUNCAN, Ph.D. (Stanford ) Professor of En,omology and Bo,;my

San Jose Sta" CoUegc

STANFORD ,' [ ER ITY PRESS STANFORD UNIVERSI Y. CA.LIFORNIA

tO~'l)ON, I1I1l1PIJ~&Y AIII.FORD OXFORD Ul<lVII.RS 1TY PRESS

1939

STANFORD N I VSRSI T Y P RSSS S.-rA'SFORD UNIVEll5..lTY1 CAUFORN-IA

LO DON: B MPIiRJlY Mn.FORD OXFORD NlvnslTV PRESS

Taa una ,uel) T~TLO. CaM"I'A.'H"Y

H ,I7TH AV'k!f1.l&, ~ .... vOIle

THII. Iou • .,,,a' CCJJ.tI""'W1'

TOi: ..-o., OMlCA, '(VOTOt .. KlMI

H'1'IolllH Ttri. tnI1T"nt I'fATU o. AJrottaJc:..\ .y rrA~P"OI(l)l UNIVKUITT J'a&II

PREFACE

T h present paper c;mbodies observations on the ~orth American paper­making wa.ps belO"l..mg to the sub famil)', Vespinac, of the hymenopterous family, espidae. The subfamily rna)' best be chara(:terized for the gen-raj reader by saying that it consi ts of the well-known hornets and yellow­

jackets. Whereas ail the North American species which are known to the writer have ~cn considered, ""ten';ve obsen'ati n Ita!; been pas ibl only with r '. pect to the CODunoner species of the Pacific Coast; any genernl re­mar~s, therefore, which are made in this paper, unless otherwise qualified. :ire to be ttJ1derstood as applying primarily to the latter g1"OUp of speci> aIltI only secondarily to species restricted to other parts of the country.

The wo rk is divided into four senions : an iotroduction . " morphological secrion, a systema tic .""tion, and a biological section. The morphological seerion consists 0 f a detailed rreaonent of the ~keletal and muscular !Mor­pl,olog)" with lbe exceptions noted below, of the Western yellowjackd, I"csl' u/1J prIJsy/t'(J,nica (Sauss.) . The detailed Sl rttcture nnd interrelations

j the ar tictt l"" sclerites of the wings, and the mu culature of Lb. male geni­tali:!, have been omitted from the pr 1t study. The systematic section con­, i. ts f a comparison oi the morphological and biological characteristics of the gene", represented in, orlh, \merica. A treatmenl of individual species b omitted. T he biological section comprises ext"!l.'Iive observations nn hahits. behavior, nest-buil" ing, and tife history . The immature stages of /'rsf" la pnuylvan;clJ are descri bed .

1n the course of his studies, which have extended. more or less con­tinuously, over the past twenty _ ears, the writer has made numerous ob­,erq t ions in the field , both on individual wasps aud on active colonies. His field studies have been supplemented by laboratory ludies. H e has col­IccleJ many colonies in their en 'rety and ntade ;1 careful census of their l'on.tmts.

lIe has al.o examined several thousand pinned specimens. These are c rua incd chiefly in the c llections of St:lnford Uni"usity, the California \ c:uleOly of Sciences, and the writer. Small amounts f material have been

"'1l.ilable, also, from U,e 'niversity of Catifornia , the niversity of Arizona, HriOUS other institutions, and private individuals. These" lleaions in­dude specimens from Alaska I ~.(e>cicn and from Cal i {ornia to :'laine. The wriler's personal coUectiolUl hav ' been aUlde in lifornia, regon . Washington, rdallO. '~vada, Colorado, Virginia, and Texas. Specimens (rom many other tat", hnve heen obtained by exchange or purcha!;e. Exotic material [rom all available sources has been compared wilh 'orth ,\meri-

(31 I J

4 ::-:ORTlT AMF..klCAN VeSP l:'\ E W SJ'S

can moterial in order to increase the validity of an)' conelu ions JrnwrI irorn the MlIIly .

• '0 atlenlpt ruu bt!"" made to review e .. <haustivcly the likrature of ",a5J> biology, a5 the prbeJlt paper ii" no sense a monograph. The work of oth"r has been freely used, however, in all C3SeS in whieh it ruu eemed the author'. presentation would be improved thereby. n inclusion from the literature. of facts or ideas of a specific natu re. are accredited as 10 source. In the cas<! of general or long-establisbed facts or ideas, on the

ther hand, it has not been deemed necessary even to attempt to trace them to lhdr ~ourc~

Spffial aekuowledgments are due to the f IIo\\ing menlbt!rs of the en­tomnlogil'3l tiff of talIford University : Dr. Isabel Mc rackco, under whose !(uidance the forerunner of this paper was prepared as a Ph.D. dis­scrtatiull. for numerous helpful suggestions and courtesies whicl cou­tTilmted materially to the author's academic progress. To Professor . 1'. Ferri,. fo r "niailing encouragement, and for setting an example of high achie \'en1cnt.

11l addition, the allthor eJCtends his thanks to Mr. E. P. Van Duzce, of the . !Jeornia Acadc:my Qi ciences. for extending the privilege of e~nm­iru ng the w~allh of material in the AcadcolY collections; to Dr. Joseph

equaert of the Har rd . ehool of TropiCllI 'M 'dione, for the identifica­tion of doubtful specimens aud for valuable suggestions; and to tbe folio \\'­ing ind"'id"a1s for th collection of much material and data : F. M. Gaige,

f the L'n i"ersity of ~1iehigan: Professor M. R. milll, of the Agriculrure an,l ~lechan i ollege. Missi sippi ; Dr . J. c. Bradley, of Cornell Un;ver· " t) • . Dr. l l. I::. Burke and Dr. J. C. Chamberlin, of the United tates I Urcan "f Entomology; Dr. T. H . F ri on . of the Illinois tate Geological and 1\ " tural History urve)' : Dr. Karl S. Hltteltine, of San Jose State

ullc),"e : and I3righton C. Cain, Fric J. Spruyt. Tom Rodgers , Mrs. J ohn Bre.ck, and IJroir.ssor l'. W. Walker, of the l'ni"ers;tyof l'lorid •.

To the author's wi ie. \' c,'a C. Duncan. much credit is due for patience ;md e.IlCuuragc.:.mCIll during m nths or concentrated £to rt, and for indispen­sa"le aid in the tedious tru;ks of preparing lbe bibliograpby, mounting and laUcling iIIu [ nations, and typi'lg manu~ ripl.

TIw writer i. indebted to Dr. ayle Pickwe.lI of the . an Jose State College, enthusiastic colleague and fine fr iend, for the greater number of c cellent photographs r produced on Plates.' 'V to LrV. Figures 197 to 202, 20S to 207, and 2~ to 242, on Plates 'XX I , XXXV, ' X , ' I, and XLVIII. were taken by Lester Brubaker of an Jose tate College. (Figures 231 :lnd 35 on Plates XL \ . and XLV I were taken by the author.) Me. BrtlLake r made al l of the prints.

Tlte rlrawings (Plates I to XXXllI) are original f'n maicjng them a

14 1

PR£PA~C~E~----------------~---- 5 ---------

camera I"cida was used to oLt:liu <orrel"tI1es., "f proportion. The drawings were then completed withollt Lhe camera lucid", many observation being made from various Ill1g1e to insure accurnCJ and completeness of d"t:til • fost oi the drawings have 1-.. made from specimens presuved in alcohol and di ected under water. \\'hene"er desirable. howe"er, balsam mounts and .keletal preparations of material boiled in K H to remove the soft IlIlrts were used also. The scale of the drawings varies.

• lOBI, c.u..,oaNIA Oetolkr 15, 1937

I 5)

C.O.D.

TABLE OF CONTENTS

1. INTROD enos II. TUE MORPHOLOGY OF V.spula P.lIsylva.ica (Sau .)

Tue F£IUJ.E TU E i NTEGUMENT

VESTITIJRE ">10 SCULPTURING .

TKE HEAD AND ITs ApPENDA ES

The Head psule Sutures of the cranium . The lent rial pits T ho crania] areas T he tentorium Additional cranial true tures

The l:Iead ppenda es and Their Musculature The anleno". Genernl featur o f the mouthparts The mandibles T he labrum and epipharyu.,< The labio-maxill ry complex The labium and associated structures The salivarium . The maxillae

The Upha.lic tomodaeum TBE T HORAl( ,\ ND ITs A pPENDACES

The Thoracic T ruuk . Gene".1 features P rotboracic sclerites

The pronotum The pro!"'ctu. The prol)leura The prOSlernum

T he muscles of the prothora.x T he first thoracic spiracle ?fesothoracic sclerile Th~ scutum T he scutellum The mesopostphragma The mesopecLUs .

The muscles of the mesothorax

[ 71

11 l3 13 13 13 14 1-1 1-1 16 16 1 19 19 1 20 20 ?2 22 23

29 31 34 34 34 36 36 37 38 40 41 47 48 48 48 49 51 54

( 7

!\ O ATH AlJ.:EIU CAN " PI NE WASPS

T he ~<ond thorad<: spirncle . The seler-ites 01 the mctathorn.x

The mctafiotum The. metaposLnolum . The rnel"-pectus and propodcurn The propodeurn .

TIle muscle 01 the mNathol'm< and propodeum The propodcal spiracle and its muscles

T he Wings • The Leg

The antellna-cleaning device or strigil Th~ wiug-cleaning device .

TIlli AsDOI4&N A}/ O LTS ApPESDACF.S •

The Abdominal Segments The. Iusculaturc of a Typical Abdominal Segment The Abdominal Spiracles and Their Musculature . The . ring "nd ssociated tructures

T U C \\' OIlKliR

T UE MALE .

The l l LA D T HE TuoaA.~

T n p. . \ 8D MES

The U ale Genita li a

ITI . . YSTOIATJC CO."~IDEJl.AT10;':S . llARACTF:1l IST1CS OP Trrl: V spillQ.f

G li...'V1.I • I'ui''' I..J"NAEUs

lruetuml Characlers Tl iological ,aract. rs .\Ja\crials

GL'ItlS I' es~!"D C. G. TBoMsoN (S.L. ) lrllctUra! Otaraclers .

Biological Omracters Ct;NUS VU~lda C. G. TlTo.,sos (S. Sir.) .

~trtlctural Characters lliological Characters ~ fateri"ls

Goms Dal;c/rovDspuZa ROIIWER

LrutlUra! Characters Biological Characlers ~ [ate.·ia.]s

OSCLUSION .

[ 81

5 50 59 60 60 61 62 66 66 70 71 71 72 72 7+ 77 78 79 80 80 80 80 81

85 85 87 87 90 Q()

90 90 9_ 92 92 9+ 9·~ 95 95 96 97 97

T ABLE O F COS'TE.NTS

IV. BIOLOGY OP THE "£SPINE .

SEASONAL H,STORY 01' A WASP OLON"

H IUF."RNATION

Foo lJS AND rEF-OrNe BEnAVlOR .

Food Materials ense. l:sed in Locating Foods .

H Ullling Methods and Behavior T reatment of captllred prey .

Crilization o f Foods by Wasps _ Food o[ adull~ . Food of larval wasps

unllibaJisDl IhrrLDJ ·c ACTl\'fTIES .

The Huilding Materials The raw materials . The making of paper. T he fini hoo product

Building the Nt . Basic feantres . . leering tile nesting site .

The "queen nest" . Modification of the nesting ite Enlargement of the combs 013nges in comb ami nest uspension The fate of old combs .

hanges in the nest wall . Location of the entrance hole . Olanges in the general shape o f wasps' nests I rregularities nnd abnom1alities in nest construction

S ize ttainable by Wa.<p C lonie • . L I VE HISTORY "X D METAMORPBOSIS

11,e Egg . R rood Production Egg-Laying by Workers TI,e l ..arva .

Charncter i'il ic.s of Ihe mature larva pinning and transformation .

The Pupa Tlte New "dull . ensle Differences Proportion oi Males. New Queens, and 'Vorkers .

r 91

.. "" 98 98 98

100 103 1 ()I) 110 111 113 113 ll S 117 120 120 120 122 123 125 125 126 130 13.J. 137 1+3 148 150 152 15+ 157 158 160 Ir.o 160 161 161 162 164 165 166 167 167

10 S'OR.TIl AM ERICA~ \"I'.S P[ NE \""'ASPS

The Autumn Crop of • ew Queens and Males . £mergence Dates of Males and Queens

MISCKLLANEOUS lItainlenanc< 01 Comb for Brood Praduction

"ccession of Hrood in the Combs . Color Pattern in V pine W. ps

Color paltern and phylageny . olor pattern and caste differentiation

Teratology BIDLIOGRAprIY .

LIST O F J LLUSTRATIONS

p , TES I-LTV (\vITll KF.Ys)

I NUF.K

L 10 1

L68 L69 LiO L70 171 173 175 175 176 177 187 192 257

A CONTRIBUTION TO THE BIOLOGY OF THE ORTH AMERICAN VESPINE \VI ASPS

1. INTRODUCTION"

T he vc.-pine wasps (fanl il. .spidae, subfamily Yespinae) are repr~ scnted in orth America oy lhree well-differeotiated taxon mic groups which arc rcgar<k.d ... genern or subgenera according to the point of view. T he pr~l1t writer holds them to be vatid genera. The groups are as iollow

V rs;pa Li nn ., the true hornets. represented by a .ingle spcr.ies. Ves~a a abro Lin n .• the Brown Horner of E urope, which has bem introduced illlo the Atlantic seaboard states, whore it has become natura~zed and is sparinlily presell t. The true hornets have the llead swollm behind the eyes anJ the occlli remote r!'Om the margin of the head (Figs. 166-1 ). The Brown Hornet wmally nellts in hollow trees. though oc:oasional nests are bllilt in the ground or in aerial situations.

Do/ic hovesp"l. Robw. , a genus which includes the so-called Bald Il ornet or \\1,ite-faeect Hornet, certain spe ... of yellowjackets, and some wasps "f other col ration. Well r<presmtcd in the humid pans of orth merica north of exico. Head not sw"lIen; ocelli close to head margin; 10-m, la r space long, antennae of male · usually with tyloids; a transverse enrina on the prono urn (Fig ·. 128. 156-158). These wasps, with rare or mor or less acciden tal exception. choose aerial sites for their nests. hang­ing them froro trees, bushes, rock surfaces, and tne tiJ'nber of bridges and Luild ill~ (Figs. 205, 206, 215-221).

V cs/,,, I. T hom. genus c nsisting mostly of typical yellowj.clcets, thou,! 1 sOme . peci .. aTe black and white. Repr ""ted by several species III I:.oth humid and arid regions. Head essentially as in Do/ic/,ovts/,lIla, ex­cept that the oculo-malru- space is quite small and the antennae of the males do not bear t ,",aids. T a carina on the pronotllm (Fig . 1-6. 38-42) . The lIlem eN of this genlls are typi cally subterranean in nesting habit, gener-

• All the f;ldJ; re-porled io thil pa(lfi" concernillg the Itrudure and behavior of \\'1Ij~. or cl C!'~ling with other upecu of ~~p biololYJ except thos,e definitely accn:dited t other ~n;am. have b«n observed by the iluUtOr. Rt(er~ to the Litc:I'ilmre afC

nWlierou.1, hn \'ing been included wherever it ba.!i setmerl the poipu would hi! Imtl'rov~ th~ret.)' , hut £ailure to accredit. an observation to own does not necessarily indieate t h~ l the obscnwon i. beiui f ll:porltd here for the firit time::. It is belicnd the reader will be ::able. from U.le COlJtext. to diJi\in i!h betwee:n inIarmation which iJ distinctly new _Ild th'll which has loll&' been eS1:ablishtd.

[ 11 1 [ II

12

ally choosing a des~rted mouse, gopher. or squi~rel burrow. or some crevice,

h ' ( Lh ' I .hou ... h in ve~' rare mstances cboosmg a hollow as I e .. te 0 e.r ne> • ~'J 30) log or Lhe dark ioterior o[ a smnJI building (Figs. 223-226, 2~2 •

Fernald (S8) ~ desig1l3tes all tru-inhabiting sp<!Cles (D{)I~/II)w$pllla) d all bt·_n~H s"""ie.s (Vcs/,uw) as yeUowJ8ckets ; but as hornets an su ~ .~.~ .. -- 'ali

this cl:u ification is neither justified by the [act o[ wasr bIology nor v -

dated by cQUunoo usage. . . ' M t [the specie ' in the su family exhibIt three well-defined castes,

ai, OSc °ti'l [ __ 1_ or queens' and infertile felThl.les or workers. BoLh

m es. er e =-, . " h 'm 'a! Vrspufa and DolkhO'lJup"fa, however, contal!1 speCIes W I are SOCl

rnsites and hav~ nO worker C3.'i l . The queens of these spec.es usurp " ,tablished nests o i olher species. kill the nghl!ul qu~ns. and depend on the adopted workers to rear their brood for Utero.

• The numbers in partnmeses following citations of author! rt:(er to items in the

bibtiography, p. 17i,

[121

II. THE MORPHOL OGY O F VES PULA PENSYLVANICA (SAUSS.) ·

THE FEMALE

THE INTE.GUMENT

The int"!rUment is neneraU)' th ick and heavily sderoli,ed (see Ferris and Chamberlin . 59) , Ill" tllose parts whim must b. Lhin to llennit move­melll being membranous and Oexible, The head and lhorax are mOte h"" vi i)' sderotized than tJu:. abdomen, being, in fact. quite rigid and box­like. T he rigidity of these parts is correlated with tI .e fact Lhat they serve as foundations (or lhe powe rful and ""ried oiUscl .. wruch move the mouth­pans and locomotor organs,

T he ouly eJ<ten.<ive membranous areas in the integument are the in­le rsegm mil merobran ' oi the abdomen. the membranes connecting tit" mouLhparts with Lhe head capsule, the cervical membranes , and the mem­branes connecting the pleural a nd sternal pans of the proLhorax wiLh the r "' t of the thoro Figs, , 14, 43, 44, 114,117,131), The intersegmental membmnes of the a"domen pro"ide for Lhe telescopic movemenls of respi · r' tion, for the distension of the abdomen when engorged wiLh food or filled with eggs , for mo\'emenls in volved in oviposition , in balancing. oK: .. and for the usc of the sting. Th C Mcal membranes and the membranous basal rC!(iUIIF of the mouthparts in<:reru.e the latitude of mO"ernenl of Lhe head nnd of the mouthparts. respectively, The membran aroWld the pl eural and ~tt rt ",1 parts of the prothorax penn it a much more varied use o f the foreltogs than is possible for most insects, In addition to the foregoing membranes, lhe , "tur between the mcsonotum and the scient .. whim surround it arC sufficientl. membranous to permit tbe up-and-down movements of Lhe meSl/nOllllll requisite for Lhe operalion o f the flight mechanism.

VESTITU'.R ANO SClJLPTURlNG

T he vt:$titu~ consists primarily of two sons oi setae, Vcr)' Illinute, dosely app r sed setae cover all parts of the body and appendage except t be compound tyes, A soft pile also covers most of Lhe body, The pile is ah; l!II t [rom Lhe compound eyes, the anlennal flagella, and th tibiae and ta rsi, " "d is only sparingly present on the abdominal sternjtes, except aloog

• Duri the ~aind~r of tbi paper, rept in the. systematic 5et.uon. and in thoe. icc tion on color pattern and ph tog~ny, abbreviations o{ aumon' mune. will be n "n ed {r m the citlltiOfll or specie! names~ This will el iminate needless and uselCh ~pt:tition.

( 1.\ I t il

I

14 SOftYR AMI::II.l(j\, · VE$PIl'iE Asrs

their caudal margins, whcr~ it farnlS a delinite fringe. Various kinds 01 highly 5Jl':CiAlized ""Iat are 10 be found on the mouthparts, genitalia, ett.

detaikd study of setae is omitted. V,spula !,<IIsy/~'lI,ica is devoid of mark~ sculpturing, though much of

the body is finely punctate. The punctures arc: moderatc!y large on the abdominlll .temites but are not crowded; elsewhere they are distincliy <l11aller and are often close Iogt'ther. The punctures on the mandibles are eJoug;ue, I ,ding to produce a striate or slightly rugose condition. The punctures on the clypells a re coarser apically. The punctures on the . y­peu he gwae, the mesothorax, the mctatho r;lx, and lbe femora an! sub­equal ill -ize and medium 6ne. Elsewhere the punctures are quite small Or indistinct.

THE HEAD AND ITS APPEN DAGES

Tru: H£AD CAPSOLE

The general shape of the head i shown in Figures 1-6 well CIlough to make " detailed descriplion unoecessary. [t is of the hypognathous type, 111 , ·hich respect it !lilly be cansidaed primitive, since such relati,.ely primi­ti,'c insects as grasshoPI>ers and ston flies are hypogna thous; but in prac­ticall \" eVC,· · other characteri tic the bead is higtlly specialized. The he.~d (j\1)'~le is highly ri iei The clypeus (CII'> is immovably incorporated inlO Ihe head cap IIle as a result of the arching upward o f the cpislOrnal 11-

[ure ( cs). The head surfaces generally are onve.1y cur,·~ and are able, [her for '. I resi l gr t stresses i rom without and the pull f strong 1ll L\ cles from within . Moreove.r the h d wall is amply supplied with sec­ondary braces in the form of strong r idg .. d •• eloped internally along the principal head sutures and lines of grealest "train. The foroltw, lIIolT"u.m (I·ip. 4. , 20; For i, small, being closed below by a postgenal bridge (P!]e8r) inlerp<>sed bclw L'CIl the foramen lU1d the seleritcs of the labium.

. uturn of I"~ crall;'''": Ih~ rl'icrallial su/t/re.-This uture i appar­l'lI th' absent. al tllough a fa int impressed line (Fig. 2, fs ' ) which extends d(}\\~ tlle midd e f the face [rom the anterior ocellus to a point midway between Il,e anlennae, and two harely perceptible ridges (fs") which di­,erge abruptly fr m thi~ impressed line and e.'<tend laterally to the antennal <CIcritc ( . ..! ), rna' be vestiges of lb [rontal sutures. 1£ lhis be true, the frontal sutures are (used in their upI" r pan.

TJu occipilal .I1Ilure (Figs. I , 4, 5,6: 0<1).- This, as commonly inter­preted . is represented by two lines which extend fr m points immediately anterior to tile JlO!'terior mand ibular articulations upward between the lateral and caudal surfaces o[ the bead. They tum onto tbe verte,. but fade ut before meeting. As far as the I vel of the upper end at lhe com-

f 14 )

Tne. 1I0RPU GV ., " VESPl1L.\ P"NSYLVA.·I,,' · (SAlJ"SS.) 15

pound ey these lines are impresse and are margin~ posteriorly by minule carinae.

Th. posloccipilal Slllu," (Figo. 4, 7, 10, 18, 19, 6J; ,ros).-This forms the dorsal and lateral parts o[ a groo,'e which completelv encircles the foramen mag mllll but is somewhat obscured by secomlary' devc!opments. Be[ween the posterior tClllorial pits and below the foramen lDlIb'Tlum the hal,·CjI of Ille groove ~"Ur\'e vent rad to meet in the poorly de/in~ SUN" or the postgeoal bridge (Pg~8r). Tbe parts of the "postoccipital suture'" be­t\\" en the posterior tentorial pil (1'/) are doubtless, in pan at least, homolo­gous with the gular .ulures of such insecls as beetles and, the refore, with parts of [he hypo! lomal sutu.res of geoerali%~ insects. Internally (Figs. 7, 8) th postoccipital suture is marked by " postoccipital ridge (POR) which i · low alld indistinct dorsally but is high laterally and ventrally, where it is cont inuous with the posterior roots of the ten1orium.

Till' .rulurBS and ridges of Ih. pm.rto",.,.- Secondary developments make il difficult to homologize in their entirety the sutur s and associated structur surrounrullg the mouth with those of a generalized insect such as a gra! hopper. The [ollowing conelu ion., however. seem warranted : The ~ti.<lo",al su/"r. (Figs. 1, 2, S : ts) fonns the dorsal and late ral margins of the dytcKS (Cl,,). Through a part 01 its course it is confluent wilb the margi ll of the compound eye. The arms of this sutu re run ventrad from the margin of the eyes in such a man.lll" as te> cut off the cranial condyles ()[ tlot m.wdibular articulation (C) from the dypeus. The "um'os/omoJ SUI!" . Fig . 1,2, 6; f's). as identili~ by the cou.rse of the .rubgelUll ridg, (Ftg, . 7, 10, 1 1; SgR), partS from Ihe epistornal suture where the latter I .",,,,,, Ihe eye margin, follows the ,'mind margin of the eye, and then passes d' ''f:ollally backw..1.rd and downward to the posterior mandibular arricula­ti<'l'- It thus fonns the boundary between tl,e oculo-maJar space (OM) and t·h. gena C.). The hY/Jor/o,"oJ slllllrc (Figs. 4, 7, 14, 19) is rufficult [ 0 iden 'f}, though it appears to be represented in part by a groove (lis) wh' ch arch ,. r the ral fossa n the back surface of tlle bead just ex­ternal to Illt! "-,,!'OS/OlHal carilla (HC) which margins the fossa.. Above the [ 0",,, this "hypostornal suture '" meet its fellow of the opposite side. and the two nre continued upward :1>' the median suture traversiug" the l ost­gtlll i bridge. Abov" this they diverge to the po terior tentorial pit •. as ha, Leon noted abov/!. The pleurostomal and hypostoma l sutures together or,' ,Oll1m 'lly r rerred to as the SllbgNloJ .ruJttre (Fig. II. gs) .

The el'islO'IIoJ rid!]. ( "gs. 7, 11 ; ER) is low and ruund~ in its dorsal purtinn but rises high lnterally and becomes con inuous with tile brand Illl/(rior lentorial o,."'s (Fil.'5. 7,8, I , 11; ,-IT). The subgeoal ridge ( gR) IS well de\'c!~ped on the pleurO!\t lla! suture, is replaced by a simple inllee­flon of the crolnial ma rgin vL1Ilrally beneath the gena and postgcna, ruld is

r 15]

16 SQRTU AW~CAX V I': F' I N'E WAS.PS

not developed at all un the p'.rt ,,( the hypootomal SUiure which arches over the oral (0 sa.

Tile OCilla,. SIIllIrcs.-The margm of each compound eye i.o Ilrrounded by an ocular suture (Figs. 1.2. 5.6 : as) along which the utid" i. infolded 10 [oml a high knife-like oCilla1' rid9~ (Figs. 7, 8, 10. I I; OR) internally.

T lte 01.",,"01 SIII"rts Figs. 2, 5,9; os).-The base of each antenna is

surrounded by groove. the 0111<11/101 SU/UT' ,' but there is no prominent internal ridge developed along the ulure, such as is described by Snod­grass (130) [or insects generally. What appears on the ""tal aspect of the craniwn to be such a ridge (rigs. 7,9. 10, 11; AR) is the wall o( the anten­naI socket fanned of the inflected antennal sclerite.

Tilt slIbonlmllol Stl/Ilres (Fig. _, sas),-Thes cxtl'.nd from the antenrol sutu r downwanl to join tbe cpistomal uture. Internally the subantennal ulures an, marked by in flectiollS of the cuticle. the subantcnn.al ridg

(Fig. 7, SaR), which buttr . the r ts of the anterior tentorial arlll . The I 1Ilorinl pils.-The cmnial udele is invaginated along the entir

latcral ma rgin of the dypeus (Figs. 7. 10. 11) to form the anterior nrnlS (.IT) I the tentorium. The in\'llgination, however, is open as a de!uule aM.n", "'II/or",1 pit only at the dorso-Iateral angles of the c1ypeu (Fig. 2. aI·· . I II comparable omnner the entire lateral portions f the po.toccipilal , lI LU re are continuow; internal ly with the posterior roots of the tentorium (1 ';';(0. 7, 8, 10), bUI recognizable /,os/trior lelllorial "its are limited to the ,·enrro-later.1 ;lOgles [ the postoccipital uture (Fig. 4. (>I) .

Til, rrQlliaJ artos.-The distinct areas of the cran ium, as definetl by fllnd.mental morphological fealu r s, differ in many respects from those commonly recognized by the taxouOI11i t. !!Ven when the names are usee! fll r areas occupying approximatcl. homologous po itions. Tn the fol ­lowing treatment the crani I area will first be characterized in l 'mlS of f undamenlal morphology. Then in such cas as merit pecial treaune" l the areas as cormnonly recOj(1lized hy the ta ' Ollomins will be defined.

TI,,' [rOIlS (Figs. 2. S, 6; Fr ).-In Vespliia pVlIS)'/vollirlJ. owing to the "b, '11' of r'C gnizahle f r lIlal sutures, the la teral limits o{ the t rue irons are undefi lled. It includ the a.rea about the anlerior ocellus and that be-1"\,"""., the antennal bases and the dypeus: but how much more is tUlccrtain. If the median impressed li ne . f.r' , and the (aint ridges, fs" . are vestiges of the {rontal sutures (see I" 14). then much of Ihe frons 35 it exi ils in g""­era li l-ed insects i. here absenL Ii the impressed line, fs' . and the divergent ridge, t.<" . are secondary (catur , then the (rons is doubtl " mOre exten­sive and tile [ronlal ntures are entirely absent. I n ta xonomic litenllllre nn wu "J.lS thc term " (rOll!!" ;, commonly applied to the broad and laterally un­bounded median facial area ext~ding from the c1ypens upward b tween the antennae to the anterior ocellus. A part of this area. fann ing an ill-

116 I

"t llE 11 KPlIOLOta: or " V.&SPO'LA P£lo..'SVLVASf .. (~1.: :. ) 17

defined triangle (Fig. _. 5, ial) with the apex directed downward between the antennae and the id marked by more or less evidc:nt ridges. is ele­,'at«l .'ligbtly above the general level o f the face, eope.:ially below. and is oilen r fc:rrc:d to as the inlrranl''''lary tritUlg/' or the i"t,raJl/rnna[ area.

The cl.l'P'"s (F,g . 1. 2. 5.6; CIP) , which i8 well shown by the figures. is markedly com'c.'<. The apparent ventral border, which bears two promi­nen t teelh. i not the true ventral margin but a pordon of the disk that IlroJt:CI. rlownward over the ma rgin so as to conceal it completely and to ,'ol'cr tile broad basal part of the labrum , well. This condition develop •• olCCOrding to Kirmaycr (SO) . during the prepupal s\3ge. at which dme the lahrum of the forming pupa becomes drawn up beneath the lower pan of Ih,' e1ypeus. The labraJ s"I,,,, (Fig. ll, Is). as a result. runs behind the c1ypcu> just below tbe level o( the compound eyes.

Til . parielal scl.rillS (Figs. I. 2, 4. -. 6).-TIt<:se exhibit no special ieature- in V"/' lIla e:<cept that, o\ving to the absence of definite ronal ,wd irontal suture ', they a .. continuoWl dorsally across the vt:rtex (V ) ami anteriorly aCTOSS the frons , c.xcept where limited below the amenn ... , hy the subantenoal . utuJ'es. The spatial rc:lationships of the bounding and includ:d 5t ructures, however. di"ide the parietals into several areas which bear di,tinct names and are taxonomically significant. The area on tlte ver­lex hearing the close-set ocelli (0) is called the oallar Iriangle. T he vrllu (Gel i, limited to the comparatively narrow strip OIl the side of the hea,1 behind the compound eye. The whole of this strip or sometimes only part "f i immediately adjacent to the eye is often r 'ferred to as the pos-1,'/'io1' orbit. The small ocu/()-molar space (OM) (sometimes called the (' h"okl below the compound eye is often considered a pan of the gena but apI'ea act II) -i f the writet 's idtntiJication of the subgenal ridge (Figs. 7, 10, 11 ' SUR) is correct-to be the ,,1.II,os/omll. On the face. the part of the rictal. adjaCCJ1/. to the compound eyes and filling the emarginations of Ihe Inuer are called the Olll.rior orbits. Betwe"" the anterior orbits and the intera tennary triangle of the frons are broad. shallow. trough-l ike de­pressiolJ.; which run diagonally upward and outwa rd from the antennae, becominf: hallowet as they d so and di ppearing before the compound OJ'CS are reached . These omy be called the Olll.' lI lIaJ slilci.

E xtremely narrOI oclilar ulerilts (not hown in the figures) and ' um",,!t:ll wider o.nd carinate allloIH.al se/mlcs (AS). which are reflected int', Ihe antennal socke to make bearing urfaces fo r the bulbs of the an­I/'uuae, are cut off from the parietals by the oC14iar s"'''''~J (as and the 0111<·" ,,01 sutures (a..r). respectively.

The /los/occipul (Fig. 4. Poc form a ring completely encirclin the [01 ameu magnum. Above the £orn.men it is reduced to a vcry narrow rim, lout Iaternlly il broade. and bears the rounded occil'ilal cOlldylcs (Oa).

I 17 I

NOR-Trt Alif..RfCAS' VESPINF. WASPS

The Inn below the foramen i. nol truly pootoecipit'll in c.haract.:r but is homologous with the b3se (at least) of Lhe gula of .uch in>CCt ~ beetles, and with a pan of the neck membrane of generalized insects.

Til. occip,tal Qrel, (Figs. +, 6) .-This arch, comprising dorsnUy the occiput (Oc) and laterally the pos/gena (Pgc)-these region however being continuous B1ld nOI 5ep.arated by utures-is greatly expanded and forms rno t oi tile ca.udnl ,uriace of the head. The upper P;lrt of the oc­cipital arch surrounds the postocciput. The lower part embraces the oral fossa nnd is excaval .d by the [ossa almost as tar :to the postoccipitaJ ring. nle oral fossa i. a larg" roughly cmicircuJar pocket on the back of the ranium that receives the Jabio-ma,,;Uary comple" wben the complex is

folded up. Between the oral fossa and the postoccipital ring. mesal exten­,ions of the postgenae mtet in the midline to form the posl(J~lIaJ bridge (PgtBr). This bridge, app.arently because of its po.ition relative to the moutllp.arls, is called the hypostomal bridge b)' Snodgrass (l.17); hut as it onsi t cbitOy of po tgenal structures, though it does indude a hypostomal

cnmponent, the term postgenal bridge seems nrore appropriate. In an earlier paper (130) • nodgrass ~a1 ls this bridge the llOstgenal bridge.

. \ bove the oral fossa the caudal surface of the head is moderately conc:,,'e and adapted to fit over the front tlId of the tho"" . Immediately >urroundin!( the postocciput the head ""rfa~e i mQre det.'Ply ronca,'e tI"," ehewhere. This p.art is set off by cun,ed dotted lines in Figure 4, the hnc:> following ra ther abrupt changes in degree of cOllca,-ity. For a ,hal' t di.t:t.nce along the dorso-Iateral boundaries of this morc sunken central arc:t there is 011 ~d, side a definite but shallow sulcl (Fig. 4, LaS ).

TI,r s-ubgr>laJ orCIJ$ (Fig<;. 1. 2,4,6, 7, to, I 1).- 1'he /,lruroslo'lId, as "Ino:ttly I' iuted out, forms tbe SO<aIJed omlo- n,aior s/,aa (Figs. 1, 2, 6 : o . J). The hypoS/olI/aia (Hn) are reflected internally and cephalad from the CllUdal surface of the head to form the lateral and dorml walls of the oral fo!>"". The <Iorsal wal l of the oral fossa is continuous along its caudal umgin with the processes of the postgc:nae which [ann the po t­genal bridge. Arou nd the margin of the oral [ossa, just internal to the hypo"tamal Rumre, the hYPDstomata are produced to form the arched 1r_'PoMoRlal cariM (He) .

Tlrl! /(II/ori",,, Figs. 4. 6, 7. 8, 10, Ll).-The tentorium is compl .. " in foml and highly specialized. Certain details of its structure wiJl be pointed out in the discusRion of the various mu cles which find their origin on it, but the general features may best be given at this time. The tentO­rium consists almost entirely of the coomlo"sl), developed anterior arm , since the /Mlorial bridg. (TB). whith develops by the fusion of the posterior arms, is reduced to a very slender arch located only a littl"

[ 18]

THE AtOKJ'llOLOCY OF " VESPlJLA l'E.NSVl..VANICAo

• ( AUSS,) 19

way in front of the forame n mal(l1um. Dorsal arms, if they are to be considered as pr ent at all, are .-presented by tile triangular processes ending in .Iender weak fibe", located on the tcntorilll = iu,! ce~halad o i the tentorial bridge (Fig. 10, DT?) . V,ewed trom below (Fig. 6) the tenlorium fol'lnS a truncated V the apex of wbich is directed back­ward. From in front (Fig. 8) it appears as a broadly rounded but

le<liall .. incomplete arcl , on which re"ts a smaller and more slender arch :In<le ~f the tentorial bndge, and from which lateral buttresses, the I ten. r tentorial roots, rise upw~rd and b,1ckward to the postoccjpital ri dge (POR). In lateral ,-lew (FIgs. 7, 10) each arm of the tcntonum Ilan, .. [anwU;e both anteri rly and po!>teriorly. is narrowest a triAe tlVer one-half its length from the clypetls. and appears as if twisted so as I rolal one of its ends through ISO" The junction of the tentorium with the epistomal suture e"tends from just above the anterior mandibular art KuJauon to just below the antenna'! base (Figs. 7. 10,11). The junction WillI the po.tocclpital suture invol\'es the entire latern l part of the suture (FI);>. 7, 8, 10) .

. lrfJiJinna/ era";"J siruclllru.-The labrun l. ille epipharynx, and the hy-1'''1'1 It)'nx, though developments of th: hea~ wall proper and not a~pen~c­ub r in nature, are so inti lflatel)' assooated 10 &tructure and lunctlon WIth lilt lIlouthp.art5 that a discussion of them \\il1 be postponed until the mouth­part; are considered.

TlI . lIet.D PPENDAGES A, 0 TUEm MOSCULATt!1lE

In the treatment of the musculature of the wasp in this pape.r the nm d es arc numbered consecutively for convenience of referencc_ (The Iluml"'r, a signed have no other igoificance and do not correspond to th. numb.", employed fo r insect muscle~ by other authors.)

Tlrr un/nmae (Figs. 2, 3, 5, 6, 13).-The general iorm of the 12-seg­lIlent ,I antenm and the proportions of the segments are apparent from an examillalion of Figure 3. T he ullJ of the scape (Sc) fits into the socket­like Ulfltlltwi ulmtc AS) to constitute with ille latter a all·an<l-soeket !"int. \ -entro-lateraUy Ihe rim o[ the antenna! bulb bears an inwardly !lir,· -tl'fl, >talked, ar/i,"/ar cup (Fig_ I ,ArC) that fi . over an awl-shaped p,."cc_. of the antenna! sclerite, the an/c,IIIi!" . Fig. 7, 1\, 14; 1If).

Th antenn3 as a whole is moved by four muscles F igs. 13, 19, 20, 21. ~3). These nre an U'ltrnal d.pressor (rad[l])- and an in/eNlal de-

• The: arabie. nu~. in brac.ket3. which follow the abbrmations reJurinlf to the dr;a.winp of thr: muscles, form ~ C0I1UnUOt15 series. each n:ferring to a particular nw It.. These numht:ra do not correspond to lhe numbers etnl>loyeci by 3ny other :ImhOf (rw the desJ~ l1.3tioo or muscles. They are included in wil .PRper iOlcly lor con­vtniWct or rderence.

[ 19]

20

prusor (iadL2J) inserted in notches in the rim of the antcnna! bulb jl1.5t laterad and mC$ad, respectivdy, of the articula r cup, and an erlernal Iln'Olar (nal(3J) and an inlt!rnal levator (ia/(4J) inserted on the upper part of the rim of the nntennal bulb. All the antenna! museles have their origins on the tentorium. That of the ute mal depressor spreads fa ll' wise over n leg of the tentorial bridge and the mesial surface of the , adjacent posterior tentorial root. The external le,-ator originates on the hinder portion of a mc:saJ extension of the tentorium (Figs. 8, 20) close to tlte po.terior tentorial rOOt and just eOlrad of the origin of the

ternal depressor by which it is partly surrounded. The internal de· preo:iOr originates on the upper concave surface of a mesal ly dire :ted Aanl:" of the tentnrium ju·t ventrad of nnd anterior to the origin of the external levator. The internal levator originates on a lat rally di rected flange of the t ntorium about midway between the anten r ,nd posterior roots of the latter.

'Ole mo~e01ents n13de po 'ble by the mode of suspension of the antenna nod the arrangement oi its muscles are numerous and '·aried. Any one muscle acting alone produces a rotation. Movements of le,-atioo and depression, promotion and remotion are accomplished by combina­tions of the rou 'des actillg in ullison.

C,'IIUal ,"alllres of Ille ",oulilparls·-Th~ mouthparts of Vcsp"la (Figs. 1. 2. , 10, 11. 12, I 7 inclusive) ub..,rve two di till Ir dif­reI' t sets f junction., a fact whic\J is reflected in their structure as learly as it ib in their operation. T he mandibles, on the one hand. fUIlLilon

primarily as tools and are only very indirectly concerned with alinlenta-lioll. The ma,''lillae and labium, on the other hand, are concerned almon solely witl, a li llleLltatiun alJd are surrounded and bound together by folds And . he·1s of membrane (Figs. 17, _ , __ , A , 26). T his membrane COIl­

,titLlles wha t bas been called by t:lric\J (J48) the "mouth-fidd-sack" (.; !JIJ'df~ldsack) . In this paper it will hereafter be referred to as the " raJ fif/d. The oral field i' developed from the oral margin between the ""Ulriibles in suc\J a way as to make maxillae and labinm quite indcpend-nt of the mandibles. In consequence the mandibles play no part in

bounding the mouth open;ng or in foml ing the cl1anncl through which food is brought to the mouth . [nstead they funct ion almost as if they wer serondary structures added to the head after the organs of inges­tion were completed.

TIl • • Ila .. dibl S Figs, i . 2. . I , II, 14, i8, 19. 20,33,34, 36).-The

• All rdenncu to position or d.in:cti n r palU in thi! section (cephali~, dorud. <1<.) •• sume the mouthparts to be extell<!ed b<nc.th the h ... d all<! the head to be in il:l normal yutic:aJ potition.

[20 )

Tn:~ JdOR PllOLOGY OF "\' &sPULA P£NSYLVASl CA" (SJ\ USS.) 21

strtlct ll r~ of the mandibles (Md) eminmtly fits tbem for their functions. • tool they are used for cutting Or scraping vegetable fiber [rom wood !urfaees and weed stems, for cbewing this fiber mixed with saliva into a pulp, for fashion ing the pulp into paper, and for atlaching titis to the n .. !. They ar~ used for carrying objects, as weapons for the capture of pre)' 10 be rcd t the larvae, and as implements for cuttmg up the prey and converting it into a sort f h.-unburger. They are used as clippers ior tearing away tl,e inner nest layers as the nest is enlarged, for ' rim­ming tile ragged edges of eUs from whic\J young adult. bave emerged in preparing the cells for new occupants, and in cutting off leaves, Slicks, and roots whic\J are encountered in enlarging the nest. They are used also a digging tool in the "nlargemcnt of the underground vities in which the nests arc buil t, for with them mud or earth is scraped loose 8wl moulded into pellets to be carried away, and pebbles are pried loo.o;e i rom their setting .

Owing 10 a diagonally truncated apex and to a mther abmpt cun'a­ttlr around a diagonal axis running from the anterior articulation down­\ 'llI'd and had.-ward to the postero-inferior angle, ac\J mandihle appears roughly ' riangular when vi wed from in fro nt or from the rear (Figs. 2 8, 11, 33). In lateral aspect it is trapezoidal (Fig. 36). T he truncated

,in . the cui 'ng margin, bears three teeth and 11 notch. B hind the.., mnrg inal or primary teeth. and alternating with them (Figs. 10, 11, 3+), arc tw secondary teeth and a molar shel f. The two mandibles, when bcinll' used, meet in a ertical line beneath the e1ypeus. In repose. one is folded OV«:.r the other (Fig. 2 .

T he articulations consist of an acolabullll" (Figs. 33, 36 : Ac) . located anteriorly and fitting over a condyle (Figs. 6, 10, ll : C) at the lowe.r an~lc Q\ the clyp us, nod a mandibular condyle (Figs, 34, 3 ; MC) 10-cat",1 posterioriy and fitt ing in to an acetabulum (Fig. 6. Ac') on the c"miunl at tI,,, lower end of the occipital suture. T he hinge membrane Ixl ll"'on the two articulation is concealed from view, wing- to the in­fl ection of the cranial wall below the gena and the O<: lI lo-malar space Fig . 8, I I, 21 ).

The mandible arc operated by two muscles : an enormous "'alldib"lar ,,<.I,/lrelor ", ,,sci, (adtlld [S]) and " mther small ""mdibular abductor "'"sd o (abmd I6 ] . The adductor nmscie is two·branched. The inner arlll smaller branCh is again divided. The fibers of the inoer and mailer branc\J originate on tbe vertex and the mesal por 'on of the occiput (see FiR'· 1 .20). The outer nd much la rger hmnc\J is lobed, T he origin of it, fiber extends over all but the lower fi fth oi tht. geM and postgena • over the latera.! pa rts of the occiput, and Ol'{,r tlte posterior surface of the o<:ubr ridge margining the c I11 pound eye. The tendons of the two

[ 21 I

22 SORTO A1UJUCAN \r5S'PINE WASP

branch .. 01 th~ mandibular abductor unite to lonn a large common telld"n (Figs. B. 11, 33, .H, 36, apd 5) which is inscrtw ioto a cotcl, in a mesal bulge of the inner rim of the mandibular base lhrough the inter­mcdiary 01 a small accessory scleritc (Figs, 33, 35, AMS). The mandibular abductor originates on the lower 6fth of the gena and postgeDa and is in.erted by means of a =11 tendon into the outer rim of lbe mandibular ~e a short distance in front of the posterior articulation .

Till labnm' alld cpipiJaryllX (Figs. 10, 12, 16, lB, 21. 23. 24, 25, 26),­The la.lm",. (Lm) is small OJld i. largely concealed behind the dypeus. It is about hnIl as broad basally as the dypeus, though Lhis broad ponion i. very 'hort. and it narrows abruptly into an dongate spatulate mWian proct:S. that is carinate medially on its anterior surface (Fig. 16), This pmc .. , projtClS beyon the clypeal margin (Figs, la, 12) and is visible an­teriorly \I hen the mandibles arc opened, The mandibles close in front of it anu concl:al it at other times, The base o[ Ule labrum, except a marginal strip. is ut weakly sclerotizw, or even membranou, The marginal strip. like the median process, is heavily sderotized, The Illbrum f",s no mus ' e:5.

The ~/>if'lutry"x (£pl.y) consists of n membranous fold arising just bdUnd the labrum and at its base. The sides of the epipharynx form rtllmdw. wing-l ike lobes that cover the angles of the moutil. Its middle portiOlI is produced into an elongate , rapering, and pointW prece which lies just behind tbe spatulate process of the lalJnun.

lletween tbe lateral angles of the labrum and epipharyt1l< on "ach side there i, a !iffiall I riJlllguL1r sclerite whith may be called the labral Iriangl,. ( Fig , 12, 16, 24. 26 ; LTr) , It is thickenw and strong on two wges uut IJ;l$,!;es into membrane On the otheL One angle 0 r the triangle hooks uncler the end of the marginal strip on the labra! ba.;e ; One extends laterad in mem!)r"ne to near the anterior articulation of the mandible: and the Uurd exlends downward in the rounded lateral wing of the epipharyn..'.

Each wing of lbe epipharynx contains in its posterior <Jl" ventral lamina a slellder, curved , mesally tapering, weakly sclerotized strip, the , />if'llIITyng.aJ bar (Fig . 12, 21. 24; EBa) , which, when tbe mouthparts arc appro»imated , fits down against a similar weak sclerite. the IIJ";"iai. bar (LcB, see p. 31), in the Iscinia. At such times the epipbaryn.'C com­pletes the roo! Over a temporary passagewny along which food is brought to the mouth, the rest oi the roof being Conned by the laciniae and the galeae, Li ke the labrum, the epiphal)'tL'C is devoid of musculature .

T/w labio-tllllXillary complt$ (Figs. 8, 10, 14, 15, 17. 18. 20, 22~2, 35, 37) ,- [n Hymenopttra the remainioj! mouthparts, the maxillae and the l.billlU. inslead o! developing and lunctioning more or les. sep.1.­rately. are united into a complex organ of mutually adapted and co-ordi-

[ 22]

23

nated part. that is "'pablc of extension anti retraction as n unit. This compound structure is ealled by Snodgrass (137) tbe "maxillolabial ""rupl""." but the present writer prefers to use til~ term " labie-maxillary cmt1pl.,,:' for lbe sale reason that it is more easily pronounced. The labium, as commonly identine.,] in II esp.tla, is incomplete basally. both the mentum md submenrum being lacking and thcir places being taken hI' membrane (Figs. 6. 28, 30, 31, 35, 37), T he maxillae are joined t~ lh siues of the labium by strips of m",nbrane, At the ba.~e o{ the trm,clltulll (Pmu) , the only well-developed a."ial sd<!orite of the labium, t he maxillae "'"c closdy bound tcgetiter On the midline Ilj a small patch 01 tough membrane (Figs. 8. 28) connecting their carcline-stipit.11 binge'S \\~th the prementum, It is this feature which insures that in all movem.enLS oi exlellliio" and retraction the ma,<illae and labium . hall move together. ,\rticulation of the labia-maxillary complex \\~lb the craniunl is eft'ectL-d solely through the maxillary cordill,S (Cd) .

Tn this connection attwtion should l>e callw to the fact lbat the basal sclerites of the maxilla (cardo and stipes) and those of the labium (sub­m"ulum, mentum, alld prementum), as has been pointed out by Kir­OW'-(!l" (00 , Snodgrnss (130). and others, are not consecutive segments in the axes of appendages but are merely sclorites developed in the rall, 0 [ tl,e organs coocernw. The basal parts 0 f the maxiU .. e and

labium 31'e largel)' non-appendiculnr outgfowtlts f the cranial wall to whkh tltey are broadly joined. Their interio ... , therefore, communicate i reel~ wilb tho main head cavity. T be true appendicular structures iD­\'ul:w in the maxillae and labium are representw chiefly by the palpi,

The latera! parts o( the oral 6e1d (Mulldfcldsad of ulrich, see p. 20) femu two folds. one IYUlg on each side of tile mouth and bounding lat­rnl ly the preoral cavity. Whenever the laLio-maxillary complex is ex­

tended for feeding, these [olds of tile oral field arc drawn !on\'ard. by a mechanism presently to be described, so as to e.~lcnd the preoral cavity apprecinoly,

Tilt labi"", and oss(JciaJ~d stn.cttlres (Fig,. B, 10, 18. 20, 24, 25, 26, ~, 30, 31, 32, 35, 37) ,- The principal derile (Prml) of (he labium ha. l>een variou,ly interpreted in the past, In taxonomic literature [""­trail)" it has bee" reierred to as lbe mentum. Tt bas also been interpreted as repreo;en ing the mentum and submentum h.sed. The faint grooves n its

urface (Figs. 26, 28) which divide it into a median, more distal area, and a broad latcral band batt ","ouod the lnat'!,<io of the former, have been interprt!le(1 as the vestiges of the utures along which fusion took /1io<: •• the median :rrea and the lateral band being , respectively, the sup­l><>set! mentum and submentum. Recently, bowever, nodgrass (137) has d.'innel), shown lbis sclerite to be the premcntum. as its posterior dorsal

[ 231

2·~

margin serves for the iru<ertion 01 the paired posta-jor labial adductor mwc/e.r (Figs. 18, 20, 21. 5, 26, 28, 32, 3S; plad [7 D. muscles which are ruways inserted on the prementum. And Liu (84) has shown that lhe submentum in many of the higber Hymenopt~ has been reduced to membrane and has thus disappeared a. a sclerite. The grooves on the outer surface o t the prementum (Figs. 26, 28) are merdy exterru>.1 indications of lhe boundaries, on the inner surlnce, between the areas of origin of U,e anterior and th' posterior fi.,.'Cor muscles of the ligul.a (Figs. 25, 35, 37 ; afli, flfli) . The posterior labial adductor muscles are the innermost of three pair of muscles (Figs. 2. 14, 21 ; plad, Irm, fist) moving the labia-maxillary complex which origiru>.te 0 11 the dorso-Iateral parts 01 the dypeus. From thcir origins the posterior labial adductors p:u , backward through the head cavity, one 00 each side o( the saJifH1f'y duct (Figs. 8, 18. 20, 2 • 26, 30, 31, 32 ; SID). and converge to the single stout tendon by means of which they are insened on the premen· tum. T he insertion i located in a deep median notch.

T he prcmentum is a stout sclerite having the general hape of a grocer 's , ugtlr scoop or an inverted sunbonnet that has been cut away hehind to form the notch in which the tendon of the posterior labial ab­ductors i' inserted. T he dorsal. or pro. 'maI, lIlargin on each .ide is joined to the max.illa by a broad strip of membran". except anteriorly. where the membrane is largely replaced by a small, irregular, pivotal cleri!:e wh icl. m,y be ternled the lalrio-1HQ,ri/lory illo"'" ( Figs. Ii, 26 ;

I. ",J) . T he latter ap~rs to have a fourfold {unction: Its outer margin nr ti rulates ,,~th the stipes. its inner margin with the prementuD1. Its stipita-clistai angle articulate with a small pivoLal scierite ( Fig. 17, /.(P ) , supporting th base of the Iacinia internally, and a point on i r curv distal (or anterior) margin articulates with the lower and mOre poster ior end of a slender, slightly 5-cur"ed, rod-like sclerite (Figs. 17. 22. 2·1. 26. 32; LaTr which may be: called the IIWral tractor. This litt! . I 'rite r llns upward and iorward (rom the labia-maxillary jugum through the lateral iold ' of the oral field (Fig. 24) to the lateral angle of the epipharynx, behind the labrol triangle. When the I~b i o-maxillary com­plex is extended, the prementum and stipes pull on the Iahral tractor, which rotates the labral triangle forward. thus IHting the pipbaryn and pening the mouth and at the san,e time extending the oral field . Cateroily the labral tractor articulates close to its upper end will, a curved. somewhat spoon- haped, weak scierite, the Iroeloral suspeltsor (TrSp) , which runs laterad through the oral field to the vicinity of the anterior mandibular articul:ttion. It appears that when the labia-maxillary complex is extended the lractoral suspensor i ' warped and a strain is developed in it. When the labia-maxillary comple.'" is retracted, the ten-

124 J

-rITE ",oRPuor. G O F " ~OLA PP.NbV I.VAI'lCA" ( ... ,,55.) 25

. ion in the tractoral suspensor aids in restoring the structures around the mooth angles to their original configuration.

laterally the prementulTl is '" tended cephalad in a broad 51l3tulate process Figs. 20, 25. 26, 28, 30, 31, 32 ; SPrint) braced at its ape __ .: by an inwardly reflected margin (Fig. 32) which serves as the surface of origin fo r tbe muscles of the sali • .'ori"," (Figs. 18, 25, 30, 31 ; Slu) . An­t riorly t.he p remcntum is roofed over by the hypoph.ry ..... (Fig. 18. 25, 26, 30, 31, 32 : HpJry) , the lightly depressed and trough·like anterior , lI ,{ace f which is quite membranous from the tip backward to just I .. d 01 the spatulate lobes of the prementum. Along each flank of the hypopharynx runs a band-like sclerile that is only moderately rigid in it' posterior half but is well sclerotired anteriorly, and which bears on th boundary between these halvcs a comb-like row of long, stiff, mesially directed, appressed setae which may be: called the hyfJOfJlrar'J1lgtal 1'~~lffl (Figs. 26, 32; HyP). Behind the apex 01 each flanking selerite tht're i another, nnd less di tinctly bounded, lateral sclerotic portion o f Ihe hypopharyngeal waU. T his ne. which will be more fully desenDed later, is invaginated at the tip to form an internal, button·like structure, the h)'popha,yngtal bulIDH Figs. 25, 30. 31 . 32 ; H)'B) . th.t rv for the insertion of a long cr.miaJ muscle (fpgl which functions as a flexor of Ih. parag/ossll.

The anterior wail of the hypopharynx at its proximal end i in vaginated immediately below the mouth to fo rm a more or less globular gnathal t Olfe" (Figs. 10. 12. 18, 21 . 23. 2; . 26 ; GnP) termed by Wheeler (161), 5ince it occur also in ants, the illf ,abuccal (/1111>11>", and by Janet (75 the poch. gna/hal • . Its lining cuticle is lightly sclerotized and there lore bas sufficient strength to hold the pouch open . The mouth of the pouch is a nor w. transverse opening (Figs. 12, 18, 21, 2 ). The gnathal pouch eryc:> as a receptacle for dirt, detritus, and . olid matter stra ined out of

the iuO<.l by a fringe of setae, the oral peclen (H gs. 12, 2S; OrP), pro­jCCl ing 0\ er the slit-li ke true mouth. It probably receives ruso dust and d. bris which th wasp removes lrom its body during cleaning opera­tions. Presumably the wallp b· some m"""s o f everting the pouch oCt:aJ!ionaUy and ""'ptying it; but there are no muscles attached to it and the writer has POt di,;covered the mechanism which will empty it.

Distally. at it. lat ral angles. the prementum bears the four·segmented labial pnlpi (LbPJ . Each palpus is moved by n single, thin. Clpering, lan­like mllscle (Fig. 26, dlbt [8) that originat 00 the spatulate process of tile prementwn and is inserted on the lower or posterior margin of the pnlpal IJ,,-'ie. It is therefore a d.p,usor of /ire pal/,flS. The paJpal base is . us(>Cnded by its upper r anterio r margin from the 'p f ~l slender p=ess oC the prcmentum projecting over the palpal socket. When the

( 25]

26

paJpu, i. d.prc~L-d by its lllU5Cle, this process i bont : and when the muscle relaxC!. the elasticity of Lhe proet. ... rai.e. U,. pall'us to ils former po.itian. Distally betw«:n the palpi. th pr",nenlum bear U,e liQula . Th · ligula is undivided basally on its ventral or po5l,,"or surface (Fig . 28) ; but on it! anterior surfac. (Fig. 32), and apically throughout, .t is dil­(crenlillted mto well-defined lobes. a large median lobe. the lill!l"a (Li,,) . amI a pair of slender laternl lobes, the paraglossal (Pgl). The lingua i. fnnher parted apically by a deep notch into divisions that are commonly interpl'ded as U,. glossae (GI), though Kinn.1yer (SO) hold. this mter­prelation to be unjUhtifiL-d. and sa)'s the di\'isions an.", during the ontogeny of the mouthparts, as purely ondary developments on an originally Slrictiy Ullpaired structure. Sinc~, however, tile lingua could hardly have evolved irom anything oth.". th:ll . the glossae of primitive insectS. the use of the tenn glossae for the lobes of the lingua sccms quite permissible.

T he base of the ligula is covered both anterioriy and posteriorly by l\ elJ.<le\"eloped sclerites.. On the po terior surface a broad, modCJ"'ately sderotizetl ate, which may be called the ligular I,j.g~ piaU (Figs. 24. 25, 21), 28. 30 ; UHP), articulates proxi,naUy II) the rn.argin oi ch~ premen­tum :lIld pl"Qds laternlly acrOSs the entire breadU, of the lij,'llia. Distlllly this sderite passes by a rather poor!) defined border into a di k-like sclcrite. (PLiP . with ouhvardly reflected laternl margins, that over the base of the lingua. Th' may be temed the "os/trior lingllal pial.. n' teriorh' tbe Hegual base hears the an/trior lingual tlale (A LiP ) . T he linguai Jllat~ articula~ with each other laterally by a narrow process of the anterior plnte which passes around the lingua, in the cldt b [wee" it and tile paraglossa. (1:il.'5. 25, 30) . Proximally the anterior linguru plate i,; cnntmuous with U,. floor of the salivarium, in the .ide walls of which the proximo-lateral angle> of the anterior lingual plate arc upturned into hun truncated lobes (Figs. 30, 37; TL) which serve as insertions fo r

some of the r~tra<:tOr muscles of the IiI:'t1a. The base of each paraglossa bears anteriorly a hairy, lozenge-.h.1ped

puragiossal scl"";/" (Fig" U. 26 • . 31, 32, 35; PUiS , thaI curves lalerally to a 100";: :lrtirulati(01l Wilh he ligular hinge plate. J~t distad 01 the amenor paraglos,a1 sclerite is a small, weakly and variably developed , second sclerite. On its mesially directed sudace. hidden in the deft be­tween it :mel the ling ua, the paraglossa bears an elongate triangular scit!rite. the parag/assal 1,.;'11Igl. Fig. 31. Pgrr) . T lli. parallcb wiU, irs anteriur margiu the anterior paraglossal sclerite, anictdat ,in a proct:SS from its I'ost~rior angl~ , wilh th" sclerotic process connecting the ant~rior and posterior lingual plates, and articulates, in a process 1 rom its proximal angle, with the truncated proximo-Iaternl lobe of the anterior lingual plate wbere tbe latter join the salivari om.

[ 261

THE .A.lORPIIOl.UGV OF "''ESt'ULA PENSVLV,\NICA" (SAUSS.) "1.7

Both glossae and paraglossae are almost entirely membranous, c,,,­cepl basally. The lip of each, however, iJears posteriorly a roundish , or rough l . triangular, cuticular thickening c:alled an aeroylossal bUl/o" (Fig •. IS. 23, 24. 2 ' , 16. 28, 30, 31, 32. 35. 37; A(8) . The acroglossal buttons have been .tudied crit ically by Liu (84), who tales that they are dovel· opt.'tI in typical form ollly in the Ve5l'idae. The nnttnor surfaces of the glo. sae and parnglos:.ae are beset with numerous transverse row ' of truncated spatulate setae with upturned tips (Fig. 82). The . setae with­out (Ioub! bave a rasping function. They must furthenuore help to retain by capiUaritj' any liquids whidl the wasp laps up.

The muscle; opcratin~ the ligula are the following : broad unpaired muscle (Figs. 25 . 37; tfn [9]) arising baSld on the 8.o0r of the preme.n­tum and inserted On the posterior lingual plaIt. This musde undoubtcd.ly « pre'ents the (used flex" .. of the glossae of generalized insects.

pair of large muscles (Figs. 25, 35. 37 ; afl; [10)) arising basad on the lal ral part o f the prementum an inserted on the anterior lingual plate beneath its upturned proximo-lateral lobes. These muscl are al ­nlo>t certainly, judging from Snodgrass's 12 ) figures for the bees. the h",lIOlo "'. f the Aexors of the paraglossae; but they do not function as -ueh In F~/>"Ia. Instead they pull directly on Ihe base of the lingua and

1 ul fle" the entire IiguL~. Tn view 01 the fact that both of the ligular nlrucle., /ljli and ajli, produce a flexion of the entire ligula they may rightly be designated as the pos/,riar jle%ors and the anterior f1~xors, respectivcly. of Ih,' ligula.

It i possible that during feeding these muscles may contract imul­taneonsl)', or they may contract independently. Contraction of the ante­rior l1 exors alone bends the ligula forward . and if the mouthparts are apl 'Tf sed to any surface the glru;.ae and pnraglossae are bent downward

rin~ it r lapping liquid from me smface. On the ocher hand. as Ki rma~ r (SO) pointed out. contraction o f the posterior Ooxon swings tIll' ~nLire ba.e of the ligula. on the anterior margin o f tbe premelltum as n hinge line, through a wide angle until the ligtdar hinge plate a nd r ", PD'terior lingual plate make a right angle with the noor of the premWlulTL This draws the glossae and paraglossae forward and proxi­Jlll!!1 beneath the o\'erlapping maxillary lobes so far that only their bpS

proj ·ct. At the same time the anterior lingual plate is forced back against th. hypopha.rynx, causing this to "'rinkl~ nnd fold inward. lid . extreme r"traction of the ~Iossa and paraglossae serves io bring into the food pa..~ge tormed by the fl oor of the hypopharynx, the overlapping max­illary loi>cs. and the epipharyn. .... any liquid which has been lapped up. T he quid may then he su' k..tI intn me mouth. (See also pp. l Q8.-9.)

The actual f1~xors of I", "Maylossae ( Figs. IS, 20, 23 . 25. 26, 30, 31,

[ Z'lJ

«UKI II i\: J:RT • ESrl .. F W 51'S

32, 3S : IMI \ 111) are a pair oi long stmp-like muscles, the longest ill the head, that are homologous witb th. anterior adductor. of the labium as the latter are sbown by nodgrnss (137). TI,cse muscles arise on the posterior cranial \\'all dorsn-hUerad of the po terior tcotorial root., p3ss downward between the tentorium and the articulations of the maxillae, between the posterior labial adductor. (plad) and the fl exors 0 f the stipit~ (flst) , aod between the paired mu!!des of the salivarium, to be inserted on the bypopharyuueaJ b,llioll (HyB), an invagination on the distal ""d of the more poterior lateral !!derite of the hypopharyro:. This sclerite begin low on the flank of the hypopharynx and, narro v at first, widen rapidly : then it is abruptly invaginated to form the hy­popharyngcaJ button . the surface of insertion for the Bexor oi the para­glo,-'<3, after which it narrows abruptly .and ben~ do,":,d to the up of Ihe proxi"1a1 ang e of the paraglossal tnangle, w,th wh,ch It articulates. W hen the Be,,-or o f the P'lraglossa contract., it pulls on the bypophar 11-

geal bUlton and tbis in lum pulls on the pataglossal triangle. r eking the laller dor.ad on the tip of the process from its posterior angle, which is articulated to th lateral process or the anterior lingual plate. The result is a forward flexion of the paraglossa.

The so i.'ari"", (Figs. 25. 30, 31; tv) is a ilattened . more or less trian!:ular POUcll Umt is in ag ;nat~ between the apex of the hypophar­,",x and the base of the ligula and inlo which ' he saliv.ry duet SID) dis­. h'rg . 1t is transversely concllve aoteriorly. th anterior lVall being infolded again t the posterior wall and between the upturned prol<;mo­la tcral lobes (Fig. 30. TL) of the anterior lingual pittt.. The "nterior wall is '0 lightly scleroriud as to he almost membranous; the posteri r wall is imilar in characte.r basad of its junction with the anterior lingual plate. T wo pairs of SLrap-shaped muscles are inserte 00 the salivarium (Fig.<. 18. 20. 2- , 30, 31, 35, 37), Both pairs an"" laterall)' and close together within the inflect<d ~pex of the lateral SP'ltlllnte proccs>; of ,he premC!11tum (SPNnl). The anterior p.-ur are plainly dilators of Ih. sl11.-1'nr;U1II (dms/v [1 2)) . From their origin they run distad, anterior to the fl exors of tl,. paraglossae, and are inserted on lhe median ponion of tile ,"1erior wall of the salivnrium. Th. functioning of the posterior pair \1.3 J is not SO obvious. From their origin they pass distad, posterior to the fl exors of the paraglo sae, and their fibers are inserted. P'lrtly 0 0 the upturned proximn-ittteral lobes of the anterior lingual pint • . and pnn ly on 'he mernbranou. margiu of th salivarium behind Ihelle lobes. Some of the fibers of tbis muscle. tllen, pull directly on the anterior lingual plate and there! re mer~y reinforce the action of the anterior fI~xors of Ihe ligula. The other fibers. inserted on lhe membranous wall 01 tbe ~Ii"arium, may, -ince the Boor of the salivarium i continuous with the

[28J

29

,ulIerior lingual pia, , have the same effect. n the other hand, they rnay conlnoutf! to e dilatation of the silivarium, or they may act as anlagoni 1:5 to the anterior muscles. Hence it is better to call the muscles oi tlus econd pair simply the posl.-ricr ,"lUci" of tll~ solivan.", (pmsl") t~n to attempt to take account of thei r function in naming tbem. The salivariu l1l i e,-idently firs t a pump for drawillg sal iva from lhe salivary duct and then. through the ~aslieity of its walls which collapse whC!11 the dil.tor muscles relax, a pres. for forcing U,e saliva our u(Jon the surface of the ligula.

Th. ma.rilloc (Figs. 8, 10,14, 17, 18,20,22-24. 26-29, 32).-Tbe cardo (Fi~. 28. Cd) is an e10n te tr~~ngular sclerite tbat narrows proximally to end in a foot-like tllickening. the "heel" f which rocks on a pointed proceSS (Fig_. 6- , 10: a") of the cranial w31lthat extends cephalad from " • durso-Ialcral portion of the bypostoma, where Ihe laller is inflecled to foml the oral fo,,", (Figs. 4, 6, 8 ) . From tl,e "toe" of each cardo. which extends cephalad from the cranin-cardloal articulation into the had caviN, a bort stout muscle, the prodUClqr of Ihe cardo (Fig •. 17, 20. 22, 13: 24 ; flcd [14». extends backward and upward, spreading fan­wi, e O\'Cf the postgena just above th dor' -lateral angle of the ral f ,S:t. This muscle rotates l it cardo ventrad and cephal d and thus, \\·itlt il3 fellow uf the olh", max.iUa. e.~tends the labio-ll1axillary om pie." frum 'lit oral fossa. Oi tally the cardo broaden and then .bruptly bends JUSl heiore meeting the stipes in the cardino- tipital binge. "'hen the mouthparts a retracted into the oral fossa lhe bent-over end of the carda i ' all uC it tllat is vis ibl '.

Tit· nipcs (St) is elongate and roughly triangular in cross-section. h tIm:; vresent three faces. an inner, an outer, and a l)Osterior. Its inner inc" ( Fig. 17) is moderately sderotized, except fo r the di<tal ."d, whi Ii is 11I0re or less memuranous, and is directed antero·mesally against the , ide of the prementum. Tbe anterior margin of this face i. connect~ \\~th th< dors., 1 or proximal margin of the promentum by me:ws of a broad .trip oi membrane, except distally, whEOre, as prev iously noted ( p. 24) . the 'nemllrane i. r placed by the labio-ma.:cillary jugum (LmJ).

'pon the anterior margin oi th inner face oC the stipes. a little prox imnd of its midpoint. is inserted the tendon of " large £an-shaped muscle. lhe fluor of Ihe stipes (Fig. 17, {lsi [ IS]) . Tbis muscle is the middle "ne of three which have their origins on the dorso-Iateral part of lh. c1ypeus (Fig. _) . The tendon of this muscle rllns along it:! anterior margin. the fibers being inserted into it at an angle, Ihus incre:lSing its effect,veness 3.' a flexor muscle. Becau oC the linking of the stipes and the I·r 'mrntum by means of the labin-maxillary jugum, the flexor of the s ipes ",rve:; "" a retractor of the entire labio-maxnlary complex.

1291

30 '!\'~ORTU A.lI..EJUCAN VltSPIN'E. \r PS

The po.tcrior face of the ,tipes (Fig. 28) i. Inr~e and more h~avily sdcrotized. Between it and the inner face thero is d'wcloped a high carina (CSt) . ",bid, continu .. as a low sharp ridge around the distal end oi the stipes and widens into a much shaner but equaUy high carina (CSt' between the posterior and the outer stipital (aces. From the ridge around the distal end of the posterior stipital face there extends a slender process (Figs. 22. 24. 28) that curves around and supports the base of the palpus and pro,ides one of the twO points of articulation for the base of tlle galea, In other being the distal end of a :unall e1onga'" scieri le (BSLc) Ilpportillg lhe base of the Iacinin externally (Figs. 22. 24, 2).

Proximally the posterior stipital face onds in the ",mhoo·stipital hinge li ne. in which it is produced internally into a looth-like process (Figs. 27, 29) On whicb is insened the tendon o[ the large • ...tensor J1",-,,/r of rlw ",a.riJla (Figs. 2. 14. 17, 18, 20-2+, 26, 27, 29 ; '.nn [16). This is the ,)ULer of llie th r~-e fan-shaped muscles originating on the clypeus (Fig. 2). II .• tendon runs along its poslerior margin and receives the fibers diag· olL'lIIy. thus having its effectiveness increased also. The e>o."tensor of the ma.'<illa is probably lbe homologue of one of the adductors o( lbe carda as cl." ribed by Snodgrass (1 37) for generalized insect'S, the il.,crDon or which bas migr.ned across tl,e cardin,,·, tipital runge onto th~ stipes. In £ ullctianing. however, the ",,,tensor of the maxilla "'n'es as an extensor of tile eruire labia-maxillary complex.

1'11 posterior raCe of the sopes is continued ID its proximal hoi ( into ~ broad 1 be (Figs . 22, 24) t.hat curves cephalad onto the outer stipit,,1 face . Aside from this sclerotic lobe the outer rae<: is chiefly membranous. In its distal hall it bears 0Jl irre!,'lliarly lozenge-shaped. weakly developed, hairy sclerik. Proximally the outer ~tipital face passes directly into the membranous oral field IVhich suspends the labio-ma.xillarv complex rrom the oral fossa. -

T he galea (Co.) w d fadllio (Lc) are oval, flattened, more or less foliaceous, and but weakly 10 moderately sclerolited (F igs, 17, 22, 2+, 26. 32). The lat"inia is supported by two small sclerites that are continuous in a narro,,' slrip around its base. One is the pivotal stlerite (L<P) re­ierred to on page 24 as articulating with the Iabio·ma."'Uary jugum (Fig. 17); the other is an dongate scierile (BSLc on the ",~temal faco of the b'ti]les at the base of the ladni" (Figs. 22. 24, 3_). Th~ galea articulates with the distal end of he stipes in two points, ne prol· ided by the small r encircling the paJpus, the other by the lip o( the sclerite (8SLc) sup­porting' the base of the lacin;a externally. Both galea and lacinio. project mesad over the labium Md bend along their basa l hiuge lines in " mOve­ment Ol t]",'lion toward the an t rior Iabinl surface. Both the flu.ror "'''sci. of 1/" gll/,a and the fluor of II" ladnia (Figs. 27, 29, fgo [17]

! 30 I

-r nE 1I0H.l'1J L GV OF ·'''BSPUl.A PENSYI.NA:-;I CA ·· (SA Ss.) J I

and /1< [I S]) originale in the proximal half of the stipes, the latter muscle being internaJ to the former. There are no extensor muscles [or the galea and lacinia.

Both the galea and the lacinia bear numerous long settle di.tributed over their surfaces Or arranged as marginal £ nnge.. The lacinia has in it> "utcr IrunilLl " wcaJdy del'e1oped, tapering selerit., the ladnial bar (Figs . 17. __ ; [<B), which. already noted (p. 22), fits against the -imiiarly haped epipharyngeal bar when the mouthparb are approxi-111.11<.'<1. The galea "haws twO poorly defined segments and two thin margi nnl flnng.:s. Its thickness varies considerably and its surlace is marked off by the changes in thickness iIllo several well-defined areas. I 'orl ions of the inner suriace bear numerous cuticular sense organs (Figs. 15. 17 ; S oBa), which appear to be of the type styled "sensilla hasiconirn" I, . Comstuck (32). Projecting mesad over these sen"" organs is a comb­ItA · P l " f long etae wruch may he called the galoa/ p.clen (CaP). As ha> heen .tated pre,iously (p. 22) the galea Md laciniae usually exteJd IJlesad o,'er the labi um to form with the trough-like anterior lVall of the lo, popharyo."C (Fig. 32) a passageway through whicb food may be drawn ill '" ~ tlo mouth. When in this position tI,e galea! pecten fits against the ""'''parable. mesinlly !lirected, hypopharyngeal pecten on the Bank of the b~'P'lpha.ryn" .

T Ilt' tIIrLri/lary {la/pus (M.~P) is moved by two slender, Bat, tapering 11111,<,", I" Fig>. 27, 29; admp, pdmi' [19 and 20) which are inserted at h~ ,ame point On the margin oi the paJpal base. Both act, therefore,

"" depressors of the paJpu. , tllougb one (adl/l.p [1 9]) is doubtless a lIluscle Ihat , origillally " lo,,,tor, but Ihrough a hilliog of its irLSer­tinn h.', hecome " depressor. This one origill3tes on a ridge across the I'nlX'imal end of the posterior fae of lhe stipes. T he other (pdmp l20J) uri0natc. in the lobe of the posterior stipital lace that curves onto the ollkr 5tipital face. Rotb muscles pass to their insertion. between Ibe nn ur of the galea and the O",,,or f the lacinia. ll1<~SC muscles may be callef!' respectively, the BJlkrior drprusor of th. ma.riJJary pall' liS and the (>fI.r/criar deprrssor af th. 1nll.lilloyv pal",os. . -

TBE C;:PltALIC ST01l00A£1J1d

That pan "I the stomodaeum which lies withi n the head (Figs. 12, 16. I , 20. 2 1, 23. 24, 25 , 26) consists almost entirel" of Ihe buccal ca"ity and the pha rynx, which together form such a hi ghly specialized >uckmg pump that thei r illdi"idual lim its arc not Mfinitely dcterminnble. J udging rrOln certain r"",arks concerning the rgans o( itlgestion in 1T)'II1et1optern made by Snodgrass (137), it is possiLI" that some elements

r 31 I

32

in the slicking pum" may even ~ oC cibarial origin. I n addi tion to the ucking pump the anterior part oC tbe esophagus lies in lhe head.

The uclring pump is divisible roughly into two "riocipal sectiollS. The nrsl (Fig •. 12, 16, t .21,24, 25, 26), which for d wptive purposes at l"-'lst may ~ called the allifrior pharyll..r (API,},). has the Corm of a broad. flattened pocket extending tePha13d from the mouth for a short distance, then turning upward, narrowing in it. upper balf, and contin­uing to ~ levd of Ihe antCllnae, where il i. continuous with the second section. O n tile lenl of its junclion with the latter, the anterior pbarynx it produced at illl dorso-Iateral angles into a pair of conical, pouch-like diverticula, wbich may be called the piwrYIIgeal div6TIiCllla (PhDi) . T he second section, Or 1'0Slmor phary" .... (Figs. 12, 18, 21, 23, 24, 25 , 26; PPI,y) , i. a tube of elliptical cross section, the flattening being from ide to side in the anterior portion and in the dorso-ventral portion where the tube crosses the ten t r ial bridge to join Lhe esophagus.

The functional mouth (Figs, 12, 18,21, 25 : Uti,). which may r may not be honwLogous Ivith the m uth of generalized insects. ' a narrow traru \'<:fSe lit between the base of the epipharynx. which functions as an anle.rior lip. and the membranous fold ~tween the mouth and the gnathal p lI ' h , which [unctiOn as a posterior lip. Kirmaycr (80 call Lhese ups t be "inner lips, " designating the epipharynx the "fiord"" / nnf>,/if>P"" and the hypopharyngeal fold the "hilltl'rt 'tltwlli"P8, " both apt! descriptive tel"Tl " though the fonntr should be applied not LO lhe entir<: epipharynx uu unly to the basal part adjoini ng the mouth, and the latter is less a t rue lip thm it is just a curved su rface againsl which lhe antenor inner Hp fits .

The lin iIl g cuticle o f the sucking pump "aries in thicknes , as the fig_ u reS show, and in sclcrotization also (Fi . 12, 16, 25) . That ronning the anterior pharyngeal wall is bllt little sclerotized, e"en where quite thick, and is therefore Quite R ·"ible. In the posterior wall the derotiza­tion is more pronounced and a Jar!; platc. the p"aryJ1gral ""'It (Fig. 1_, PIIPI) is de eloped in the ventm l half f the waU. From tbe dorso­la era l angles of this "late a pair of slender rods. the dorsaJ ft/Jarytlg.aJ b rs (DPhB) , extend toward the lateral wall. of the pump to tbe np,ces of the ph:1 'ugeal diverticula. The pharyngeal pla Lc and the dorsal pharyn­gcal bars serve to keep the a!lterior pharynx wide-spread and flalten!!(\ .

shorter, tapering, :utd much les. pronounet.'CI pair of processes. the ".lIlraJ plraf)'ngeol bars (Fig. 12. VPhB), extend [rom the velltro,L'lcral angles of the pharyngeal plate dOIVOIvard to the ang! f U,e mouth, hold ing them wide aparl and maintaining the slit-like character of the llIouU, . Both th ' alUerior a nd the poslerior pharyngeal walls , the latter more extensively and DIOrc densely th.,n the forme r, are beset with nu-

I - I

THE .wOIlPUOLOGY Po " ESPULh rENSYLVM1CA" ( css.) 33

",crous transve,."., rows of finc setae for SOtH ' di,tance back of the mouth . These seUle point toward the mouth. The ant rior pharyngeal wall, 31

it>; oral margin, is strengthened by a very narr W sclc:rotized .trip of cuticle ),earing a comO-like row of long dose-set setae (Figs, 12, 25 '

rP) . This may be called Ihe oral peel,.." , T he setae of the oral pecten arc likewise d irected outward , and form a sort of fmc grating o\'er the mouth, Ob. iously lbe grnting and the setae on the phar)'nC"-'l1 wall. within Lhe mouth serve ta prevent solid particles from entering .he mouth anti pharynx.

111. muscles of tbe sucking pump arc everal in number, Jme paired and "jfn~ unpaired, and they fall into distinct sets, according to whether th'" ,)riAin.te on the c1ypeus, the [ron . the vertex, or U,. tentorium. ' ll1~ lI !Uscles Bre as foUows:

An unpain.'" muscle, here <:alled the elyp,.1 dilalor of Ihe mOtllh (Figs, 2, I I , 16, L8, _5,26: cdlll/it [21]), originating in a transversel , elliptical a rea on the center of the c1ypeal di ' . ' ternally the origin is often

tarked 1,)' a ltlack do t. Th i ' nmsde spreads to a thin fan of fibers in­serted 'In the anterior pharyngeal wall immediale! above the mouth, It evul rnlly .. r"es to dilate th ·· moulh and may ~ a hom logue of the d,lalvlN cibarii o f generalized insects .. descr ibed by Snodgrass (137),

. \ pair o f muscles (Figs. 2, L1, 16. 18,24, 25 . 26 : dlb, [22]), originating un th,' dypeal disk a litUe below :utd laterad ' of the preceding aJld in ,

rted un Lhe ucking pump just below it> midpoint. These Illu>cles <I re

hrn~" beLts of ftber f slightly arcuate cross tion. They are powerful ,lt lator. and are p ssibly Ihe dilator .. bllU,dis of nodgrass. Their origins are oit"" indicated by "Icrnal maculati ns on Ihe clypcus.

. \ pair o f short SlOut muscles. the dll.Jor~s pltaryngis irOlllall's (Figs . ~ , II . L6, 18,24, 2 ,26; 1 dip" (23». or fro"laJ dUDlorJ of Iii, allll'rior pllw"· ,,.r. T hey originate in flattened ellipses on the frons ' lW""'1 the .nt ",no and 5pread to broad inst.ruon. in the trough on the anterior ",;til OJ ! the pharynx between Ilud below the bases of Lhe pharyng I diverticula.

\ pair of slender musdes. the dilalor~s poslplto.ryJlgcoles (Figs. 18. 20. 2 L, 1.; , A : 2 dIp" 124]) . or dorsal dilalar" of Ille poslerior I'Jlaryn,~ ,

"ng"lll'lting on the vertex and iDS<:rtcd on lhe posterior pharynx where it wid... sLightly abovt Ibe tentorial bridge.

A pair o r slender/aural phor)'H(J.alllluscits (Figs. 19. 21 : 11"1111 (25)), o f uncerutin homology and function originating on peg-like processes (Fig . 7. 8. 11. 19-21 . ppJ alongside the ventro-Iateral angles of lhe el)!",U and inserted on the upper ends of tile dorsal pharyn~""'1 bars. I I « "' tratting, these muscles must spread the bars Or at 1e."lSt their III''''''" ~'ll .• and also the pharyngl!:ll dil'ertiCllIJl. The), may produce other 11lQ"Cmellts a wcll .

[ 331

J4 NORTTI AWBJUCA..'." VESP1 !'\£ \\'ASp'S

An Wlpaired ponu;c, dillJlctr of Ih,! on/rrior phar) .. x (Figs. 12, 18, 21. 25. 26; ~ dlph [26]), rigioating in a long slender lend n attached to a poinled (orward-<lirected spur on tlu: middle of the tentorial bridge. and inserted on lhe hinder median ponioo of the pharyngeal pL,[ •.

An uIl{XIired vtlllral dilator of II" "osl.rior pharynx (Fi • 18. 23, 2-l: 4 dlpl. [27J , omposed of .hort fibers arising on the traIl Ve= por­tion of tbe lentorial bridge, where they surround the origin of rhe pre­ceding muscle and spread [0 an extensive insertion On lbe under side of the posterior pharynx, from directly abo"" lbe tentorial bridge forward to a point more i.ban hal f way to lbe anterior pharyru .

A mas!! of intrinsic muscle fibers. tbe olllerio,. j7l/ri",sic fJharj' lIgeal "III.cI. (fi gs. 16, 18, 24, 25, 26; ali", (28)) . extending transversely on the nnterior pharyngeal wall between the dorsal pharyngeal bars. They are of doubtful function . thougb they may acr as antagonists 10 the various ph. ryngeal dilator mu des already enumerated.

A imilar but thinner, less extens;"e mass of intrinsic 6berIi on the upper half of the posterior waD of tJ,e anterior pharynx, lbe posterior j,,'rinnc p'.aryngtal muscle (Figs. 12, 18, 21 . 24, 25, 26 ; plion (29J .

lotrinsic muscle fibers which run lengthwise in the waDs r the pharyn­geal dinrticuln, the inJrinsic 11Iuscle oj tho "hor),1I9'01 dit'6I1ic.J" (Fig. 12, dim [30]).

A pair of rather large intrinsic muscl (Figs. 12, 16, 21, 24, 26; I'dlllll. r31]), originating on the posrero-lateral portions of the pharyngeal plate, curving forward and mesally o\'er the anterior sudace of the pharynx and spreading to a broad . thin insertion on the thickened part oi the anterior pharrngeal cuticle close to the mouth and alongside the c!ypeal dilator o f the mouth r l\o. 21 J. They dou bLless function to some "" tent a /J1tary'1{}601 dilators of II .. ,,,(}JIII., though their action i not nearly so pronounced that of the clype.aJ dilator of the mouth.

THE THORA ' AND ITS APPENDAGES

T n!! TIlORACIC TI<UNK

Gmtral ttaJuros.-/'.s in the case of all the higher Hymenoptera, Ihe tboracic trunk r a vespinc wasp consist!. of four segments. Ihe usual prothorax, mesothorax. and metathorax. and. in addition, the propod""m (Pr) . which i. dcri"ed from th · abdomen (F igs . .38-42) . TIle propodeum is somenm .. caned the "UldiClII st{Tm~tll. a designation which was given to it by I_~treille. finitely a part of th .. abdomen in the lanai stage. the propodellm i. transferred to the thorn x during the tran.r"rmn.tion 1(1 the pupal tage. J t becom"" firmly and immm-allly fU!led wilh the meta-

[l.q

Tn" IoIORPROLOGY OF "VESPU ~A PENSV LVAxrCA" (SAUSS. ) 35

thorax in all the higher Hyn'enoplera. In some gonera (Gast,""'/,lioIJ . CapiJDflius, Figil s, Rllodilts. rl nl.) lbe intersegmmtallinc between meta­UlOrax and prQpodeum nearly or quite disappean., bur it is plainll marked in the Vespinae. It is generally bei.ievcd that the lergum only of the propodeum has rerained its sclerotic character and that lbe sternum has heen redu~ed to membrane. The remaining abdominal segments become ",paraled r rom the propodeum by a de"p COil tri~tion. They constitute the [Jan" (Figs. 43-47, 114-117 . Functionally, the gaster is th~ abdomm oj the adult insect , and it is so-called bj' most temati IS.

he thorax of th .espme wasps is broadly and a1mo>t perfectly oval, a f rm which confers upon the thoracic walls the high degree o( strength nec<k'<l to withstand the g reat strains put upon it by tbe powerful leg aml \\ illg muscles which nearly fi ll the thoracic intenor (Figs. 43, 44, 74, 77 ). 0nJ a sphere could provide greater t reng-th, and in cross section the ·a. p thorax sesse UIC propert ies of a sphere, for its ~ross section i", clrcular.

To the mtthanical advantage ascribable to hape is addl'<1 rigidi ty oC conHr uction . With lbe ""ception of the pleu ral and sternal parts of the 1'f<ll h<,rn:x. and o f the cutum, the scutellum. and the metallOtum, the th"rax o f lhe wasp (orms a ~ompletely rigid box. Interlocking mechanisms hel \\'""" pronotum and me;opleura. between mesopleura and metapleura, aml between mesosternum and metasternum secure rigidity at the inter­, bO'" " tal boundaries even though lines of membrane still ~xist along theio junctions. Me>opleura and me 'ostemum are fused imo a single cleri te without obvious trnCe of sutures between them (Fig. 40) . The

. ,ort, diagonal. impressed lines at Lite approximate location oC lbe pleura­, l ~ rnnl utures in generalized forms are purely superficial. The metapJeura, 1ll01"'ttrnulll, and propode.aJ tergum are fused likewise into a single scierite.

. till other features which increase the rigidity of the rhorax are> I rc-ent. The margins of the proootum " re abruptly bent inward to form, r"'I,..:1i, Iy. the a7llmor. dorsal, and /los/urictr prollolal iuj/ectillll! (Fig. 46, IITI . DITl , PITl). Broad, knife-like. marginal apodcmes ;11" OC­,'e oped anteriorly on mesopleurn, metapleura, and metasternum, and pos­ttriorly a well 00 the nl opleurn (Figs. 47, 74. 77. 79, 88. 92; PI2A I, T'll,/1, T V Fl, PIZAl). TI,e anterior mesopleural apodcrnes are continu­ous v<'OlraDy with the base of the mcsosternal apophysis (Figs. -1-7, (2). and the lurc;d arnr of the laU"r (.111'2) are firmly bound b. the furco-tltural ' · ... scl~ (fitI'll) to the "~SOl'lturai rid!]" (PI2AZ) and adjacent parts of the hody wall (Figs. 4S---l7, 51-54, 88) . The marginal apodemc of the Otetast rmun becomes medially the Irollwcru vorlical platt of I"e .. ,eI4-sl~",al apopl.yns (TrFl). Laterall)· ti,is apodeme is mntinuous with the suolldary lIIetaPI~.r.1 .p.dclllr.r (FllA2) . Finally the furcal arms of

[351

36 :-;ORTU 1\ \1' ER ICAN VESPDIE WASPS

tile ",,,tastltnfaJ apophysis (.'IFJ) a .... rigidly fused wiLh the broadened upper end of the secondary fIIelapleural apodemes Figs. 79-81).

It would be difficult to concei"e of ~ stru<turc beuer adapted to with­s=d stresses and strains than the thorax of n . spine wasp. And this is well. for Lhe Lhorax of n oLber insect . not even of ants or Ihe honeybee. .s so S<!Verely taxed by the I.,bors of its po e;sor. . 0 other in eel per­forl1lll such a variety of arduous Iasks.

Prolilortlcic sclent.s: Ihe Immot"". Figs. 38. 4O-4{i: Tl).-The pro· nolum arches O\'er the proLhorox J rotU on. (ore <oxa 10 Ihe nLber. lL is deeply .. x cavilled on Ihe dorsum by the mesosculum. and in consequence is divided ill to two roughly triangu lar side pieces which are <onnected anteriorly by a transverse, narrow. <ollar-like strip. Owing to this and other modificauons, Ule side pieces are so oriented that each possesses a dorsal margin. an ,mtero-ventral margin, and a postero-ventral margin. T ile dorsal and anterio r margins. respectively. are ontin uous aero !l.e lOp to form U-shaped 6gur when seen from above or from in front (Fig . 38. 4 1 . The dorsal margins tegeLber represcnt the middle Lbird of lh primitively posterior pronotal margin . the antero-ventral margins ~re equivalent to the primitively anterior margin, and the postero-ventral m rgin.; represent Ihe lateral thirds of the primitively posterior margin. The primitively ventral margin has been either eliminated or in 'orpo­ral~d into tile postero-" l!Ilt"ra1 margin.

The anterior pOllotal influlioll (AID), referred to n page 35. is developed along the "ntero·ventral margins of the pronot"1 side pieces. T lti inflection . in addition t bracing the pron IU IlI , provides a bearing u r ace directed in its lateral port ions alfolinst the lateral faces of the

propleura ( ee p. 38), and iII its upper nod middle portion. against the projecting po.loccipital rim f the foramen magnum (Fig. 63. Poc) . . ve il a bearing surfat · i neces itated by Ihe lT1ovemcn~ of the head and the foreleg. T he anterial' Ilronotal inflection is produced externally to "orm thc antfrior Immotal ranna (TfCl) . T he dorsal pronolal infttcliqn (DITl ) is developed on Lbe dorsal margins of tbe prnnotal side pieces. It is directed toward the anterior and I.teral margi.lS of the me! osculum. with which it is connected by means of a strip of membrane (Fig".[ S) . 'n le membrane provides for Lbe up-and-down movements of the mesoscu­tWll during Aight, and the posterior pronota! inBecti n provides a bearing surface ior the mcso:;cutal margim.

Each postern-venITal margio of the pronotal side piece. shonly belo,,; i13 upper "nd . bounds Ih slightly convex Pos/frillr lob, F ig . -12. LT1),

nealh which lies the lirst thoracic piracle (Figs. 45 . 46, 65). Above and b"low the post rior lohe the postero-ven.ral pronotal margin hrors th' posterior /,rOIlMal ;"j/rrtioll (PITJ) . Aero the po'lerior lohe the poste-

(36)

TH& MORPHOLOGV OF " YES'U!.A PENSYL"",I[ .. (SAUSS.) 37

ri r pronotal in fl ection is separated from Lbe pronotal margin and modi­fied to form the arch of tlr. spirocular 1Jfstib"l~ (.4Sp2j . Immediately above Lbe posterior lobe Lbe pronatal inOcctioo IS produced externally into " small triangular looth (Figs. 4 1, 4_ tha t o\'erlaps the margin of the IIIcsoplcuron . Scarcely higher till . Lbe pronolal inflection is produced internally in lo a somewhat larger triangular tooth (Figs. 42. 65). which tUldcrlaps Lbe margin of tile mesopleuron. Thus Lbe edge of the meso­pJcuron above is held fimtly ill the groove berween Lb""" t wo pronotal t tho T he II ppermost segment o f the po lerior notal inflection above the ( W I) teeth is concave posteriorly, forming a li ttle fossa in which the /o'9 111u ( F igs. 5- . - , 5 . 69. 70. 74, 77,88 : Tg) fit.. The free margin of Lbe po>terio r lohe is r.cun'ed internally and hooks Qver a carina il ti,e meso­I' _ural Im rgin . Just internal to Lbis canna the arch of the spirncolar \(' tib li le fi ts into a hallow .margination of the mesoplcuron. to which it is I 'Jllnd b' n strip of membrane (Figs. 46. -17 ). From Lbe poslerior lobe to h. at/ach",£1t1 peg for the ocelusor '" "sci, of tlte first lI,oracic spiracle Fig, 46. G- : "s~P and 2osp) Lbe post rior pronatal inflection presents a

grcI<wed m faee. which recc:i"es a low ridge on Lbe mesopleurru margin. The iolkctiofl rapidly narrows as it approa~hes the attachment peg of the oe I",or mu de of the spir"de. Below the peg it is reduced to a very narrow ,trip. which, II w ver. p rojects as a tiny carina Lhat bends back-ward over t11~ tu~ pleural margin. Qoscly paralleling this carina anteriorly is a groove which is developed internally as a low ridge tbat abuts against tlte anterior me ')pleural apodeme (P12Al) . At ils extreme lower end the posterior pro­not'll infl 'ction widens abruptly and joins the anterior .08ection. From the jllll'lion o f the two a long, triangtliar, apodemal tooth (Fig. 46, VPT) "mjte inward to lap aero the anterior me>opleural apod"",e shortly be­r rt the latter turns inward nod becomes continuous with the hori::oJllal /,1111 ~f the IIIcsotlerlllJl apopJ,.vris (HF_). At thi saDIe point the ventral apc: o f the Mitrd anterior and po lerior pronou" inflections allut against tht end of th flange-like all/ nor muoslenw carilla (Figs. 38. 40. 42-47, 6<'. ;~. 92: lei) projecting (rom the an terior margin of tbe meso tunum. Tbi. abutmCllt completes the interlocking meeh:uti.m between the prn­nOHlIn and the Dlesopleuroo.

( In each side p iece of Lbe pronotum a short di t:lnee above the coxl!. i, f .. und Lbe forono/aJ pi/ (Figs. 38, 40, 42; T1~) . This is t he mouth of an it l\ agination I rom which a I IIder rod (Ro) projectS inwardly and c<1.baJad to the anterior pronotal inHection F ig. 46) . This rod appar­'ml)' rves to brace tile pronotum against the pull f a muscle. lhe SfcoJld fo0S/frior 1Iotal "ro/rac/or of tire prop/,,,ron (Fil!£. 46, 61, G4; I rIll6) .

Ti,. ~ropect'lS.- s in the Hymenoptera generally, the pleural and ster-

(37 J

38

naJ p"'"'-' of the protho...,·, colJectivcly tenned the /'ropul.u by Snodgrass (127), • re separated from the r .. t of the thorax by extensive membranes and constinl!e an exceedingly moLile 8uspen..,rium tor the head and front legs (Fig. 38, 40, 42-45, 61~, 66, 7, fB, 73) . This stmcturnl feature makes possil>le the versatility in the use oi the head and forelegs that is so characteristic of wasps. bees and ants.

The propllura (PI!) make up the larger p"n of the sus[>en1lOrium, but the Pros/.mum ( 11) and its apophysis are not inconsiderai>le. Each pleuron consists of" ingl~ sclerite whieh is SO bent as to pr~t a lateral face and an antero-ventr.l.l face. The !atem! face is commonly co"ere<! by the ide piece 0 r the pronotum. oong exposed only when the pleura are extended forwnrd (Figs. 38. 40, 42). The boundary betweeo the lateral ;md the antero-ventral iac", of the propleurao is mversed by a minute carina, which is well masked below but fades out after attain­illq nbout two-thirds 01 the height of the propleur:t.. Ventrally this carina cunli nu around the lateral angle of the antero-entr.l.l propleural race anrl e 'Lcods mesad nearl), to the midline. The antero-ventral faces of the twn proplettr.l overlap the presternum nud meet in a straight line mediaUy "'. far upward as the neck membrane (Figs. 38, 40, 62). Above this IC\rl they are excavated so as partially to encircle the neck The angu­laic f>OrIlO Il of each p rop!euron immediately below the neck is curved cephalad so as to be visible from the side (Figs. ~2. 73). The jwctaposed \11'-.;11 edges of the propleura are thickened and fl3uened (rigs .. 43-15. 62 CH) . and constitute a sort of a rticubtion. but they are not joined to each oUler. They are connected instead by infolded membranes with II;· lIIargill5 oj tbe presternum which they o,'erlap.

F r III the anu:ro·dorsal angle of U,e propleuron " stout process. the (i(ritUal proctss of Iltr tro/,/"uTI7rI (Figs. 43-15, 61. 63, 64, 67. 73 ; OP), =end>; ",phalad. slighUy dorsad. and mesad . to articulate with the Co opitul condyle of the head (Fig. 63, Occ) . The principal ruo"emenl of the head on the two occipital processes is an up-and-down mOven\CJIt.

tnce. howevO!f, the right and left pleura are independent of each other. and the illlerposition between them of the complex pr05l=1 apophysis prevent. any signifieant amount of rigidity, torsion movements of the heau are enoily brought about also.

\.o,ide from tile contact between the ccipital condyles oi the cranium anti the occipitul processes of the propleura, the junction of the head and thora.'( is effected by the c<!rVical mtmbrane (C.If) alone. Certain portions ,'f this membrdne. howe"er, are greaUy thic.kened and are ligamentary in ""ture (FIg. 63) . ne of these. lhe 'Ventral csrtJicolligaJ1lml (CLI). runs from the ventral margin of the foramen magnum to the UlIUrWr process of Ih. prosllT"''''1 (Vi. 76, .JPP). The others, the laUrol cervical liga-

1.lS1

39

mtllls (eLZ) . run from just below the occipital condyle to the bases of the occipital processes of dIe proplcura.

Inte.mnlly au apodemaJ />ost~rior ralllus of II .. occif'ital proc.sr (PROP) (Fig. 61) extends uackward for a sbort di tance and provides urfaces of insertion for four of the prothoracic muscle. From the base o[ the poste­rior ramus a bracing sscondary prop/~"ral ridge (J RP) extends diagonally downward and mesad across the propleuron (Figs. 4~5, 63, 64. 67). Ex­t.:mall), the course of thi ridge is mark~ by a faint groove (Figs. 38, 40 .

The poslero-dorsai margin of the lateral face of the propleuron bears an invagination wrucb forms tbe marginal apode~ of III' ~op/n"on (Figs.. 61 . 67 ; MAP). T he apodeme is broad below and ",mo,~s to ~ po~nt upposite the occipital processes of the propleuron. Just bclow lIS nudpoltl t the apodeme widells abruptly to foml an angulation on wbich a mu cle, tbe O/Ilmor olld inn" 110/01 prolrac/or oj lit. propltJ4ron . is inserted I [' ig. 67. [tIllJ) . In front of tJ,e insertion ot lhe muscle a curved thick­ening of the cutiele forI!l.'l a IraJlsv<rst /"o/,/l'1trol bra", (Figs. 45, 64, 67 ; Pl1 8) tbat cr the apodeme and ontinu .. down the inner surface of the lateral face of the propleuron (Fig"'. 45, 67). This brace: strengthens the propleuron against the pull of both tile protractor muscle n:ferred to abo\'e and tbe pleural ~olraClor of ,,,. fore cora (Figs.. 64, 67 ; '1m2) \\ hiclt ari< On the under suriace of the apodeD1e and on the adjacent pltural wa ll just behind tbe brace. t\ short distance above the pleural ar­ticulation with the fon: coxa the marginal apodeme i. produC<.'<l still rurther ill \\'ard M the proplet4rai """ (Fig. 45, 61-64, 67, ;3, PIlA), a long. taper­ing invagination with recurved tip that runs backward and mesad a long the (m'N" {rest of the first furcal arm (OCF1). The bas of tile propleural ~rm i, bound to the crest of the [urcal arm l>y a .hort f"rco-~/cllr'" liga­;" ", (fig. 61, FPL) . which is cuticular in nature, since it is not broken down 11,' Ron.

nodII'''''' (127) ca1Is the propLeural nrm the "'/,;.ntral ar ... ," and says it i, "apparently not the homologue of the pleural ann of ther segments, .;nc~ it d<>e not arise from between the pleural plates." The tenlhredinid, .1",. p., i~ tlle only hymenopteron which Snodgrass cites as h.wing a well· drfined epimeron. Io this genus the "epimeral amI" is said to be invagi­nated irom the posterior angle of the epimeron, but in Hymenoptera gen­mtll)! "when the epimerum is absent this ann appears to arise frolTl the ~pi.t mum." Tbe lauer appearance is the logi""J expectation if the absence of recob'll izablc epimeron is due to it.s ~luction . RedUClion of lhe epimeron would carry rhe "rpimeral arm" forward liU it would come to It · all the pleural utore and would become virtually equivalenL to the true pleural arm. \ Vhen this smgc is reached Lbe "epimoral ann" would "ppe r to be borne by the episternum. I" Vespula (Fig. 73) a suture extends

[ 39 1

40 NO&'TH AMEJlICA..~ VESPlN£ WASPS

from the articulation Wilh lhe coxa up into the propleural ann. This may be the laIr.u suture, in which case the narr w strip behind it is the much­reduced epimeron , and the r",,1: of the pleuron is lhe episternum. Interven­ing between the end of the sutu re in question and the co"a, and sharing in [he coxal articulation , is a minute accessory seleril, of III, for. 'Oral

(Figs. 61. 73, erS). From the foregoing paragraph it i e\'ideot that the Cundanlental

morphology of the prol,leural arm in the majority of Hymenoptera re­maim yet to he po itivcly established. The same i true oi vinually the entire prop[euron as the latter is here defined. Berlese, according to Snod­grass ( 127 , on what seems to be rather in~ure grounds, holds it to be larl{cly sternaL in origin. Crampton (35) rcg3rds it as a iusion product of the propleurol1 and cervical scI.riles. calling it the "cervico-propleuron." Snodgrass (127) and Weber (153) contend that it is strictly pleural in natll rt. .\lore rettnlly, however . nodgrass (137) recognizes th possi­biJi~' 01 fu..ion betw..,n cervical and propleural sclerites, though the illw.tration he g ives is hal of POl/orta (o..."."ludinis and not a hymenop­teron. The fact that the occipi tal process of the propleuron (p. 38), r it , lUI va! nt, is borne by a discrete lateral cervical sclerite in mo.~ in­sects , tends to support Crampton's interpretation.

T he p'roslrnr"". (Figs. 38. 40, ~3. 44, 61-63 . 66. 67, 76; Sll) is sma1I and relativcly imple, though the presternal apopb)·sj· is large and Co.1111-pi x. The body of the prosternum is Ii roughly oval plate of which the po_Lerior nwrgin and adjacent l>ortion o( the side margins are rcllected \'~ntr"d to rono a stout rin>. Anteriorly the presternum hears three proc­esses. pair of 11l1"al prousscs oj l/r, {>roslrr .. ",,, (LPP) extend out­ward in iront 01 U,C [are coxae, and a parallel-sided . round-tipped allltr;or frau ...... of tire prostmlltlH (APP) Fig. 61 cxtefl tlq forward to meet the ven­uul ocr vical ligament. The oval body of III pro temum is appro>timately horizontal in po. ition, and is readily isible: the proreJ; -bearing portion of the pI' ternum i' hent upward and is o\'erlapped by the propleura. The bod)' of the presternum is invaginated medially and around its po terior margin to form th prosternal apophysis, the locus of tl,e medial invagination being vi ible externally as the "'tdian ster"al uroer..'t ( I/Isgl ). The groo\'e is marked .. nteriorly by the ",oslrrnal pit (f/'l). [0 the mem­brane between ."ch lateral process of tl,e prosterrunn and the corresponding cOXllI rim there often occur. a tiny weak sclerite (Fig. 76, Tr). This \\'ebcr (153) hnmologizes w.lh the trochillltin 01 genernlizecl insects. ~ nod­g~ (127) calls its npparem equhalent in certain other Hymenoptera the "accessory precoxal plale."

The ba>c of the pro lemal apophysis is quite strong. being formed of til united vI'1'l;cal I'lal t of In< proslrrllnl a{>ophysis (Fig'!' . 43. 63;

[40 I

Tnt: ORrHOLOGY OP "VESI'U LA I'£SSYL.VA:-"C,A" (SAUSS.) 41

VPFJ) , invaginated along the medillll sternaJ groove, and U,e curved . /,orl.rior v~rlical fllaJ. of th. prosl,mal apopi.ysis ( Pip. 43. #, 61-<i3 . 66, 67 : Fl), invaginated around the posterior margin of the body of the prosternuLll . These two pIa . honly combine and give rise to the hon:onlal pkUe of tI .. prostmuzl apophysis (Fig. 4 , 44, 6L, 63, 66, 67; HF1 ) and the furcal arms. Each furcal arm seen from above is some­what coop-shaped. with a median trough (PT) and elevated side and rear margins constituting. respect ively, the f)'lIla, iuner. o .. d posterior crUls of Ih. first furcal arms (OCFl, tCF1, and PCF1). The inllC!' crest thins and broad ns anteriorly into the spatulate i1lntr "ro, ... of tI. first furcal arm (rigs. 61. 67 : IPFl . T he out r crest ends in the <lInrt out., "rocess of tl.o firsl t"real arm (Fig. 67, 76, OPFJ). The trough o r the furcal arm ends anteriorly in all angle (F igs. 62, 66, 76). ilnd is crossed just c pllAlad of its midpoint by a pronounced ridge, tl,e IrOllrl'I'1'sr ridgr of tllr jirSI f"r(ol arm (Fig • . 67. 76, RF1) .

Ire 1I((iSdts of Ihe prothora.r.-ln describing the muscles of lbe thoral< of Vts/>fllo pfnsylvonico the sy I"'" devised by Voss and used by Weber ( 153) will be followed generally. though the symbols employed to desig­nate the indh-idual muscles "ill not always be the same as those in Weber's p.'lJl<' r. 111e rea.500.S are two: First. Weber distinguishes ymbolically be­l\\' ." dorsal. ventral. and dorso-"entral intersegmental muscles. Tn the "r<'>Cnl work nI). aile b ic sY111bol will be used for all intersegmental 1l111>cles. econdly, the present author's observations and interpretation .,e Ulu!;Culature do not coincide witll those of Weber. It i. quite unlikely tlu l lhere are any very significant differences belween the mw;cies of I'"'/,a crabro. the species investigated by Weber, and those of Vl!Spuia t. IIs),h'IJJlictJ, which is the object of the present research; yet several serious di ,crepancies exist between Weber's findings and those of tbe writer. Tbe cxphn, tion "'""" to ~e in the difference in technique employed in making 4 )1.l t:tef"VatiollS.

\\ 'eber worked chielly with ections of 60 microns in thickness, or of a thickness between 300 and 400 microns. These were cut in celloidm. the thinner ones first being stained with haematoxylin-eosin. [n other rases he used sections "1/4 so did: als d.r gaMt Thorax" mounted in balsam. In the"e the cuticle was 1I1eached and the muscles were tained w.th carmine. uch sections are entirdy too thick for accurate observa­tion under " binocular . especially whell the xtremel)' thin tendons of . orne of the smaller muscles are ncemed or \ hen one must determine rhe course taken by muscle tibers that are ut diagonally r must de.ter­mille the nature and locus of their attachment to cuticular structures. T he present ",nter has used the method of di secllon under ",ater through­uut. employing w II-preserved material for all studies of mu.cles_ Both

[ ~l I

42 .NORTU AMBRlCAN VSSP L."4£ WASPS

alcoholic specimen. and specimens macerateu in KOIl have been used in working out the deta.ils of skeletal morpholo"ry. Every muscle shown in the figures and thereafter described Ix:en followed Crom origin t in",roon and observed from many angles. Everyone has been iso­lated a a unit from adjacent muscles, its fiber' have been separated from their surface of origin, and in all cases in which the structure of th" muscle pem.iued, the general action of the mu!<tle has been determined by actual manipulation. lL is believed, therefore, tbat the descriptions given here will be found to be entirely trustworthy. and that any d iffer""ces observed ither will faU within the range of normal variati " or will rep reseot abnormalities .

Each muscle has been given a name, the tl:Ul,,, being basoo on its appar· elll chid function . There is no intent, however, to sugge:. t that (unc­tions oth"r tllan 11,0se implioo by the nume may n<lt be engaged in by any o f the muscles named. The effect of a muscle contraction depends not only on the structural relationship between its origin and its inser­tion, but also on the action of other !ll1IScies concerned in the movement of th" same tructuraJ units. Following th· name, figure reference. and symbol given for each musde is a number; these numbers together form a series, begun in the section n the morphology of the head, by which. for convenience of refe.rfDCe, the muscles a re enumerated throughout tltis paper. In the symbols used in the text: and as labels on the figures. any number preceding a letter s}1l1bol indicates the segment in which tll muscle or other part occurs. Any number following a letter symbol indicates which muscle o f a p:1rtkul:tr group is referred to. For ex­ample [II",] is the third leg muscle of the mesuthorax. Intorsegmcntal tllU. Ie!! ree"i"e the egmental desigrmtion of the segment which precedes. [ ism is Ule first intersegmental muscle described as occurring between prothorax and mesothorax. Symbols for muscles between head and pro­thorax are begun with O. E numeration of the ml1>e1 .. follows.

Pleural elroalor of ti,. head (Fig . 43, 44. 61, 6 . 64 . 67. 69. 73; O · 1 [32]) (Weber' OWlu).-This is a broad, fan- baped muscle with a markedly concave inner su rface (Fig. 67), the fibers of which origi­nate partly on the lateral face of the pr pleuron below the marginal apodeme and beneath the pleural protractor of the fore coxa (11,"2) , and partly on the ventro-Iateral face of the propleural ridge (lRP) . T he fibers of the pleural elevator of the head converge 10 a stout tendon in the

eck membrane laterad of the cipital prace of the propleuron. The tendon is inserted on ule poslOcciput a short distance abo"e ti,e occipital condyle.

Furral ,I"'olor of lire !trad (Figs. 43, -14, 61 , 63. 6-1-. 66. 67. 69. 7J; Oi$2 [33]) (Weller's O,: ..... Z) .-Thc fibe rs of this mw;c1e tak · origin in

[ 42 j

TUE I[OR.PKOLOey Of' fjVES"'PULA PENSYLVANICA" (SAUSS.)

:t .trip along the inner surface 0 1 the outer ",:til of 'he trough of the fu rcd nnn. alont::' the outer crest o f the furcal arm. and on the outer process

f Ihe fur al arm. They converge to a stout tendon whiell is i=rttu n the r im of the foramen magnum immediately laterad of the tendon

to; the pleural elevntor of the head. The 'WO tendons panially coalesce at their tips. Weber says tllis muscle ari e on the hinder end of ,h" pnlplcural arm; but he doubtless confuS<!'; its origin with that of the first posterior '101al prolrtu:lor of the proplcuron (lpm5). . I irsl dcprussor of /lI6 1rtad (Figs. 43, 44, 61, 63; Ois3 r34]) (Weber's O~·(IHZ).-Tbe fibers of the firs t depre.w>r of the "ead arise partly on the t rough PT) of the furcal arm behind the transverse ridge (RFl) of the furcal arm, partly on the under ide of the inner process (lPF1 ) of the i ureal a rm, and panly on the furcal am, imnlediately below the inner pr"(e,,. T h fibers converge to a slender tendon inserted. along with Iii . lendon of the folio, ing muscle, n the lower rim of the foramen llla<; l1um a , hon distance laterad of the midline. \Veber stat" that this mu It- ari;es o"er the entire inner process of the furcal arm, in which respect . ii the musculature of (' ,spa and Vespula "'" essentially alik", he i, in error; as the tip of the proces and its upper surface serve a the origin for the fibers of the prollloracic turco-pl.u,al "'Iucl. (lfP).

The rigin of the second deprtssor of /1.6 head (Figs. 43, 44, 61, 6.\ . ("' , ": Ois~ [35) (Weber's O'l/lm1) includes aU the upper sur­i ICC vi Ihe furcal trough that lies cephalad of the transverse ridge. TI,is i, a I1ll1eb thicker muscle than tbe first depressor of the head, but its li" .... < are shorter. They converge sharply to form a stout tendon tlmt join the lend on v E the first depressor shonly before tbe rim of the fvram ' magnum is reached. Weber (153, Tar. 3, F ig. 8) shows the tend"n of this muscle as bcing inserted on the foramen magnum far to the 1<1< of ti,e tendon of the first depressor. 10 V Ds/",Ia P6'1S)'/t'a/liclJ the Iwo muscles might wi th equal propriety be co. idered subdivisions nf n ... ingle muscle.

T Ii_ rrolNOI'lJcic furco-pleural ,"lIseie [o'igs. 43. 61, 63, 64, 66; If.- 1.16]) ari . panly 00 the upper surface f the inne.r process of the iurcal arm (fPF1 , and partly on the very tip f this process. Jts liLtr r rm a Hm. strap-like mu c1e, which narrow' but slightly to its itl-"n ion IlIl th .. post~ri r rnl11us of the occipital process of the pro­~ 1 "lIron In some individuals the fibers arising on the tip (the inner )lrocc'" of the furcal arm are separatoo from the others. They then {"ni l a s.pamte muscle whicb co""erges more than the upper group and i. inserted On the under side of the posterior ramus of the occi pital I'r e.s o f the propleuron dose to it~ base. Weber's prothoracic "Gaild­srilrIUluukrl." !I,m. the de. cription of which most nearly fits this muscle,

[ 4J j

XORTn AME kI 'A,N VE.SP[NE WASPS

appears LO be a compound mu .. l ~ which owes its "characteriStics" to incorrect observation. T he description and figure £ ~ origin gi"en by Weber cxnctly til the origin of the rurcal elevator of the head (0i.s2 [23]), but its insertion is giveo as the anterior end of the marginal apodeme of lhe proplellron (MAP on wbich, s r:.r as the writer can di.cover in r'esp,,,,, /, trJs),lvullicQ. tbere is 00 muscle inserted.

AII/rnor IIO/al .lll1Ia/or of tire !>roplcuron and l!tad (Figs. 43, 45, 61. ,69, 73 ; Ip",1 [37]) (Weber's I"ml) .-This i a tiny, slender, ribbon­

like muscle which arises 00 the inner sur face of the pr notum and is inserted on the Lip of the posterior ramus of the oecipit:l.l procc of the pr pleuron .

Lal,.,.aJ !/O/al e/ll1IOJor alld rotator of /Iu pro"lcur(JII (Figs. 43-46, 61,63. 64, (J} . 73; Ip ... 2 [38]).- Thi ,one of the largest of the proUloracie muscles ill Vcspu[o p~'tryl,//(lIIica, i neither described nor fi~'UI'ed by Weber for V <spa "..a/;rO. It is broad, lIat, and fan-shuped . It "rises on the side piece of the pronotum in its poster<>-<lorsal angle and narrows to a !lat tendon, by which it is inserted on tbe outer rim of the tip of the posterior ram US of the occipital pr= of the propleuron .

. ·llllaior and i"" .. r 110101 pro/raetOf' of Ihe pr"p~"r()11 (F igs_ 43,44,46, 61. 63, 64, 67. 69, 73, [pm3 [39]).-Thi. and the. next mMtle (1" ",4) npP<'ar 0 be equi "lllent. logether of eber's' pull, I pm3. and I pl>l4 . I p",J "ri~ on tbe pron tum between the out"" and inn« pronot:l.l de­pressors of the scutum (Figs. 69, 74; Jis1 and lir2), and runs diagonally ,·tn u"w and caudad. passing just mesad of tbe lateral nOlal elevator and rnwtor i Ule propleur n. [t makes a slight (wi t and is ioserted Oil the allgulation of the marginal ap den1e f the propleuron immerliately be­hind tli " traDSverse propleural "race (Pl1B) .

The jn/rnnediaJe nolal "rolractor of tile proplc"ro" (F igs. 46, 6 1. 63, M. 67,69, 73; fP,1I4 r401) follow closely the course of Ipllla nod is of the same character, Hat and ribboo-like without a tendon. 1t is broader lltan l p",3 and lies laterad of II'm . It is inserted 00 the marginal apodeme u f the propleuron where the latter gives rise to lhe propleural arm.

T he first Pos/t,;or lIoloJ protractor of 1}1$ "roplel/rOtI ( F igl!. -I3-4<i, 61, 3,64,69, 7.3; '".nS l41]) Weber's 1f!m5) i Oalti!b, parnllel-sided, and arise<> 0 the amero-ventral border of the pronotum. It runs caudad and is inserted ' lemally on the ti].> of the propleur.l l arm (PlIA).

The stco,1d posterior tlolal " rotrac/or of tire "roplcllrol& (F ig . 46, 61, 6·1. 69. 73 ; [p",6 [42]) has its rigin on the ventral an~le of the side­piece of the pronotttm just below the rod (Ro) in"aginated from the prono I pit. It is II.otrened, e. ... tcnds diagoon.lly dorsad and caudad, and tapers 10 its insertion 00 U,c propl.-", ,,,I arm close to the preceding muscle (/"",5) . \ cber apparently failed to find thi, muscle. T he

[-HJ

T If~ fOltPliOLOG Y OF "VESPl:T.A P ENSY1.VA~"'1CA" (SAUSS.) 4S

muscle to wh ich he applied lhe dcsignati n, 1/'.H6, in VuplI/a ptr,syl,xmica is inserted on the rim of the fore CO"" close behincl the coxal articulation and serve either 10 tilt the prolhorncic SUSl"'OSOriUUl forward or as a redactor i the foreleg. Th · origin £ this muscle (f 1>",6) as described and figured by Weber coincides exactly with the origin 01 the coxal muscle in uestion 11 ... 6 of this pa""r , q.t'. p. 79) .

S/enla/ .. otalt" of tire for." COra (Figs. 44,62. 64,66. 71 , 72; Ib,.l r -131) (Weber's JI,," I .- The origin of this muscle covers the enti re lateral : uriace o f the vertical plate of the prostemal apophys is (VPF1) . The Jlluscle i thick, and sOnlewlta.t fan-shaped. 1t crosses the coxal bare and is ins"rted by a short t 'ndon on the outer rim of the coxa weU ill front oi U", coxal articulation .

PI~lI raJ /,roduc/or of I"l' for. co~·. (Figs. 44, 63. 64. 67, 71. 72; 1/ ... 2 [44 j). - This muscle and the n"xl (11 .. 13) appear to have been inl<:rp <eted by Weber as " single muscle (his fb",Z) . \ eber describ 1/11/12 in the hornet oS h"" ing its origin 'pread Over the hinder half of the illiler sur face of the proplcura. In ( 'csp,t/a pensylwnica the pleural prodllc tor f the forc OX" arises on the under .ur iace of the marginal ap, )tlemc f the propleurnn between lhe transverse propleural brace (I' ll Bl and the ba of the propleural ann. The musde i. triangular in IT"'S =lion anteriorly bU l has a thin wing of fibers l>ehind. It tapers to ., tendon and i inserted on the OUler co><al rim a little behind Ilmf.

Th" ple"ral depr/'ssor of tllr !-rocho"'",,, and primar). addllrlor of II.e far.,lrf] (Fig. 45, 6 1. 62, 64, 67. 71- 73; 1/,,13 [45 J) is the second of the two I''; 1U1Iscles to arise 011 the ptopleuron. I t is a broad, curved, lu n­shaped muscle f some, hat irregular cross section. Its fibers take <J ri -in 011 the proplC>I ro ll beoenU, the marginal apodeme and behind I PIItZ. 1m the under sur fnce and on the OUler edge f the propleurnl arm. They couverge to a ,lender tendon which traverses the lumen of the coxa. where it is reinforced by fibers originating ill the CO><3.. and is in­, .. rtell 00 tbe inner rim of lhe trochamer. This and the next muscle te­gelher appear to be tbe equiv(llent of Weber's Ibm3, which is described '''' having itfi origin 0 11 the forward part of the outer surface of the prof tI rea I arm. and to be inserted partly on tbe bind margin of the coxa a l1ll l'arUy 00 t.Iro troch.ant.. . T he outer edge of the propl"u at ann "", ily may be c llfused with the "forward p.~rt of the outer sudace at lh,· pro furcal arm" unless it is carefully scruti nized and separated (rom the furcal arm. \ "eller probably d id thus confusc it. Hen"" lhe part of hi. TII",3 wb ich i. described as traversing the coxa to be irt.erted On the t roellanter is n whole o"IUsde. and the p."t rlescribed as being inserted on lit" coxa is a difTerem mu de . the 1I",s of thi' paper. which docs originate fln lhe furcal ann.

[ 4S I

NORTU ~td£.1JCAN VE$PrNE. WASP

The furcal rOlolew tifllo. fore coxa F igs. 43, -14, 61, 63 . 66, 67, 71-73 ; 11",,/ [46]) is a very stout. curved, fan-shaped muscle that originates on the under udace of the posterior and outer crests of the profurcal arm. il runs ventrad and laterad, converging as it does so, and i inserted on the rim of the coxa a short distance behind the coxal articulation.

11te suondor)' adduclo r of III. few, coxa (F igs. 62, 64, 71. 72 ; f/ • ..s (4i ) ) i ' a very slender, w~k DlI cle 01 roundish cross section which a rises ben~th the tip of the propleural ann n the inner side and is in,erled by means of a thread-like tendon postero-inten1ally on the coxal rinL Weber makes 0 meli llon of this muscle.

Pr01lola! addlU/or (!) af Ih. fore ,a.re (Fig . 45, 61-{)4, 67, 69, 71-73: 11,,,6 [48) (Weber's {bm4).-This muscle originates 00 the side picce. of til pronotun l a short tlistancc cephalad f the midpoint on its poslero-ventral margin and not far from the fir5t thoracic spiracle. 1L i. more or less parallel-sided above but tapers below to n cord-like tendon that is inserted on the outer coxal rim close to the coxal articula­tion. It is t.he origin o r thi muscle which \ .ber appears to have con­fused with the origin of the second posterior notal protractor of the pro­pleurou (lpm6) . Hc de.cribes {t",() (his f bm.J) as arising on the pro­notum, bu t does not say where, nor does he figure the muscle.

Th diagonal rolaJor of Ih. f()r~ co.m (Fig . 43, 6 1, 63, 64, .7, 71, 72 ; /1",1 149]) ( \ ber's Ii,,,) is a spindle-shaped muscle which arises by means of a thread-l ike but strong tendon , on the tip of the posterior ramus of the occipital process oi the propleur n, and crosses diagonally through the prothorax to b ' inserted on lile Quler run of the coxa on the " de of the thorax opposite (1) its origin. T he insertion i. between those of {/ml and 11,"2, This mu~le Weber refers to a. the "Dorlllmuslul: ' and 'ays thal it goes "von cI .. r Propleura .,,; ch 1 d..., Iuskeln OwllK I a nd {blitZ [= 0 .. 3 and 111,,1 of this paperl aus und schief naeh vom ohm zur Speiseriihre cmpor, die er herab- und tw;lS zuruckziehen. wahl aucl, ausdehnen mnn.' , In a de.cribing this muscle \ Veber iomlUiates conclusions on \'cry incomplete evidence. He must have observed the muscle always in specimens cut U'TOugh the sagittal plane and hence have failed to sec the origin of the muscle. t\ ~"Ut in the sagittal plane always o( necessity severs the tendon at the origin of the mllscle, and i f the ob~erver s~s only s\1ch a preparation hi interpretatiolUl are almost certaiuly destined to be incorrect. The tendon of the origin of this muscle passes just under the esophagus, but it does not attach to it-

The i,m"r /'rollola! r~lrotior of Ih. sculu,,. (Figs. 45, 46. 69, 74; lid [50)) (Weber's [JIm) is a small, thin, intersegmeotaI muscle, out­lined somewhat like a seal ... e tnangJe, which arises a ll the inner sur­face of the [\3.rrow, transverse. collar-like trip of the pronotulD imme-

[46J

THE MORPHOLOGY OF uYESPULA PHNSYLVA~fCA~' (SAL:SS.) 47

diatdy beneath the anterior n tal elevator i the propleuron and hend ( /tHln· It runs I .. teratl, slightl), dorsad, and cauclad to i i.menion on [he Oluter angle of the mesol'rephragma (PIrI) . It acts, along with the next f is2) , to retract the scutum to the rest po ition following a COD­

"" rioll of the indirect depr or 01 the forc wing ({{dlI). T he oale!' prollola! rclr(Jl;lor of lire sculu". (Figs. ·r , 46, 69. 74;

(i.,] [51]) is extremely thin, but is much broader tban the preceding and is fan-Shaped . It arises anteriorly on the sidepiece of the pronoLum and it fibe ... £pread over an area extending I rom the insertions of 11'111) and {/,,,,4 to · point above the insertion of Ip,nJ. Weber mak no m""tioll i this mllscle. B«ause of its thinness and the fact that it is overlain

wn h a layer of fat, it i easily overlooked. T ite /IIuontrTnal ' 6/rM/or of lire proptelus (Figs. -1-3 , 61--64, 67. 71-73 ,

&': l is.f [52J) ( Weber's ivlHll is a stout, tapering, iutersegmentaJ muscle Ot oval cross section that arises on the borizootal plate of U,e meso­'lernal apophy is (H F2), run cephalad and laterad, and is inserted on tit. 'Juter rim oi the fore coxa just behind the pleuro<oxal articulation. \\'dJrr ,tat lhat, in the hornet. the insertion is on tbe posterior margin of the pro ternum. but it is quite likely that he failed to see the actual I"i," of msertion of the muscle.

TIle first n,u()furcol r~lroclor of lil~ pro"ccllos (Figs. 43. 6 1-63, 66. ('7, 73 : {i.,-/ 1531) (Weber ' (vllll2) is a stout. thick, intersegmental

u"l:le that ari.es On the meso furcal bridge (ArF2) , and is inserted on the hod)' of the prosternum at the lateral extremity of the marginal im"llgination that gives rise to the curved po lerior vertical plate (CFt) .,i th.e prosternal npophysi . P osteriorly the filJers of this muscle are

lie! Anteriorly they converge to the tendon by which they are in­,ert£d. TI~ fibers spread at their origin over the bridge o r the meso­,tern>1 apophysis (rom the midpoint of the nlcsofu rca.l bridge AF2) to the anl" rior prot ' of the mesofurca.l amI PF2 . Mo t of the fihers "rise IOn the ventral. ur face of the bridge, but a few of the lateral fibers are ' wcad over illi upper surface. ~ The Sf co lid ",esofurca! relroclor ()I Ihe I'ropsel.,-, (Figs. 43, +I, 6 1, d: lis5 [54]) (Weber's Iv"lI3) is also a stout intersegmental muscle. It oril<inates on the upper sudace of lhe anlerior prouss 01 I"~ muo­I"prn/ or", (PF2) , rUDS cephalad and dorsad, and i. inserted partly on t l~e apex of the propleural ann (PitA) and partly On the adjacent crest "~ th~ profurcaJ ann. The mllScle tapers from its origin to it insertion. 1 he fil>l:TS are inserted direclly. there being nO tendon dev loped. Weher tates tha , in the hornet, it is tnserted 00 the hinder part of the aliter

""riace of the proapophysis. The firSI t"oracic spirarle (Figs. 45, 46, 65 : S 1'2) is located in tbe

( 47 1

SORTll ~CAN VESPfNE WASPS

membrane lin ing the spirtJCtllCII' flls/ibll/e. " little poci<et beneath the poste­rior lobe (L Tl of the pronotum. BI!J1""th thi5 lobe. a!I pre\·iously Doted (p. 3n. tbe posterior pronolal inflection (PITJ) departs from the edge of tbe pronotum and forms the bracing arell of /h6 sp;racular vestibttl. (ASp2) . The vestibule is invaginaled between the arch and the pronoturn. The piraclc itsel ( is a curved 'Iit adjoining the arch of the vestibul~.

which serves as nn inner lip to the spirade. T he Ouler lip i the thiclc­ened margin of a ave;, cap-like operculum (CS!,Z) , wbich projects illlo the: spirncular vestibule. T he outer lip of tl,e spiracle is cOlltinuou witb a sclerotic thickening in the wall o[ the tmcben internal 10 the arch of the vestibule. O n this thickening the closing mu e o( the 5pirncie (20sp f 55 J (Weber's [sl./I) is inserted. The muscle takes its origin 0 11

a peg-like process (OSpP) {rom the po terior pronota! ioliecti n" hort dbllUlce below the: spimcle. There i· no dilator musele. the spi" ele being open d b)" the elasticity of the closing apparnt IS.

MeSOlhoraei. scltfTiles: I"" sculul/I (Figs. 38. 39. 41-45, 48. 49, S. 56, 58-60. $ . i O, 74, 77. .92: SCI) .- T he scutum of VespuJo. haif-oval ill utlinc. is not entirely equivalent to the scutum ot generalized iosetts. The po tero-lateral angle I)f the primitive scuttUD arc cut off by the Iralls.>c,,1 I 111"" (IS). which forms the posterior boundary of th defini­t i, c rutlUtI. The severed angles are incorporated into the s<uldlun! ( <II) as th ' Intern! sCl/I.Uar segmt:llls or =iU"" (S<uSc). The trnnsscutnl su­t ll re coincides with tile primitive sCllla-seltldlor slIlurc (vsJ except whcr~ i t LOlllld the lateral utelliLr egOlen .

Parapridal furrows Of) "-'<laid cephalad from the tran •• cutal suture to points a little anterior to the wing b: e, partially separating the parap­sides (Par) [rom the scul.al disk. , ~otauli are absent, though Caint de­pressions ( 1I0wing the course taken by the notauli in other subfamilies ui the Vespidae are sometimes di~erniblc , and til anterior margin of th ' srotllm is sinll"te at the points where it is attained by notauli in genera which po . ' thenl. In its anterior h.~ l f the scutum b.r a median nalal .m/llr( (ItIIIS) .

. e parapsidc:s bear posteriorly the posllegu/o., (Fi~ . 4 1, 42, 56; PTg). smal l. erect, Lruncat scales close behind the ba.!.es of the front wing .. The o"l~,ior IIOtal willg processes (Fi~. 41 , 55, 56,60 ; ANP2). also. are borne by the pampside.s. Immedintcly behind the anterior nota! wi ng processes are the /o/trol rmargin/dioll' of tilt scululn (tl..2) in ",bicb the neck 0 f the first axillary sclerite (1 A.r) fits.

The scutal mnrgill, (:Xcept its posterior scl,'lllent an ti the portions in­volved in the wing articulntiofl, is turlled downward strongly and bcars a narrow opolltmal fold (~IFS). At the front of the scutum thi s told i. produc<d inward to foml two small I bes which constitute til II>I!SO-

[ 48)

TI1J! ldOR.PHO LOG Y OP " Yl·.$V1JLA rE.N5YLVA~(CA" (SAUSS. ) 4

/, rcplrraglllo (Fig. 44, 45. ,0. 74; Ph 1 . The down-lumed scutal margin auO"e the apodemaJ fold presents a surface that bears against the dorsal in Rec ti II of the pronotum DIT.!) during the movements incidental to Right. The apodcntal fold of the scutal margin ends posteriorly at the , ing base, ' here it is continuous through a strip of membrane with the body of the first axillary. lnuned iately anterior to the axillary the apo­dernal iold bears tbe legu/a (T9) . The posterior scutal margin is inflected , lightly at its junction with the scutellwn.

Tlrr seultllum Figs. 39,41-46. 48, 49, 55. 58. -9. 77, 92 ; Scu).-The deliniti, e scutell um of Vespula comprises the primitive scutellUJil and the lal "ral segments noted allove as having been derived froiU the scutum. T he nn~rior and posterior ma.rgins of the scuteJ lar disk are turned i11ward n the lmlerior alld posterior ;nf/8clio"s, respectively, of the scutellum (. J/S. PIS) . Behind the posterior inflection a thin fl ange projects backwar l over the edge o f the metllllotum T3. Along the C 1II>e of the primitive scuto-scutellar sulure there is developed the apodemal elllodorsal ridge (VRJ . T his forms a werful brace that crosses the: <cutellurn laterally along the boundary t the 3-xillae. It then become c"incident with a second apodemal ridge developed [rom the posterior inlk cti n.

T h axillae are bent forward beneatll the posterior angles of the ,c UlUID. where their truncated ends are bound limlly to the inner urface

i Iii ·tiTSt axillary. The lateral ends of the scutellum, therefore, hare 3.~ intimat I)' in the wiog articulation as do the anterior ootal wing proc­c" c., . Externally the lIxillae bear high, thin rests. the sCIlIt/Ia,. rr~sts ('f ). These are rcaI l)' ' xpaosion~ of Il,e marginal Hange tliat o"erlaps the metaootum medially. From the midpoint of e"eh scmcllar crest a r i<l~O!'likc <. ... rina, the lrallSscillel/lJJ" can'/lo (CSCII). cro:lSeS the scutellum tCi il s anterior margin . Ikhind this carina and above the scutellnr crest Ihere i . 3. deep poci<et, the lateral set/leUar pocklt SeIlP). The scuteilaT cr6 roject laterally between the fore and hind wings aod end in blunt I"li nts. to which are joined the a.riIlCll')' cords (Fig. 60, AxC) of the [ore win!:l;. T here seem to be no true POS/nUl,. notal «';/lg ,.rocesst. developed . unless the posterior angles f th" truncated tip_ of the axillae are \\~Dg rrrX'csses. On the midline the scutellum IS longitudinally grooved .

Beneath the scutellar crests the posterior margin of the scutellum turms a t riangular lamelliform proces (S",L) that fits into an emargina­ti"n of tlte anterior lanlina of the me.o;opostpbragma (PII2). The process i joined to the lrunina of the phragma by a memhra/lous suture. Laternd of the lamell iform process tltt cutclluru and the phrngma are more fi rmly united. though the line of union is sti ll flexible .

The mMoposlpJrrt1gmo (Figs . 43-46, 48-5 . 54. 59, n, 92: Pir ) is

( 491

so NO.KTIl A1I.lEiU AN \'ES1~JNE WASPS

an ~nonnous 5truC[Ure that ardles backward from the "1:lrgins of the scuttllar a."i llac almost to the wall of lite propodeum and extends ventrad nearly to the leg bases. It i · attadled by it laleraJ rallli only ( F ig. SO, PhZR), having no connection with the extenlal cuticula in its middle portion (Figs. 59, 77). Its continuity anteriorly "'ith the scutellum nds at the inner edge of lhe triangular lamelliionn processes (Sell/.) of lhe latter. It is notched beneath to allow the passage of the hean. Its lateral rami project forward under the wing base (Fig. 45, 46, 50, 5+) and end immediately beneath the mbalart (S ,,2) .

The tip of each ramus of the phragm.1 is partially detadled from the resl ,~f the phragma by a line of flexibi lity so as to foml a dilinct iclerite (PIo2') . Tbe true morphological narure of this sclerite is in doubt. ""cber identifies it as the fourth axillar)'. This, however. is cert" inly an error : for the genus Prpsis, of th~ famil Psannnoclt3rid.,e, possesses a wcll-devdopcd lou rth a"illary in addition to the scleritc in question. )Ioreover, the fourth axillary is a sclerotizMion ill the dorsal lOt!rllbrane of th~ Idng base, and the detached tip oi the phral,'mal l'tl.lnllio is W1'lues·

i(lllalJlv pleural in po ition. It is probabl: a derivative of the epim~ron. Sllod~ 128. p. . Fig. 3-1) figures the homologous .clerite in tru. honeybee, but marks it r and does not ventur~ an opiniun concerning its nature. The honeybee also has a well-developcd f ounll a."ll<Llj". Vest'"la SeetD! to I.~ck a founh axillary unless it is repre:.cnte<1 by , minute brown

dot in tIle a."illary membrane. 1.1"",<1 of Ihe' detached rip. the phragma l!ears on i anterior surface

an elongate, me.ially d irected proc'!». (MPPIo2), Ihe anleri", proeou of 'he meso{1osl/>/trag1l!o, the Lip of ",hidl S«l"Ve as the insertion point for a muscle I hich arise on the meso [urcal arm. I n the honey~ this process is separated i rom the phragma, though Lound to it ill a 8exible articulation. The later;tl end of the scleritc thus produced is cios'y ass0-

ciated with be fourth axillary and Snoo!tf3SS (128, p. 88, Fig. 3-l, and 127, p.84, Fig. 19, and PI. 15, Fig. 70) tenns it the ac=solj' .deritc. of ille fourtIl axillarv. In an undetermined cimbicid e."amined by the wri ter this structure is ;ev~ra l rime larger titan it is either in tbe honeybee or in V.sputa and i unquestionably an integral part of the phragma. 1n P~psis it is mucb the same as in Vespuw, though n little larg~r. The eon­r1ition in ~ honeybee. thereiore. would seem to be a en;e of specializa­tion, and the union of be process with the phrngmn to be the generaliz~d condition. Weber (153) describes th~ process simply as a pointed anterior [Irncess 01 the "mesophragma," but in his I.e."t figures £' and F' depicts it" part of the fourth axillary. His dl3racteriutioo ill the text is the

corr~ one. On the posterior surface of the phragma lere is on each. ide an oval.

[ SO 1

THE fOUBOLOO", OF H 'E.SPULA PENS.\"LVA.~JCA'· (SA17SS.) 51

\\ iog-like laleral (resl (Figs. 50, 59. 77 ; LPPh2), on which is insert~d a muscle, th~ rolrat:/or of tire ,neso/,oslp/iraglllQ (/lldl) .

Dorsad o[ the lateral crest f the plUagn13 and anterior to it there is " 'trip o£ membrane conn~cting the posterior lamina oi the phra~a with an tI,er small sci 'rite of somewhat dubious character (Figs. 47-50, 5,3 . 5' . i7, 79. SO, 94; T3'). Weber call it the me.sopostnOtUll I. which it cannot possibly be. A postnorum i. a continuation of the anterior lamina of a phragma. whereas this scieritc i. connected on the one hand with the poste rior lamina oi the I1lesopostphragma and all the other with the ante· rior margin of the mctanotum. This relati nshi[l may b<! seen dead, io a section cut diagonally through the thora." so as to P"-'S througb the scutellum, the phragma, and the sclerite in question (Fig . 48) . The in­\agilt3tion of the mesopo tphragma is open in front of this scleritc and " bristle or needle tip Ot3y ",,_ily be inserted intO it. The sclerite appear · t" the writer to be a detached piece of til ' metanotlun . for it bears IWI> t<l<ltlt-lik1' processes between whidl U,e neck of the fir t axillary oi the hind wing fits, exactly as Ihe first axillary o[ the froot win!: fits into the laleral C!lllarginatioo o[ lh scutum. The anterior o£ the tWI) teeth mu t, tl rdote, be the all/mOr II olal ,ui1l9 ,"DeISS ANP3) . The notch loetween th, teeth is equally dearly the lateral emargillation of the ><:utllm. Tn the n <".inner of thi, paper the sd~ntc itself will be referred to as tI,e ""'"fral .rrlml. of llot "'~la'lOt'''''.

TIo' ",#'Sop cllIS (Figs. 38-47, SI-5-l, 59,69,70. 7-1, 77,88. 92).-The IIIw'f'lrura (PI2) and the ,""os/ernum ($12) are complercly [wed. there heing not the slightest trace of pleurO,lern 1 ulures betweell th."", A (lair oi Rhort impresed lines are present veotro-Iaterally on the meso­pcelu . . but these ha\'C ~v~ry appearance of bei ng secondary in origiu. They are entir.Jy super6cia1 nnd do not show on the inside.

Th~ mesoplell ron is sbortened on its dors.1 margio to only ,~bout

h"lf ib widtb lower do\,,". Almo. t the entire dorsal margin is concerut'(l in the wing articulation. Anteriorly the do~ Ot3rgin presents a small n ll1cale process (Fig. 42), behind which is a small emargination in

which Ihe bo.raJar~ (Figs. -16. 47. SO, 51. .;-l, 55. 92; Hal) rocks in and " Ut. Behind the emargination is the pl'ICral filing prauss (PII"P). Tllls IJrc:ent· various elevation. and depressions adapted to contacts with the "illaric.- o[ the wing base. Behind the pleural wing process is a addle­

like ia.sa in which the hird axillary turns during movements of tllC

wing. I n the memhrane connecting the inner edge of this [os· with the wing, tbe mba/arr (Figs. 45-47. 50, 51, 54, 55, 58, 60, 88; $u2) lies. It is <[uite small and hidden irom view. A tin)' u!,r.unent connects the , "\'"lare with the third axillary (Figs. ro, 88) .

The fossa of tlte third 3xillary i. margined ext<foally by " strolll(

lSI )

NO.TII AllEUCAN \~ESP1.sf. WASPS

subalar ,,,,,so"'~ural ridgr (Figs. 38, 39. 41, 42. 74, 77. 92: SuR) . Ante­riorly the ridge bends UP' c rd before the fos<a . broadens, and ends on lbe pleural wing procts5. Po teriorly the ridge broadens, and ends at the ma ,in of the me.opleuron just in front of the puilrcmt! (Pn-l) of the ~ond thoracic spiracle.

I ntmediat~y anterior to the bend 01 the subalar ridge is a deep pit. the swbaJor 'Hcsopl,ural pit (Fig. 42. sup). This pi t is tbe meeting point of c group f sutures that traverse the mesoplcuron. The prinlllry mor­i>hological identity of these sutures is unCI:l"lain. One of them, here rolled the Irans~/."raJ nllurs (lpI2s), arcl>es upward from a point on the anterior mesopleural margin just bel v the posterior lobe (L Tl) of the pronotum, to tbe subalar pit. An apparent continuation 01 this .ulure cxtdlds C3udad f rom the subalar pit along tbe under side of the .ubalar ridge. j\ second suture descends f rom the pleural wing process and skirts the ante rior edge of the ubalar ridge to the subalar pit , where it appears to turn caudad and to unite with the post rior rontinuation of t h~ Lrnnspleural sutme. Th · resultant suture turns downward when very close to the po terior l11esopleuml margin and continues along this margin t the leg aniculation. A broad porlmor ,nosoplrllral apodNllc Figs. 51 - 54. 77 : PU.I3) i invaginated (rol11 tlus suture along the po ttr ioc Ilw""JlI ~ura l margin. branch of the cond utur. descnbed above n !llS forward I rom the pleural wing process to the anterior mesopleural margin. T he third uture (1'/2s) of the grouP. meeting at the 5ubalar pit, descends \ ertkally i rom the pit in a broad cun'e. until it nearly attains th. poste.­flor l11e5opleural margin at about the midpoint of the lalter.

Each of the SlIlures just described is marked i.nternal ly by a w~l­de\'~l oped apodemal ridge. That on the thi rd suture (pI2.<) is developed illlO tlte higli , thin, .ICo1ld ,nrsopl."rai a/lod.IHq (vigs. 47, 51-54 : P12A2), T he ridg 011 the oth"l' sutures are much lower but are thicker.

. om portiou of the system of sutures just described doubtless repre­, nlS the primitive ""sopl~"ral SI.lur.. Rohwer's comparative studies of b . according to Snodgrass (128), inwcate that the CUT.., of the Ill ' opl"urnl suture is dcviou . The suture on V upt.la, which Sd!111S 010St nearly equi"alent to the mesoplcuroj . uture in the honeybee. as the latter i .i' ribed by nodgrass (128) on the basis of Rohwer's studies, is the .ccond described above. It begins on the pleural wiog process, skirts the subaJar ridge In lbe . " balar pit, and bends backward beneath the sub­alar pit to the poSt"rior mesopleural margin . where it turns downwa.rd and COllti nues as a ubnla.rginal groove t the coxal articulation. All the othe, uturcs of the group concerned, then, are second."y. If this inter­pret.tion is correct, tbe S II alar ridge, the fossa of the third axillary, and the exceedingly narrow strip ""-tween Ule suLmarginnl groove and the

152 1

THE. MORPHOLOG Y OF ~lVJ::.SPI.1LA l'E,SSyLVA....'HCA ·· (SA05S .• ) 53

posterior mesople\1rnl margin constitute the mesepimeron. And the poste­rior mesopleural apodeme is homologous with the mesopleural ridge of generali .. d insects. Some support for this interpretation is found in the presence in VtsPultJ (and in Prpsis) of a structure bere called the 11'U<>­

'''oracic f"rco-pl~rdl muscle tlnd ligtllllent (Fig. 77, Ilf1'12, and p. 58, bela .) T he ligament is a very thin, elongate, triangular organ that extends inward from the tooth-like, in-turned, lower end of the posterior mesor leural apodeme to the tI"" of the t1«'sorlt",al dpo/lhysis. I t is quite possible that this ligament is a vestige o( the mesepleural arm of gener­alized insect. On the other hand , Rohwer ' interpretation amnot yet be <ollsidered as proved . The author, therdore, following common prac­tice among systematists, refers to the sulure which descends from the 50' alar I)it and arches backward to the midpoint of the posterior meso­pleural n"'rgin as the mosupillurai sulllre (pI2s) . The sderite cut off from the mesopleurnn by this suture becom then the mesepimeron.

The anterior mesopleural """gin (vigs. 46, 47, 69, 70) is practicaUy .t" ught for a borr distance. It then exl,ibits a hroad. rounded emargi­nation, which receives the posterior lobe of the pr notum. B~ow this the margin bulges trongly forward. The manner o f its articulation with the pronotum has already been described (p. 37) . A broad apodeme, the onlerior tHIso/lI ... ral apodeme (PI2AJ) , is developed along the entire an­tcn ur margin 01 the mesopleuron (Figs. 47. 51, 54, 74,92). This, as preyi­omly n ted (p. 35) . continues ventrally aCrOSS the skrnum, where it , .-p"nd. ill ward and is contilluou with the "0,;co1l/a1 plale of III" tIIeso­st.·mnl apo/l")'sis (HF2) .

The sternal margin of the mesopectus is produced anteriorly, in front of the marginal apodeme, LO form the 40181ior tncsostrmaJ ca,;na (vigs. 40, 43-47, 74, 92 : Mel ) . The invagination of the combined marginal npodo!n1e and borizontal plate of the mesostemal apophysis occurs along the Lound:rry ~tween the carina and the mesostemwn. As this boundary I'I'roach the nutlline it curves backward alKi becomes continuou with

the median sternaJ groove of the Olesothorax (Fig. 40. ",sg2). There is Il<l apodente or carina on tbe po terior margin of the mesosternum. short di ,tance in front of the middle coxne a lratossltlrttOl suillr. (vigs. 40, 42-i: 1st) marks off a precoxal scleri te (Pres) 0 11 each side oi the median

lill' ·. Each precoxal sclerite is produced backward between the coxae t form a pointed //Iuos/mud tooth (Figs. 40, 4245, 47, 77, 84; SI2T). The inter egmcntaJ line between mesO !ernum and metasternum follows the I""terior margin of the middle coxae to near the midline. It then bends i rward abruptly beneath the mesosterna! teeth and, where it crosses the midline, pas es througb a point that is distinctly anterior to the CO""" ( Figs. 43-45, 47).

[ 53 1

~ORTH 1\ I EKtCAN VES PtNE WASPS

A ma.rked median sternal groove (msOZ) occurs On the m""OSlern um (Figs. 40. 74). and marks the invagination of the lIIosos/rrHOi OPOpl,yN. The vl'f"licol piau 0/ II,~ »I"sos'l'n,al apophysis (Figs. 43, 47 : V FZ) ri~ directly above the median terna! groove. its two idmil1ae joining the Iron­"0"101 plul. (HFZ) of the apophysis, and forming the ventral lamina of the latter. The median .ternal groove ends anteriorly in a shallow pit. tbe 1>Iuo.rurnol pit ( Figs. 38, 40, 74; /p2), This is not the furcal pit, for the latter occurs posteriorly between the bases of the mesosternal teeth (Figs. 40.43 : fpP). A pore leads directly from the furcal pit into the fureal base, where iL bifurcates. the branches passing into the furcal amlS.

The furcal arms (.-I F2) of Ihe mesoslt!1'1,al apophysis arisc as diverg­ing tubes from the horizontal plate (Figs. ~ 3-I7. 77. 88, 92) . pass diag­onal!:- upward. and then tum outward to end io thin, fan-shaped tips directed toward the "epimera" (Ep",Z) between the s<!CQnd mesopleural apodeme ami the posterior mesoplcural apodeOle. The tips of the {ureal arm. are attached t the "epimcra" and to their bounding apodeme5 by a pair f muscles, tbe turco-plEural mllsclcs ot tI'6 HluolhortJ.I· (Figs. 45--47, j}-54. 84; llfp/I).

A transverse mesofMeal bridge (ArF2) connects the fureal arms at the level On wh.ich they turn outward toward the pimem. The brid~e is broad. ",Iati\'ely lIat, and horizontal. It i somewhat bilobed (Fig. 92). and eacll lobe bears anteriorly a parallel- ided, blunt process, the anl"rior process of /lID f!'J'So/iircal bridgo (PF2) which i turned down at tbe tip. Each o j the processes is supported beneath by a lugh median carina which i ' cont:inuous below with the edge of the hOriZO lltal pL~te of the meso­ste rnal apophy is (Figs. 4+-47). From the base of each process a low carina curves backward and outward ar.ros~ the surface of the bridge to the furcal arm. T b.is cari,,,, is the inner l>olUldary of the "rca of attach­tIl""t o f lhe second mesolureal retractor of the propectus (Fig . • p , lisS). The midpoint of the anterior margin 0 f the mesofureal bridge is some­what salient and angulate, th~ angle being upturned slightly and its mar­gin' carried bar.kward for a short distance as low carinae on the upper StlrraC!! o f the bridge. These carinae bound tlle area.q of origin of the ftrst mesofureal retraclOrs o[ Ihe propectus (Figs. 43, 74 , 92, liN .

Tire muscle. of Iho 'l1ssolhora.r.- T he i .. dirccl olld PrillcipaJ dtprtsso .. of lire ffJTS willO (Figs. -13, 70. 74, 77 ; lfdll [56]) (Weber's IIdltlll) is enormous, being rnu h the largosl muscle in the body. rt is a paired, longi­tudinal, dorsal mwel •• the members of the pair lyinR' 00 each side Ot the middorsal line. Ameriorly the fibers of this muscle are attached to the SCIl I~m and it> prcphragma from the free edge o[ [he phragma back oearly to the tranSScul1l.l suture. Laterally this are.~ of insertion extends to the line marked oj' the notauli in forms where these are present. T he

[54 1

TK£. MORPHOLOGY OF "VESPULA PENSY.LVAlHC4\' · (SAess.) 55

",rSQPrp!,i,ragf!Ja provide but n small fraction of the area required for the ~ttachment of this muscle. T he fibers of lfdl1 are attached posteriorly to the l11esopostphrngma, the area o[ attachment coveriol; all the phragm~ except a narrow median strip .nd ti,e phragmal rami. As this muscle serves to arch Ihe nOIUI11 it can hardly be said to have an origin and an insertion in the usual sense, Both ends of the filius arc origins or insertions , accord­ing to the way one chooses 10 regard them.

The enonnous size of thi , muscle and its vigorous activity during flight necessitnte a tracheal supply of unusual character. The muscie fibers are ':paroled inlO n large number o( disti nct bundles. The divisions shown in Ihe figures represent Ihe actual number and rclative size of the ftber I.lundles in Ih~ SllC"imens from which the drawing's were made. Jjetwe~n Ihese fiber hundles runs an elaborate maz of tradleal branches, each fiber buncUe I,·in\: completely enswllthed by interlacing branch tracheae.

The iud'r",i and prif!cipal cln!alf>r of I/.e fora .uing (Figs. -13, 44. 70, 7-1. 77. 88; IIdvl [57J) (Weber's lid"",), abo a paired mu..<cle, is the 'L'Con<i largest in the body o f tbe wasp. It ~es latcrnd of the preceding and ;, al,o broken into fiher buodles that are enswathed by traLheae. The fiber 1llllloles are inserted mostly on lhe laternl parts of the scutum, l>ctwtcn the depressor of the fore wing and the pru-apsidal furrow (/If). None. how­e' er. roaclles the anterior fifth 01 the scutum (F ig. 44) . A few ftber undies ron " 1I,,~hl'<l ahove t the scutcllar segments (Fig. 45, ScuS~) . The indirect clevll!ur of the (ore wing i. 113rrower anlero-post<:,;orly at its lower end but i, thir.ker transv~rsely . Its fiber bundles take origin bel01Y on the mcso-1'""ltl.;. from a line on the "ternlUll in front 0 f the middle coxae, lmerad nnd upward 0 11 the pleura tIS far as the atto1ciunent of the direct muscles of th~ for ' wing (lIp1ll1 to JIPIl1S) . Th. fibers of the indirect elevators of tl,. wi llI{ pull against the ootum, bending it along the transsculal suture, and Werelly deprcs 'ing ;L T his in turn produces the upbeat of the wings.

The p.rcaJ r~traclor of t/l. 1I1uoposl/,"'OO'/IU (Figs. -t.J.. SO, • 2; lId" 2 [S Il (Weber's lfistn) i. a relatively .mall, f"n-shaped muscle that ari . 0 11 the meso furcal anns and converge upward to " tendon tll"t is i sen.d 00 the tip of the anterior process of tll. mesopostphragma (Figs.

1-4 , 54, 92 . MPPhZ) . T his muscle resis . the tendency of the indirect depre. SOr muscle of the fore wing CUdl1 10 Ij(t the mesol1ostphragmll, alld helps to retract the phmgma to the position nf rest following a con­traction of the indirect depressor muscle.

T he basala, ,/II/.Sc/. of Ih~ tllcSolMrar (Figs. 45--47. 50. 51 , 54 ; I rpm! [591) (Weber's Up".1 is a thin. (an-shaped muscle that arises 0 11 the mcsQpleuron just behind the anlerior me50plcural apodeme (Pll .. J1) and cunverges to a long. slender tendon that is attached to the basalare CBaZ). T ip contraction of the basalar muscle turns the basaI" r" in its notcb on the

[55]

56 NORTH A)t£lUCA. VESP I l\:£ WASP

dorsal pleural margin. T ills in tum pulls on the wing base and draws !he ving fOrw:lrd into !he position of flight. T he ba.salar muscle is a pronator f t.he wing.

The sutmd and third pl.llral "tuscJu of the mesothoro,ic third a..-illary ( F igs . 45-47, SO, 51, -4; Ilpm2 and IIp,"3 [60, 6 1)) (Weber's Tl pm2 aod I f ?",J) arise On the rnesopleura close to the origin of the basalar muscle. The fiber o[ IIp ... Z arise on the pleurnl wall a linle higher than those of II pm1 and beneath them, that is, they are overlapped by the fibers of 111',"1. The origin of I1!> ... 3 occupies the same level as tilal of III'",1 and i immediately caudad of the on gin of 111'",1 . II P"tZ and 1/1',"3 are inserted close together on the margin of the thi rd axillary ( ig. 7) . \Veber sa)" the tendons f these muscles are inserted on the membrane between the first. third, and fourth axillaries. ( Jbviollsly, then, he <lid not find their real insertion.. T hey, like 1/ Im,l, are thin, fan-shaped mu cles ",bicl, converge to slender tend ns by which they are insuted. They func­tion as flexor of the wing.

T he pl,,,ro/ depressOT of til. ""t.//u", (Fig . 45. 46. 2 ; [/1>,"4 [62]) (W eber' lIP"''') also i ' a thin, fan-shaped muscle. Tt arise on the meso­pleuron just caudad of J I pm3 and is inserted by a tendon on the 10Wl'r angle of the tip of the scutellum. Since tbe tip of the scutellum is bound closely to the nock of the firs t axillary, the depressor n ( the scutellum exerts an almost di rect pull on the axillary :ldd therefore on the wing base. lL functions as an levator of the wing, reinfon:ing the action or TldU. I t might well be cnlled the direct and secondary tlwator of th" torr wing.

The illtrinsic ",usclr (or first pilUral musd6) of the third (Jj,'j//ary (Figs. 54, 57 : ",JAr (631) (no designation by " . cber) consists £ a masS of short tiber that almo t completely fill the cavity of the axillary. Th y converge immediately to a iny, cord-like tendon that runs cephalad 3 short distance and i attached to a peg-like pronliocnce On the inner postenor edge of the pleural wing proct! . .- ince the peg-like process i fixed and the axillar)' movable, the tendinous end of the muscle i ' its origin and the fib rous en within th axillary i its insertion . This muscle functions. in con junction " 'ith Jfpm2 and iIpm3. as a Bexor o f the wing. Weller failed to see the tendinous origin of this muscle, describing it as attached to m=brane in immediate proximity to the third axillary.

T he subalar ""<scI. of Ille ,nlSotliorax (Figs. 45-47. 51, 52, 54. 77,84; 11(>",5 [64]) ( Weber's ll/'t.rS) is a slender muscle that arises on the lower pan o f the pleural wnll, behind (caudad of) the meso£urcal ann and close t the posterior mesopleu ral apodeme. Some of the fibers arise on " peg­like process from ~ mesopleuron immediately above the pleural articula­tion of the leg. The fibers arc convergent but are of dirrerent lengths. T bey a", inserted on the outer 'ide of a long, .I .... der tendon that glides over

( 56 ]

THE MORPUOLOGV OF " VESPULA P£I<SVLVANICA" (SAUSS.) 57

tile mesofurcal an n and is inserted at its upper end on the tiny .ubalarc (SII2) . The subalare is joined by a short ligament to the third aKillary (3.'1.1'). In pulling on the third axillary, IJp",s also (unctions as a flexor ui the wing.

Th . produciOT of tlte middle cora (Figs. 46, 77, 88, 92. 5 ; IJIml [65) ( \-eber's llb",1) i. a stout muscle with fibers arranged more or less l' inlk1td) (Fig. 5). It is convex bdow and trough-like abO\'e. The fihers arise partly On the median vertical plate (VF2) of the mesostemal apophy­. is. partly on the adjacent sternal surface. and partly 0 11 the barizonml plale (H F2) of the apopbysis. This muscle is strongly convergent and . in; rted lac.rnlly on the anterior rim of the middle coxa by a short, stout tendon. It turns the leg forward on ule c xo-sternal hinge line.

The outer rcductor of tJ .. ,Hiddh CO~'a (f'ig ·. 45 . 77. 88, 92, 9S, 111m2 [(6» (WeLer' 111»>12 is a slaut mU$cle of somewhat rhomboidal outline. It is much smaller than 1/1 1111 . JI lies in the trougb of 111",.1 and arises partly on tbe verticnl plate and partly On the horizontal plate of tile meso­Hemal apopbysis . Its fibers converge slightly to a short, stout, Bat tendon that is inserted on the posterior rim of the coxa. T ills muscle turns the leg L ckward.

The IIIuofurcol depressor of the trochaHtcr w,d adductor of 1116 lIIufdlc ICrT Figs. 43-47, 51-53, 74, 77, 88, 92, 9 " ; lll",3aab [67J) (Weber' TI b"," and IIb ... 4 ) is a two-headed muscle. One of the lUusc.le heads, 11I1n3a. is a broad, thick group of fibers that a rise on the ,'mtral ur face of the anterior process of the meso furcal bridge (PF2). T hey converge ,lightly and are attached to the inner edge of a stout tendon . The other h= 1 of the muscle. Ill,n3b, nsisb of a broad band of fihers that arise on the under surface of the mesofur""] arm. The two ets of fiber. are slightly t om'ergent and are attached along the side of n tendon. g roup 3b distad nf group a . which traverses the coxa and i. inserted on the inner rim of

trochanter. Weber interpreted the two divisions o f this muscle in the h,'mfl as two distinct musc.les. [1I,n.311 corresponds to his llb,n3, which he .'ay is ill rted on the hinder rim f Ihe Coxa. as a resul t o f which its action « infor os that o f 111m2. Illlll3b c rrespond to Weber's llbns4, which h describes c rreetly :u; being inserted on the trochanter. Weber appar­e11l 1)' oonfused the in$ertioll of lJ/mJa witl. that of lIl",". . The j'lII~r redue/or Qf lI,e middle cor a (Figs. 44, 77,95; {fI",1 (68)) 15 " ""'all . tapering IIIU SCJ~ that arises on the posterior end of ti,e "crtital rlal~ of ti,e mesosternal apoph)"sis. rtm5 diagonallv backw;lrd .nd outward, ; IItl i, in.~ed by a , hort tendon nn the po terio~ rim of the middl coxa Ill". ad of 1I/m2. Weber '<'elll! not to haye identified this muscle at aU, Ihough he e" idently <3\0 iL I t appears to be represented in his lI oll lencJ.~­ture of mlLSc.les only by the in~rti n of his Ilb.,sJ.

r 57 J

58 SORTU AMERtCA. · VESl'TN"E WASPS

The d~f>rcmtr of I"e Itl/uta (Fig. 56. mut [69]) (no designation by Weber) is a liny, <;hort ribbon of muscle Ih .. ~ arises n the apodemal fold of the scutum (AFS) and is inserted on the ,nner surface of th~ .tegula . . It pulls the legala downward. It is opposed ollly by the elashc.ty of the

articulation betw«n the. cutmn and the tegula. . The lIIfso/lloracic f"rco-pl,ural .""sci. (Figs. 45-+7. 51-5-l-, 88: IIfpli (701) (Weber's /f:;",) is compo""d of a mass of .fiber, thai radiate (rom the lateral euds of the meso furcal am IS. They are Inserled on the posten r , "rface of U,. second me opleural apodeme (PI2.1!). on the "epimeron" (£1'",2) . and on 11,. anterior surface ot the posterior mes<>pleurn! "i;'O' dt:ll1e (PIZA.J). Thi. muscle douhtiess opposes aU the other muscles whIch "n.e on the mesofu rcal anns and tends to correct nny defomlal1on of the me,.,furcal amI <nused by the action of such other mu..,les.

The lIIe.rolhorocic {«rerr-pl,l/ral ",,,.rcif and ligm"cnl (Fig. 77 . IIfpl2 [i ll) connects the lower part of the me."'fu~ca1 arm WIth. the pleural wall immediately ,thove the coxopleural amculanon. It COnsIsts mostly of a " cory thin triangular sheet of connective tissue and mlnlerOUS shon fibers. T ile lendon-Iike apex of the triangle. presumably the ongm of the struc­ture. is attached to a uoy, pointed, apodemal process 01 the mesopleuron in nginated at the lower end of the posterior mesopleural npodCIDe (PIlA3). The b.~se of the triangle consists of shorl fibers inserted beneaili a narrow c;1rill ~ n lhe mesofurcal arm below the bridge of the meso.tema! ~poph)'. ,i. . n ,e fuoction of the mesothoraoc fureo-pleural muscle and hgrunent i, nr>t d ear : hut it.cems probable lhat, like the preceding muscle (llfpl1), il I " tOr(!. the meso furcal ann to its Donnal form after its defonnatioD In th. action of other muscle, which anse upon it. • T he rdractor of ti,. sclllellllm (Figs. 43-47, -3; lIisl rn]) (Weber's

Tfdlln2) is a mnll, fan·shaped muscle, which, if its function is correctl? implied in lhe name given it. arises in a tendon on the mc~nonun ~nd IS

inscrted b\' lh tips of lhe muscle Iibers on the entodorsal ndge (V R) of lhc .iCulell~m. Presumably it contributes t the retraction of the mCSOl1olum foll')\I'ing each con traction of the principal depressor of the fore wing, Jfd/l. and thlli aids in producing the upbeat of the WlOgs ..

T he illl.rfwrcal ,,,,,,cle (Figs. 43-47. 5 I- 53, 79-81; IJrsZ [73]) (We­her' 11.'1",) consists of a narrow transverse b.111d of shon. longitudinal fibers which connect the mesofurcal anll with the metafureal arm.

The soeond tlloraei€ tpirade (Fig. 68) i located beneath the base of the hind wing 011 the inte egm~ta1 line between the mesothora;" . and the mctathorax. 1t is protected by a small triangular sdente. the /,mlr.",. of Ihe ucolld thoracic s/,;ra<{p ( Figs. 39, 41 , 42, 51-53. 68. 77; ~.r.J).

The corners of the peritrelh ' eur.'e inward arolUld the [Hracle. The closing mechanism is imilar to that of the first thoracic spiracle .• \ cap-

[ 58]

THE 1dORPUOLOCY Uf' "n';SPULA PENSYLVASJCA" (SAtl'SS.)

like operculum (CSp.J) is pulled down by a minute, fan-shaped muscle to close the spiracle. lts opening is accomplished by the elasticity of the lllech.1.Oi. m. The fibers of the occlusor muscl (figs. 68. 77: 30.rp r74]) (Weber's liS/III) arise on the posterior surface of the posterior mesoplcural .pod.me PIZA.J) close to the >l'irncle.

Tilt scicrilu of Ihe ,ntlathorlJ:r: lilt ,,,<,Iallol,,,,, (figs. 39, 41-49. 52-4, 77. 79, SO, 94: T3 and T3'}.-The metanotum is a transverse sderite

Lhat extends across the dorsal part of Ihc postenor declivity of the thorax. It s anterior margin is comparatively straight when ,-iewed from behind r nly slightly curved, but the posterior margin is strongly angulate in Ule

middle where Ule metanonlm is produc<'<I to fit into an emargination of the propodeum. The margins of the metanotum are inAected to r rm, rospcctivcly, the allicrillt' 'lIelalloloJ i"j/Bcl;on (AI.lf) and the poslcrillt' In tunolal i"f/eclioll (PIM). T he anterior inflection is much the larger. L,tcrally eath ramus of the metanotum extends forward Dlesad of the base 0; the hind wing (Figs. 52. 53). Each ramus is squllrely truncate but bears <1 11 its lower an~I ' a curved, spoon-like pr~s CPT3), which curves be­neath the firSt axilla ry (IA.r) sclerite to which it is bound . This process is \'Cry flexible, but there i no suture between it and the m.tanotal ramus.

n th latera l parts of the metanotum almost directly above the pro­p<"lo:al "pirades are val, depressed areas (Figs. 39. 4 1, 43; OT3) which are b unded laterally by curved tTOIwndollo/aI ,ariMoe (CT.J) and me­dially by cUrI'cd /rollsm",lano/Dl .... ,"res (I'~s). The transmetanOtaI sutures ,ir the external e,;dellce of invaginated cuticular folds that (arm powerful ",do ll%l braus (Figs . 47, 52, 53,89; T.JB) . The ends of the metanotal ral11i laterad of these sutures and braces may be called lhe ' llelanoial Sl'!Jm.lI1s (Figs. 42, 47. 52, 53, 79,94; N3S). Eacl1 metanotal segment is nnnected anteriorly with the small elongate sderite (Figs. 47-50. 53, 59.

i7, i C, SO, 9-1 : T3' ) partially described on page Sl above and there ""111 d the 1",,"~roJ scleritc of Ihe ",clulIO/um. The oUler end of dle hum 'ral scleri te is thick and harc\. [\ projects freely beyond the edge of the mdanotum and bears the anterior notal wing process (ANP.J) and tbe lat,'rn! scutal cmargination into which the neck of U,e fir t a:<illary sdenle ti ts. The rest of the sclerite is thin, flexible, and seOli-<:ylindrical, the trough of t1,e semi-cylinder being directed upward. T he edges of tile t IIgh Ilre joine<l by membrane with the po tenor lamina of the m opo t­Ilh ragma and with the intern! segments of Ole metanotum. The metanotum, like the mcsonotum, lacks a delinit. posterior notal wing process. The aXIllary cord I joined simply to the posterior margin o[ the metanotum.

netween the transmelanotal sutures the anterior border of the meta­n<ltum is traversed by a rub",argillni .mlcu:s (Figs. +3-47, 52. 53, 79 : T3s) , mIl) which the posterior flange of the scutellum fits . Just above each lrans-

! 50 I

60 NORTH AMEIUCA..N VESPINE WASPS

metnnotal.uture (fig, 53, 79) th" ubmarginal sulcus desunds aero the anterior m.tanotal inflection (AIM) and becomes continuous with the trough of the humeral sclerir.. of the metanotum (T3') described above. T he sulcus, therefore, does not ~end onto the metanotal segments. Em:h metnnotal se!,'filet1t bears anteriorly two elevations, the more lateral of which is usually the larger (Figs. 52, 53) and is bere called the "",talIOloJ

prom ill"'" (T3P). . , The ",elapo.fI"olulII (Figs. 39, 41. 42. , 49,51,80 ; PN3) IS lackmg

in many of the higher Hymenoptera. Snodgrass (127) lists Odolliaulacus , S.'mJotlwspi.r , Colo/arclls, CoCCOPho.'l"S, Dilll.~ocki", RI.odilu, TekJlo""IS, ""d Al'is as complete.ly lacking a recoglli7.able metapo matum, In VrsllI/la the meiaposmolUm is represent"<l by well-defined lateral remnants, These have the f rm of urved sclerit forming crests that unllount portions of both the metaplet1ra and the propodeum (F ig, 42). These sclerites may indu e both posrnotnl and epimeral emenlS; but if so the elements are c<Jmpletel.l' {used , for there is no trace of a suture on the postnotal sderites. 'n,e mctapostootal sderites are fused rigidly with the propodeum, but the , uture ioetween them and the propedeum is still well defined (Figs. 48, 4 ) , \ \ ' ber [:tiled to recognize the metapostllotal sclerites,

TI. r mtloprclll4 and ftropod,,,,,, (Figs, 3 -49, 5 I-54, 79-81).-The "'tIO~/'" "0 (PU) and tn,l4slt"H"K (SI3) , like th "of the mesothorax, nre united into a continuous sclerite, the metapectus.

The meiapleuron is much smaller and simpler than the mesopleuron. It i, divided externally into two sclerites by a diagonal suture, the 11I4/a­plt1<roJ stcondary suillre (Fig. 42. flUs) , The lower of the two sclerites i, Jightly mrg",. than the upper, The upper sclerite is separated from the propode\lm by 11 groove-Uke suture along which the lateral margill5 of the thi rd phragma are invaginated, ,\t the lower end f this 8uture there i, a snl.,11 but deep pit, the first ",,,Iatltllral pil (Figs. 39, 40, 42 ; PI3p). I mlllcdi:udy above the hind coxa, the mClapleuron toears a smaller uCOHd

,"til/pilI/rill /,il (PUp' ) which mar a,e inv",,.ination that foml the con­dyle for the plenral articulation with the 0"", Below the first metapleural pit th metapleuron is bmmded posteriori}' by an obsolescent utllfC that t3kes the form of (l. slight carina. The meta pleural secondary SUlllre curves diagonally cephalad and vent:rad from the fi r·t metapleural pit to lbe ante­

rior metapleural margin. Dorsally the metapleurn are much like the mesopleura, the same

proees:,es, ernarginntions, and fossae being present. though of course smaller nnd somewhat simpler. The ilasalare (Figs, SI-S-t. 79, 81, 94, 96; 8 a3) is nearly as Inrge as that of the mesothorax. but the tllbalar6 (Figs, 51. 52, 96 : S"J) i~ much "OIai ler. It lies in a pockel of melnhran~ invagi-

ted between the wing base and the rnetapostoOtum.

[60 I

TilE MORrnOLOC;V Of" '" ' ""ESPlTLA P"I!NS'i LVANlCA" (SA{;SS.) 61

·n,. m.tapleuran bears on the upper two-thirds f its anterior margin a wide. thin apodeme, the lIIarg;1Ial apod,,,,,, of lire mttaplMrroll (Figs. 51-5.J., 79,81; PI3Al) , s6colld ""tap/~ra/ apod.",,: (Figs. 49. 53, 79. ro,

I ; PI .42) is invagi nated along the metapleural secondary suture (pI3s). _-\ r the lower eI,d of th is suture the apodeme continues across the meta­.t mum to become the IrallJVC1'St 'Vertical /,/01. of Ih~ 'Helasltmal apopllysis , Figs, 43-4 -,47. 79; TVF3). At its upper end the second metapleural apo­deme '-' continuou -, across the invagination f the first metnpleural pit, with the third phragma (Fig. 4 ,PII3). The upper half of the second apodeme is produced medially into a broad t runcate process (Figs. 89-91) that over­laps the fu rcal ann o[ the mdasternal apophysis, The apodeme and the iu al am, are fused throughout the area of overlapping,

T h melas/.r.mlll (Figs . .39, 40. 42-47, 5 I-54, 79-81 ; S13) is sunken o\'er the arc':tS covered by the mesocoxae to ud, an extent that the median purtioll fonn a high carina in its anterior half (Figs, 4<l, 42-44, 47 . T he depressed areas are outl ined on eacb side of the sternum by a series of small diag nal wrinkl (Fig, 4<l). The mediall sll'rnoJ groOlI. (msg3) follow' the crest of the metasternal carina, and the midline bebind the carina. Oose to tho posterior margin of the sternum, the medjan sternal groove broadens into the f~lasl.Ylwl pil (fp3) , The posterior ma~n of the .ternwn Bares laterally, overlaps the pleural coxal articulation. and partially surrounds the co~ base making w;th the propodeum a sort of socket for the coxa.

Til onelaslmlai apophysis (Figs. 43-47, 51-54, 79-81) presents ""Ie­riorly the trOl'.l'!IcrS6 vertical plale (TVF3) nlready referred to above. which i an apodemal fold acr the nmrgin of the sternu m, On the mid­IUk tlle apophysis rises as a /oJllPrud;'J(J1 vertical plale (VF3) that is ill­vagi""led along the median sternal groove (msg3) . A narrow I,ori':ollial /,/111,' If 1'3 extend.. along the crest f tbe longitudjna\ vertical plat .. Fr the junction, anteriorly, of the three plat ., the furcal arnlS (AF3) IIf II" fnelasllr1lal apophysis diverge. Passing upward and outward the iurt-aj arms flatten and fuse with the poste.rior suriaces of the t runcate proces,;es from the second mdapleural apod meso An open passageway e.'teuds from the metasternal pit Up3) diagonally forwa rd and upward to the ba { the furc:tl arms, where it bifu rcates, a brancb extending 1111" each of the fu rcal arms. T he f ureal arms bear several very narrow IlIw carinae \ bieh increase the sur face availa tole for muscle attacbment.

T he propodC1lon (Fi~. 39--49. " 1-53, 79-81; Pr) covers the lower three-fourths of the posterior declivity of the thorax and .. '<tends laterallv to the sides oi the tborax. I t is emarginate dorsally where jt joins th~ metanorum, T he ventral border is reduplicated (Figs. 39, 45-47) , uter­ally , n eb side, the reduplicntion shares in the formation of a socket ror tho hind coxae. Between the co,,-nl ockets and the midline it nares to fonn

(61 )

62 NOIit.TIl AWERICA,N VESPI~·E WASPS

the al'ica/ scol,s of Iht pro/,odsum (Fig •. 39--42.80; ASPr). Dorsally the reduplication is produced ventrdd into two 11:lJ"d. strong tC!eth. Ille propodeaJ leell. (Figs. 39, ~O. 43-47, 79; prl). The vertical plate of Ih. abd,m,;nat p~rJicri (figs. 43-47,79,8 1. 116, 117; VPd) glides between the propodeal teeth during movements of Ille abdomen. TI,e tips of the teetb slide in a pair of grooves on the abdominal pedicel alongside Ille verei",,1 plate of the pt.-dicel :md are prevented from slipping off the p<!dicel by a transverse groo"< across the anterior margin of the ""wcel .

Dorsa- Iat rally the propodcum bears the rropodral srirocies (Figs. 39-42, 48. 75, 78, 80; Sp/) . The openings to the spiraculnr vestibules are "'ongal< slits 011 lolV elevations.

. \cross ih dorsal margin the propodeum bears the third phragma of the thora."'. • ince this phragma is inseparab y fusecl with U,e propodeum it becom the pr0ftod.al (1reph,agma (Figs. 46-4 ,:i- 53,:md 79-81 ; Ph.!). Laterally the phragma conl;naes down the lateral propod<'3.1 margins as fur :tS the first metapleural pil. where it joins the 5~ond metaplcural apo­cleme.

TIle phragma is almost nonexistent on the midline, owing to th! deep emo.rgin:;tion of the propodeum by the metanotllm. On its anterior surface "ach hal f of the phcagrna bears a curved ridge-l ike carina (PII3C) that partially bounds the surfnce of origin of the retractar muscle of Ill~ nl 0-

postphragrna (Ill dl). The tnu.rc/es of the mclall'O,QI and /"opod~I"".-The relraclors 0; 1/",

"",s0ftostp/J,agma (Figs. 46,47. 53, 77,79; Wdf (75]) (Weber's Ifldl",) arc ,horl, .. tout mllscles of elliptical cross secti n and parallel fibers that arise on the anterior surface of the propodeal prephrngm;, and are insened 011, t.he lateral crest> f the nlesopostphragrna (LPPlr2) . T hi p.lir of muscle strengthens the me.<opostphragma against the pull of the principal depressor muscles of the fore ,,-ings (Ild/]). and retract. lhe phragma to the po irian of rest a£ter the contractions of Jld/l .

The baso/or muscle of /I.e mc/olltorlU', 0' prollator of II .. Irilld wi,,!! Figs. of I , -I-54. 79,81 . 94. 96; lll~tHI [76J (Weber's mp",1) is

an asymmetrical. fan-shaped muscle that ari es partly n the anterior sur­iaee of the transverse plate of the mebstemal apophysis (TVPJ), partly On the lower half of the second metapleural apodernc (PI3 .. IZ) , partly on tbe Darrow slrip of the nletapleuron in Cront of Ihe second metaplcural apodcme and below the marginal metapl.ur-..J apodeme (PI3Al) . and partly 011 the metastemUlD behind the transverse vertical plate of the ",etasternal apophysis . Its fib~ conve.rge to a long. slender tendon which is inserted On the basaJore (Ba3) of lhe metalhorax. Its COntra tion pulls the hind wing into the position of fligh t. In this act ion, however, it is helpecl by the basa1:lr muscle of the mesathorax (ffPllll) . The pull of ([pm I is lrans-

r 62)

TUE YORPllO.L('H::\' OF ·'v·, PUL,\ PE.N5YLVA..~1C.,,·t (S .. \I:SS.) 6J

miued [rom the fro nt \\;ng to the hind wing by the hamuli or ,r",wl hooks ( I: ig. 87, H) on t.he cosml margin of the hind wing. These link the fore and hiud wings together and insure Il,ei .. moving at the same timc. It is interesting in this connection to note the curious misconception of Janet, '''Ilressed in his Obse",,,'wns sur '.s 9"~PtS (75), to the effect that the hind wing of w s is devoid of musculature . peaking o( Vespa crab,o . he say the hind wing are "completmlml depou rvues de loUIe espeee de museulature." As a matter of fact, except fer the large indirect muscles, and disreg'urwng differences in size, the bind wings are as well supplied wit.h muscles as the fore wings.

Tho plCi'TOi "'"Sc/<S of the ,""to/l'l1racir Ihi,d ru:illary (Figs. 45-47, 51-54. 79,81,94,96 ; mpm2a&b [77a&b]) Weber's IIlp,fJ2j.- Thesc Olll~c1es might be interpreted as two muscles in~ertcd by a common tendon I.,r a." onc muscle with two heads . Weber reports only il single muscle for , ', spa crabro, but it is po -wle that both are pres~ot in crabro and U,at he failed to wfferentlate between them. Doth have their ori!:irl on the posterior .,mface [the rnarglnal meta pleural apodemc (Figs. 79 and 8 1). 1I1"".2a is , maller thM /lfpmZb, ties between rlf/,IIl2b and the apodcme, and arises a lil ll higher than IITp,H2b. The eommon tendon of lIIp",Za&b is in­•• rted on the tl1ird axillary .c1erite--not, as Weber says fOT the corre­,pnnding IlTp",Z in VrMpa crabro, on the membmne between a>.-illaries one. two, and three. rrrp",Za&b function as flexors of the hind wing.

The pleural ",Usc/.s of Ille $lIbo/ao" (Figs. 51, 52. 79, 94, 96; IT!pm3a&b (7&&bJ) (Weber's fI/p",/) ar' in general similar to 1II" ,"2a&b. They taper more abrllptly, bowey r . and are more or less pyramidal. whereas III p",ZaErb are elongate and thcir fibers arc inserted " I',ng but one side of their tendons. fllpmJo arises partly 0 the anterior

etnpleural apod"me below 1I1 " ... 2b Fig. 79) nnd partly nn the adjacenJ. p!eural wa ll. Both are inserted very close together by very sho n tendons On the subalare. The subalare is connected uy a ligament ,~th the third axillary. Hence muscles fflp.n3a&b function as fl exors o f the wing.

The drf'ressors of II.. ''''''ana'.''' (Figs. 45-47, 49. 53, 54. 79-81 : I ffl'ml [79]) (Weber's IlIp.n3), among the l:)r!.'CS{ f tile metathoracic. muscles, arise on the anterior surfaet'! of the second lTIetapleural apodcme above and behind the origin d[ the metathoracic basal .. r mllsde. The area o f origin covers nearly the entire apodeme, including its broad truncate inlier proce ' (F ig. 79) . These muscles are rather complex. the fibers run­lI ing in several directions. some converging toward the origin , some toward Ihe iw;ertion, and others being approximati!.ly p.rallel. The fibers are in­' ert~'<I On lhe n1emnotn l segments and 011 the spoon-shaped process from the I~ ""er angie 1 the tip 01 the metanOlal ramus. The contraction of the ncpres ors of the metanotum pulls strongly on the metanotal rami; alld,

[ 6J I

64

.inee the rami are bound finnly to the firs t a.xiIlary tbrough the spoon­shaped process of the lower :Ulgle, these muscles must e.'<en a downward pull on the axillary and thus ontribute to the upbeat of the wings. 'V.her says of these muscles only that they hold the metanotum steady.

The (o.ro-mbalar mustle of lite 1IJ"'aI/,orax (Figs. 46, 47, 51. 52, 81, 93, 94, 96, 97 : II 1pm5 [SO]) is a long, slender, spindle-shaped muscle that ari"", in a tendon on the outer rim of the hind coxa ju.t behind the co."<o­pleural articulation, and is inserted, by means of a long, slender tendon, on the subalare close beh ind the insertions of Iflp ... 30&b. It functions as a flexor of the hind wing. The tendon by which it is inserted slides in a notch between the truncate proces of the second mempleural apodeme and the propooeal prephragma, Weber apparently did not find lItis muscle in VUpIJ <'robro.

The sl~",al prod'lClor of Ilr. hind (0.1'0 (Rgs. 40, SO, 93, 97 ; Illlml [Ill ] (Weber's Iilb".I) i a tout muscle with spreading fibers that rigi­nates partly n the posterior surface oi the tmnsverse vertical plate of tIle metaste.rnal apophysis and partly on the metasternum. fts 6bers converge 10 a stout tendon that is inserted on the out~r margin of the hi nd coxa close to the pleural articulation. Th is muscle swings lb~ hind leg forward ,

The rtdllc/or of 110. hind co.Tt> (Figs. 43-45, 79, 81 , 93, 97 : II/1m2 [82]) (Weber's I ff/ml in pan).-This is a broad, Aat, fan-shaped muscle arising 00 the under surface of the furcal ann from its base outward ahout two-thirds of lhe distance to the. metapleuron . .mall purt 01 the origin extend onto the transverse venieal plate (Tr-F3) of tbe met"tl.­srernal apopll}"'is and a few fibers may arise on the longitudinal venieal plate of th apophysis (V F.l) . All the fibers converge to a stout tendon that ' inoerted on the posterior rim 0 E the hind coxa well beltind the coxo­pleural articulation . Thi muscle swings the leg bacl.:ward.

Weber d c.ribes the presumed homologue of {1I1",2 in V <!Spa crabro ItS a complex, three-beaded muscle. lIf/,,,? in Ves/,,,Ia seems to correspond with dh isioo 2 of his f flbmZ. T he other two divi ions of Weber's IlIbm2 are i olessegmental muscles that are inserted on th~ abdominal pedicel. Weber simply failed to follow their tendons to their insertions ,

T he d.prBsSOY of 110. trochanter alld adductor af Ih. hind leg (figs. 45, ~(j, 79--81, 93, i; fff/m3 [83)) (Weher's fllbmJ) is a broad mu cle that appears fan-shaped when viewed eith~r from before or behind, but is V­shaped in CrO section , The stem and one branch of the V arise on the under surface of the outer three-fourth.! of the furcal ann. The other branch ari os on the poste.rior .urfac" "f the second metapleural npodeme. This muscle cony rges to a tendon which pa ses throllgh the co"",, nnd is inserted on tile inner rim 0 f the trochanter. T t s'rves to ext~l1d lhe tro­chanter and therefore to adduct the leg as a whole.

[ 64)

TlIE )OIOR-puOLOQY Or HVES Prrt.,,\ FE SYLV,\N'tCA" (SAUSS.) 6S

The pl ... r<11 praduc/ar of tlu /ri"d COxa (Figs. 4i, SO. 81, 93, 97; 1[[1 ... 4 [84'1) ( \ cher's l adlm3 ) is somewhat pectinate. [ts origin extends over parts of both the mernpleuron "nd the propodium between the propodeal spiracle and the propodeal prepbragma. The tendon at this muscle is Ill . f rted on the ant"-rior rim of the hind coxa JUSt lat:ernd of the insertion Ol /I lImI and immediately in front 01 the coxopleural articulation. The nalure of this muscle appears to be erroneously interpreted by Weber. The mU5Cle of his listing whicb coincides with Ifll,"" in form and in rlri!(in is by him called I adl",3, and is described as being inserted on the a!>,lominal pedicel instead of on the leg.

The pi, ural d.prtSSDr of Ih" .bdo,""" (Figs. 43, 44,53, 31; [Ilis1 [85) (\\ eLeT's IJJbm2 in part 1) is an dongate. thin muscle that originates on [h~ posterior surface of the tnilltal.ed process of the second mernpleural apodem. (PIJA2), abov~ the intersection of the apodeme amI the furcal am I. The pleural depr. sor of the abdomen narrows downward to a <lender t~ndon that is inserted on the second abdominal sternum ( 11/ . the propodeal 5tem um, If, not being developed) near the Outer angle of the latter. In CODjunction with oth r muscles it bends the abdomen down­ward, When contracti ng alone it brings about a twisting IDo\'enlent. T his Illuscle corresponds to the outermost group of fibers of \\' ber's IIIblllZ l>art I, r was overlooked by him completely .

The IIIcla/ureal depressoT of tI,. abdomlJt. (Figs. 43, 4-1-, 53, 81, 92; I/l i, _ [86]) (Weber's IIlbm2, part I) is a uroad, thin, {an-shaped muscle, the rigin 1 which e>.1ends from the longitudinal vertical plale (V F3) of the meta.temal ap ph}'si., laterad along the posterior and dorsal suriaees

i t.he meta(uTcal arm for its entire length and onto the truncate process of the second mctapleural apodeme beneath the origin of II/id. [ts fibers com'erge to a broad tendon tha t is inserted on the edge of the second ab­dominal sternum between the ins"ni n of If lisl and the mid ine.

The stlmlel torrio" muscle of Ihe abdomen (Figs. 43-45. 8 1; flf is3 1Il711 (Weber's II Ivl ... ) is a triangular, thickish muscle that ari ' . on the

IUII <; it udinal vertical plate 0 f the memste.rnaJ apopby is, and is inserted by a cord· like tendon On the lateral angle of the s cond abdominal sternum. I t brings about lateral Or twisting' moven,ents of the abdomen . alul when Ih" ahdomen bas been eleyated by muscles fadl/ and fadl2 pulls it down­ward to the normal position.

T he tropodtal cltValor of tho abdo",,," (Figs. 43-47, 79, 8 1; fadl1 ]) (Weber's ladfml) is the nly cievator muscle of the abdomen , It i

hro Cl d, thin. and Ian haped, Mises on the propodeum behind Il,e phrngma, and i. inserted. b means of a broad. flat tendon, in a depression on the abdominol pedicel (Pd) just behind and btuad of the vertical plate of the abdominal pedicd (V Pd),

[ 651

66 NORTH A.M£JUCAN VESPrNE WA PS

T he propodeal torsion ...... cI. at ti,. abdomen (Figs. 43-47, 79, 81 ; Jodi? (89)) (Weber' 10&",2) i thin and fan-like, but is narrower than the ~e\"ator of the abdomen (ladlI) , It arise laterad of 10dl1, partly on the posterior surlatt of the propodeal prephragma and partly on the pro­pod!!lll11 proper. It narrows to a tout tendon thal is inserted on the lateral angle of the ,;econd abdominal tergum where Ihe lergum IS boood to ~e sternum on Ih side of the abdominal pedicel . Like Ihe sternal tors.on muscle t JlisJ) . Ihe propodeal torsion muscle brings about t\ iSling move­

ments of the abdomen. T ile propodtal spira". alld jts musc/IS (Figs. 39-42. 46, 47, 75 , 78, 80,

81 ; ospl. dsPJ [90, 91]) (\ eber'. fa.thn=dspl).-The occlusar mechanism of the propodeal spiracle (Figs. 75. 78) i of the type called by Snodgr.1Ss an internal 'Iosing apparatu having a single valve. The opening {rolD the piracula r vestibule inLo the tmch ..... j surrounded by sclerotized bars. A

rigid 1m, termed the bow (8) forms one . ide o i the slit-like opening. Op­po.ite lhis is a flexible bar, the closillg wi ... (Cl' ) . From the ends of the bow there project a hort dorsal bar (DB ) and a longer o;:clliral bar (V8) , '1'h :e are flexihlv articulated with the bow. Between the ends f the dor .J and ,'enl ral bars' runs a short oecillsor muscl. (Olpl) . Irs contraction com­pre S the closing valve, cnusing it LO bend loward the bow and close the spiraclII"r opening. From the tip of Ihe ventral har a long, slender dIlator muscle (d.,'pl fOUlS venlrad and caudad to irs rigin on a peg.like invagina­tion (ds!,P ) ju t abo"e the eket nf the hind coxa. T he contraction of tlt i, muscle rdeases tbe stroin on the dosing ,'aI,'e, stroLi,ghltlls the valve. and o)lCns the spiracle.

TilE WINes

Th wiugs of V u/,,,I" pe'lS')'lvamm (Fi . lOS, 109) exhibit all tbe characters common to rypical hymenopterous wings. Tboy are held flat along the back in repose. The fore wing i much laTgl'r than the bind \';ng . Tn flight the wings are held together by the I.alllll/i Fig. 87, H), r fre nal hooks, a eries of hooks on tJ1e costal margin of the lund winl: which en­gage the recurved posterior margin . or frenal fold (MF) , of the fore wing basad of the tip of the first anal vein (1.-1). This link ing of the fore and hillel wings insures their peration ' "unit and thu ' increases lheir effi­ciency as organs of Hight. It i correlated also with lhe facl thai the great indi rect !light muscles of th thorax operate the fore wings only . The hind \v;ngs, th refor •. since they :1fe operated nil' by the small nirect Hight muscles attached LO the axiIIa.ry sd.rites. would be ineffective if they wore not linked to the fore wings. The wing membrane is highly transparent, strong, ancl glassy. nd ;.. devoid o[ allY large hairs or scales , but is overed, iairly closely with ~hort, microscopic. fixed hairs (Fig . 87) . T he wing ve-

[ 66]

TUB MORrncH.O(;Y OF "VESPULA P"ESSYLVASfCA" (SAUSS. ) 67

nation is highly moru/ied, the vein being reduced in number and their urses consid 'rably changed Irom the primitive type. T he co~ta1 margin

of the fore wing is marked by a /,lerosl;9I11a PI) . A ptero ugma is absent in t.he hind wing.

Tn .drution to the above characters the fore , ing of V . p~llS')'lvaniea is folded lengthwise in repose and in the manner characteristic of nearly all members of the Vespidae. The hind wing is entirel)' without an anal lobe, the axilla ry e.'<cision apparently having migrated campI lely to the base of the wing,

Vei .. alld all 1Io.""nclaJure.- T he n menclature of the wing veins and cel ls of H yrnenopte.ra which has been most wid~y used by systematists, at least unti l ,'ery recently, is lhat established by Jurine more than a hundred year ago. Figure 107 represents a wing of V ospa crabro with the veins and c lis named a~'Corrung to the modification of the Jurincan system adopted by Rohwer and Gahan ( 11 4) . The advantages of the Jurinean sy>tem arc it relative simpliciry and the fact that il has been employed in the paSt by ma ny workers on Hyrnenoptrra and is therefore the system most com­m .Ill ly met \\~th in all but the rna t recent li terature. The objection to a c nlinuation of its use is that it takes no aCCOunt of the homologies of Ib veins of H ) nenoptera wilh those of the wings nf other orders. It there­for ' pro,·ides no foundation fo r the comparative sturues SO essential to a determinati n of relation hips and hence of phylOlteny.

The omstock- eedham system of vein and cell nomenclature (30), the main features of whic.h were worked out more than a third of a cen­turv ago, accomplished the general homologizing of the principal vein , and laid a ecure foundation for ' mparative studies. I:n applying their system to the wings of the Hymenoptera, however, Comstock and Needham 3r· ri ,'cd at condusioDS that have c mplicated rather than simplified the prob­lems of ti1C studenrs of tili order. The most serious defects of the system aJ! 6r<t propounded by its authors, and as daborat cI later by MacGillivray (86), :Ire !lIe concept of s rial veins and the explanation of tl,C mode of r luction of the principal veins :Inti their branches. The cature of these defce is so well known that 3ll explanation of them would be superi\uou h reo ne example rnay, on the ther hand . be profitably examined . Accord­ing to the interpretation3 oi omslock and ""eedham, the tip of the most

sterior vein in the fore vein of I ' <spula should bear the designation M3 + M4 + CIIl + CII2 + 1/1 + 2A + JA . Such appellations of v~ns are obviously too aunbersome to lend themsel\'es to practical USo1.ge, ~nd mnny hymenopterislll bave been discouraged from attempting the use of the Comstock-Needham nomenclature. Figure I 12 shows a wing of V (S­

tula .w/pill/rca labeled accord ing to tile Comstock-j\:eedham nomenclature modified by Bradley (20) .

[67 I

68 N'ORTIi AMERICAN VESPINE WASPS

In eflt years three notewonby attempts have ileeJ, made to sol\'e the diffi<ulties in-'olved in the application of the Comsto<k- eedham syst<m iI$ it 113$ bt!en heretofore interpreted. Forbes (64), Tillyard (144, 145 , and Ross (115) have advanced newer and simpler interpretations, all of which have many features in common, though arrived at by different lines of reasoning. These three author agree in the ,·iew that Com ta<:k and Needham er red in the basic assumption on which their inteJ1>retation f hymenopterous wings was founded, namely. that simplification in Lbe ven­ation of Hymenoptera has taken place through a 10 of cross-vein, through changes in the ourse of the orancll. of the principal vein, and lhrough the apical fu iOI1 of many of these branches in the manner in which ins Clll and 2A have fused in dipterous wings. Forbes, T illyard, and n hold that much less shifting in the cou.rse 01 ma in \'eins 3Jld mucb less fusion have oc<urred, that many of the branches o f Ule main veins h:n'c been lost, and that a c asidcrable number of cr ·vein. have been preserved.

]7orbes's interpretation was uased on a comparison of the wings of Hymenoptera and Coleoptera. Sin , howenr. any re/'ltionship between tile, two order is plainly trernely remote, any im,larities between Uleir wing patterns could have no great phylogenetic significance. Other things being equal, 'herefor , Forbes's system could have no more val idity than that of mstock and. ' eedhanl, however useful it nllght be. Hence it will nO! he COn idered bere exc pt to point out it< principal features. Accord­ing tu Forbt!s, Radius in the primitive hymenopterous wing i three­bra ll clied, Media is uobmnched, nnd Cubitus is two-branched. The n'i"" wliich, in the Cornstock-:..reedham interpretation (Fig. 112) are termed R.f, ru, the fin.t abscissa of JfZ, M J + 4, and M4 + CI/I, are con­sidered by Forbt!s to be cross-veins . The vein commonly temled M4, 'the fi rst abscissa of the free p"rt of 313 , md the ",edial eros - ein are inter, p ... ted as consecutive segments of Cu. The !>a'Oond ab .. issa f the fr e part of .1 /3 is interp reted a CIIl.

Tillyard 's nomenclature i ba>ed on a rudy f certain fossi l insects chidl)' from the P rmian of Kansas. These were belie,·ed by hi,n to have bt!cn nn estral to the T:1ym nopt m, and were made the basis for the de­. cription of a new order, the ProlohYlIIl!Iwplcra (Tillyard, 144). Accord­ing t(l Ross ( 115) , " arpenter (L930) has shown fairly concl usively that TiUyard was in error ( I ) io the original interpretation o f the venation in the P rotohymenoptera and (2) in suppa ing the Protohymcnoptera to be all ied to the Hymenoptera, since he former arc apparenlly close rel'bves of the Megasecoptem."

r.cording to illyard, Radius is usually unbrJ.nched but may be two­branched, ledia is two-brJ.l1thed, and ubitus is unbran<hed. R4 . R5, tile

[68 I

THE "ORPl! LOGV OP "VESPULA P£NSYLVM<l .. (SAOSS. )

firs t absci of ill 2, M3 + 4, Mol +Cu1, and the secont! ab5cissa of the f rff pnrt of M 3, as tIlesc vein. are commonly interpreted, are considered oy T illyard to be cross-veins. 1014 . the first abscissa f the free pan of M3, and the Uledial eto -vein , as commonly interpreted, T iUyard bolds to be co cutive segments of Cu. Figure 113 haws a fore wing of DO/id,ovts­",,'a wenana labeled according to T illyard's interpretation.

The interpretation of h)1Uel10pterous venation by Ross 115) is based on the research - of numerous students of insect phylogeny who have at· empted to determine the ancestry and relationships o f the Hymenoptera,

and on a critical compari on of the wings of mecopteroid insects and Lbose of the llymeuoptem, many indications pointing to the mecopter id insects as the group to which U,e Hymenoptera are most closely allied. T he inter­pretations of Ross, therefore , in all probability, have greater validity than any others yet proposed. They have been adoilled by the present wri ter as Ihc basis fo r lhe systematic section f this paper. Figures lOS and 109 ,how the fore and hind wings of V IS/mla pensyltlClIlico witll lhe veins and cells named according to the interpretations of Ross.

The main point of Ross's interpretations are as follows : S ubcDsta, when present, is o f U,e u ual type. Sci fuses with C, ScZ with R. Ab ence of' ubcosta may be due to atrophy or to amplete fusion with Radius.

Radius is tw branci1ed, Rl forming the terior boundary of the ftigma and the an lerior boundary of the wiug beyond the stigma. The other bral1ci1 of Radius is Rs.

1 edia is uobranched. At its extreme hase it is often fused with Radius, LUI it sho rtly leaves Rad ill as the vein commonly interpreted as the cubital , (cm. Tt is soon joined by Cul . UlOugh in most IIvmenoptera the free part of C,d, basad of its union with M, has atrophied. Media and CuI soon ,ell:U"'4 te again, Media running diagonally toward Ihe stigma as the vein t"mled ,"-e" in the usual interpretation, ",-cul in T illyard's interpretation , 'lJld I:!asal ein in the J urinean sy tern. Media then [uses with R.s for a .hort distance. or, rardy . is connected to Rs by a cross-vein "--11/1, or it tll<;e,; ", itl! the radial stem ior a short distance. Media then separates from it, radial accompaniment and rull . direct!) to the outer wing margin .

Fr m its point of "'paration with :\ledia, CuI runs directly to the ouler wing mar!,';n, eJ< opt for a jog in the form of the vein commonly inter­prded as the first abscissa of M3. ,,} bifurC;'ltes at (his jog, the usual ,",ond abscissa of M3 being interpreted a Culb, and the usual serial vein m& lIZ. being C .. ) a..

There ar iour anal veins, lA and ZA being two-branched. Only 1.1 rcm:uns in the wings of Vesllllia .

Veins R4, R5, the first abscissa of the free p,ll't of 1Z, and M4+Cul, <J f th" usual interpretation, ar interpreted by Ross as cross-veins.

[69J

70 NOllm AltHJllCAN VIISPIN£ WASPS

The detailed structllre of We axillary sclerit ,their relationships wiw the thoracic sderites, and the mechanism of fl ight wil l not be considered in this paper.

T HE L EGS

The legs of V . l>c"s),lvo"ica (Figs. 98, 99, 101, 102, 104-1(6) show comparatively little pecialization. Thcir functions are rather generalized and consi t chieRy of walking, of clinging m vertical or overhead suppons, and i holding prey. The function of walk ing is exercised during We .earch for animal prey o"er ground and beneath vegelation . nnd ill lbe performance of "arious lluties about the nest. Tbe clinging function comes into p y during the gathering of material ior paper making md while thi i being incorporated intO the nest . during the preparation of animal food to be f to the larvae, and during Ihe actual feeding of Ihe larvae. [n addi. t.i ~n to being used for walking and clinging, the log . as in other Hymen­optera, arc used to clean Ih various pan of thr body. In adaptation 10 the d eaning function, th fore and hind legs. on whidl the work of cleanin{; largely devolves, show certain speci" lizations, as described below p.71) .

The leg, of all three pairs a.re moderately long, thougb not especially slender. The front legs are the shortest nnd We hind legs the longest. • 'J he COXIlJ! (Cx) of all the legs are lnrgt and more or less conical.. pro­]ccung frerly from the hooy surface. 'The fore cO>CIe are the largest and the hi nd coxae the smallest. The difference in size between the fore and the midtlle coxae is quite evident, that between the middle and hind coxae is slight.

The Irocha."t'rs (Tro) an: small Md closely articulated with the femora Dut exhibit no noteworthy modi6catioll5.

The ftmora (F.) are subcylindrical. and are arched lightly outward, so trot \ hen 8exe(] they fit snugly against the unded sides of We thorax. T he upper surface of CAch femur where it Ii against the thorax is Bat­tened. The same i true of that part of the lower surface ngaiost which the tibia is 8exe(] . Each femur is somewhat enlarged toward the base so as to appear cluD-like. The greatest diameter of the ntiddl· femur occurs very n~r its proximal end, that of the hind [emltr a little further di5bd, and that of th [ront femu r n good third of it length from the base. The ,:,iddJe femur is the slendemit, the hind femur is moderately stout, and the lronl femur is still more so. The hind femur is sl i h Iy longer than the middle femur, which in tu rn is slightly longer than the front femur.

The l ibia~ (T') are subcylind rical, Battened on the surfaces which flex agninst the femorn , and gradually widened from nnrrow ba.aJ articulations with the femora to broad truncate apices. The [ore tibiae differ from tile

r 70)

T ElE )lOIU'1l0LOOY O P " VEsrULA PENSYLVANICA " (SAUSS.) 71

others m being thicker in the middle. The fore tibiae are two-thirds the length of the fore femora, the middle tibiae are between three- fourths and iOllr-fifth as long ' the middle femora . and the hind tibiae slightly exceed the hind femora in length. Each fore tibia is provided with a single large libial spur (TiS) attached nearly a third o[ the way back from the distal end. The middle tibia has two spurs, which are barely subapical in attach­ment. The bind tibia also has two spurs, and these are distinctly apical in l)Osition.

The tarsi (Ta) nre comparatively unmodified. The tIIetolarsi on all the legs are much longer than an: of the ther segmt!ll!.s, being as long as, Or longer than, the next lhree segments combined. They are subcylindrical . The front metatarsus is stoutish; the middle and hind m.tatar.i are slender. The fr nt metatarsus is scarcely more than two-thirds the length of we i ront tibia, lhose of the middle and hind arc progressively longer, though still &h rtet than the corresponding tibiae. egments two to four of the raT' i on all legs are flattene(] do~-\'cntraJJy, bilobed apically, and fringed distally by &tiff, spine-like setae. egtnents two to fOllr are subequal in length . The tenninal tarsal segment i half again as Ion as any of the pre­cedin!: three . and is provideu with tw simple claws Md a median pad.

TlI6 an/t"na-clcani1l9 dn;ce or slngil (Figs. 98, 99. 102).-The anlenna cJcaner consi ts oi the spu r on the [ore tibia together with the proximal half { the metatarsus (UTa . The spur is large and curved, the in ide of the curve being directed toward the melatan,-" and provided with " win, knife-like scraper (SCT) with a finely serrated margin . The metatarsus is excavated ba.sally on the side toward the tibial spur. This excavation is lined with a brush of fine hairs. When cleaning the antenna, w e pr lC imal part oi the antennn is plnc.ed between the metatarsus and the t ibial spur, and the Clrsus is Bexed until the antenna is held tightly. The anlenna is thc:n drawn thr u"h the cleaner and freed of dirt, pollen, and ther foreign maher by the scraper on the tibial pur and the brush On the nl"latarsus. The debris which i collected by t.he antenna cleaner is then removed from he btl"r by drawing the forelegs through the mouthpans . The debris is

probably temporarily stored in the gnathal pouch (p.25) . The tuilllrcloa"j1l9 devicc.-The hind leg also constitutes a cleaning

npparntu . The hind tibia is provided on i~ lower and posterior surfaces with an extensive brush of short hairs. The metalaJ'liUS also is provided with a brush o[ hairs. T he tibial spurs are broad basally and slightly bent. Tn cleaning the wings, the log di lad of the femoro-tibial joint is pushe(] backward over the wing surface. Entenl30 ( 52) d · cribe wasps of the genu Polisles tIS using the fordegs also in cleaning the wing bas . The old book, Naill" Display'd (93) , Slates that the tint use of tl,e hind 1"1.'" by a newly emerge(] wasp is to wipe off the wings.

(71)

72 SOIlTU Allt!:RICAN' VESP I~.E WASrs

THE ABOOMEN A ' D ITS APPENDAGES

T HE \anOlllNAL S£G .. £"TS (Figs. 110. 114-126, I3l)

The abdomen 0 f the female of V"tmla Ptll.1)'/t,atl ira is elongate-conical, pointed at the apex, slightly narrowed basally before the middle. and then abruptly and squarely truncate at the extreme b::tse, joining the thorax by a narrow and extremely short pedi<:.l (Pel . T he abdomen is lighlly com­pressed vertically. mewhat flattened ventrally. and strongly arched dor­sally .. U de from the propodeal seg111ent, which . though really:ilidominal, is more closely associated witb the thornx, there are six vi iule segments. T hese are segll1<OJlt two to Seven of the rnI abdomen or segment One to sill of the g;c.tcr.

Each rypica abdominal segment is ring-like and consists of two large ' clcrites, a krgu", (Tlf to TI'/J), and a sl67'lfu", (SI[[ to StVIJ) con­nected laterally by an infolded membrane. the conjullcliva (COli). There are no pleural seleritcs in an abdom.inal segment.

In all tbe exposed segments of the abdomen, the tergwn is mucb larger than the sternum, overing lilt do rsum, he larger part of the ides, and a litt le of e ventral sur face anteriorly on whi h it is reRected. T he t.rgum is narrClw "r on i posterior margin than on its anteri r margin. As a result the lat ral margins are diagonal.

In the second segment the tergum is even lar~r pr I'o rtionally than in . m' Is three 10 si .. It presents three subdivi ion : a very small anterior

portion forn ling lhe dorsum and sides of the short abdominal pedicel: " large ver tical face overing the truncature of lhe abdomen; and a ring-like portion continuous abo,'e and laterally with the margins of the vertical face. and coveri ng the dorslIm, the sides, and most of lIle venter of the segment. This t.rgum (Fig. ltO) i. bluntly angle<! " t the truncature in .uch a manner that the vertical face is rather rhomboidal in shape and pre­sent. a dorsal and two lateral, 'lightly umbonate elevati ns (u). TIle sec­ond ,ternum (StU) is reduced to a narrow transverse triangle (F ig. 115) .

In segment sev tbe tergum and sternum are not o [ the ring type just de cribed but are in the form of triangular plates which meet along their lateral edges. This form has been derive<! from the nom1a1 ring [arm by extellsion caudad f the ",edial portions until the postero-Iateral angles have been eliminated, and by a reduction in the amount of lateral overlap­ping.

At tbe point of ar 'culation with the thorax, the anterior margin of that portion of the second tergum which covers the pedicel i .. recurved upward to fonn a transverse gToove into which hook the propodeal teeth (Figs. 39. 40.43-47,79 : prl) (1'.62). Tnterrupting this mal'!{inal gToove medially is a short, longitudinal ridge, the verlical rial. of the (Jodicel (VPd) . which

In]

THE MORPHOLOGY OJ' .. 'SSP LA PENSVLVAlH A" (SAUSS.) 73

extends back along the midline of the pedieel to a pit at the base of the Irunenture of the 5egnlCUt. The pit behind the vertical ridge serves as the i rti n (or the tendons of the propodeal elevator muscles of the abdo­men ( Fig. 81 , Jadl1 ).

Segments tw and three articulate laterally at the ends of the transverse second sternum ill a close joint which permits up-and-down movements only. The third sternum bears a deep transverse groove (Igs) extending between the articular points. Anterio r to this groove there is developed an articular surface. the 11"lerior jlange (FStTrr , oYer which " somewhat silllilar l'on"";Dr lU'licuillr jll1>19< (FSIIT) o( the second sternum slides. Then, is 110 close restrictive articulation between any of the otber 5e!,'T11ents . AU the contrary t.hey arc joined only by eX lensive conj uncti\'ae (Con) so that "ach of them may be telescoped into the Olle preceding. The articula­ti ons between the t.rga and the sterna of the other segments, however, a re 'Iu ite restricted. Little if any vertical .l<JlIlnsion and contraction of the sel(l1lcn ts is possible. As a resu ll. therefore, the expan ion and contracti n of til . abdomen necessitated by breathin~ is accomplished almost entirely by the alternate extension and telescoping of the segments.

!\ typical tergurn. such as occurs on segments four, five, or 5;',<, when flattene<! OUt (Figs. 114, 13 1) i roughly a trallSverse oblong. Its posterior and ~.teral portions are refle ted internally and clo y appr~ to the dorsal portion to form an exten ive redup/iCaiiDn (Figs. 117, 13 1 ; RT). T ltis strengthens the tergum and pro\'ides a scltronzed bearing surface to glide o\'e.r the b::tse of the succeeding segment. Th anterior margin of the intcr<;egmental conjunctiva is attached 10 the edge of the reduplication (sh WII in tbe figures by brok<!ft lines). Each reduplication shows rwo btoarl. ronnded emarginations. T hese provid~ for the origins, on the ento­d rsal surface, of the intertcrgal ·tensor muse! . (iu). Immediately allterior to the lateral limits ot the reduplication the tergum is crossed by " st rong internal ridge, which is presumably the anle,-oslal ridge (Ae ) . Cephalad of the antecost3.l ridge i a narrow transverse strip, the precosla (I'il(. 13 1. Pre). to the anterior edge of which tbe posterior margin of the cnn junctiva is joined. The lateral angles of the precOStae are invaginated to fo rm broadly triangular Iffgal apoelelllcs (ApT ) . The precosta is mar· gine,l anteriorly by a ridge, e anlmor I.rgrn brac. (ATB) . Dia!1olllu Irrgo/ brace. (DTB margin the tergal apodcmes internally and cross the precosta to the aoteco ta. LaieraltergaJ braCts (LTB) margin the tergal apodelOe5 externally.

A typical sternum (Fig. 114) is lik · th tergum in most respects, exhibiting a posterior "duplication (RSt) , an aHlecosla/ ridgo (Ac ), .sl.,... "al apod.,He .. (ApSt), an alllerior .·Itrfl. / brace (ASS ) , lateral sternal braces (LSS), and diago',,111 stl!1'./lJI brIJCe; (D B). si de from differ-

[73 I

74 NORTU A)lEIUCAJI'i V P. PINE WASPS

ences in outline which arc clearly shown by the figures, Ihe principal differen~'C:s nre the following : '!be precost i. reduced in width medially. The sternal apodemes are proportionally longer, narrower, and differemly shaped, having distinct necks. The anterior sternal braces in some indi­viduals thin down almost to the point of disappearance.

T he rerga and tema of segments two, three, and ~Yen differ from the others for obYious reasons. The external features of tergum 11 haye al­ready been de:;cribed. If! addition. it lacks the cmarginations of the redupli­cation. as there are no illtenergnl extensor mu.cles between terg:! II and Ill. Sternum /I exhibits on the pedicel a transverse groove and an aCCOm­pallying ridge which cut off a small basal sclerite. Posteriorly stemite Ii i lIenl dorsad, and the.n cnudrul to fonn the articular fiallgc (F 111) pre-

iously rderred to (p. 73). This Bange is probably" modified redupli· cation.

Terguru III e"bibilS no I)()stcrior specializations. Anteriorly it is spe­ciali,ecl in several respects. The precosla is of uniform width, and lack the tergal ajJoderncs. There are 110 anterior, lateral, or diagonal tergal braces. The precosta, the anlecostal ridge. and the adjac .. nt part of the POSICOSIa arc shaped in section like an I-beam, the upper trans"er~ member of the Ie IUtituting an articular fiang, (FTIII) "hich bea ... again .. t the redupl i­cmien nf tergum IJ.

lCrnu01 III is typical posteriorly .. \nteriorly it is narrower propor­tionally tban the following SIerna. In place: o [ an ante<:osW ridge of usual type. Sl rnl1m J II presents inlernall a broad, rounded ridge. vhich is the internal evidence of the presence of 0. deep rounded sulcus (Igs) . Th., Jl r ~-..: ,,.\::I of sternum II I i widest medially, narrowing to prnctical blltera­tion laterally. T he prccosta func lions as af! anterior arlicfl/ar fiallg"

FSII[J) which bears against the posterior articular flange of sternum II. The sternal apodemes are long. slender, and tapering, and they project mesad rather than lalerad . as is the ,se with the other Sternal apodemes.

Tht tugum TI'IT) and sternWT1 (StUI) of lhe sixth externally vis­ible segmenl of the ga. ter, with two exceptions, exhibit no structural modi­Il .tions excepl peculiaril i of outline thaI ar" corrd.atet.l with the func­tion of these sclerites in dosing the abdomen posteriorly. These pecl1liari­ties arc well shown l>y the figures. On sternum /'11, bowever, the pre­costa is complelely interrupted medially, and the apex exhibit. a median notch (StiG) which sen' as a stillg g14ili •.

TUE Muse LATURE Of' A TvI" AL ABDOMINAL SEGYENT

Segmen four, fi ve. and six have idwtic:al musculature. All of the uscles of the abdominal segments tw t 5e.Ym. incidentally, • re compara­

tively lhin and ribbon·like, though thei r oblique course of len result,; in

[74]

THE MO RPHOLOGY P "VESPULA PE...'iSYLVANICA " (SAUSS.) 75

their origins or i~rtions having an elliptical outline. The individual muscles of a segment are as follows :

Dorsal ;,,'.rl"'gaJ r,lraclor (ditr (92». - This OlUSc\e in Vcspula . ~,,~ept in segments two. six, and se ven, is di"ided into two portions, a

1;. narrow inner portion and ~ much wider outer portion. In ~Wt5 tw and !Ii" it is undi ided. The dorsal intertergal retractor arises dose ~h i nd the antecostal ridge of its segment and just m d of lhe diagonal t<rh"u brace. It is inserted clo~ to the medinn line on the ant"rior tergnl brace of the tergum following. 10 ~gment two tbe dorsal inte.rtergnl r,,­tractor is quite broad; it originates on the truncarur o f tbe segment and j , inserted on the antecostal ridge of leTgutn II I. Segmt.llt seven lacks ,lor..a1 ill tertergnl retractor, unless the muscle labeled litr' in figure 117 L a dorsal ill tertergal retractor, which seems unlikely.

Latrral illttrlrrgal Tclractor (lilT [93 ] ).-This muscle arises along a diagonal line ,)11 the tCTgum laterad f and partially underlapping the dorsal intertergaI retractor, It rims diagonally caudad and laterad, crossing, on it» dorsal surface, the apical portion of the ttrgal apod=e f the succeed­in!! , cgmrnt, to be inserted on the apoclemc do e to the antecosta. The iotertcrgal retr'detors act ing alone, r in njunction with the lDtcrsternal ext.'bOr (iSt) , straighten the abclome.n and elevate its apex. In conjUllc­tio11 wi th intcrs lcrnal and tergo-sternal retractor muscles they bring about a tc1c;coping of the segments.

I n ~ .. nts lhree to six the wd of origin of the lateral intertcrgal recr.CtOT is slightly bilobed. In segment two the muscle is separated ttc."lrly to its insertion into two masses of fibers. In segment seven the IIl llscle labeled lit'" in Figure 117 appears as th probable homologue of lhe lateral intertergal retractors of other s""menlS. It arises, howeve r, on the t rgal apodcme and not 00 the usual place behind the antecostal ridge.

The. jlltmergal '.T"tC1lSor (it. (94» ari5e. n the entodorsal 'urfnce in the emargio.,tion of the tergal reduplication, extends cephalad and I. t­,orad. IlDd is inserted dorsally on the apicnl portion of the IUgai upodeme of e sue ceding tergum. The intertergal extensors acting alan', r in c"njunction with the interstemal retraclors (lisr, 1II;ff) , curve the abdomen downward . In conjunction with inlerslernal and tergo-slernal extensors thty bring about the e.xlension of the abdomen.

In segment two llle intertergnl xtensor is lacking. Because of the do,~ union o f t 'rga 1/ and II I and sterna 1/ and III. the points of union . erving as f ulera for the movements between segments two and three, e " " mt ~ undergoes movements of elevation and depression only and

i, not extended, a whole. With such a relation lip existing, an inter­Ic al extensor in segment two would be sl1jl<!rfl uous and incapable of fu n tianing.

[ 75 J

76 NORTH AMER ICAN' VESPIXB WASrs

M~d;'", inlersternal rtlrlUl"r ( til"" (95]).-This muscle. except on sternum lIT, arises rn1the anterior sternal brace just mesad of the diagonal stemal brace, runs caudo-mesad. and is inserted close to the midlme on the anterior sternal brace of the succeeding sternum. The median inter­slernal retractor appe4rs to be lacking in segments two and seven. though it i possible that the muscle labeled lisr ' in Figure 117 is lhe homologue of the median intersternal retractor of ther segments. In segment three, !.he med ian interstcrnal retractor arises on !.he sternum behind !.he rounded transver ridge that replaces the ante<:osml ridge.

The IIJUrai inJerstrrnai relraclor (lis>' [96]) arises on the stema1 disk behind !.he junction of the diagonal 5t 'rnal brace and the antecostal ridge, runs caudo-Iatemd, and is inserted laterally on the ventral urlace of Ihe sternal apodemc.

The lateral interstemal retractor is lacking in segment two. In seg­ment seven i~ apparent homologue (liST') arises on the sternal apodeme. In segmenl three the origin f!.he lateral intersternal r tracto r is bounded

by a bon diagonal r idge. . . . . . Th intl!rStemal retractors acting alone. or tn conj unction With the anter­

tergal extensors. Hex tbe abdomen_ 1n conjunction with int rtergal and tergo-sternal retractors they telescope the a1xlomen . e lack of both median and lateral intersterna! ... tractors in segm.nt t\\'o is simply ex­plained. egment !.hn:e does nol shar' in the telescoping oi the abdomen. and the .maU amount of flexion between lerna J I and r r r is easily hrought about by !.he weight 0 f lhe abdomen alone.

T he j,,/crslerlUJi e ... ·I.".$u .. (ise (97) ari~s on the entastern,.] surface in the emarginalion of the sternal reduplication, rum cepbalo-Iaterad, and i, inserted on the inn.,.- half of lhe apical margin 01 the sternaJ apo<Ien1e of the succeeding sternum.

'rhe interstema1 extensors, acting a10lle or in conj unction wi!.h the intertergal retractors, str.ughten the abdomen and elevate its apex. In Con­j unction w,th the interterlr,,1 and tergo-sternal extensors, they eJ.'knd or lengthen !.he abdomen . The intersterna1 extensors of segmCDt two, bow­ever. contribute n.ly to Ule straightening or elevntion 01 [he segments f the gast.,.-.

The oul'" 1~,ga-strnlaJ rolraclor (alsr (98) in segment fO ll r, Ii" • and six a rises on the tergum a sh rt distance dorso-c.a.udad of the spiracle, run cephalo-ventrad to the temum of the same segment. and is inserted on the lateral sternal brace just caudad of the nte<:astaJ r idge.

The outer tergo-stema1 retractor is laclcing in segments Iwo and three. Tn segment seven its rigin is typical, but its insertion is located laterally on the sternal apodeme instead of on !.he lateral stemal brace (Fig. 117, olSl"' .

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THE MOflPUOLOGV OF " Y&sI~U J.A P B="lSYL AN"ICA" (SAUSS.) 77

The outer tergo-sternal .. tractor raise the anterior portions of !.he 5t.,.-na and thus contribute to the flexion of the abdomen.

The in~cr fn:go-slcrnal rel'tUIOT (it.rr and jiST' [99]), in segments four, n,·c, a lld SL,< . anses on the tergum between the origins of the lateral inter­t <gal retractor and th · outer lergo-sternal extensor, runs cauda-ventrad, ami is inserted on !.he edge of the sternum of !.he same segment immediately allterior to the sternal reduplication.

Tn segmell ts two and four to se"en. the inner tergo-sternal retractor is cionglll fan-shaped , and comparatively slender, and is inserted by lI",an of a hort tendoll. In segment three it is broad and rectangular and I,," no endon.

1e inner tergo-sternal rerractors lift the poslerior POrtiODS of the ;[('rna and thu contribule to !.he traight.ning f the aildomen and the ob" ti u of its apcx. Acting in conjunction with th~ outer terge-sternal reI raclors. they compress lhe segments.

Th trrgtrslcmal .. rlms(1T (Ise' [100]) is a short muscle that arises On

Ih~ Ia ttrJJ part of the te rgwn in the angl . between tbe rergaJ reduplication alld the anrecostnl ridge. runs ccphalo-dorsad, and c.QO\·erges I its insertion on the lateral angle of Ille slernal apodt!lne of the sternum of the sallle segment. I t i lacki ng On segments two and three.

\ cting :a.lone or in conjunction \\~th the intertergal ~traClor and inter­sternal extensors, the teTge- r mal extensors contribute to the straighten­aug 01 the abdomen and tht: <'Ievation of its apex. In conjunction with tbe illllTlergal and int.,.- tern' e"tensors they bring about the lengthening of tLe abdomen .

• \ ,ide fr m the movements accompli hed by the differem muscle groups a~ O11tl rnLd an I b~ ~receding paragraphs. movements of lateral bending and ,,. t" r"-lOn r tWlsbng are produced by Ule co-ordinafcd acti"ity of \'urious oj rhe abdominal muscle .

-\U IJOlll"'iAL ' l'IRACLES (SplJ TO pVfll AND THEIR MUSCllLATURE

. ~ "en of the segments of th gaster bear spiracl laterally on Ille tccgn. 1 hOlst n segments IWO and !.hre. are "I 3 YS exposed to view: those on

<'lJllImr eight, since it hares in Ille comlXlsition of tbe sting mechanism (h '. I I , IJ9, 121 , are always conceal d; those on segments si. and . evt n ar usually concealed . but are exposed during extreme extension f the aGclornen : and tho. on segments fou r ,md five may be a1lernatdy ex ,ed and concealed during respira tory movements.

The spiracies of the gaster ar essentially like U,ose oi the propodcum; hence a desc . t' . I' . ., np IOn I~ unnecessary. !oach IS dosed b)' an occillsor musde (Qs/, lf to "sf/Viff 1.101]) and is opened by a diialor m"sr/r (dspll to ds/'Vrrr ( J02]) . The origi n of the dilator muscle is On the lateral angle of

en I

78 NOR.TH "lfRR..ICAK VESPINE WAsrs

th~ sternal apodeme internal to the origi n of the terg sternal utensor muscle.

TilE STINe A>I"O AsSOCIATED STRUcrURES (Figs. 118-126)

Segments ~ighl. ni~, nnd ten of the abdomen ""' muc.h reduced in siz~. are highl), modified as parts of the sting mechani m, and are om­pletely withdrawn into the seventh segment by the e.xten i.e invagination of the seventh i n ter~""taJ ' nj unctiva (Fig_ J 18). The terg-d and tema of segIlJenrs eight to ten, however. retain lheir norm31 relative

positions, bllt their modifications are. e.~ten.sive th.,t their identity i not apparell! .xeept in the light of comp:trativ. studies. The cavity formed betw.,.n the tergum and stern um of Lhe seventh segment as a result of the invagination of the seventh intersegmental conjunctiva is called the sting rccrss (SR.) .

T he eighth tcrgum is represented by two large Ia/eral lobes (LTV//i) . ~"c.h of which bears a spi rae! (SpVlll) and a narrow dorsal ttrip (DTV I II) connecting th~ lobes. The eighth Sl mum i represented by a pai r of small somewhat triangular or lI,r6c-an"6d sci<!1"iIU (TSV/ll , the :ulterior (morphologically ventral arms of which are continued into the . rting lallc6t.· (first gonapophyses) (SL) .

The ninth tergum is repr ented by a pair of roughly StJlliLircuWr sci.riJa (LTTX), which, like those of me eighth t~rgum , are connO!Cted across the dorsum by a narrow, sclerotizcd dorsal strip (DT/X ). The ninth segment has DO spiracle. T he semicircular sclerites of the ninth tergll m correspond 10 the "quadrate plales" of the honeybee as described by Snodgm (1 ) . Each articulates anteriorly (Figs. 120. 124) with

tic. of the arms of the trianglliar plate a! the eighth sternum. The ninth sternum i represented by B pair of roughly <ntal or oblong "Ialu (PSI) . T hese (tre produced anteriorly imo a pair of .talk!, the slalks of til" "inlll sl.Nlt/1Il (SSt/X . the extremities of which bear the arms (ASS) of Ih. sting slo~~/' (second gonapophyses) (S ' ). Posteriorly the plates of the ninth t.mum bea r the hai ,finger-like, .lling "alpi (SPa) . which are the third gonapophyses. e median portion of the ninth emum below the .ting palpi is quite devoid of any membranous lobe such as Snodgr ... describes for the bee.

T he teDth segment is largely membranous and cODsim of two flap-like lubes (TX and SOC). between the apices 01 which is the anus (An ). The Irrgal lob~ (TX) exhibits a pair of slightly oblong plales which ar~ ap­proxima tel:l baslLlly and fused to some extent wi th the narrow 1!"3l1!!\"erse median bar uf the ninLh tergum. TIle tergal plates of the tenth segment are weakly sderatit e:<<:epi for a pair of disc:U. areas which "rc more strongly sderotited. Caudad of the h",,,cily scleroli.erl aJ"ea..', the tergal

[78 1

TUE 1I0Il..PHOt.OG\' O!l' "v PlTLA P£NSYl.\'A...''flC,Ah (SAUSS. ) ; 9

plat hear a numbB of loug hai r . The sl/!NOal /Qb. (SIX) of th~ tenth se),'I!1efIt show only the fai ntest sclerotizatiOD-

The .ting is formed of the modified first and second pairs of gona­(lOplly . The second gonapophy5eS. rne by th oblong plates of the lIinth lemum, are fused in tbeir distal halves to form U,e sting sheallo SS). The sting sheath is roll d vCDtrnd along its margins to form an

in<nmp~ett: tube, w~id l is o~ beneath. Th~ proximal end of the stmg '~tath I expanded IntO the sun bulb (SB) . The dorsal posterior margin

I tbe 5ton~ bulb hears a paIr of heavy arms, the "'''''' of t/oe tling bulb (.·Jr8) . which curve upward and caudad , c nverge, and fuse 10 foml a . trong V- haPf:(! structure, to which heavy mUscles are attached. The proximal halves o f the second gonapophyses form the slender upcurved ,Irm> o f . th~ sting sheatl: (.-ISS). These articulate anteriorly, as already n" lcd, WIth the extremlttes of Ihe stalks borne by the oblong plates of the nill'h . t mum. The first gonapophyses, borne by the triangular plates r~preH'llt ,"g the sternum of the eighth segment, are modified into the long. . ender sling lanetls (SL). U'e proximal hal,'os of which slide along the afl lls of. the sting sheath. The di tal halves of the lancets enter the sting ",.,11. at It · base and tend thsough the sting heath to near its tip. Tbe stm~ bnee o f the wasp are simpler in Structure than those of the bei! . . n,"lgrass (128) describe' the lanCl!t:!! of the bee's sting as having valves all ~chcd to that portion of them contained within tbe . ting bulb. The aCl lun. o f the e valves, as the lancets are worked back and forth thr ugh the ' tong sheath. pumps the venom out t.hrough the sheath. There are no M Id \'alves in tbe sting of the wasp. The barbs (bs/) of the sling la"rol of the wasp are also smaller, proportionally, than tbose of the bee.

The PQison sac (PSa) of the wasp i a large elliptical c lying just ~~1t" roor 10 the snag apparatus and to the left of Ihe middle of the segment. I he leftward displacen1ent is due to the presence of lhe rectum Rr) in Ihe mIddle of the abdomen . The poison duct (PDtI) is a stout tube ex­lelldong in a sinuous cOUrn! r r 10 the p sterior end of the poison sac to th ('ase of the sting bulb. [t is with li t obvioll. musculature.

THE WORk'ER

Tbe worker V. ImlSyivall;ca is practically identical in external nl0r-hoi . 'U th f rtiJ f "In I\~ lee e emale r queen, the only obvious difference being

that the. worker is considerably smaller. Various internal differences in the proportIonal development of the ali mentary canal. the reproductive y tem " lid lh~ fat bodies of course exist . but" discu.sion of these lie ' olltsid~ the "mIls of this paper. There is also a difference in the ranee of varia­to On Ot O,e color pattern in workers and queens, the yellow element in the

[79 J

80 NORTlJ AM£R.lCA.'t V£;SPINE WASPS

pattern being less developed in the average w~rker lhan .in the a,:erage queen . This subject is more fully discussed In the sectIon on bIology

(pp. 175-176).

THE MALE

The Inale wasp, wbile agreeing in general with the worker and queen, exhibit.~ a number at stnlctural peculiarities. These, as is to be expected. are (ound chiefly . though not entirely. m the parts accessory to the iune­

tiun of productio"-T he male is intem,ediate in size between tbe worker and lhe queen,

but is distinctly more slender than either. It also has longer and 51 ndercr

anlennae. THE HEA D

T he bead of the male (Fig. 100) i about the same in proponions as that i tbe female. The clypeu i very nearly squarcly truncate below, in <tead oi being sinuate-emarginatc. and the diagonal lateral portion. of the lower margin are more nearly straight than in the female. The. d?rs:u Ula rgin of the head is not depressed near the ocellar tube~c1e as It IS I!\

the female, but i slightly raised . Tbe lnter-antennary area IS less elevated than in the (emal • . less or not at alltubcrculate Uetween lhe antenna! bases, and the impressed line running do\ nward (rom the anterior ocell (fs') IS les developed than in tlle female, being visible iu only the upper hal!

of its tour e. The aJltennae ( F ig. 103) are 13-segmented, instead of 12- egmentro

as ill the female. T he scape, including the bulb, i a little shorter than the I,edice! plus the first flagellar ""&'ffi<'nt . whereas in the female it is dis­tinctly longer. T be fi t.! flagellar segment is f about the same propor­tions as tl,at oi tbe female. Tbe remaining flagellar segments are longer proportionally than those of the female, ,.arying from one-half to two­tltird as long :u U,e 6rst, instead of frorn one-third to one-half as long. The terminal segment i·· subcyliodricaJ, taper lightly, and bas a rounded apex. There are n 1)'loWs (Fig. 128, Ty) on the ant nM of the male

I . pClIsylv,micll. THE THORAX

The thora>: f the male 3~rs to be mare nearly globular than that of the queen and wurker, otherwise it i identical.

THE BDOMEN

T here are scven exposed .~'Il1ent in the abdomen of the male (Figs. 127, 13l-137) a5 opposed to six in the female. These are all of typical (oml

lllO]

Tlffi ~ORl'ROLO<;Y 01' "VESI' lILA PENSVLVA.\, I CA " (SAUSS.)

'><cellt lhe lut. which is the true eighth abdominal segment. The tergum ~r the '''Chtll ,;q,'lTIcnt i. narrowed abruptly at about two-thirw. the disl3Jl~e i rom its base and is then producro into a rounded lII~diall lob~. This me­dian lobe i thickened considerably. From lhe under surface of the sides of the median lobe. two thin scales (ScV III) project. T he exposed portion

i the eighth sternum is broad . transverse. and shallowly sinuate-c",argi­noHr'.

The ninth tergum is rep resented by. narrow, transveroc, membranous and dorsally, but exhibi15 two c!Qf1gate sclcrotizcd plates (LTIX) in it.

Interal portions. Th~se plates articulatl!, by means o( slender lateral arms, \\ ith the fu sed eighth and ninth temites.

.\5 was determined years ago by Verhoeff, Kluge. and Zander, accord­i l1~ to 1.I0uiange (19) . the ninth sternum is [used with the ·ghth. The fu>iull i3 not evident except on di section and examination of the resullani l~lI l1pound plate (Fig. 137). The anterior margin of the compound plate exhihit, Iil'e proces '. The middle uJlpaired process (S,,,) and the extreme late ral processes (ApStI X) belong to the ninth sternum, and tbe interme­diate lateral processes (ApSIV fll) to tile eightb sternum. These processes c.>rre. pond to U,e oomlal .rtcrncU apophyses of the other stemn. The me­,li311 unpair;,d process. Or spiculum, which was so named by Kluge, ac­N rding to Boulange (19) . and was called the "opopltyst prlomita/i' hy n .. ul.nge, i a special development. It project! far (orward inside the body <a" ity antt:rior to the eighth sternum and serves as .. base for the insertion o i III",d es which run to the gO llocardo {GC of the genitalia.

The tCDth segment is dC"eloped as a triangu la r lobe (SgX) , which Lea r, lh 011110$ (An). [t is bidden beneatJI the media.n lobe of the eighth tC l'I:um. The tl!rgal portion of the tenth ~ellt is entirely membranous Iillt the sternum consists of a wcll-develcped plate. The lateral portion~ ro r thi plate extend forward in the body cavity above the genitalia (Figs. 1 I. 14l) and provide attachments for muscles.

'flIE MALl! G&'\,ITALIA

The genitalia of the male (Fig. 127. 13l-149) are developed from the st.ntum of lbe ninth segment. They appear to ari"" in the membrane I""wtell the runth and tenth sterna, and are normally almost entircly ' .. ncealed in n 1l",ilaJ ch4mblr comparable to the sting recess of -the female. Th. iova,,';natioJl which ha given rise to the genital chamber oi the male '''"mr,; between the eighth and ninth obdomi nal segments. whereas the im':lJlillation wbich bas produced tbe sting recess o( the Cemale OCcurs lJ<:t ween the seventh and eighth segment .

T he genitalia cOllsist of ""e selS of parts. Two of these, the gOllucordo ( ';C) and the "tnis (Pen), are unpaired, and thr e. the gntloslip •• (GS)

181 I

82 Noa'fU A14£IUCAS vesPI~£ WASPS

(gono.rtipiIU, plu!a l), the ,·olIdla .Vol), and the sagill" (So). are ~r~. The unpaired comlilioo or the ~IS appear. to be secondary. It '" on­,idered to rep resent tWO originally separale processes fus~ more .or less completel) along the midline. arious names have been applied by dtfferent au tho to the parts o f the genitalia 01 male H ymenoptera, but no allempt will be made here to elucidate their synonymy. Boul3nge ( 19) presents io the fom l of a comprehensive table a comparison of the oomenc.1atures app li~ to these tructures_ Bequae," also 14~ briefly presents the terms to be met with in the literature whIch are eqUIvalent to thos h employs in ru5Cll ing the male genitalia. Reader arc referred to these. papers for further data. The significance of the tentlS applied to the gerulallll on the present work will be clear from the 6.~r",: . . ., .

T he [Jonocardo (GC consists 01 " flng-like p,ece which IS scleroll 7. d "entmllv and laterally but is m"",branou ' dorsally. It surrounds the ba

01 the gonostipites. . . The gOlloSlipilts CS) are heavily sclerotized structures whIch artIcu­

late with ch ther b lIy on Ihe dorsum. by me3JlS of a palf of rounded I bes (BL ) wbich meet in the midline. These lobes will be referred to her fter a~ the btUoJ lobes of 110. gOllOI lipes. T he margins 01 the gona­stipes are refl ected " , ad SO to end ~ar~e cOI~ca\' i ties. Together the gonostipites foml a s rt of oblong box. wlthm which are found the \'ol-"'IIae, the sagiltae. and the pen is. . .

The ilme.r wall of tbe free porlion of each gonostipes bear a hean ly cluoti .. d marginal strip (1"i '. 143, 144, 148) . Inside thi , dorsaUy and

apically. is a zooe f membrane. T his is wider in its dorsal than in its apical pornon. T he balance of the inner \\,,11 is replaced by the vo1~ella (V?l) .

The gono tipes presents three free margms-a dorsal m"rgm. an apIcal IIIll rgill, and a ventral margin. T he dorsal margin F ig. l39) fo rms a bisinuate curve whid l eods proximally in a deep. round sonus oext to the ba al lobe • . Distally the dorsal margin e"ds in a shon truncate process, the dow u lermillal process (TP1) (= the ""/lPsr inner edge" of Be­quaert), which is di rected "iSlo-mesad. The ventral margins (Fig. 140) o f the two gonostipiles are subparallel for a short d,stance from the base. They Ihen diverge strongly to form " V-. haped figure, the arms of whIch a re slightly inarched. t the point where th ru ,'ergence begins each ven­lral margin bea rs a rounded lOOI!" The apex of each ventral.:nargm .ends in a hlunt-pointed 'IIt lllral lernunal prottIs (TPl) (= the /tl'wer """" .dgc" of Beq'Illert). T he apical margin ("ou/er .dg[ of Bequaert) lorms a transverse arch between the twO tem,inal processes.

F rom the marg;nal sclerotized slrip of the inner I:onostipital \Va:1 !llere an es distally an elongale process which . a(ter Dequaert ( 14). IS here c:illed the sqlUJllla (Sq) . The .quama i broadened and comcal basally.

(8Z I

THE 'MORl)UOLOGY Of" " VES PULA P£"'~SYLVANrCA" ( SACSS.) 8J

slenderly elongate and fi nger-like distally. fringe of "'tile extends along the bali<! of the squama and around the apical mar in of the gonostipes to the dorsal terminal process.

The flo/s./la ( Fig. 145, Vol) is a large. roughly trapezoida l scientc which is s trongly inarclloo and closely appressed above to the inner waJJ of the gonostipes. with which, ve.ntra lly, it i continuous. The free dorsal ponion o f the volsella i. na rrowed to half Or less than the ventral width and is bent sharply mesad to form a heavily sclerotized lobe, the dorsal lob~ of 110 . volsella (DLV . Tbis is visible from above in the distal sinuation of the dorsal margi n of the gonostipes (Fig. l3 ) . The dorsal lobe of Il,e volsella I~as a 6~ely serrate margin and i. fringed with setae. T he fri nge of setae I conbnued around he apical margin of the volsella. T he mesal surface of the volsella is cloth~ with fine setae (Fig. 143) . At its proximG­ventral angle the volsel is produced mesad into a broad, more or less flat­wpp<.-d, "entrallub"c/e of 11,6 ,;:olsella (VTV). Around the dorsal, proxi­mal. and w~ltrnl margins of the tuber Ie tlle IselIa is deeply im'aginated to lorm a process that projec into the base of tbe gonoslipe ·. T he rela­tion of the tubercle and the invagination is such thal wben iewed from witbin ( Figs. 143, 14S) the tubercle appear to be the summit 01 a stout peg which 6ts into a pocket.

The sagilla (Sa is a c mplex process that is evaginated from the proximal ma rgin of lbe vo1s lIa. The sagitta. exhibits two portions, each oi wlllch makes up ahout hall of the ent ire structure. The basal portion is Ratt cd..: nical, and is heavil), scleroti.~ except for about half of its III I sur ' ace. The distal portion i finger-l ike, is slightly bent about a third 0 f its length before the apex, and is only lightly. lerotized. its axis is offset slightly mesad I rom Illat of the basal portion, SO that the two appear Irom within t be overlapping. separate struaures ratber than continuous part 01 a sin Ie organ. At its ba.~e on lbe veotral side tlle distal portion bears a back wardly directed Iagittal 5pillc ( as). The sagirta is hberally supplied with setae which are minute and sparse basaJl but long and ,It-nse at the apex, where they form a dislinct tuft.

Th author differs in his use of the terms \' lsella and sagirta from Dequaert (1 4) . whose brief dis:ussion of the morphology of male genitalia " the roo t recent to "ave appear~. B quant tcrlllS bOUI volseJl" and S:l<:illa a~ here employetl , the " t'al~Y1' illiernac.' · and restricts the term ""I sella to the process here called sngiua . Ilc applies no specific tcrtll to the , ,,'-'diu, as that term is used in the present paper. The usage of the pr enl Wflt er was determ ined after studying the comparison of nomenclature apphed to male hymenopterous genitalia that is given in tabular form in the paper to ' Boulange, to which reference has "Iread)' heen "",de. Accordin~ tn tlu ' ble, whenever aUlhors, in the past . ha"e IIl3de a distinction be-

18J I

NOItT({ AM£lUCA. V£.'iipr,Nl:t WASPS

tweell ,'ol.clIa and sagitta, it has most frequently been made in the manner

followed here. ..' Th . (F'g 1 '6 1117 Pan) is a mediaD unpaired strllclure whIch e penl !; 1. "1 I . , '

'culares basally with the ba5aI lobes of the gonostipe by means of a ::11, di3k-like pl3te, tho bluallobr of rho penis (BLP). The base of the '. i broad and p},o{orm, the middle porllon IS elongate and slender. pem. s . ' I

and the apex is dilated into a spoon-shape.d P'~' The I t IS strong y convc:>< vc:ntr:tlly and concave dorsall r . ~t IS part13.11y scpa~ted Into two lobes. Fr m the pyriform penis-base a parr of lrongly d l v~gmg apode",u Qf the pm;" (AP?) extend into the bases of tlle gonosopes, wh.re they

serve for the insertion of muscles.

[8+ 1

III. SYSTEMATIC CONSIDERATIONS

The systematic portion of the present paper will be limited to a COD­

sider:ttion of the haracte .. o f the super-specific groups. All characters o f proved diagnostic value that are known to the writer wi ll be given in tlle generic diagnoses. ther characters which give promise of having sy !ematie value but which need f urue .. study will be discussed also.

CHARACTERT TI S F THE VESPINAE

The subfamily . l'espillae, is " Irroup oi social, normally monogynous wasps which has been defined by Bequaert ( 13), th~ !>test reviser of the group, as foUows:

YJlCUS bro:tdly truncate. at the ~ojCLI margin, which is as a rul~. slighlly em;;J.rginat~ or ends in twO rounded. latei'd! teeth. Maodibltt. short and brOiild, foldcd Q\'e.r ~h other beneath th~ anrr:r-ior margin {the d~us, with a bro;ul, sharflly " XI,h<d, .pic:!l, <:UItinlJ .<!g.. W ial p.>lp; of 4. maxillary p.>lpi of 6 oegmm... An· h:llnae with 12 Stgml!nl.l in ftmale and worker. with 1J sqmcn in the: tmk, the ap"cal one normal Tars31 tl ... W5 impl~. Middle tibiae with two !Spical spur.!. \Ving \"emtion of the t1!uaJ diplopterou$ type': three dORd cubitaJ cdl,; radf21 ceH elongate, pointed towOlrd th~ 3.llC:X which lies c1os.e to the c:ol121 marvin i hind wings entir~ Ilnd rwTO\v at the base. without IlIl1LI or f1O!tt:rior Jobe, but 1.he preaxillary aC15icm \'cry d~p: fore wiJ1¥S tlistinctly "tailed Ion&itudinally, ~f~QIIOtt1m withotu lr:u:e3 of notauli [paraptidal fumlWS of .u<hOl'S; , .. p. ~I. Mcset>imerom comptetely :lleparaled by a ulure from the llleteplSternum; but the mer.epl.Jternuru tlC\'er iUb­div ided by a median obUqu~ su t.ure into an uppe:r and a low'cr IJlate and nevcr with a prnpectal sutu~. Hind mugin or the f)OslSCUtellum produced in the middle, ionnins a long trianguh.r lobe w"dgtd in the upper port o[ the propodeum. I'1r5\ abdamirud sqmmt sub~si.l~, not forming 3 podton; t~ tera'ite. IlS a rule. wrtb a rh,-tinct eds:e Or roundrd angle: between th~ anttrior. \tertia!, and the postt.1"ior, hOrltOctal iace. Exttn50ry muscle or the abdomen inscrtrd in a broad, oval valvula br:tw~u the apiC11 1CU1e.s of the- propodeum.

The V tSfHnD. arc c:lclined Uf il combination r characters rather tlum by Clnc r tn rl" peculia titiet which they alone: e..~i t. Thcti .. m08l u:dulive {amra appear tu be: th lIb5ence or UIC anal lobe :lnd thc deep preaxillacy excision of the hind wing.

A!! defined above, the subfamily f/cspi,ule comprises evera] super­spocific groups which ha,'e been con.ider.,.;1 by various writers as consti­tuong from one to fou r genera.. 'TIle oldest view, uniting all the species in one genus . esf'O, is nO longer adhered to by any student of the j'espidac, Ulough it is followed by many nonspecialists. In Lhe opinion of the leading modem students, among wh 111 B j llaert is the outstanding worker in this country, the treatment which most nmrly represents the truth is that which groups tbe species of V.spillac into three genera-

1851 [85

86 NORTD A)(ERICAN V£SPINE WASPS

PrlJfJupa Ashmead, V.s/>a . Dnaeus, and V .. pula Tbomson-and which ..,gregat"" lrom Vupu/Q a subgenus Dolic/,oves/wla Rohwer. The wri ter agrees in general with this grouping but believ"" that Dolicl.olJ.,s/",4J should be recognized as a dislinct genus.

Only r' rspula s. str. and Dolich()fJcspula contain species which are indigenous to North America. V.spa is represented in this couotry by tbe European V.spa crabro U nn., which has become naturalized along part of the tlantic seaboard. PrOt'Dspa is .trictly rienta l.

V'spa, V Dspolia, and Dolichw.s/",Ia agree rather elosely in most par­ticulars. T he detailed description [ the e><ternal morphology of Vcspula />~"s)'I,:a"ica Sauss. given in the preceding section of thi paper wiU apply in the main to all three. nly the points in which they differ, and which are therefore diagnostic, need be di,;cussed here.

The main event in the nomcnelatorial history of the V DSpilU'" are given in the lollowing paragraphs.

T he genus V espa, as originally described by Linnaeus ( 1758, Syst.ma N nlurae, 10th ed.) ( ). embraced all the then known species now in­cluded in the sublamily and a number which have ince been placed c1. where. Some of Ihe latter were not even "espoid Hymenoptera but belQllged in the Sphecoidea.

Henri de Saus,ure ( 119) ( 1853-1 5 ) recognized the possibility o[ subdi"jcling Vespo l>Ut did not venture to do so. ncuning the oculo­malar space, the length of which iorms the basis for Rohw r's separation of J)olid rovcsp"la from Vespula, Saussu re said :"~ ractere pem1ettrait pellt-erre d 'etabli r une coupe, il n'est cependant pas sans fTrir de trnnsition."

T he first defini le subd ivision of I' .spa was made by C. '. Thomson (1 42) in 1869. He restr icted Vespa, proper. to the large spec; in which t'he head is dilated Ix·hind the eves and the ocelli are remote from the

sterior margin o f the head. au"d de ignated JI csp,. crabro Linn. as the ty I'or the remaining ·pecies. which he treated as a subgellus of V <s/la, T homson proposed the name Ves/'l.la but diLl not designate a type.

Ashm ad (2) named ( 1902) Vesf>uilJ /U.slriaca Pamer as the type of Vcs /Juia.

R. uu Buy on (24) recogn ized ( 1903-1905) the tw groups segre­galed by Thomson. stating that they are very n"rural ones. He did not, howe\' 'r, appear to attribute as much importance to the di tinction as have other writers, fo r he aid. "En debor ' de Ia conformation de Ia tete, les caract ere. qu'il indique pour 111 distinction de c deu" grOllpe' sont purement iI1usoires ou inex.acts."

Rohwer ( 11 3A) (l916) subdivided V'SP"/4 still f Li rther. separating f rom it as a subgenus an assemblage of species cltara :terir.ed by a I ng

(86 1

- YSTF.:),IATI C CONSrOEB.ATIO!olS 87

oculo-malar space. He called thi subgell us DolichovcspuJa and designated V.spa macula/a Linn. as the type .

Boqune rl ( 13) ( 1930) reviewed the classification of the entire sub­family and presented a much more extensive exposition o f characters for separating the super-sp~ific groups than bad appeared before. I n this t rcalm nt Vespa and Ves/>ula are considered as genera, while DDlic/IOTles­/'u la iB held to be of only subgeneric rank.

BiscltotT, in " paper (193 1) whiclt has nOt been seen by the writer, proposes a revi ion nf the classification of the subfamily which, to judge i rom Bequa rt's ( 1+) review 01 Bischo ff 's classificati n. seems without adcquat justification. Bi~choff r~ognizes three g nera : .If acrov.s/>a Dalla Torre == Vespa Linn. of this paper ): Vespa Li nn. (== I'upuia

homson. minus the parasitic workerless species); and Puudwespa .: hmiedeknecht (= the parasitic speci remo\'ed from Vcs/,ula ) . His I' u P" is subdivided into V cspa proper (= V.spula of this paper minus the parasitic 'pecies ) and Dolicho<'Mpula Rohwer (= Doli<:htrJcspula o f thi s paper minus the parasitic species ) . Pseudot'cs/,a of BischofFs cia si!icalion com pri es two subgenera : PSC'll4ot'espa prope r, which in­cl udes only P. austriaca Panzer) : and PuutJOfIcsfrnia Bischoff, includ­ill~ on ly P. adullerill(J (R. dll Buysson) . Since P. ails/naco mo I no:arly ..,,,,,,bles a typical V t.rPIiIa s . . tlr ., except in lackin a worker ca te, and P. adlllt.ri,1O most nearly resembles Dolich!n'cs/",Ia, except in lacking a worker caste, it oems quite arbitrary to remove them from ti les< genera and place them to!:ethe r in a different gen u •.

J3eq uaen (14 in a synopsis ( 1931 ) of the AmcriC;l!\ Vesp;,IO. , ad­heres to the classification used in his pI' eding paper of 1930.

GENUS VESPA Ll NNAE S

TRUCTtTRA L -aARACTERs (Figs. 107, 15 167)

I Lead 'wollen behind the yes. This enlargen'ent onsistS of a length­ell ing and a d rsal and late r~ 1 hulging . . \5 a result the genae (Ge l arc "" ry broad and in the f male and worker may be seen laterad of the eOll' jXJund ey,,- when the head i viewed in cephalic nspect ( Fig . 166) , and the vertex (Fig. 167. /I,) is unusually long. Thi character i not as pronounced in some species, (or example, 1 '~Sl'a b./l icosa nuss .. as it i in most, but it is ahV'dY' unn,istnkable. fl is less marked in the male o( V. crobro Linn. Ulan in the queen and worker. Whether chis difference between the male and female is true also of other species of V <spa is not known to the author, as he has se n the male f V crabro only.

Posterior ocelli well in fronl of ti ,e postorbital line. usually at or

[ 87 1

88

n.ar the midpoint of the dl'lrsal lobes of the compound eyes. and ~ to 6 ti a. far from the occipital margin as from the ey"', Ocelh '" a rule small. exceptionally somewhat wollen. the posterior ones usually closer to each other than to the eyes F ig. 167) .

I nteranlennnry triangle (F igs. 166. 168: ial) carinate laterally. Be­quaert (14) in hi characterization of the genus fails to mention this character. thougb it is quite obvious on aU the spec,es seen by tbe author anel is di rinctl}' not so on any of the species of Vespula aDd Dolic/lOt·cs-

""ill which ase available. . .' Gena (Fig. 167) separ.ted from the postg""" hy a canna, wh,ch IS,

how."er, incomplete above, the vertex and occiput being conAuent. (.Ik­qu.,ert usea the tenn posterior orbit for the genae, and the tenn OCCiput for the postgenae.) The carina in question and the impressed line imme­diately ill front of it mark the course of the lOWe! part of the ann of the

occipital suture (aes). . . etae on head mostly rather tout, frequently bnstle-hke. The setae

on the clypeus aFe restricted in the queen and worker to a muslaci1e­like fringe aeros the lower portion. etae largely absent from the genae except along the posterior margin, especially above. In r'. "ol1ro the gen:ll "etat are absent nly in a narrow anterior strip along the compound eye. In the male of 1'. ,rabro all the setae of the head are fine and silk'Y ;l1Id the genae and c1YJ)C\IS are completely clothed with setae, as they are

in I' cS/)]I/a and Do/icho'US/'tilo. Oculo-malar pace (Figs. 166. 168 : OM) either I ng or hort, but

not in any pecies known \0 the author as long relatively as in Dolic/lO'Vcs-

I'll/a. Clypeus as wide as long, or slightly wider, .. peciall ~' in the male. Third segment of labial palri (3.lleast in Ihe female amI worker ) with

one or two strong. stiff setae on the inner side before lbe apex . Flagellar ,egments (e><ccpt U,e firs t ) of the antennae of the males, .

fa r as known to tb writer, provided with narrow, ..Jongate, raIsed en50na On the posterior su r face ( r ig. 165) . There are ~.o sensoria per segment.

II ()Ccu r io linear order :liang the antenna. This type of sensonum has been termed a tyloid {Ty by du Buysson (22) . .

Pranotum pro\-ided with a wcll-<levdoped .econdary carina (F,g. 164, nCZ), which is low and rounded dorsally but high and sharp on Ule ides. This carina follow the posterior margin of the short J oo:al sec­

t; n of the pronotum but trave.rses the lateral portions distinctly lD front ( their midpoint and runs t the pointed ape" of the lateral lobes J~'t

above the fron l coxae. A short distanL' above each end of tlus canlla there is a pit (TT fI) . arowld lhe anterior edge of which the carina arches. Debincl Lhis pit the surface 01 the pronotul11 is olten marc or less rugose.

! 88)

S\~STEld.ATrC CONSIOE.R.ATJOSS 89

Posterior Jobes or humeral calli (LT1) separated from the lateral lobes 01 the pronotum by a woill-<leveloped ,ertiary cdtina (TICl) .

Wing (Fig. 107, according te. the nomenclature of Ros : see Figs. lOS, 109 relatively narrow. Rs departing from Se + R very far from the stigma., the distance ncarly three times the length of the stigma and mice or more the length of the first section of the free part o! Rs. Cell lR1 + 2R1 shorte.r than the distance between its apex and the tip of the wing. (In the Onental gtllUS Provcsp<l, c..JII R1 + ZRl equat. the t1istanoe from it apex to the tip of the win .) Cell 1 Rs broad. broader than high, witb it. anterior vidtb about half its pos terior \\;dth. S~ond medio-<ubitai ro. s-vein (,,, - cu2) bent definitely outward, sometimes alntost angu-

lat ·. its ;lDterior end attaining cell T Rs just before the middle of its posterior margin. so that the part 0 f vein .If fanning lhe anterior margin

f cel l 2.1' is about ne and one-hall time as loog as the section of v .. in J! that forms the apical boundary of ceU 1.11. Cdl ZRs higb and narrow. ,lightly narrower posteriorly than ant.riorly. Apex of cell 1 M oblique. \ . ~i n5 At and Rs not approximating a straight line, but separating from Rs + JJ in uch a fashion as to fonn a well-marked . though obtuse, angle between them on the outer side, thi angle distinctly though only hA"htly smaller than that between the correspond ing veins in wings of

I' , s/,Itia and Do&Jrovcs/,ula Fig'. 108, 112, 113) . Stigma (PI) small ,, [ten inc.onspicll us. Hl1IIluli of hind wing bq,';nning before the poin; at which R + Sc attains the costal margin .

• II tibiae with long, erect hairs on the upper face. First abdominal segment trune"e anteriorly, the tergite mOre or less

angular between lhe vertical anterior face and the hori~ontal posterior face. As the genitalia 01 the male of but one species of V tsf>a.. V . crabro

I. im!. . have been studied by the \Vrit~r, it \\~U not be possible to tate with ert:linry the gClleric differences in he male geoitafu between V cspa and

tht other g,mero of Ves";II~. An examination of Figures 159-163 will, I""'·ever. show lhat the genitalia of I',sta crabro differ markedly from Ih«<e of any species of VespII/ll or Dolic/oo'llcspula represented in orlh

merica. T he foUowing characters in the male genitalia of V . crabro are note­

\\·nnhy: Penis 'Prll) distinctly shorter than gonostipes (CS) . comparatively

hrnad. '''.'d of "bout the same width throughout, though slightly wider I.~I It, mIddle I)(,rtion. Ending in a pair of divergent, spatulate lobes. ~"mpletely sclerati . ed abo,'e, but wilb a broad. men,branous strip below. I he '"ternal <l!vergmg apodemes of the penis (A f'P) arise nearly half­

way out [rom Its base. Gonoslipes (GS) in lateral aspect distinctly narrowed apically, squama

! 89 )

90 NOR1'H A.AI.£fUCAN VES r- INE WASPS

(Sq) short, searcdy eXce<"ding the spatulate ventral tcn ninal process

(TP2). Basal lobes (BL) of gonosapes narrOW. . V lsella (Vol) small, n t or s"","cely developed dorsad a ( the sagttta,

without a ventral tube.rcle. but with a pointed apex (Vol'), whioh projects distally beyond the sagilta.

Sagitta ( a) ql1ite small, its apex squarely tru~cate and remov~d from the apex of the gnnostipes by a ciislaDce equal to Its own length d,stad of the sagittal spine (saS) . Basal, more hea\~ly sclerotized portion of sagitta poorly developed, narrower than the termi~ 1 membr:anaus portion, and not ~hilJ itiDg along its dorsal margin a heaVI ly sclerobzod strIP .uch as IS fOlmd in some species of VespulG.

BIOLOGICAL CEl IIACTERS

The allthor is not sufficienlly familiar with the habits of any sp..'<:i of Vespa to say whether or not they are of any value in making generic

diagnoscs.

MATEIIIALS

Material of the following specie of Vespa s. sir. have been examined in the cour e of this study: V . flIlIJldarillja F. Smith., V . dl!Usla Lcpel~ V. ,il",'a Fabr., V. rrabro Linn .. V . onell/alis Linn., V . bel/ieosa Sauss. var. -<c",peri Buyss .. V. fcruida Smith., V . ",ongoljea Edm. Andre. and

V. parallela &1m. Andre.

GENUS VESPULA C. G. THO MSON (S. 1-)

ISM (= VESPULA S. STR. + DOUCHOVESPUUl ROHWER)

TII .... CTUII.AL CUAR,ACTEIIS

(Figs. 1 , 3&-11, 98-106, 108-1 13, 11 5, 116, 127, 128, 132-158)

H ead of moderate size. 11 I ,wollen behind the eyes; the genae (Gt) of normal width and the vert.". short (Figs . 1-6, 100,157,1 58).

Posterior ocelli dose to or tangent Ivith the postorbital li ne and at most as far from the occipital margin as from tile eyes (usually closer to ~.ach other than to the eyes) .

[ntex-ulLlennary triangle (iaJ) not carinate laterally, though usually well ddinecl by rounded margins.

Carin.' and groove marking course 01 the occipital suture (ocs) ?etwcen frenae and post genae varying in e.'<tent 01 development, absent 10 some

species. [90 I

S't"STE:l1ATIC CONSIDER.ATlON 91

Head co\'ered almo t entirely with long, soft, hajr-like setae. ?\o bristle-like sel;)e present.

cule·malar spact! (OM) Inng or short, in ome species practically a sent.

r'roportions of dypeu, variable, generally , wide as long; distinctly wider than leng in some species, slightly longer than wide in a few s 'ies of Dolje/tOfJupuJo.

Third segtnell! of labial palpi generally without, but occasionally with , a 'ubtcrminaJ seta. Bequaert ( 14) calls attention to this character in the following manner: "Third segmenl of labial palpi without :I strOllg, stiff ><Ia. sometinles witb " small hair: ' From thi statement it would apJ'>C<,r that there is an important distinction in character 10 be made between th ' ".trong, stiff setae" found n ('he labial paJpi of the species of the genus Vespa and the "small hairs" found 0 11 the labial l)alpi of spedes of T' csp"/a. far as the writer mn see there is no striking difference uetll c-en the two. The species of Vespa are large and robust and their pal pal setae are correspondingly strong. The species of Vespula are much .maller and lighter in build and their paIpa1 setae are correspondingly slight. The only distinction of importance which can be made on the basis o i the oharacter of the paI",,1 setae. so far as the writer can see, i that their development is cansta.nt in V.spa and is variable in Vupula .

Flagellar segments o f the male antennae (Figs. 103. 128) with moder­ately developed tyloids in some speci (Dolieltovespula), in other speci (VupII/a s. sIr.) entirely dtvoid of tyloid .

Trnnsverse carina on the pronot llm omplete (Fig. 156), indicated in th · lo wer ponions on the sides only. or entirely absent (Figs, 38, .JO....42) . Pron tal pit (TJ p) near lower end of this carina variable in development. sOlllct i:nes much r duced. Surface of pranotum behind late ral pit some­time, striale or rugose. often unsculptured. Posterior lobes of pronotum. or humer,,1 calli, not separated from r ' t of proootum by a carina.

Wing. (Figs. lOS. 109. 112. 113) reL~ti vely broader than in Vespa. Rs (nomend ature of Ross ,separating from Sc + R closer to the stigma than in r·upa. the distnnc. being less than twice the length of tI, .. sti a and auout equal to the free section of R.' h -t.wecl1 C + R and loI . Cell ll(] + 2Rl 'horter than rhe di. tance between its apex and the lip f the wing. Cell IRs broad, but not Mticeahly hl'O(\der than high, its anrerio~ width half or Ie limn half its posterior width. Second medio-cubital er.. vein (m - m2) sligbtly ""rved outward, but not bent, attaining c"!llRs either at (Ve"ptda s. str.) or we.1I before DolichovuplI!a) the mddl of .t5 posterior margin . 'ell 2Rs high and narrow, either of same wid,h tht'oughollt Or distinctly narrowe.d posteriorly. lJeX of cen 1M eilher square.ly truncate or very slightly oblique. eins M and Rs at

[ 91 1

92 • 'OIlTli AYER1CA:i v£.SPINE. WASPS

their point of divergenCl: making only 3 slight aug.le on ~he .outer .ide, at least B definitelv smaller angle than the corr,,"pondmg vems In V upa. an~ tbus approxin";ting a .lraight line. stigma (PI~ well developed. Ham.wI of hind ' iog (Fig. 87. HI beginning at tht. POint wbere R + Sc attams

lhe co,tal margin. . I First abdonlinal .egment truncate anteriorly. tbe terg!te more or :

angular between the vertical anterior face and the honzontal 1>0 tenor

fac"-ine male genitalia of the speci.,; ind uded in the genus Vesp.J<J s.l. ariable' but the variations are of ouch a cbaracter as to

ore very v , V J I d Do permit the easy segregation of the pecies into Up" a s. s r. an -Ii Irov.s~ula Rohwer. Tbe genitalia of Do/~chtr~DSI"'la resembl.e those of the mal!S 01 Ves"a. a smning the gt.tlltallll of V. crabro Linn. to be t 'caI more than thev do those of Vtsl'lllD s. sir. The d13racters of tlle YPII . ·talia thercf~re constitute one of the I110St cogent reasons for

ITh"li! g<:.m I ' . . '"

recogniring Dolic/rov8sfH'/0 as a valid genus Instead of Lreat1.tlg It ,IS a subgenus under I' •• /lula.. In "iew of these facts til: characters of dIe male genjtaJi,,~ of V.sp,lIa s.l. will be given under the dISCUSSIon o{ V.spu ta .<. sIr. and DoliclJovcsptda Rohwer, rather than here.

B<OLOGICAL CIlAIlACTERS

AccDrding to Ihe location nf the nest and the details of its . onstru~on the species may be separated into two groups . the limits.of which COinCIde , .u. ' th those of the g1'OUPS V ul'ula . ,tr. and Dolrcl.oc'tsprda. as de­""Be. ) WI f Ii th b'olog> termined by morphology. It will be hetter, ther< are, to (.cuss e I -

cal characters o( th e groups separately.

GENU ' VESPU VI (S S. , .)

STRU TU\t.AL CIlARACT£ll5

(Figs. 1-6.38--42,9&-106, 108-113. 115, 116, 127, 133-149)

Head (Figs. 1-6. 100) onl)' a trifle narrower than. the tho,:"",, , . . lntc:rantennary are" (il1l) di tillctly mangular, Its margms ilivergmg

trongly dors.~d, broad and distinctly flattened . _ . Oculo-malar space (OM) short, at most hall the length of the penw"­

mate antennal :.egrnenl (in female and worker), olten absent. the eye then

touching the base of the mandible. _ . . ypeus (ell') never long.r than wide, and often distlllctly WIder than

long ,,"pecially in the males (fig. 100) . • ccipital suture (ocs) generally moderately well developed. at least 011

192 1

. YSTe.acATTC COSS1DRRA'TlONS 93

lhe sid .. of Ibe bead. often omplete bela, and vctending to the bas" of UI. mandible.

F1agella r .egments of mule antennae (Fig. \03) without tyloids io aJ.I cases, so far as known.

~routhparts of the foml shown in the figures illustrating the mor­phology of 1'. p.nry/~Vl1Jjco (Figs. 1,2. 8, 10-12 . 14--37), moderately short and stout, not especially lengthened. Cutting portion of distal margin of mandible wbich lie b hind the apical teeth considerably less than tw ice as long as the portion which bear. the teeth (Figs. 33. 34, 36).

Secondary pronatal carina generally enti rely absent, faintly marked at the lower ends in some species. Lateral pronatal pit.. TIP) .nlall. in o.ne ,pecies ve.ry much reduced. Surface of pronotum behind m..e pit.. smooth.

clllRs o( fore wing (Figs. 108, 112) broad posteriorly and narrow d anteriorly. its anterior width UJiuaUy about hal f its posterior width, in sante ')l<.'Ci· · (F ig. 112) slightly 1= than half the po teriot width. Second llIedi<H:l1 bital cross-vein (m-eu2) attaining the celllRs at the midpoint of it , poste.ri or margin, so that the part of the vein .~[ fomling tbe anterior 1II;lfgin of cell 2M is appro:omatdy twice as long a. the section of vein M that fom ls Ihe apical boundarv of coli /.It. Cell 2Rs high and narrow, in Ill,,,t pecie about the same width thr lIghout ( r ig. lOS), but in some peei " distinctly broader ante ri rly (Fig. (12). The latter also have the

cdl JR., less than half as wide ,mleriorly as it is posteriorly. T ibiae as a r ul e without long hairs on the upper {ace. (They are pres­

ent in ". u.oslriaea Panzer, tlle type of the genus.) 'rhe following are the salient characters in the male genitalia (Figs. 133-

149 J .

P nis (P,,,) distinctly longer than the gOllostip.,; (GS) . either mod­u atcly broad or v"ry slender. but always wider basally than in tile miudle ('<m ion and ,ometimes ,'ery much so . Ending in an expanded . spoon-like I' i~cc. which mayor may not show a slight subdiruinn into two lopes. C mpletely sclerolized above. Membranous strip below broad ill the species wi th broad penis, extremely ,I."lrrow in species with lender penis. T ntemal <Iiwrg ing apodemes of the penis (ApP) arising from the broadened basal part.

. (;OIlOStipes (GS in la teral aspect rather blong, only slightly narrowed "'I"a1ly. Squama (Sq) pointed or blunt and moderately short to long. but 11\ all species oi which males ban been seen by rbe writer it project.. be­),nllo. in some cases ver}' much beyond. the tip o{ tI.e ventral t rminal pruccs (TP2) . Basal lobes (BL) of gonostipes moderately narrow to rathu wide.

\',;I5Olla (1'01) ,ariable in .i"e but proportionally larger than io Vrsta. In species with slender penis the volsella i developed dorsad of tbe sagitta.

[ 931

.sORTH A)t ERICAN V, Pl!'tE WASPS

&> thnl it ocoupies nearly the whole width of the inn r face o[ lhe gono­

stipes (Figs. 143, 144, 145) . Sagitta (Sa also variable, but its basal, more heavily derolized por-

tion and the t"nninal, weakly ""Ierotized portion always sharply differen­tiated : tbe fornler distinctly broader than the latter and olten produced inlo an espl'cially heaV11y sdcrotized lobe (DLV) along its dorsal margin. The apical portion variable, finger-like or lapl' ri ug, not attaining the apex

of the gono tipes, or much exceeding it.

BIOLOC ICAL C HARACTERS

'esls nearly always os 'entially subte.rt":lllean : begun usually in a rodent burrow, which is enlarged by the wasp as the ne t grows (Figs. 197- 202, 208-211, 226. 228-230 . ombs, "xcept fo r a small conical a rea around th main suspl'o.sorillm, olten a r~ essentially plane (Figs. 227. 246, 2- 5) . Each comb uSP"fid I rom the preceding ( t he first f rom the n t wall ) by a large number of small , cord-like suspensoria (F igs. 229 , 246), T he nest wall compoSMl of numerous, relatively small "shells" or scales 01

paper, in which the minute arcuate strips constructed from single loads of paper-pulp are concave on the downward ide, and in the last-made por­tions of the scales often completely circular So as to produce an appearance like that of the growth rings about the umbo of a molluSCMI !heU (Figs. 226, 236, 237). Nest wall ultimatel ' not added to above and on upper part of sides but only extended below. ppe.r part of nest waU thu ulti­matel)' constant in thickness and not ,'c r. thick. Inner su r face of upper part of nest wall often plastered over with paper pulp to produce a reln­li,'ely smooth surince. Doo rway through ne. t wall in ld n 'Is well up on . ide of wall or even on top of Ihe nest (Figs. 228-230 . As the tlest is enlarged , additiona l cord-like suspl'osorin are built between the nest " all and the roo f of the nest chamber ([:ig. 229).

Colonies enduring generally until late in the [all, often not breaking up

until frosts come. 'i oung males and females u nally not produc d earlier than September,

and often. in the wannu sections of the country, 110t unti l much later.

MATE IALS

The following species of the genus V upula bave been studied by til. writer: V . ol<slrioCIl ( ramer) , v. sqllamoso ( Drury), V . su.lpJlllrea (H. de

aussure) . V . I'll/a (Linnaeu.) , V . ' lifo var. (OMob,illa (R. de Saus ure) , V . , ufo var. vid,Uj (H. de Saussure), V . Tu.fa \'ar , alropi/osa (Slnden), V. g.-nll,mica (F abricius), '0. ",acu/i/rons ( R. du Buysson), V . p<lIs),/­val/ieo (H. de Saussure), and 1'- vlIlgons (Lirmaeus) ,

19011

SY$TENATr C CO NSIDE.R.ATJONS

GENUS DOUCHOIlESPULA ROHWER t916

SnuCT RAL IMRA=

(Figs. 113, 128, 132, 150-15 )

95

H.ead (Figs. 157, 158) di stinctly narrower than the thorax, in some specjes considerably so.

lnterantenna ry arell (iol) not distinctly triangular, and not definitely flattened. but havmg ,he [arm of a broad low rounded'dg h' h . , . , . n e, W lC 1$

sam 'what WId r above than below. Oculo-mala r space (OM) long. nearly as long as the penultimate anten­

nal ' cgment or longer. ClypeU5 (Clp) usually about as wide a it i long, sometimes a little

longer than it is wide. . Occipi.!,,1 sutur ~ocs) usually reduced; well developed only on the

mIddle third o f the Ide of the bead al,d obsolete bel .. . 1.1 . I b I I 0\',. ! occaSJona \' en-tift y 0 so ete. .

F lag-cUar segments of male antennae (Figs. 128, 132) with well-de­\'eloped tylolds ( T.v) In a f~ species; in other speci .. the t),loids are small a nd poorl)' developed: III sl111 olher pl'cies they are entirely absent

:II uthpart d i tinctly elongated and slcruler as compared with those of V. pc>!s.l'lv"I1lc4 . Cutting portion f eli,tnl edge of mandible. which lies Lchl~d lhe three teeth c1.early twice as long as the portion bearing the teeth.

~econdary canna ( T1 C2) o r pronolul11 well devciopl'd on sides and u, ually d "clooped on dors ulll ; occasionally obsolete on dorsum. Laterdl pronotal 1m ( Tlp ) wel l d velojX-d . Sudace of pronotum beh' d th '

h. . m e prt

511100\ . ·trlale, or s~ghtl :, rugose. Cell IRs of the fore wing (fig. 113) a liltl broader tha h

O

h b d 1 '1 I ' d . .. D Ig , raa tK,lIJl( at; , qUI te I~arro~ antenorly, It · anterior width not more than one-.~lrd of Its posten Or WIdth . etond medio-cubital eross-vein attaining the , ost nor .margll1 of cell I Rs one-third of the I\'ay from it. proxilnal end. thus makmg tl~ ection of vein M that fornls the ant' rior bounda ry of cell 2M "pprox1DlateJy equal to the section o f M which forms the apical n~arl;lll of Cell ,7M. Cell 2Rs high .. narrowed posteriorly, broad anterio rly. H !1 ~. t.en~r Width no over two-thirds it · anterior vlidth.

I J ,b,ae 10 all ca ' . with long, ereCt ha irs on the uppI' r face, especiallv on

t Ie lund legs. 0

( ... Peni (1'",,) shorter than the gonostipcs (GS) , in dorsal aspect mod­l' ,Jlcly na~row basa1~y , broadening gradually a its midpoint and then

~,,;~'Ptly WIdened 10 I~ greatest width; mpcriog from midpoint to apex I ere .' t ends In a pa1T of clongale. pointed, inctlrved lobes. Conlpletely

:: erohzed .~ove except for a .hort median strip eXlending basad I rom be­en lhe apIcal lobes. Entirely membranous below [or one-third 10 two-

[ 951

96 ~ORTJI :\ lL.EJUCAN V£.sPINe w..J\.srs

third. of its length (rom th~ base; <;C!uotized laterally and ~IIIbr:mou, mesall), on thr apical portion. Internal divuging "po<ienles (A/'P) ari ing from the middle portion. whete the p~n is is widesl.

Gonostipes (GS) in lateral aspect oval.- dbtinctly oarrowffi apic:o.lJy. Squama (Sq) pointffi and short. but always projecting a little beyond the apex of the ventral lenninal process (TP2). &sal lobes (8L) of gono­stipes narrow to mod~rately wide.

Volsella (/ ' 0/) .mall . not or sc:o. rccly developed dorsad of the b:o>e of the sagitta ; dorsal lobe f the volsella .illlall and hidden behind base of sagitta : vo1scl1a without ventral tubercle.

Sagitta (S .. ) moderately small. it,,; apex broad and usually ohliqurly truncate. about equaling the tip of the gono,tipes but not exceeding it. Basal. more heavily clerotized portion {each sagitta poorly developed. narrower than the (emiMI membranous portion and not developed along the dorsal margin into a heav il. sclerotized strip such as is found in some species of V.spu/a.

BIOLOGICAL CHARACTERS

l\; CS . (Figs. 203. 205--207, 215--222) . always essenti"lIy aerial, though occ.:I. .• ionalJy built so close to tile suriaee of th ground Wt the nest soon touches the ground and spreads out 011 it, developing a Rat under surface ( F igs. 221, 222). Combs (Fig. 216) . except for a small conical area im­me li:uely around the central suspensorium. uSllillly tumffi up at the mar­,;n5 :lS to be concave above and convex belo" . Eacl. comb suspendffi from the preceding uy a rew suspensoria and these! in the (om> or thin walls or buttres.<e:I (Fig. 2~O). The nest wall composed usually of a rela­ti \ ely small number of quite larg layers of paper. which are at first ar­r.wged as a serie of concentric spheres or oval globes. Inter more reduoM in size and overbpping somewhat. [n nests of Dolic/latcsp"la ",anI/ala the rooi of the nest is oftell made of l1umerOIlS overlappim; mall ,heets of paper so that it has n cellular appearance (Figs. 219. 247). The minute strips of paper constructed r rom single loods of paper pulp nearly Straight and running horizontally around the nest (Figs. 215, 21 7, 219, 238). New layers added to upper p:,n of nest as well as on sides and bottom . though in old nests om.what (ewer are addffi above than below. The new layer. addffi to the upper part of the nest .oon corne to intersect the support f rOIll which the nesl is suspendffi. They are then attachffi 10 this support or started from it. thu providing additional susprosion for the DCSL Sec­ondllr)' suspensoria or the usual type are not built for the ne.t as a whole between Ihe De t wall and its support. Inner surface o( upper l>art of nest wall eommonlv trimmed. all r.1l.ogcd edges left by remo\'lI1 of inner layer, fastenffi dow~ and all chinks fillffi . hut interior not smoolbly pla.sterffi

[961

"\'ST EMATIC CONSIOE~ATION' 97

o"er with paper pulp as is don~ by SOme specie.; of VOlpu/a. Doorway through nest wall ventrdl in youllg nem; often btrral in old neslll but never high on lhe nest.

. C~lonic;s genemlly o[ relatively.short durat i n, diSSOlution often begin­mng ID lDld-June. and many eoloDles completely di pe~d by the end 01

July. SO.me colorues elld~re until autumn, but probably few if any persist on an acbve oDdlbon uotd destroyed by the advent of winter torms, as is true of many colonies of V"pllia.

MATERIALS

The fo!lo wing species of the subgenus Dolichovu/,uIa bave been stud ied by the w~ter: D. adullen,,:, var. arctic", Rohwer (=V. borcafi.r [Uwis not of Kir~y l , D. arenanD (Fabriciusj (= V. diabo/iCIJ [H. de Saus-ure]) . D. "",clliat .. (Linnaeus) D. ""dia (Retzius) d D .

(F b .. ) , , an . tlDl'W4f11ca a n clUs .

co CLt.:srON

, F~om the elnta presenu.d ill the foregoing pages. it is ob"ious that U,e 1 eSf,noe, though constituting a compact and relatively hoonogeneou sub­farndy, are clearly divisible into well-mark .. l genera which differ not only 111 a~u lt morphology but also in such biological cl1.lracters as choice of nestrng Sll<'~ character of nest constructed, nnd dltration el f life of the col­Ony. Addmo:w study will uodoubtffily brillg to light many other differ-nces Of . \'ary~og degree which will contribute to a bettet understanding of

Ihe rel.allon hIp of the genera and oi their constituent species. It IS rcason~le to expect that some of the wfferen es yet to be disco _

ercd ar~ to ~ ~ound in the immature sblges of the wasps. The writer has flot ma~e a cnocal compar:llivc study o( allY immatnre stages. but a cursory

mpanson of Y,e la rvae of V.sfml .. PClIsylwllica ( auss.) and Do/icho-" .. pula MenanD (Fabr. ) shows these two species to be di tinct in la rval head charncter. as they are in adult head character .

[97)

IV. BIOLOGY OF THE VES PINAE

SEASO,lAL HI TORY OF A W SP COLONY

\ ith the exception of the socially pa=itic 5peci.~ all normal wasp colonies develop in the same gene ral way, . hibittbe same genc~1 charne· teri.tics, and undergo the same sequence of developmental changes. Each olooy i founded in the spring by a single queen, which has passed the

winter in hibernatioo a ltcr having malc-ci tfte preceding summer or fall . The queen selects the. nest !i e, Collects vegetable fiber with \" hieh to make pa.per, and constructs the first few cells ef the nest. he also constructs from one to thr~ paper eIl,elopes to en 'Iose lhe cdls (Figs. 197-203, 205-207, 213, 214) . The lest in this early Illge, . ,nee it is entirely the work of the queen, is often cal led a "queen nest ."' The. queeu lays an egg in each cell, then cares for the larvae which soon batch from t.he eggs, Ceeding them da ily with freshly killed insects. All these larva and most o f tbe uccecding broods until midseason or later mature into workers. These assume all the dutic f the colony except egg-laying. The queen rarely leaves the ne t aiter !.he first workers mature and e,'entually not at all. The workers stendi l)" Qdd n w cells to !.he comhs and new layers of pap«r to tbe nest wall. The inner layers ate lorn away to make room for comh cnlargt:ment as new layers are added to the outside wall : The rate of en­largement accelerates with the maturing 01 addition' I workers. SO that within a few week or mouths the colony becomes quite populous and the nest becomes th romId or oval structure that i' 0 well kno vlL '!'owwd the dose of the sea on cells f a larger me are buil t, and ;n these are reared new males and queens to a number varying from a few do:r:en to several huudred . The queen mother, £oUllder 01 the colony, now quite worn ou t, dits. The new queens wei males lea,'e the nest and mate. The neW' q U<r05 then enter hibernation qUMter· . each selecting her own hiber­nacuJum and occupying it alone. The males feed at Howers for a few days or weeks and then die. The. workers remaining i ll the n t die a iew at a time and the nest population steadily dwindles . During the Ia t few week> th . n::maining wo rk r9 develop erratic habits, oftel1 becoming cannibalistic and feeding on the remaining larvae and pupae. Finally the last worker dies, the nest struclure i left to the merc,- of wind and storms, nnd once more the species is represented only by hibernating queens.

HIBERl\ATJON

As a general rule only queen wasps hibernate but there is some evi­dtltce to indicate that ill rare instances a few workers al.o hibernate. Fol-

98) [ 98 I

gfOJ..QC\' OF THE VESPINAE

som (6.3) sugge.ts that not only workers but also males may hibernate and that such hi bernation occurs in the nest; but he doc not present !.he data on which his suggestion i based . There is evidence, on the olher hand, that workers sometime hibernate av.",y from the nest in the general man­ncr in wluch queens usually bi\>tmatc. F or example. dead workers (lJ'e oc­casi nail to be fou nd beneath loose bark on trees, a it often c.hosen bv queen \v;"ps (or hiocnllltion. Some of the dead workers are doublless i;­dh'idual which have been caught and killed by other predatory insects or I ,,· opiders. Some. probably, are ind ividuals which, enf~bled wi!.h age, have .t",yed into the paces heneath the bark and have died there. A few, how­ever, are apparently aewally hil.!ernatiog. On December 17, 1922, willie collucting hibernating queens of V",pllia pC/1.Sy/wni<"a from beneath loose eucalyptus bark -on tile Stanford University campus, U,e writer found One worker in n fresh and apparently normal condition. UnfortunateI it was se 'erely I, inched il"l pulling oII the piece of bark beneath which it was foulld . I \\ ru; not possible, therefore, to revive it by warming :U1d to determine PQ,itivcly if it bad been alive. On January 18, 1931, while dnving ;n the hill s 'vest a( Stanford University. the write r took a live worker 01 Vcsf",/a t " lIt.\'h ,iea wh icb fl ew into nis cur. As the mooth of December 1930 had illdud d the longest unbroken cold spell n record to that dale for the di -t,.i c!-fen consecuti ve days {freezing weather-and as the therrnOlnete.r had registered below- freezing temperatures severn] times during the firs t h"f[ <If January 1931, it is certain that the worker in question had bccn dor­m ant. if not in a definite hibemacu lum.

\ \'hetber or not male "'. p hili.rnate is more unc<!rtain , bot at Stan­(unl l 'ni,'C ity 00 a warm afternoon of December 22, 1929, a oumber or mal o f l' cs/, .. /a prllsy/va/l;ca were taken in Bight o,'cr a seepage area and on vel:ctatio l1 nt-.tr by, It is qui te po sible, howe"er, that these were .imply male" which had matured very late in the fall aJld had not yet died or been killc by cold. T ney may not have been in hibernalion at all. M"le wasps an' o~casiollruly taken during the spring mon hs in regions of mild dim.,t • . \ 11 tl'ost ",hidl have come to the writer's att .. ntion, however, have been

"mng adults which m"tlIre.d in the spring ill which they were taken. The)" have all been spring produc of vigorous colonies which successfully Q"er­wjltte red and c(lntilluecJ into a second season (see pp. 141 :u!d 159 ). and n,,1 individu;l.1 which had hibernated. OverwinterinJ( wasp colonies doubt­Ie,; du occasionally contain males that pass the cnld season in a dor­nmnl condition along with workers and queens : but positive evidence that the ' do rtlTUlin< yet to be secured.

Conlrnry to popular belief ~nd to most I)ublishcd smtem<nts on the >uhjcct the dormancy of queen vespine WllSps is probably IIOt typical hiber· nation; ior cold weather does not appear to be necessary to induce hiLerna-

1991

100 NOIlTn Ar.tERI . V Es-p r X'E WASPS

rioo, oor dO<.!i the period of dormancy n cessanly incid. with WiOleT. It does. howe er, include the "vinter seasOIl. ue~ dormancy seems to bt. brought on by a particulnr phyoiological Slnt~ ~.thout regard to w~~ther conditions. In thi respect it appears t be s.m.lar to tbe so-alled oblI­gate domlancy" reported by Roadio ( Ill ) for ~edl'~ius ~.T~,,"aJUS. The ,ucens apparently enter hibernacul3 and beeom~ macbve wlthm a few days '~ flcr lea"ing tlle parell t colony and mating. Th~ o~ly publ~sbed .tat~enl to thi d£ect. however, which has come to the wnter s attentton tS conlnmed in I: , i\1arti " Duncan's 011' i nsect Fritndsand Foa (51). In certain ca~, t t o f Doije"I1IJupllta arennria in central California, for example, young queenS may begin hibernation as .arty as the first week of July . A hibernat­ing queen was taken by T om Rodger~ i? midl;ummer in Slnte Redwood rark. Big Basin, Santa rut. County. CalifornIa.

Indirect evidence to the effect that queens enter hibernacula before being forced to do so by cold weather is fOllOd in the fact that, disregarding queens taken in hibernation and young 5peC.OlOIS captured on the parental nest,; before leaving tbem for mating, most speolnens of queen wasp found in insect collections prove to have been taken in the pring, not.in the lall. D uring the fal l of 1929 the author kept a carc!ul watch for flym queen. Only a hal f dozen all togeth r were seen ; but large number of mal ,'ere tak~n . e"en thougb no special effort was made to secure them.. Yer ord~ ­na rily new males and queens are produced by was~ colonoes 10 approJ(]­m.ately equal numbers. Coloni e which have 10M tbelr queen mothers or to

which queen mother. ha\'e beeome tOO nfeehled to funcuon do, howe"er, produce an excess of males from infertile eggs lai~ by. workers, but ~s exce S appears far too s",al l to account for rhe dlspanty between R)'mg male. and '1u ns du rin the fall, It seems probable. lherefon:, tha.t early hibernation is th rule among the Vespinae; but much morc e:<leDSIVe ob-

sen'ations are needed to clear up the matter entirely. . With the advent of old w.ath"r the "obligate domtancy" ot the queen

wasp: w ldoubtedly beeOlnes a tYPical hibernntion. During such cold weather dormant queen s are quite torpid. T hey may be roughly handled and put into strong alcohol or killed by cyanide . witho~t manifesting the 'Iigbtest reaction. n the. other hand. tbey readily re~"'e on wanll days and mo 'e about or threaten t sting if uncovered and dISturbed.

In the sdection of hibernacula, the only important consideration.s seem to be eonce.,Iment and a certain amoull t of shelter. t\ great vanety of place ' a rc chosen, but there is 11 0 evidence Illat the queen ' abibit the striking intelligence th..,t. at times, has been altnbuted to them. They not inf requentl ' choose hibernacula that re entirely inadequate to protect th",:' from the rigors of a 'evere winter or from the abruptly changmg condI­tions of even an ordinary winter. The author took one queen wasp but ty,·o

( 100 I

BIOLOGY P' T U B V£S PI NAE 10

ioebe. {rom lhe outer surface of " large rock. in a crade so nearly vertical tltar it could scarcely ltave kept out the raill even during mild showers. Some of the hibemacula are quite dry. Othen, such as decaying logs per­meated by {UIl!.'U. hyphae, ""'y be quite wet. Wasps choo ing hiberrulcula of lbe latter type are sometimes ki lled by the fungi present in the wood. The thiclcness of the bask which covers hibernating wasps may be as little as one-eighth of an inch or as much as two inches, or tlle wasp may be

.ered with as much as ten inches o f wood and ha rk. Protection from dew and frost is not sential, in the rngion around Stanford University at lea t; ( r the :luthor has taken hibernating queens there which were covered with fr t, yet they revived upon being ",anned and appeared to be no"nn' in every way. Tn some species (Vu~ula "mrs/vanka) certain individuals hi­hernnte in cracks in rocks in open fields where they are exposed to larKc tIaily 8 uctuations in temperature. Other individuals of the same species eboose reiati,ely uniformly cold . damp ' Iuaners in shaded galUes.

ctnal hibernacula recorded for various American species of espine w ps :lrc as follows : Queens of Ves"ula !,CllSst, Qlljca have been taken under loose bark on eucalyptus. willow, cypress, alder, and various oaks.

bviously there is no preference for particular kinds or types oi trees. Til y have been taken between baled shingles, in leai-and-twig debris on the ground, in the spaces between stides of stacked cordwood, between rhe iolds 0 r old clothing and sacks hanging in a shed, between stored book , in cracks in a pille log. deep in beetle burrows and ther holes in rotting logs of c1iffe rent sor ,and in erades in boulde.rs exposed upon a hilltop. The author has taken V UPUt4 voJguris beneath the bark of a decaying log of the Blue Oak, Qtteretls douglasii, and DolichO'lJts/'Ula ""u;u/ala deep in a log ill the last stages of decay. Phil Rau (107) records finding two hiber­nat illg female of D .• naCluaJa three feet apart under a log. Specimens of Do/;ehO'lJ.spula. a'N/aria, from burrow' in a rotten log. Were sent to the amhor by L. G. Gentner of the Michigan Agricultural College. Gentner also sent a specimen f VespoJo ",orollif,olls Inken while sifting dead leaves. Z. P. Metcalf, West Raleigh, 'orth Catolinn, sent a queen VespllltJ, sqUll­tI,osa which was taken "in earth."

DUring the period o i dormancy th wasps assume a very characteristic pose (Fig. 2-4). The anl""""" afC held against the head. witb the scapes extending along th lower edge of the ernargillation of the compound eye and with the OageIJa bent downward along the occipital sunorcs. The distal end of the llagella arc curled mesad behind the mandibles. The abdomen i <:u rved downward : the segments are retracted slightly and, except on "'ann days, are held practically motionless. On waml day obvious respira­tory movements may occur. The legs are fl exed closely again t the sides, the hind tarsi being held against !lIe undersurface of the tip of the abdo-

[ lOl J

102 NOa:ru AMERICAN VESPfNE WASPS

m~n; tb. middle tarsi are extend<:d backward under the sides of the ahd mm nt it! base. and l h. front tarsi are doubl<:d oockward along the side. of the leg bases. The wings are folded. carri<:d backward between the hind femo ra and the I>ropodeum . and tuck.d beneath the sides of the abdomen. with the co tal margin of the wings dir<:cted outward and the wing lip overlying the hind tarsi. In addition some hibernating queens tightly grip a small pledget of wood or bark with their mandibles. I f pulled loose the pledgets of wood or bark will usually give way without the wasp opetling its mandibles. Other indi';duals eilher ding but lightly by the tarsi alone or lie quite unattached . sapponed simply by the W'~IJ of their hiber­naculum. Figuier (61) refers briefly to the hibernation attitude, comparing th appearance of the bibematiag wasp to chrysalids. ,\nd Westwood ( 157 records having observed a wasp " in a frosty morning in October. hanging uspended by its jaws to a curtain . with its wing folded up be­tween its legs. and upon i . brca..~t, " but he does not state whether or not it \ 5 a espine wasp. Memb= of certain otber genera (Po/isles and Jfisritocyllllnts) assume the ame attitude, not only during hibernation but " bo during ordinary repose at night on their nests.

The length 01 tbe dormant period diff~ with the species. the locality, autl 'lCasonai conditions. 1t may be as short as two mOnths or as long as eight. A t low ele'~dtions in California. and doubtless in other regions of

ild winters, th domUUlty of the \ 'p: lMy be interrupted by warm days ill \\' inte ... Onuch day ' hibernating wasp may hift their positions or e ytll , ally Corth on the wing. After thu. venturing fortIl, the wasps appar­t'nlh' u tilize any convenient uew shelter in which to continue their hihema­t ion~ T here i n e,'idence to show that th'y ev 'n attempt to return to the original hibernacula aUer their sallies. They ha\'e been observ<:d to 8) a waY without manifesting any particular concern ior the location of the shelter which they have jllst felt. H.ibern.~ting queen. on severn.! occa.sions I VB been found in places where inspection had shown there were none a weck previous. With the merging [winter and ~pring, warm days and ex­plorntory flights by queens become more Crequent, and hibernation gradu­ally breaks up. The queens feed at earl.. pring flowers, and ~s soon as sui table conditions are sufficiently continuous and food supplies are de­pendable each seeks O\lt a ne ling s;te and founds a n w colony.

The following data c ncerning spedmens takm in hibernation pertain to specimens in the author's collection (except where then";se noted the collections were mad , by himseli). V.spula ,noell/'frons: Agricultural Col­lege, Michignn, December 19, 1923; collected by L. ,. Gentner. V~spu/a squaillosa : West Raldgh. ~orth arolina, F"bruary I, 1918; collected by Z. P. Metcal f. Ve.r/,,,/n p .. o.ry/vollim: Stanford University . California. lXcember 28.1 920; December 17. 1922: December 2-1. January 14, Janu-

t 102 'i

Ht OLOC:V 1-' "tUE Vr..5PISAE 103

ary 25, January 28, and February 1 . 1923; December 28. 1924 ; early No­vember. ~ber -, and D~cember 14, 1926 ; November 11 and November 1- . 1927; November 20 and December 28, 1928; November 16 and 17, and February 2, 1929; January 1, January 13. and Febru.1rY 21, 1930. At ' ,. no. California. December 26. 1922. Dolich01Jtsfrl//a or6,UJria; Agricul-

tural College, Michigan. pril 20. 1923 ; collected by L. G. Gentner. Doli­d'Oflesp"/o 'flDeli/olo; Washington . D. C. . April S. 1919 .

FOODS AND F EEDIKG nEll V[OR

Tn any discussion of wasp foods or interpretation of ieeding- bebavior tll'O jacts must be kept io mind. F irst. fo raging workers collect the food supply for all the wasps remaining a t borne in the obt, for the queen after 51 , becomes confined to the nes t, fo r all larvae except th se of the first Lrood whose food i5 collea<:d by the queen. for ther workers which rna be engaged in caring for brood or in other duties about the nest, and for )' ung males and queens which have not yel left the parental home. Sec­ondly, for anatomical and perhaps otber reasons ad ult wasps (Pi>. 32 and 33) eat only liquid s ubstances, thougb wasp la.rvru! read ily ingest solid food . '\ ny lid foodstuff, therefore. wbich a wasp m~~' be observed to coll= art destined to be carried to the neM and there fed to lan·ae. Lack of ac­quaintance with these facts h35 led m.·my persons to draw erroneQus con­clusions concerning wasp foods and feeding behav ior.

FOOD ? IATElHALS

) f diff~rences in larval and adult habits be disregnrded, wasps may be de>CIibed 35 practically om nivorous, for the foodstuffs whidl they on urne ar~ l1um~US in kinds and diverse in character. Howe\'er, \Va ps do not, except aCCIdentally, I cellulose in any form. :-rost of their foods fall into two well-<leli lled classes-liquid foods high in sugar content ; and meats. 1<')Od5 of the fonner class are primarily encrgy-}~elding food and are eaten ?,ainI by adult wasp. 1l10se of the latter class are primarily Ii ue-build­mg. foods and, except for juices whicb m3)' be sucked from them, are eaten en ti rely by lar at wasps. The general character of the foods consumed. therefore, is correlat<:d with the physiological Deeds of the larvae and adults. respectively, which consume them.

Energy-yielding food of high sugnr content are obtained chiefly [rom plonra, tile most imPOrtant source being tl,e nectar of 1I0w rs. Honey also I< eaten when~ver available. An old book, NaJ",~ Dispwy'd or L. S rulaele de la Nalllrr, publi heel in 1733. says that wasps will enter beehives to plu nder them of the honey. and \ ariou8 authors credit the Eu ropean homet. Vespa crabro. with robbing be<'hives. This the writer has not wi tnessed.

[103 J

104 NORTJr AlLEllICAN Y'HSJ1'r.;~ WASPS

Th~ author has t:.ken W:lSpS f various species at the ftower of Amp.topsis quillqlle/olio (\ irginia CJ"e~er), Bacchoris douglasii (mule fall, .Chr},so­thom",u sp. (rnbbit brush), CI~Offle sp., Dallcl4s corola (carrot), E71ogo/lum umb,lltJIlltn, E. fosciclIlaj.",., and other species ( wild buckv.'healS), Foel/iell­IUIII <'Ulgau (anise) , GrimJelia sp. (gum plant and rdatives) , Heliolro/'ium cura.uCltlin4", ( wild heliotrope), Up/'ia 1I0diflorll (Iippia) , Scrop/lulorio califorl/ica (California bee plant) , Solidago SPI'. (goldenrods) . Sltrculia (8ro<h)'c";lolI) ditJl!1'si/olia (Australian bonle tree), and Symp"oricorpus racemosus (snowberry). A careful investigation would greatly extend thi~ lisL

With the exception of H eliolrop;",,, , 1.i/>/>ia, and of the composite, 8acehnris, Clor)'sollram"us, Grindelia, and Solidago, all the plants listed bear flowers with rei,ltiveiy broad and shallow ooroll.,. The nectaries are thus well exposed . a fealure es enti.~ 1 to any fto ver in which the nectar is to be reached by the mparntively short and broad mouthparts of vespine wasps. The crowded florets of H tliolro/'iu'lI, Lip{>ia, and the compo i~ listed have coroUas which, through not brood and shallow, are quite short, and are uffic iently open to accommodate the mouthparts of a wasp. "'hen visiting the urn-shaped Howers of SIn-culio dit'usifolia th ' ''".lsI's thrust their heads dC('p into the corollas.

Honeydew iront any and all sources is eagerly sought by \Y85PS. TI,e author has taken thml coUecting honeydew [rom Abiu conca/or (lowland fir), dahlia, Cirsi",n (wlfeo/aJ"", (bull thistle), CoHiu/II tr'acuitJIuI/I (poison hemlock), Cotoneaster 5pp., CrtJIoDgus (hawthorn) , Pinus radiaJtJ ( [onte­rey-pine), P'},roran/ila pp., and Vacd"jlllll OVall"" (huckleberry) . In most of lhese cases, the honeydew was produced by aphids. though that on the Pyraconliw was p oduced in part by mealybugs, ,md that on th~ pj","s rad;ara by the scale insect, Toumi~lla piHicola Ferris.

Manufactured sweets will be catm whenever available. Wasps are regularly captured in fl y traps baited with molasses or other syrupy mate­rials. Davis (430) has reported taking severa! specimens of Vrs{>o eraora in mol ass ... traps set for Orthopler.l near Great Kills, taten Island. Jams and jellies are pi lfered at every opportll nity. .

The juices and pulp or well-ripened fruilS are often COIl.unted III great quantities. In year of abundance, wasps may in fact, because of their fond­ness for iruit juices, become " vcry real [lest to tbe fnut grower. Worker wasps have been observed by the , riter to feed On appl and apple. peel: ings, canteloupe and watermelon rinds, gropes, mulberries, ~he {rllll' o[ Opuntia sp. (prickly pear cactus . peaches, prunes, grapefru1t. and sweet pickles. rkers have be .. n observed bUlling about jars of ~ned fruit in "shed. One was noted at a pil of discarded tomatoes, but II was not ob­served to feed. Additional fruits recorded in the literature are apricots,

I \().I I

IHOLOGY oy TUE VESNNA& 105

greengage plums, and pears. Bromley (21) includes cider in a list of foods of Vrs/,u/a "'Deu/i/rons.

\ 'arious gbuldula r ""udates otbes than nectar, also planl saps IlIld other liquids oozing from wounds On plants, are collected by wasps. The writer h ob erved workers of l' upuJa pcnsylvanica and Do/icl1Ofl~s""la arroaria Ii 'nj; the involucra! bracts of cornflower. CC>Jlourea noar"'D . On ovem­her ,1927, a worker V u{>IIID pmsylvorrica eaJll e to .. woodpile evidently at tracted by freshly cut eucalyptus and oak wood. I t hovered about the CUt ,urfru:es, and occasionally alighted, but was not observed actually to eat anything, Workers o i V . Pltllsyit'fJn;ca and fl . suJp/rurca were observed on ~erltmber 2 1. 19_ , feeding on a sticky exudation from diseased acorns of the Da t live oak (QuercllS agnfo/io) . pecirnens in thl! t.~ford Univer­.ity coUection taken by G. A. Coleman are accompanied by the note: "They ,ct llled to b· intoxicated from ipping a very hea,,}, liquid ""uded from . <c(\ cups of tile English i,'y and all were easily taken by picking up with the forcep ." Bromley (21 ) records VtS/la erabro as coUetting in late sum­mer around bleeding injuries on elm, maple, and birch to feed on the sap. n"" is (43) reco rds V. craoro as feeding On the sap of the birch, maple, popl,..., 'm<l oaks. often at w unds produced by wood-boring larvae.

~! eats, chiefly /nU$cular and glandular tissues, no matter what the source, are collected as food by wasp . Meats are acceptable fresh Or cured, muke<! Or raw. They will be taken when partially decayed, thougb th au. thor c:umot recall ever having seen wasps feeding on substances that were I"tlri d.

\\ hen coUecting the ti sues of animals (usuaIlY"ertebrntcs) too large III be carrj~~ away in a single load, " wasp a1igbL' on th · food mass, bitcs nut a roundish piece nf appropriate i,e, and flies away, carrying the food I ell · b eath it head. T£ the mate.rial is too tough to <out readily, the wasp Lut·s o r scrnpes off a number r li t1 le bilS and Ihen works these into a pcllet b ior~ carrying them away.

The cutting Out of pellets of meat is often strenuou proces requi ring tw .. or three mirll1t . The wasp cliDg$ with all six I gs to U,,, meat works hoth f rom ide to side and downward, arching its body mor and rna;'" until, when the pel1 t is finally cut loose from the larger mass, the wasp's man­d,Ll . may be cLltting close to ilS own abdomen. I f the meal has been cookl!d "" the fibers are separable. th' \V<lSp ,viII combine tearing with cutling, in OJr er to ge~ a manageable piece free, In haping a peUer prior to 8ying away wrth .t the w p sometimes brings into play the forelegs and also Ihe middle nes . Wher. large pellets are carried. not only lhe mouthparts hut .the forelegs as well are used for bolding the pellet.

. f he following foods of vertebrnte origin have been observed by the "TIler to be coUected by wasps; roast beef, roa t lamb, boiled ham, cooked

r lOS I

100 ~ORrn A).I1!:'A.1CAN VESPINE. W ASPS

beefsteak. bacon rind. cooked calf's liver. a mixture of raw i,.,.ound bee!

L• b~f lun'" ""d salt salmon froo, which the salt had b~ removed by lver.,~ ... - . • f' I .-~ )

soa.king (th e mixture was being used for tee<1mg fish r)' to a '""",ery . wienerwursts. a dead cottont.-.il rabbit, a dead field mouse (Per,nllys,ur sp.). the vi cern of freshly killed tame rabbits. fish (both ~ked and raw} • . a toad. :trld a dead lizard. Other foods oi vertebrate anglO nre recorded '? the literature. Bromley (21) lists blood among the foads of Ves/>l1ln onaClII,­frOlls. Davis (43. 46) records a dead house sparrow and a watedrs~c as food for undetennined species of wasps. and a rattlesnake ~ " co,:" snake" as food fo r "Vespllia ,orolilla" (= If. squamosa). Phil and Nellie Ran (104) list greasy chick n bones. the eyes of a dead rat, cold roth. and the head of a rooster as furnishing food (or "1'. gem,a"'''a''. (probably Vespula tlmet/lifron.r. tbe Anlerican species most commonly rOlStaken lor

Ves" .. la g.·rtnanira). . . ' . Though the preceding list 01 vertebrate toads IS mlpr,:"""e. the bulk o f

the animal foods con.um,(\ by wasps without doubt conSIsts of to ects and other arthropods, with an occasional true worm thaI IS p,cked up acoden~ tally. All species of vespine wasps . e.~cept per~p. the workerLess. soc~all) parasitic species. hunt insects assiduously. C~tlpedCS and arachOlds. ,:hieH~ spiders. are also taken on occasion, though It .s p~51ble t."at only prevlOusl) disabled individual nre s~ured. In so far as anllllals louod already dead are concerned. wasps appear to exercise liltle or no choice ~ to what shall be eat~ . In regard to living prey, aD the other hand. certato lactors. slIch as peculiarities of structure aod mode of life, tend to protect cerlaln organ­i5lllS from the fo,ays of wasps and to render others parucularly :ulncr:-ble

.

A b· ...... ,elI and burrowi.tg fonns. inaccessible because of th.If habItats. qua .5 .. __ . ' h' hI . .

rarelv become lhe prey of wasp . Creatures WIth po,sonolts or 19 y Irn-

rating secretions are doubtless con, iSlentl)' aVOIded. Certato large bc:etlc~ a nd other [onns that are too hard-shelled to be readily hewed. and nllnute

. t - " 'I to l>c at any consequen<e. enjoy emplete 100muntty. Milch spCCles 00 ,,~ . nall T ed smaller pre\' than would be expected. however. are oc:=aslO y ub IZ .

Rau ( 110) 'records "Ves/,,,Ia gqnllall;ca," ( = V. ~".c"ilfronrf) as coll~ct: ing cba.lcid wasp. T he write r has obscrv~ Dolu:"OfJ~s"ula, arClUlNlJ 001

lecting minute flies less than on~ghth ot an lOch ID length from the

flo rs of wild Inllstord. Several of the flies were taken one after another

W b . d ch and the whole lot then worked into a pasty !!U\.5S to e carne away. u . sma1I prey pr bably are taken only when very abundant aod when the"

capture is ea5)' . . b AnIOng the recorded insttt prey of wa.~ps. beetles nre conspICUOUS '!

their scarcity. The :lutllor has not encountered a singLe reference to a nan~ed s~es. i-leither has he records o[ personal obse"'ations of l>cetles !iCrYl.ng as wasp prey. Still it seems ani)' reasonable that solDe of the soft-bod,.d

t 106 1

BIOLOG\~ OF THE YF..5Pl.NAE 107

beetle. should occas.ionaJIy be captured . and complet"" observations ,,~ 11

probably ,how this to be the case. Janet (75). in France. list. fin ing the clytrn of beetles in food debris in a nest of Vespa c-rabro. Dipte ra, on the other hand. fonn a large pe rcentage of wasp food. Few if any species seem ttl be actuall), disdnined uy wasps . Adults usually are taken. though larvae are occasionally utilized. ipterous prey of wasps observed by the writer iJldude various 'Illthomyids. a bambi lid (bee fly) , Luci/ia, spp. (green bottle fi ie,). C"",""yia, macelJaria (screw-w nn fly). Phornua regilJa, CaJli/>"ora rrytltroc8{>haia. and other blowflies. Mllsca, dome.r/iclJ (housefly) . Eristalis W IG-" (drone fly). TI,trl!'l!o '!lcu/a (Th"evidae). and species of Tipulo (crane Bies) . Records in tile literature are numerous but, except io the case "f the houseAy and blow Hies. rarely Jlam tlle species concerned. Bromley (2 1' . bowever. records Dolic/,ov"s/"Ja rnat:lllaia as preying on Musca do­"".fica. StOtllox)'s cald/rans (stable fly). Pltormia regilll>, Taba""s orio" (a horsefl y), Eris/aiis saxONI1I. (Syrpbidae) , Ti/>lIlasp., and the robber Oies 11rrUIIIJoia ",,,orilla • .!lsi/us ",.yll,rooulI";lIs. and Loptogarlcr jlavipcs. He aIm r"""rds Vespllia ,,,acul'froM as captu ring Musca d01flulica, SI01M.,·\·S <di<1Ira,lS, and TabanllS lasioptltalmus. Davis 43) ci tes ''V.s" .. la caroUn~" ( = V. sqllolll<)sa) as pturing an asilid (robber By). Watson (152). in F.ngland. says wasp food " consiSts largely of Diptera. One observation 00

a Ulicny or about sh..-ty workers showed two hundred twe:nty-seven rues 10 ue brought to the nest within an hour. In another nest examined about 90 1"" cent of the fl ies brought in were muscids." . \' ery iew Homoptern are known to be taken by wasps. Gyrolla NIgam " lhe only one observed by the author. Davis (44) records an observation o i He nry Bird at Rye. 'New ork. on September 18. 1924. who saw a I.'"'ta croOro capture a cicada, Tibic",. lyri' ..... in midair. And at Arrocha. St aten I sland. _cplcmber 10. 1916, Davis. in company with WWiam . \\' right anll Edward J. Burns. "\Viln~'SSed a Living Tilliee', chlorOIlUlra (T . . ,o),i) being devoured by a craOro." Watson ( 152) records jassids ( It ... r -hoppers) as wasp prey.

V tt.rious<lther HymenopLera a.re frequently taken by wasps. Th honcy­be is reponed by Bromley (21). Denton (47), and Fabre (56) . Uau (110). " already stated. repons "1'0,,/,<,10 gCfflla"ica" (V. maw/ifrollS) as cnptur­I. ~~kids. ,\fatson (152) states that larv.e of P/erollus (N.IOm/us) nITr.w (gooseberry >3wAy) are attacked by both qUL..,n a nd worker wasps. . \ ce'"dmg to Fyles (65), . tOne r ports a winged ant as occurring among the prey of an E nglish species of wasp. Various of the smaller wasps oft"" fall prey t larger wasps and occasionally to others of their Own species. RllU (110) records ''Vespulag"''''G ll ica'' (= V. IJllUulifrollsf) as being at· tacked and killed by Dolicloot'gspula macura/a. Bromley (21) records the sallie tiling. Davis (46) reports aeeing an individual D. maculala catch sev-

1107 I

108 NORTt! A MEltICA.N' VESV'IN"B WASPS

eral individuals of "Ves~lJ4 (arolin'" (= Vespu ta squamosa) and then let them go. The author's wife on September 11 , 1928, a t tan fo rd Uni· versity, California, discov.r.d a worker Vuptda p,nsylvanica hanging by one hind leg (rom a chrysanthemum lea f and chewing on the abdomen of nnother worker of the same species. Jt seemed to make little progress and shortly flew away carrying the stil l intact abdonlen with it . [n declining colonies toward the end of the season, lan'ae, pup.'\e, nnd unemerged tenera! adulu of probably all species a r dragged f rom their cells and eaten by workers of thei r own colony. A fuUer discussion of this cannibalism occurs on pages 117 to 120.

Larval and adul t Lepidoptera are important sources of wasp food . Es· sig (55 ) includes caterpillars among the injurious insects destroy by wasps, and Rau (107) records Doli("O<Jrs~uIa ma"lIma as coUethng cater· pillars. Rau (110) al o records a red· winged moth as prey to "VIS/",Ia germanica" V . ,1t4€ulifrons?) . J. G. Wood, accordi ng to Bromley (21 ), reports a V cspa crlWro in England that c ptured the red atalanta butterfly. Bromley also list ti,. cabbage worm in this country . According to Janet (75) . in F ran e. Girard records hornets (Vespa crobro) as intedering with the rearing o[ caterpillars of AI/acus ya"'a· .... a.:, and Flllioo in einc-et- isc found hornets attnck,ng caterpillars 0 1 Allac"s 'l",lhifk which he was at­tempting to accli matize. O'Byme ( 6) in thi country reports finding" worker of " V . stula gcrm<lnico" (V. maculifrons') at work on a battered but still living Eac/es ill.penalis. The "asp entered the moth 's abdomen thro1.lgh a hole on the under side and emerged with a yellow mass larger than its head. 1I'l.any trips were made at fairly regular intervals. In Au­gust J926 the author walched two worke", o f Vts~ura ~CII.sylva"jca dis­member a cutworm which he bad dug up and killed a rew moments before.

Only one reference to a mecopteran among the prey of wasps l ome to lhe writer's nttention . Jan t (75) reports fin ding the wings of Pallorpa com,,,u,,is in food debris in a nest of V .spo crabro .

donata as \ p prey bave been recorded but rarely_ Bromley (21 ) ree rds EnalklgllllJ sp. and Losles foreif/al lts as prey of Dolichovespula

",aculala_ Among the rthoptera, Dou",s cora/ina is recorded by Rau ( 110) 3 .

flllling prey to "Vrspula Oar"",,,ico" (= V . tnaCUiifrollSf) . Rau lllso, ac' e rding 10 'Byrne 96), lists a roach among the prey of "Vespula g fr-,"att;ca." Essig (54) l i sL~ grasshoppers among the prey of wasps.

few foods do not faU under any of the ""tegories already given . Tbe author has observed species of V.sf/ufa fe ding on cooked mush at a tan1P table. also 0 0 boi led egg, on \' gembl. soup from a film relnaini";l: i~ ~he bottom of a plate, and on melted raspberry mousse (ice cream) . J n ImblblDg these liquids [rom the surfaces of the plates to which they adhered, a wasp

[ 1011]

8 (OLOCV 01" Ttl£: VES Vf NAE 109

would thrust the ti of i momhpa.rt3 into the liquid and spread its man. dibles so that thei r cutting edges were separated by an angle o f about twenty d~rees. T he mandi bles were thm alternately spread a tri Ae more and approximated at a rather rapid rate. the head meanwhile bobbing up ,!Oct dowlI slightly a t the same rate. P resumably the ligula was being ex­tc:nded and withdra wn in a lapping motion. udace tension appeared "de­tll"'te to d raw the liquid upward between tile mandibles : the oth r activ ities 31'1"" n: l)' w,:,e concerned with getting it up into the pbaryn x. (Sec p. 27 for a diSCUSSIon of the probable action of the labium and maxilla dur­ing' the imbibing of liquid-.) Bromley (21) records "garbage" amon" w;u,p foods. And adult , asps o f all species and castes eagerly imbibe lh~ -.Ih'ar.y secretion produced by lar 'ae of thei r own spede •. T his food is e. ten in considerable quantity and POMeSI>CS great socia l ignificance. I t is inlly discussed on pages 114 and 115.

' r: NSEs USED IN Lo TIN G F OODS

Both <ight and smcll seem to playa part in th locating of materillis suitabl f r food . In the captu r of living prey <sec next section) . sight seeUl. t be the more important. ight alllO is undoubtedly import.u1( in the Tl'Coh'l1ition of fl owe rs from whi h nectar may be btained. In the location of lIIuch \Ie.'\d aoimal matter and of exudates from extra· flora! glands and ir"l11 wound on plauts, mell is certaioJy the ense n which dependence i chiefly placed . . mell certainly guides wasps to the molasses or other ba.it p!:.ced l!eneath Bytraps, for the baits are invisible from outside the traps. A worker wasp, ob.erved b the write.r as nttrn ted to canned fruits in a .herl. was buzzing about the f rui ts in the early morning before there \vas hgllt cnoncrh to distinguish objec.ts clearl . When cooked meat is exposed at a picnic or luncJ,eon served ut of doors pa.sing w' ps are quickly at­trdcted. and it nJaY readily be observed lhat the wasps approach the (ood fr m "down the wind, " following tile odor- unti l their sourc is reached. A per iormance in which b th ight and smell may bave been involved ,vas reported to the author by J. . hamberl in, H e collected several specimens of /'esfruia .f'II{p' II/Tea Sauss. On the beach at ear u.k., California, wh",e the)' were "hawking OVer a ditch full f dead dry gnat.., fann ing them with their "'JJ]g5." T he gnats proved to be chironomids. It i prob~ble that in the lIoI.tom of the ditch the gnats we.re not dry, bllt were decaying and ~roclue.mg an odor. That sight may 'ometimes be Illore u clul than mell III the location of dead animal food is indicated by an observation of Rau (110). A worker " Vosp"la o~",allica" (V. maculifroltS!) , which Rau was watching as it cut up a Iarse moth. was using Rau : a landmark, and had ~~.lIicllJty in find!ng tll moth when Rau changed his position whil ' the wasp

as away carrytng a load of food to the nest. Rau removed the wings f rom

[ 1091

110 NOKTII AlL&.RJ AN v£SPINa WASPS

the moth's body, placing them a lew inches away. The wasp returned to the wings 6rst but eventually located the body again. Rau co"ered the moth body with leaf and th!: wasp had difficulty in 6nding it. Just as the motll was rediscovered, the wasp was frightened away by a dog. Having had no opportunIty to make a new locality tudy to aid it in rccogniullg lhe leaf that covered the moth, it required twenty minutes for locating its prey again.

HUNTING METIIOOS AND BEHAV IOR.

H unting wasps, according tn the writer's observations, all show the same general behavior. T il y fly back and forth more or I slowly a rew inches above the ground. beneatlt the eaves of bwlrungs, in [rant of , , lis , over weeds aud nther berbage, or a little in {rant oi lhe foliage of tr~"'s and shrubs. Whenever wtable prey is found it is instantly pounced upon. T here is usually no stalking proc . Occasionally a huming wasp will ali ht on the gr und or on vegetation, sometimes, apparently, to r~ t , at oth" times to search mar thoroughly [or prey. Wasps do not at any dOle, however, engage in the as.iduous ground-hunting th3t i characteristic of llIany o[ the spider-<:atching psammocharids.

Typi cal hunting methods are well illustrated by the following obser· .-ations of the author. On July 2- , 1923, DolicilOtJ8Sp,,14 ",acII/ala. work­er were observed catching blowflies on th' docks of a fish cannery nt Dodson Station, regan. The flies collected On the outside of the walls of tllC cannery and 0 11 the empty fish boxes tanding on the docka. Hue Ill' , sps came to gathcr provende r. Mostly th y flew back ,nd forth making udden pounces upon the flies from a distance of four to six inch . They were not very skillful, as 3 dozen or more pounces might be made before a fly was ecured. One wasp which ,vas taken imme­diatell' al ter capturing a fly had its lego around the fly and beld the Ill's thora.~ with its mandibles. Occasionally a wasp would bunt by alighting and . taIJ..;ng n fir, or by merely waiting until a Oy came within reach. On. wasp followed this procedure for several minutes without, howel'e r, securing any prey while under observarion . \V h"" afoot in this manner ~ wasp as stanled by any 8y which canle near. If a fly passed overhead \\ ithin a few inches, the wasps would quickly raise their heads and fore­legs :lS if to dnrt at tl,c l1y.

The wasps apparently did not distinguish their prey by any sort o f real recognition so much as by size and perhaps color ala n<:. Conl~'l with the background was probably important also. They repeatedly trK'<I to capture the nailheads which were conspicuous on the whit." hed cannery wall., ' 0 w:lSp was seen to mak. more th3n one attempt on the saIne nailhead, but veral pounced on nearly e,cry nailhead encOLmtered .

[110 I

o rOLOCV OF Tl'E V£.sPJNAE III

The. riter bas observed the same belmvior 00 the part of f' "p,lIa 1> ..... ,.1_ ,;amca, and ,t has been recorded by other workers for ,'>!rious species of n~spine wasps.

The wasps occasional ly pounced on one another as they did on the flies. Almost as soon " they ca.me in contact, howe"er, they separated ag in. Recogni tion, if this und"lay tllei r immediate separation, thus 'PI>eared not to OCCur umil actual contact had bc:cn made. Davis (46) r (,rd similar behavior on the part of a D. ,,,,,(ulall> worker, which " wn seen to ca tcb stveral of these wasp .. ' ( VUPII/I> sq"lJ>llOSO, reponed as "V. corD/I",") "and then let them go. The ",am/alrJ bad probably mistakc:n them for Ries."

W hen hunting prey or when cutting up food to be carried away, w".ps arc o ften very much preoccupied and can be quite ea ily captured. Five specimens f DofichO'l.respul4 ",aClllaia h1mting blowflies at the /ish cannery referred to above we,." taken with a handkerchief. As nlany as Iwellly-,,,, yellowJuckets of sev ral species have been picked lip with a forceps by the author in Q space o{ five minutes as they came to iced on cold meats s rved at a garden party. \Vben present in large numbers howe" e<, foraging wasps sometimes become :lTouscd and resentful. O~ ull" 0 casion wh"" the author netted some V O$p,tla p."sy/tm';CrJ workers c!t~'1IG'ed in cutting up a dead cottontail rabbit, other workers left the rah . it and buzzed around him excitedly, though none attempted to stiog. . r real",«,,1 of Col>lur.d prcy.- W hen prey is secured it is usually killed """,ed,at Ir· The capto r crushes or cuts the prey ,vith its mandibles. On the ther band, captured insects are occahlonally carned some distance I .. f .. re being killed. A Da/ici'DVupu/a maculala work .. , taken by th allt h r at Dod n tatiou, Oregon, was carrying an unharmed By it had ·nptured. BOlh were put into a cyanide jar, where the By died no sooner t lian the wasp.

The writer has not observed any "espine wasp sting its prey, as certain nther authors have reported them to do. For pie. Denton (47) in rl'[>ortmg the capture of a boneybee by "Vrspwa" (Dolic/,Ot'upula.) mocu­l~a :u- Robbinsville, rth :lTolina, n October 4, 1929, says : "A fter ' tlll~ng It several times between the segments and hiting it severely, the hnn'et carried it away to devour al leisure ." F. Martin Duncan (S I), in "' ferenee to Vesl",/a tJUigans, says: "It is chieBy as a means of procuring the n~essary ani.mal food [or her young that the mother wasp uses her ling. And agar.n. r:aptured Aies are eventually killed by the " repeated

.. t'~I, of the wasp' stiog." Rau (UO), also in reference t the belmvior of I 'r/,lIla gerffuJnico" (V. fHllCu}jfrOHS f , says : "Another was seen to

tlng an adult l~ras51 hopper. foll ow it in its agonized Bight, a",[ sting 't repeatedly until lost from view." These Sllltements the writer believes

[ 111 I

112 NOItTtf A.M£lltCAN "F-SPJS£ WASPS

to be simply errors ba~ 0 preconceptions rather than precise obser­

vation. Prey that has bem killed is usually cut into pieces. The wasp will cut

off a piece small enough to be carried and flyaway ,,;th it, making a locality study before leaving to insure that it hall find the remainder. The cutting and carrying , ill be repeated until the entire prey bas been removed. 00 the other hand, prey ,hich is small enough to be carried without cutting may be takm to Ihe nest whole. T hi fact has lJeen noted by F . Marcil) Duncan and by Rau as wtll as by the writer. Dun­can (51) states tha t VU/>Iua vulgaris occasionally carries flies intact to its nest. Hau (107) records having observed a Dolich01lcspula maculala

carrying a , hole caterpillnr into a nest. Captured insects and OUlI,. animal foods are always sooner or later

thoroughly chewed by the , orkers and convened into a paste; for the Ian'll., to which such food i destined to be fed, are ltn3ble, because of thei r mall and weak mouthparts, tu cope wilh large. tougb, or very bard masses [food. Tbe chewing of the prey or other food is termed "maJaxaLion" and m3Y be cnrried out immediately after capture or de­ferred until just before the larvae are fed. O r it may be partially carried out when the prey is fir t secured, and complt!ed later. The [ollowing .,.'Camples will ill llstrate the variable behavior of wasps in Ilus connection. On September I, 1924. seve.raI workers o f V rspula i',,,,sylt'OlIica w~re captur d as they return"! to their nest laden with prey . One carried the head and anterior third of the thorax of a bombilid fly . 1 burden ap­peared !lot to have been chew d at aU. AnoU,e .. worker carried a partially malnxated pellet compos..!, as nearly 3' could be made out. of a part 01 a spider. A trurd carri ed a pellet which had been so thoroughly malaxated

thaI itll nature "'as quite unrecogniz'lble. When culling "p capw red insects the wasps usually discard certain

parts. These regularly incl ude. the legs ~nd \~';ng . oboriou Iy ~use of their high p6Centage o[ heavily sc\erouzed ,"legument ~nd. their rela­tively non-nutritious c.ha. .. acter. Sometimes the head :liso IS discarded, or the abdomen. The advantage in Ih . cases is I10t always clear. If the prey be a fly of a species with 13rgc ahdomin31 air sacs. thl:D lack o f nutn­ment is doubtless the reason for dise3rd ing the 3bdom I. The head, on the contrary, containing as it doe the brain and I3rge optic gang~. is certain ly not devoid of food ' 'lIlue, though it is probably less nutnUOUs

than the very muscular thorax. . Tb dismemberment and malaxat i tI of pre)" m"y be carried out nn

the ground o r on an elevat support. The weight of the prey is oft"l

the deciding factor in is matter, but there seenr to be no other guld,ng

prill il)le. few examples will illustrate.

I 112 I

nlOLOCY OP TfiE VESPIIS'AE 113

On October 1929, a worker VultUla ".n.rylt'tmica was observetl 0 11

the ground at fenlo Park. a1ifomia. cutting off the wings of a dead Erisla/js ttna.r, a By obviou Iy 100 I3rge to be dealt wilb elsewhere.

On July 26, 1.923, at Dodson ~l:3LiQn. Oregon, a worker Do/ie/lovup"la "'Jeli/aW which had ju t caught a blowBy flew with its prey 10 th top of a box II r by. Then holdiog the 8y with its front and middle legs. il curled it> abdomen under, apparently for suppon, unt.il its body made a half dr e. As ia,[ as could be seen it did not sting the By. With its mandibles it eU ofI th lIy's wings and then its legs. After the first wing was sev­.' red the \ p bit the ny" bod)' and head a rew tim . When both wing and all the legs h3d been removed, the wasp crushed the 8y thor ughly by biting the head, thorax, and abdomen repeatedly, turning the 11 )" IrOJ11 side to side as it did so. The fl y was not chewed to a pulp, how ver. and the integument apparently was not broken open. The wasp the II Iln . away \ ith the 8y.

Bromley (2 1) records a somewhat different behavior [or D. "'1JC14/ala, "');lIg : "As soon as he \'ictim is seized, the hornet lollows a regular procedure. .It immediately seeks a resting place where it assUlll the follu"'ing g rotesque atti tude. It su pends itl;elf by One o f its posterior lel(s, head downward, and v igorously chews up its prey after clipping otT the wings and leg : ' Essentia.lIy similar behavior was exhibited by Ihe llecimen of Vnptlla PN".l"vonica observed by the "uthor's wife and rcpurtlCl above on page 108. Bromley reports anoth<r D. macula III :IS

Lli crnh ' ring a lIy on the ground . clipping off the bead. winb'S. legs. and al.<J men . Part of the abdomco (doubtle only liquids) was consumed; the Ihorax was carried awuy.

unLIZAT' OI< OF FOODS BY WASPS

Food "t alillitS.- The rood o f adult wasp, :l.S was pointed out on page WJ . colloisl;! only I liquids or ' ulially liquid ubstances. A portia! c planaliun of this iact is fou nd in the structure of the mouth an eso\>I13-~" The mouth (Figs. 12. 18. 2 1. 25) i reduced 10 a ver" narrow " ,"",verse slit. Tbi is guarded, moreover. by a comb-like ro,~ of tiny, Olll \\ ardly directed setae. rhe oral penen, which effectively prevents Ihe ~'n l nmce of any particles o f solid matter except tllOse of microscopic .ize. I he lining membrane of the phar~'tI'" aI ,for Ome di-tance inward fr '

'he l11outh, i pro,,;ded with olllwardly directed setae, these being excced-1II1:ly line ""d close et. They douhtlcss train out solid particles which • l":lPC tbe oral pecten and return Ihem to th mOllth. The solid materials 'llu . tmined [rom food subslances arc stored in a membranous invugina­"'m. the gnathal pouch (Figs. 12. 18. 23. 25, 26) . This presumably is pcn u ,cally emptied, thOUGh no specific mecha.oism for its eversion has

( 113 J

114 NORT U AlolERlCA:-: V};S.P l S't,. WASPS

been discovered. \\'heder ( 162) = d Janet (74) describe :L.nts as having a imilarly org:ullzed mouth, pharynx, and gnat~a l pouch. T!'e esophagus of adult wa.5pS is simply a slender, thrrad-hke tube whIch l r:lYerses

the thorax- . . The bulk of lhe lood of adult wasps consists of sugary liquIds such

as nectar, honeydew, honey. and plant saps-ill f~ods of ~gh en~r~y content. rn additi n to these, w:lSJls consume liqUIds of arumal onb:.n. Foraging workers often suck all the liquid nutrimenL that may be eas.ly extracted from the pellets o( m:lialGlted insecLs wb'ch they earn' back to the nest. Wheo engaged in cutting up the tissucs of dead verl<brates, worker wasps imbib freely of exuding juices. [I will be recalled aL thl~ point that Bromley (21 ) rerords blood as one of th~ observed foods. 0 1

Vrspu/a IlIOlII/ifron,. A workor Vcs/,fl la p."s)'lt'tJlllca,. olrsened dunng the ummer of 1926. sipped the juice from a bit of the mt",tme of a cut­worm that hnd just been killed by the wri ter and was adhering to a clod oi earth. From such foods wasps undoubt dly = ure enough nitrogenous lIutri ," ent to satisfy their needs and round out theU' clict_

Wheo liquid ioods are available in quantity, foraging worker some­times gorge them elves and ca rry the foods to the ne.t to ~ shared. th rough regurgitation, wi th ther. adults and perhaps also WIth lan~. Lubbock (85) record. ma rked Vup,,/a gC"""lIlic" . worker which came repeatedly. at .inten ... ls of from five to fourteen .mlOuLes, generally. to a di.h of honeY which he set out for hem. Thl conUnued fo r several d:lYs. rabre ' ( 56) stnte. that when V cst" craoro capture; bees it kil ~s them quickly, rips open the crop in th ' abdomen, and laps up the honey It contains. Whether foraging w rkers a1. collect nectar or honeydew and carry it to the nest to feed other members 01 tbe olony has not been

determined. . . The sharing of engorged food by adults may be observed r""dily w,th

captive wasps. All that is necessary is to withhold food for me hOUr>, theo t feed liberally a portion f tbe starved wasps, and lastly to liberate the remaining starved wasP" in the cage containing the satiated ne . The starved nes will pr()111plly solicit and recei l'c food from the others . Whether all liquid are thus shared by adult wasps .cannot. be stated on the lJasis [ available data, bUL this cenlS li kely. It IS pOSSIble. howel'er, that meat juic.es. being I ... abundant. are shared to a Ie ' r degree t":'" ",eets. I f so. the result would be a slight difference between the dId

of work",r. and lbat <l f lbe olbor c:astes. . ' final normal elenlent in Ihe iood of adult wasp . bneRy mentioned

on page 109, is th~ salivary secretion of the larvae. which IS said to be sweetish Th i secretioll is emitted whenever the larl'ae are stunulated and is ~erly sought by the adults. The production of secretion may

[ 114 I

o tOLOGY OF TH E VESP INAE 115

be induted artifiCIally by touching- the head of Il,e larva with a straw or other suitable implement. The stimulated larva bends iLs bead back­ward and expels a droplet of the secretion. Ad ult wasps of al l castes induce the producti n of secretion by gently biting the heads of the lan ae. Th" drople of secreLion are sometimes quite smaU but may be larg~ enough to COy", nearly lhe entire face of the larva This reciprocal lceclmg of adul~ b~' larvae has been termed I ropirallo.tis by \\ beeler ( 161) · nc.l oocolrophobrosu by Roubaud. according to Wheeler_ · The source of the secretion , which was fi r ' observecl by Janet (75 ) , ,s the labIal gland- This opens by a transve.rsc aperture (Fig. 171, SO) on Ihe lablUru f the larva. This gland is very remarkable in that it lI "d~rgoes .two di ti nct challges in function during the life 01 a wasp. At hr t salivary, It becomes, at spinning time, silk-secreting. In the adult il . regains its salivary function and also prod uces the cemcoting sulistan'

Ul whIch tlte vegetable fi bers compo ing Ihe nest pal>er are glued together.

· !lte fo od ~f /(In!!,i uosps.- The food!. f larval wasps are of two types, h, !uld and sOh.d. K Irby and pence 79) state that very young wasps are ,ed only hqulds. Other authors have presented the same view_ The nature of the solid foods is well known . They consist simply f animal I ".'!les, hewed to a pasty consiSlency by Ille workers, and sucked more or less iree of lbei r j uices. W henever the n h o[ vertebrates i used the resu ltant paste contains only soft particles that are compleLely digestible: \\'hen chewed illse<:ts or other arthropods make up the food mass ti ny pice' of indigestible cuticle f va rring degrees of hardness are 'inte; . mixed with the .oft portions. Much of Iltis indigestible Dlaterial appears to be swaUowed along with the soft er porti ns. as Ille lan ... e possess well­,clcrotized mandibles and always rechel\' the food thai i tendered to litelli. The .largest and hardest of Ilk~" cuticular food mponents are e\ ,01 ntiy reJecred, for a considerable amOUDI of finely d ivided debris of • cuticular nature is to be round 'tl times in wasp nests.

The exact nature o[ the liquids that are fed 10 wasp larvae i not n cJ.a r. All arc presumably deril'ed from the liquids imbibed by the work-r. and later regurgitated. though it i possible that glandular st'cretiollS

IIIay be added by the workers to the other liquids. Then, too, the treat­m""t accorded liquid foods before they are admi nistered to the larvae ~ , a)' vary . For example, tbe liquids may be administered in unchallged oml a fler tempo,:"ry .storage in th e crops of the worker, or they may

lIndergo partial digestion [ore being given to the lan·ae. ~'estwood (1 57) held Ihe latter to be th~ case. ft is possible tl1.,t foraging worker may regurgitate only the coUe'ted sweell;. n the other hand , the larvae llIay be red only meat j uices, Or both sweets and mea l juices may be fed

[ 115 J

116 NORTH AMERICAN vl£-SrlNE WASPS

indiscriminately. Smitb (1 ~3) implies that the liquid. fed to the larvae are of plant origin ooly . Whecler (161) also seem. inclined to tbi. ,-lew. though ho makes no !!p&ific statement to that effect. \Vhat he does say is thM the habit which ioraging workers have, of eking the juices from 1l1e pell ts of onimal food brought in. deprives the d.v~lop­ing larvae of much nutritious food. pecially dunng the early part of the seas n wh"n insect food is less abundant than it is lat"T on. In fact, WI~er pr<!Sellt_ Ule hypothesi. that this elimination of meat juices rom the diet 01 larvae early in the season is oue 01 the (actors re-

sponsible for caste differentiation . Saussure (1 19) long ago raised ~ similar ,'!ue.tion when he asked if it were not possible that the larvae 01 one sex are reared on animal foods and mase of anothe r on vegetable aliments alone. Rollw r (1130) . in a stalemMt referring to the social Vespidae generally (= Vespinae a.nd Polybiinae, me former in his classi­ficatio n i"dud.ing Poliste. as wfil as the typical vespi ne genom), includes honey and pollen amon the foods ied to larval wasps. rt i. the wriler s impression. howev~r. t"" 1 no " pine wasp collects pollen. Ki rby and

pellee (79) state that very young larvae receive only weel liquids, such as nectar. honey, and fru it juices, older larvae being given solid {bods as well.

The administration o f solid foods to la rw,e is reported by various authors to be carried out in me following manner. Holding 0 food pelkt with her mandibl ... , and perhap a1so her maxi llae, in such a manner that m O I of the l,.,]Jet is behind the mandibles. a worker wasp approacbes a group of larvae. The worker may be a foraging indi\;dUllI distribut­ing food which she herself has collected or she may be a nurse wasp tbat has obtained a food pellet from a foraging worker. From the front of th. pcllet " small morsel is cut off and given to a larva. This pr~dure is repeated unti l the entire pellet has been disposed of.

Uncon umable particles of food that are rejected by the larvae. along with discarded remnants o ( prey that arc cut up and rrutiaxated within the nest, a,'e u Ullily ca rried out of the nest by the worker . This has been established by direct obse rvation as weU as b. the fact that w:.sp nests are gMerally qui le free of superfl uous marter. Portions of me integument 01 crane Oies and of caterpillars have be.," identified by the writer in the burdens of wasps captured as they left meir nests. ther particles could be recognized only' being of a cuticular nature. That not all of the leftovers (rom fceding arc thu removed is proved by ttte fact that they may occasionall y be found on the floo r of a nest below the combs. The presMce of such debris may indicat. that the . colony. in which it is found is ;0 a derulling state. On the other hand. to colomes of l' cspula . which are buill underground. mere l11ay be enough of such

I t161

OtOl.OCY 01' TilE VTlsrlNAE 117

debris 10 meet tbe food requirements of numerous 8y larvae living in the earth below me combs. 'nle author has observed these 8y larvae in the underground nest chambers of V {spula ".nsylv(llIUo/l but has not reared the adult flies. Janet (75) records their presence beneath a nest o f Vrspu/a mta near Beauvais. F rance, and states that larvae of four species were prcsenl.

A rather anomalous situation with respect to the food supply of wasp larvae is reported by du Buysse" (22) . Tn wasp combs kept in his laboratory, starving larvae were seen to consume drops of their 0'\\'11 sal i­vary secretion. thus engaging in a1llolrophaJla.ns. It seems quite probable that this pmctic. may OCCur reguh rly in dedining colonies. An intereJ;t­ing question thaI arises is whemer such autot:rophallaxi~ could prolong lhe liIe of larvae. Incidentally. for creaUlr.,.. accustomed to being fed daily, " . p \a.n.."e exh ibit considerable ability to resist death by tarvation . On June 2 , 1928, the author cQlIected 0. nest of Do/j~lruvesp .. la ar,'1tar;a at Cupertino, Californ ia, using carbon disulfide to asphyxiate tbe occupants. _ ot a ll the larvae were killed by the treatment, and some o( these were still alive on Jul~' 12, two weeks later.

An interesting item f beha\;o r has been reported by Wheeler ( 161). The larvae tu rn their head, backward until their mouthparts contnet lhe W"d lls of their cells and then scratch me paper wim their mandibles, a performance which apparently serves to attract their worker nurses and =ure iood for rhe larvae.

CA NNIBALISM

In many. if not nil . old colonies nearing disintegration, the workers, rcduCL'iI in numbers, bt!Come erratic in behavior, cease to care properly for the developing brood. tum cannibal. and d •• ou r them iru;tead. This umocial behavior has been witness d many time and reported by numer­ous auU,or (Kirby and S~ce, 79 : ouper. 34: Figuicr. 61; Lewis, 82; Janet. 75; F. Martin Dtl n<::an , 51; and thers). J anet ( l" ) reports observing Vespa (raiJro worker kill certain larvae in me nest to procure food with wbich to feed others. orne of the earlier writers tlAively re­g'drded the killing of immatllre \vasps by the last of the workers not as cannibalism but as an act of mercy to save tbe brood !rom deam by - tarvation, and ascribed this benevolent massacring of "innocents" to

peciaJ instincts instilled in me wasps by their reato r. SUcll a view was expressed by Kirby and Spence, who say, " The old wasp drag out of the cells all lhe grubs and unrelentingly destroy them. strewing thur dead carcasses around me door of their now desolate habitation." And Lewis (82), as late as 1897. says: " Late in fall . alter al l the flowers arc gone, if t"ere are any larvae or partly grown wasps in the nesl . it is said that they

f t t7J

118 !iORT Ii AloIl::.RlCAN VI:: Pt~E WA SPS

are stung to death by the workers who see'" to realize that they would die from cold and starvation if this was not done."

That there was some justification. in view of the prevailing philosophy of the times, [nr the views e..~pressed by such o[ the earlier entomologists as Kirbv:md pence is evident from Lhe following fact. The killing and dev~uring of wasp brood by workers in thei r own nests is not simply cannibalism, but i a more complex phenomenon; for the workers, in orne cases at lea t, ki ll more larvae than arc required for their neeos

and those of the remaining brood. Iso they do drag or carry many oi the dead larvae out of the nest, which would not be done if the urge to ki ll came only froOl hunger or [rom need for food with which to feed their charge ' . tn a IIest of V cspula ,.,,,,ry/t'a,,ica, on the Stan ford t'ni. versity campus, which urvived the compamtively mild winter of 1923-24, and which did not finally disintegmte as a colony until midsummer 1924, the following and imilar occurrence took place: n January 30, when tile colony was first discovered . a worker was captured as it left the nest carrying a larva "b ut one-thi rd grown wruch appeared to have been but recently killed. On pril 20 three worker· leaving the nest ,vitb dark pellets were captured and their burdens examined. One pellet consisted of parts of the integument oi a crane By ; [he otl,er two appmred to be the heads and body integuments of black caterpillars. Thus it appeared there were still wasp larvae that were being fed in tbe nest. Four worke caprured 00 May 11 carried th [ollowing burdens, On carried the head and the first (ew body segments of a young larva. Two were carrying portion of the head.~ o[ newly transformed males. The fourtb bore the 'utieular shell of the base of the abdomen of a newly transformed male or that f a worker. The apl=J'ance of the burdens of al l but the Ii .. t indicated that the victimized young wasps had not emerged nomlaUy [rom the cocoons but had been extracted from them.

The caUSC" of this uLlSociai behm'iar 113 \'e not been elucidated. far as the wri ter i· aware ; but se\'eral hypotbeses arc suggested by well­known aspects f wasp hehavior. "one of Lhtse has becn ubstantiated . They are presented hen!: in the hope that other inter~ted in wasp bio!ogy may find opportWli ly to test th m and determine what degree o( vahdity they may 1'0_ . 10 so far as the ph,,"omena observed r'Present impl. cannibaliJim they may arise out of cither of the situations described below.

first: During the declining phase o( a wasp colon ' , owing to the devel pment of young males and queens. only a few workers mature to replace those dying of old age. The worker population diminishes. Th. amount of work in foraging for food that falls to the lot of each rem",n' iog worker, therefore, increases. At the s,une time. ill Ihe case of tho c colonies, ch iefi y V rspu/o colonies, which remain active throughout the

[118 1

Bl01.0C'~ Of' TU"'E VESt*rNAB 119

fal l season. insect prey ""come. increasingly difficult to secure because the general insect population decr~s on the approach of cooler weat her. An actual food shortage dnelops. and lmder pressure of hunger the worke" tum cannIba l. Havlllg so.tisfied lheir own hunger at I'he e.'<pense of (X11'l f the brood, the" nursing instincts become dominant and rem. nants of the dead lan-ae are red 10 the remaining live ones. The cmmi. bali m i extended to pupae and newly tranfonned bu t II.! yet un. dTlerged adults; the COCOOIlll of the latter are chewed open 3.lId the occu. pants dragged from them.

Second: Cannibalism may arise out of the trealment accorded ti" larvae during lhe trophaliaetic relationship. . s the normal food SUPPI; decreases and Lh~ larvae b.ecome more and more starved. dleir a ili ty to prod.u~ llle d~lrable sal"'ary secretion probably decreases. Work"", !IOliollllg ec.reuon may th n become more and more "iolent, until . in their " :tempts to mduce production of the se retion . Lhey may bite out pieces ot the heads of the lan'ae or even kill them., a result dley discover that the larvae are good food, hunger supersedes their nursing instincts and cannibalism results. '

The ~ng of killed and partiaUy COLlSumed young out of the nest may i.Jc SImply an outgrowth of nomllli nest·sanitation behavior. In anv n t it is customary for the workers to remove from theIr cell and car~, away any larvae which <lie from dise".e or any olller cause. In a olon; .. where workers ha"e tu rned cannibal. once the hunger o( a worke r i' "at, fied ~t IS quite possible LIlli! its "sanitation in tincts" causr it to carry the remams of it meal a, well as other brood which have been ki lled out of the nest.

rtain objection to these hypotheses are apparent . The colon i of : me species of Dolichuvespulo, tbose o f D. ar.llana in centrnJ California, I r example, often decline in early smrune .. and come to a complete end II) ' mIdsummer, whIle there is still 3D abuncl1nce of insects to 'erve as !'rcy . anJ while en"ironmental conditi LIS are still highly favorable to life III general. Workers in such colonies certainly e.>q>erience no difficulty in r. ndin~ ~uffi.cient food (or d,em elve.. They may. Qf course. because of lim: h rmtauons anel their reduced numbers, i.Jc unable to l>«u re enough additional food to " pply the larvae. Th ' iml)ul.e to feed the larvae may then be strOng enough to cau e dIem to regurgitate much of the food \\ I~cl! they have eaten. , ith the result that hunger develops and canni. hahJ;n-1 follow -. On the other hand. il is possible thM cannibnlism may not develop to the so.me degree in colonies which come to an early end, and ,Perhaps not at all . lIper (34) expresses some doubt in this con. n ellon. The autho: has observed cannibalism only in V n/,,,Io colonies. dlOugh he has not Investigated Dol;chrn·e.rfJuIa c lonies with the definite

[ I t9 J

120 NOR.'[}1 ~JERICIoN VESi>INE WASt'S

aim or determi ning whether or not they exhiLi t canni~aJism. ~pecific refer­enCes in the literature that have corne to the writer 5 attention also refer only to species of flcs/",Ia. A second difficulty in the way o{ acceptance 01

th nnsed hypotheses is the fact that the canntbal \ orkers CUt open

c pro,__ 'thj' 'r-h- Id M nd drag oul the pupae and yOllog adults w, n. 15 WOll ..,em

coc~!IS a . ' abl' ~'e11 quile UJUlccessary as long as larvae remam .avail . e tn uncapl"-u ~. The operation of some other stimulus '~ addition to hunger, .therefore" I plainly sugges .. d. Obviously, wa p bIology IS fraught WIth unsohed

problems.

B ILD ING ACTIVITIES

The building and maintenance of a nest consumes a great deal of the time and energy of vespine wasps. No olher endeavors. except perhaps the prOCH ring at food and the care of Il,e brood, are so exacting. A ,l11all amount of construction work is done by the queen at the llme ot the (ouuding of the colony, but by {ar the larger part o( !, ' performed by the w()rkers. They toil (rom dawn to usk from the ume the first .brood n"'-Iures in the spring until inclement w""Lher or ,nlemaI ~gcs on the colon v Lring their enterprise to a do -'. Bllilding :lct1,· ,tJes IOdude such di\'c~e operations as coHeering building malerials, conv.~mg these mto pal''''-, inc rpo rating the paper into the nest SLructure. deanng the nest Ite of litter "nci other obstacles, and . in lhe ca.se [ spec,es of 11 <spula, excaval­. " th-' solid earth. The versatile implements wilh which these van ed mg lll I,; A- . . . by la, " are chiet1 v accomplished arc the mandibles . ss,stance l> gwen , the fordegs and by other organs. of course, ~Ul no ~ of the \~ p s b<Kl: i, US<.'<l more frequently than the mandibles, or \.0 1' such d,verse

Ilurpa>'''.

TUE BUTLDING 1'Il",. F">u LS

Tlte rw.o' mQlerials.-The materia 01 which the paper in ves~ine wasp nests i, constructed are get:illie fibers and salivary secr~t' o,: TI:e fiber u-.,u I 3y be grouped , according to their nature or ongon, 'rUo ,t.." cia ...... lIameh' : I) the fiber i rotten wood; (2 the fibers of wcather-

., be I { f posts and worn hIll not d . yed wood, uch as can scrape! rom ence rails, (r<l111 unpa.inted buildings and other wooden sLrucrorcs, and from the suri"c!: o i dead trees from ' ruth the bark has fall n; (3) .fibers de­rived frllm the cort"" and pericycle o( the d."d stems o( \-:t:,ou n~n­woody pi n , such as the coarser annual weeds ~d perenruals WhlCh have annual tops; (4) fibers derived fr0111 the hVlDg cortex of shrubs and trees; (5 cocoon silk in the form of [rag~eots of . the caps of coroons within the nesL cut away by young wasps m emergUlg; and (6)

11 20 I

BIOLOCY OF TH E VESl'JNAE

fibers (rom non-natural source • ."eb as discarded newspapers, ""per bags, pasteboard signs, and other artificial products of a fibrous nalure.

·rtain pecific differeDces in the choice o{ fi bers are exhibited bl' wa.ps. II spa crabro, the Brown Hornet, is the only species found i;, Ihis country ,.hich is kner,vn to lise fibers from the cortex of living plants. That it actual ly does s , however. seems thorougbly established. Although Cory 33) states thaI he has sew no evidence in nests in this c uotry of the use of green parenchymatous tissue . as has been reported by £uro­pe'tn bservers. he, as wcll as other observe rs in b th Europe and America, il'om Reaumur, according to Westwood (157), to Bromley (2 1), Davis 4311), and Rothke ( 116), reports II. crabro as gnawing the bark of living

t Ce.9 and shrub _ Lilac is the shrub most frequently mennoned. It seems to be damaged occasionally La slIch an extent as to impair the h..ulh of the I'tant ~ concerned. Bromley says II . rrabro occasion:llly girdles lilac, birch, and hododendron. Bromley {mth nnore reports Britton as saying that 1 . ,rabro damages dahlia stalks loy gnawing. On the other hand, living Ioark probably (urnis),es only " minor fraction of the fibers used by '-. crabro. Kirby and Spence. accordi ng to \ estwood. give decayed wood a, the source of the fiber. and Janet (75) says lhe wood used is gen­e ,lIy more r Ie. rotten. V <spllla .mlgaris !leems definitely to prder thoroughly decayed wood in the brown rotten stllgl!. All of the sel'cral lIe.ts of t.hi specie. examined by the writer have been made from rotten wnod and have. in consequence. been quite fragile. (Figs. 212-2l4, 226, 227.236.239. A5. 246. and 255 . ~urtis (410 .tat - that V. ",liIlO-tis (in England) constructs its nest of fi bers "scraped (rom sound timber. as 0), rved by lhe late Mr. T. A. Knight" : but it is highly probable that Knight's identification f lbe wasp observed was erroneous. Most of the theT America n specie , while using fibers from several sources, seem

tOJ prefer weather-worn fi bers. Wasps of the genus Polislos occasionally lise pla nt hai collected from very pubescent plant · (Duncan. 50) ; but tillS has not ueen reponed for any espine \Va I'. The failure of wasps "enurally to usc the matwe. dry, dead bark of trees is probably due to the fac that bark contains a high percentage of cork cdJs. Thesc are non­absorbent. aod a prereq llisite of a good fiber fo r the manufacture of 11:1 r is that it b~ absorbent ,

In addition to fibers it appears that other sub ' tante' , on rare occa­,ion . are incorporated into the paper of vespine wasp neslS. Isabel ]Ok­-racken o{ Stan ford Urriversity, some years ago, in reporting to the writer

a neH of some species of Dolicl'r)'/l('splIla (p robably D. arauaria) which sh· had observed hanging from the eal'cs oi a brick bu ilding, described it as having a consideraltle quantity o( bric k dust mi xed in , illt the tibers of the paper.

[ 121 1

122 NORTH AME1UCAN VES[' INE \ \'ASrs

In gathering fibers for paper pulp Lbe wasp a~mes a position wiLb its body parallel to the ""is of the stem or the gram of the wood (rom which the fibers are to be collected. She lhen walks slowly backward, biling or leanng off the fibers as she goes. When gatbering. fiber from weed slems r >emi-sh rubby plants a clickmg sound, which may be hea rd at a distance of several f~t, is produced. The sound doubtless carnes from the breaking fibers. The collected fibers arc , orked into a ytt~e baU which is hcld behind the ",audibles. The maJcillae pr bably rud til

holdi ng the ball of fibers . As soon as enough material has been colle<:ted it is chew d and mixed with s:Ili""ry se<:retion to make a doughy mass. The wasp then Ries to the nest \\~lh its load of pulp. Upon arrival. at the nest the pulp may be carried inside (or addition to the comb~ or It may be added to the envclopes 00 the out.<ide o( the ne l. The sahvary se<:rc­Ijon hn lJ en said by various authors Folsom. 63 : Lewis . 82) t~ make tile paper water]lroof. This may he t rue in the case of some specIes. ~ut il certainly is not true of all Most of the nests observed by the wnter absorb moisture r""diIl' or nearly as readily as otllcr malcnals of a papery nature. The aerial nests made by the species of DoiicllD1.'rsp,Ja, howe~cr . " re more durable than rhe lIbterranean ne>LS made by spedes of

fle.pula. . ' . The mokillg of pop"r.-The conversion of p,up <nro pape.r and Its add,-

tion to the n t may be observed readily on an aerial nest . and has been described . usually brieR)" I>y 5e\'eral authors. The method of adding paper to one of the envelopes in the nest wall is as follows: Th. wasp walks backward and forward along lhe edge of the " IWeiOpe eeking t\ IJlnc~ 10

begin. When it has sclected a place, the wasp bol~ he bal! of pulp ag~n;t the edge of the c:nvelope and bite ' It U1 such a tashlon as to fasten It to the <llvelope. he Lhen walk uad<ward. biting ~e ball of pulp repeatedly, working it into a slender st ring ru,d attaching It throughout Its length to

the edge of the ' '''ISling paper. fn tIlis performanc~ the wasp u es ani), her nuddle and hind legs for walking. The middle legs are held more or less at right aogles to her body; the hind legs are directed backward. \ ' ith her forelegs Ihe wasp holds the edge o{ the paper to which the pull' is being "dded. one leg inside the paper, the other outside. ""ben the ball of pulp has been strung OU L and attached the wasp run.s fo~\~ard to the beginnin!:, of the pulp string and. walks. backw":,,d agam, blung th~ pulp once more, this time to Batten It. This behaVIor IS repeated unltl the string of pulp h .. been convened into ". thin . trip of paper [rom one- ixtccnth to three-sixteenths of an lOch In WIdth and from three­fourths o f an inch to onc and one-iourth ioche long. few mJnlltes then suffice for the new paper to dry. aft. r which it is an onte!{l'1ll part of

the nest. [1221

SIO T ... OGY OF Tn'S. VESPUiAE 123

'rIl' Job 01 attaching the new paper to the old is !lot always perfectly done. Frc'luenLly" string of pulp will be attached nly at interval" and ."metimes only at the ends. In such QS., little holes or lIlits are left iu the paper. Th e receive no fllnber attention and may be found in olmost {Ull nest.

W hen CQllecting fibers and making paper, wasps work assiduously. 1{l1U (107) re<;ords observing 11 Dolic/'OfJ.spul4 IIWC14/al4 which worked "'JlIlinllously during the period of ob crvatiOll (4 :21)...9: IS. presumably ... Y . .). The lapses of lime while the wasp was away gathering pulp were r' pectively 5. 5, 7. S. 6, 7, 7. S. and 7 minutes. The len~1 . of time <cupied in ndding paP<'r to the nest envelopes were 2.5, 2 .5. 2. and 2.5

Imnu te . Those spent illSide the en\'e1opes were J. 12. 5 .5. 16, II. and IS minutes.

TIlt fi"ished f>roducl.-J n any nest. though mOr so in the ncsts of cer­rain species th.n (others. L1le paper in the nest wall, and to a 1I!S>Cr e xt"11 1 that in the mbs. shows oumerous variations in color and texture. T he l1XJst obvious variation is Lh:ot of color ~sulting from the use f lil",r- col lected from diverse sources. White paper, brown paper, tan pal' 'r, yellow paper . and gray paper are often juxtaposed Lo make a stri king pattern (Figs. 23~23S, 244. 245) . Textural variations depend ·um.ewhat n the source f lhe fiber, somewhat on th· amount of in­cuq Jt,rnted >aJiva. r n general, the paper in the nest walls has the coorsest , ... ture, that io the suspensoria whicb support the combs the fin t. Ln ne. t. o f Dolic"01!£sf",/a ING,"/ala, however. many of the buttress-like aoldi tinns to the original suspensoria are composed of sheets of coa rse­tnturc I paper exacLly like that found in the nest walls.

In . tr 'Ilgth, also, wasp paper varies greatly . It may be very weak, as in Ule ease 0 f the paper made of rotten wood by I' /!Sputa t.,i{I(Jris : or it n1:1) be fair ly strong. The po.p r made by species of Dolicl'O'IJc",m[a (D. Ill<l. ,,[aro and D. armoNa) of fibers from weathered wood Or weed stems i, ,. ftcn comparatively tough. The nests of these wasps are ft.n suspended frnm the branches of trees and must be able to endure considerable huf­f 'ling. The paper in nests of certain spede~ of V .. pu/a (V . pct/sy/vanica), t""ugh II ·uaJly inkrior to that made by species of Doliclwvu/>,iJa, i of !luod quaJ.i ty ntv nbele... Oue nest of V csputa />""syit'4I1ico dug' up at SlHn ror(! university by the author provides an adequate ilIustralion . The ne:t had been b"ilt ;" ston. soil and was ery irregular ill shape (Fig. 1%. 231 ). The wasp had removed the soil from around the stones en­r"ull lcred and had attached the paP<'r of the uest walls. bm not that of ~he tombs. to the toues. All the stones, bowever. had some support I rom below, in the forn, of other stOlies or of earth . at the time the "est wa o Jug out. [n the digging tltis support was removed_ Maoy o f the

[ lZJ 1

124 ~OH.TU AMERICAN V'&Sr l NE W~PS

stones. including one about two and ne-hall inches in diameter, were then held up by the p"p"-r alone. The largt'St one, however, broke loose and fell 10 the bottom of the nest cavity before the ncst was completely

removed. Th~ pape.r composing the nest walls e.xhibi~ the. gre~test variation in color and texture, and the least strength of all the kmds 10 lhe nesL The pulp of which it is composed is in the ",,',in brought freshly from the fidd, though in Ider nests some is derived from layers at pap"-r on the inside o f the nest wall that are being tom down to make room lor en­largement of the combs. In any ca~e the w:,11 layers are compos:d ot in­numeral,lc lendet strips of paper with POinted nds that are mterlaced to produce a peculiar striped pattern (Figs . 2.>6-238, 244, 245). Each strip is ntllde of a single loael of paper pulp. The nWllb<r 01 strIPS 10 a l.riven piece of paper is thus the same as the number of tflPS tnade by the wasps in search of fib·rs lor pull'. As an tnde:x to the amount of labor in volved in the building of a large ncst, atlentt. n IS called to FIgures 83-86, wh ich ar drawings of lour of the small ",bdls" of paper from the ncst of II CSP 1<W /JenryifJOIli£a shown ill Figure 22~. ! he vertl""! hne betwec:n Fignres 84 and 85 r~presents one IOcb and ,ndicates the .,ze of the "shells" f p"per. These "sheUs" consist'..:! 01 ; 0, 3 17, and 4<i strips of pap"-r, respectively. Many thousands of these shell of paper are mllde during the building oi " single large nest. The nwnl r of trIps andd for the gathering of paper pulp. thereiore. runs mto hun reds of

thousand . In lIests of Doli,liOfJupula. pp. the component ,trip in the wall

paper are appro.intlltely .tmight and run principally in a horitontal direc­tion (Figs. 2 15, _17, 218, 219, 221, 238) . Ln nestS of I'esp"/IJ sPI' ·· as may b< in ferred from the preceding parngraph, the strips are nearly always arcuale, with the inside of the arcs directed downward (FIgs. 226, 228, 236, 237). 1n each I the " hells" ot paper in a V .s~"/IJ nesL the first or uppermost arcunte strips generally are of longer radIUS than the later 0<

l ow~r ones. They al represent shorter segments oi a circle. The last strips to be added to (l "shd.!" of len are of very small radi~.s and .. may even b< complete eireles. \\fheo the la tter LS the case, the sbells are

distinctly umbonate ( Figs. ~36, 237 .. , . , The com lis, also, "re blult up of hule trIpS of paper (FIgs. 211. 21_.

246, 24 , 249), but the trips are of much 5IIlaller ext~t and they lIr: more homogeneous in character. They run tran versel), tn the ,...ruts 01

the cells. as they are added to the cell rims when the latter are btllit ,~p . Sometimes one of the little strips will be limited r a portion of the nnl of a single cell. At other times it may begin in the wall of one cell and extend into the walls separating one or more adjacent cells. T hlS rea'

1 1241

nrO LOGY OF T il E. VESPUfAE 1"­-~

l~trc o f the comb paper knilll the whole group of cdls composing a comb logether and thus adds greatly to the st rength of the comb.

T~e pulp used ill. comb construction. like that for the wall layers, i. 'Ollletlmes gathered rreshly afield. On the other hand, it is often derived [rom the tnne~ layers of the wall that are to rI) dow" a. the nest i. en· larged .. Late In the eason, puip for the newest combs may even be ~ken 1 rom the olde r combs in the upper part of the nest. fo r these are < tten abandoned as brood combs and their rims cut away (Fig. 243) . ~us most of the comb paper is made from fib<rs thm ha,'. b«n -bewed IWI.ce or even ~.e times lind have had more tha« the usual amount of salivary ,,,,,rebon mixed in with them. T his accoants for the gre<lter ~l . mogene lty and strength of comb paper as compared with ,,-,;II paper .r he toughest a nd most h moge.neous f all nest paper, howe"er, is tha; ~ound 1.D the suspern;oria upporting the combs in Ves/Jula nests (Fig. _46) and I ~ the plasterings of paper that are sometime laid dOlvn on the fac~ 0 1, very ?ld abandoned combs in tl,. topmost recesses of large nests . (FIg. 2J . fhese S metur , appear to be built up gradually as a nes,t lD<"~es in siz . and w ·~hl. Tbe component pulp is very finely chewed. 1> added 0. bnle at a time. and often con tain s much sativarv malena l as to develop a di stinct gloss. .

ertain specifi c differences between the paper · made by various wasps have b<eo pomted out in the pr ceding [lages (pp. 123 and 124) . To these It rnay be added that the. paper made by Vrspulll ntl/>lturoa (Fig. 237), at leas tha III the walls of tlle three nests examined by the writer, though made. of weather-worn but not rotten fibers. averages slightly thilmer, contaIRs a la rge.r number of min ute inter ·tices, and has a somewhat fra iler appearance than any similar paper examined. It SlIggests a fabric ap­p~oaching the thre:r.dbar stage. Other specific differences in nest paper ' \111 undoubtedly b<come apparent as the biology of wasp is more

orough ly investigated .

BtJTLDINC THE . EST

Bosie !rBlur.s.-lt bas already beeu pointed out (pp. 11 and L) !hat ;reo? of /)ol,,"o'Vn/>u/o characteristically build in aerial ituations (Figs. _15, _17, 21 8, 219, 22 1 al\d those of Vu/",Ia in underground s ituations Fig. 226, ~~30j. Thus the charader of the nest site has gerleric (or ubgener~ cJ Slgmncaoce. T hat the distinction between aerial and ubter­

rancan SItes IS not always ftnely drawn. howev. r. is apparent from the follOWing exceptIonal examples. On J uly 22. 1922, a .ntllU nest of Ves/>u/a " tlgt1YlS was found 011 Mou nt Hood. Oregon. suspended from a small !')g, olle end of '~hich was buried and lhe other of which projected from the slope at a shgbt angle. This nest \ s protected on the ,ides hy an

( 12.51

126 NORTIJ AWERICAS' VJ::6PlXE WASps

accumulation of leavrs lodged again.t the log. Except for the shelter pro­"ided by the lodged lea.es, thi nest would have been typically aerial in poition. In late spring of the present year ( 1937) near Hoberg., Lake County, California, a small queen lIest (Figs. 197- 199) wa fouod hang­ing from a roof timber in a dark shed. Th · nest contained a dead queen VcspuJa pellSyl''''nlca. On July 6, 1922, on Mount Hood, OregoD, a nest of DolichO'llesp.M. tlrclUlrin was found behind a slab of loose bark on a fallen PseudotsuYG log. And on J uly 3. 1923, a Dest of the same sp<:cies was found built among coar e talus rock in tlie lumbia River ,,"yon in regon. T his n t had been slarted on a tnlly aerial support, but the .upport was so low that the Dest had been exl nded il\to th spaces bell' ecn the rocks below and had thus become more subterranean than otherwise. Rau ( 106) reports a similarly anomalous case in which "Vespa g~nlW"ica "

(almost certainly Vupulll lIIucuJifrolls) nested in a doS<:<! and dark tool cloSC!t at Kirkwood. lissouri . Bischoff (18) figures , nest o f T'rsfula t~dgaris that was built in a rome or shed and anached to a large beam."

uch nests are undoubtedly xc.ptioual and do not dcstroy the vnlidit)· f the general statement. tatements lhat have been made by variou_

obSt!f\'ers and uy authors of general texts t the effect that "VCSPIJ (I'u­~lIla) {}cnnanicu" (which does not even occur in th is country) nes either in tre or below grouod, bave been based on mi idenljfications. 80th DolichovtS"/'lItll tlrC'Ulria and V,sp"ta /I1aculifrlJlfs have been rq>Catedly c nfounded with T·. gcrma"ica. T he same is apparently true also of V . vtligaris.

The n lIDg sites of the various species of V u/,o are not so well known. V ujla crabro nests in hollow trees, in the alties of buildings, and apparenlly, ooc,,-, ionally ( aussure, 119), among the branches of a tree in the op<:n. or (Bischoff. 18) in th ground. Cory (33) describes a ne t of V . crabro coll ected at mtervi1le. lIeen nnc's oWlty. }lary­land. lhat was buill inside a twelv -inch-square fence post which con­tained cavity. Thus V. crab,a i more variable in its selection of ne.'t­ing sites than wasps of the llier two American genera, though an urc (119) sa)"s it prefers hollOW trees.

Sfiutill!J lhc nesl,ng si'a.-:'>lany writers on wasp biology, especially the earlie.r writers and those whose works are of the called popular cl:15s, have credited wasps wilh tbe exercise of extraordinary intelligence and foresight in the selection of nestillg sites. The old book, Naill',' Displav'd (93) . says of underground ne>ts, that they are built "always in risi~g ground, lhat the water may How dO"~l below their place or rci-

• The author', copy of Biscboff"a book was A«ured too 1&l( to ~ UJed thoroughly. tnadt!quatc cita tion of this imporunl work, therefore, though rf:gnttable. i5 un· avoidable.

r 1261

BIOLOCV OF THE VESJ'J~A£ 12i

dencc. and nnt be incommod ious to them." aussure. an exce.lIent ento-1I1ol<ogist. who laid the foundations of Our modern knowledge of the \. pldae and related Hymenoptera, ""),5 (119), "Les l.'Uepes fit ails I I. precaution de choi ir pour leu r etabrissement d.,. arbes touffus tel~ 'Iue de& ""pins, t.lODt Ie branches sont un abri de premier aloi; . .'. ." liut he realizes thm the j udgment of th wasps is not always sound. (or he ontinue : '"mais j"en ai vu qui, plus maladroites. ··.;ta.ient etablies au sommet d 'un poirier nain, cOmm. si e1les s'etaient plu :i defier les orages." 1{'lll (107), a gmerally ~ceJ l ·nt observer, says the nests of "I' cspu ( [J or;c"O'IIupflla) "Ul<ldalo" are I;enerally built out of r~ch. Coup<:r (34)

} S that "V"SP" (Dolicill)1.·esp .. la) macilla/a" locate thei r nests ne.lr human hahi tations, "no doubt to be close to localities wbere Ihey enD procure four! to rear their young ' juickly.' · The reference plainly implies tnat wi" re man is, ther. also will be /tic., and in abundance! .

Lttensive observations. on the other hand, have con,inced the writer that the !iIlpposed j udgmem of wasps is a myth based on the chance "I",:r 'arion of a few lavorably located nests. Poorly local nests are too nllmer us for the situation to be otherwise, a a few examples will demon tratc.

T h" matter of concealm""t would seem an important consideration in Ihe building 0; so large a Slructure as the nest of a vc pine wasp ; yet, a, ('\'eryone knows, many aerial nests are built in ,,-"posed locations and are 1101 onccaled at all. Moreover. they often uffer ill ·Ollsequ~nce. The r,.,,,· ,.,led nests. 0(1 tlie contrary, are rarely di~co\'ere<l or destroyed_ : ,'S! . of II ~sp"la spp .. bein!; subterranean, automatically enjoy the bene­tIl nt ronccalment, and the concealment is so effect,,'e tha l few laymm e\'en know .there are such things as wasps which n t underground, I\"hcrcn.s aenal n.,..u, at least) through the medinm of the comic strips 10 . our newspapers if not through other channels, are very fanliliar olJJects.

Protection from weath r aJ 0 hould be important. yet it is apparently " 'It 111 the least taken aCCoLln t of by ""liSps. gain Ihe ne Is of I'csl'ula

1>[1 .. b . alise of location . enjoy a large measure of protecti n, hieOy f r',10 "-IOd and from temperature changes, less [rom rain. The aerial fie", of DoIichO'llup,ull SPI). present the extreme of contras in this rr peet . Some, because placed beneath bridges, the eaves oi buildings, ~1 11f1('r large tree limbs, or ill attics, barns, and caves, secure the utmost 10 • Oth protectlon_ erg ar suspended frol11 pendulous branches which ar" readily swayed bJ' the wind and are thus SUbjected to COn iderable hulT ting. Many are exposed to U,e full fore · o f Summer rains or are " :en [/Uilt down among grass and weeds where they may become sodden \';Ith . t D I' I . motS ure. 0 U' 1000rspuia paper IS, to be Su r~. more resistant to water

[ IZ7 I

12 N'OR-rG AMERlCAN Vt5.Pl!"l"E \ ASPS

than that made by V uf"'(o. a fact that is doubtless correlated with the gene.ral diifer<!llc' between lhe nesti ng' sites chosen; but many 00/,,110-fllSpuJa o~sts are undoubtedly brought to "" untimely end by adverse weather condition .

It would seem that ample room f r the enlargement of a nest would be a sine qua non in the choice of a nest sile, but there is no indication that such i the <"OSe. T he two n ts o[ DO/;c/ICr.Jupul4 orenaria noted above (p. 126) as ha ving been buil t, respectively, ""neath " slab of loose bark On a fallen Puwdo/s"ga log and among tal us rock, were both so localed that normal enlargement w impossib le., \ nest of the same sped found on the Nevada shore of Lake T ahoe was built in a man­zanita bush about two and a hall f~t from the ground whe,. the rooked . closely placed branches a f the bush constantly presented obstacles to growth and required modification o[ buildi ng procedure. Still .nother, the one shown in Figure 221 , fmmd in A lum Rock P ark noar an Jose .. Cali fo rnia. was tacted SO close to tbe gTound th, t its downward enlarge­!nen t was definilely stoppe<! as oon as the ground was reached and Ihereafter only lateral exp.1llsion was possible. Davis (43) ropons an identical ituation in the Ca$ of a n •• t of "Va.rpa (D olichovespu/a) macu/aJa ." [t might be pr umed that the wasp would lig out the earth below and make room for the nest gTowtb . but the species of the genus Dolicl!ovcsp"la appear to lack the instinct t excavate which is so well developed in the gCllus Vespula . Fyles (65) reports a nest o f " Vo.rpa diabolica" ( = DolichUIJespu/u IlJ'lmmi<J) built in a hollow cap of a newel post on a ca irway. Between the cap and the top of the post was a small space in which the in sects worked T here was room fo r but one comb of cells.

Various writers during the pa t two hundred years have expressed the IJcliC[ that coloni"" established in unsuitable sites eventually abandon lheir nests and build elsewhere. Rau (lO7) in a recent paper infers this to be the case with Do/;,hUIJupula manila/a. T ilis bclief, wh ich appeals to the writer to be utlerl naive, in view of its inconsi teney with Lhe known (ads o( wasp behav ior, ha arisen out of the occasional finding of deserted incomplete nc ·ts. A much more logical e.'<planation of such deserted nests is t11.,t the queen has nlel a tragic deaUI and the colony has died Ollt

in conSt!quence. This explanation was recognized as probable by Sausstlre (119) and admitted as possible by Rau (107) . ctually there is 1I0t a shred o f positive evidence. so £af as the present \,rriter is aware, that vespine wasps eve r voluntarily abandon aO established nest. Whenever abandon­ment is forced as a ro ul t of the destruction of their ne IS by man or some predalory mammal. the wasps rebuild. or attempt LO do so, at the same site. ~{oreover (see pp. 153, 157, 158), the reuuilt lIests appear

I t28 J

o(or .. O(lY OF TIJt VEsr l XAE 129

alway. to be abnormal. eXhibiting many irregularities in struclure, and ar ' often decided monstro itics. Janel (75) figures a nest that was be­ll llll by a queen "V upa SaxOIl;Ca!' on a rolled canvas cunain. Shortly ,he"'" i ter, when the liTOl envelope of the nest had been completed and a sccond barely "tarted. som.,gne pulled the curtain down just <!Ilough to ro­tate the roUer on~ lurn . T he indpiCllt nest was carried rOWld the roller to n IIe w position in which it projected horizontally outward from the curtain . The queen tber<upoll abandoned the nest and built a new one beneath the rollcr in the position originally chosen . T1Us instance., thougb a case of Itelluine abandonment, is hardly the kind to support the hypothesi that poorly chosen nest .ites aro abanrloned . Such parallelism as does exist, in ial't , sel' ves to strengthen the opposite view.

; \ mple rOOm for the traffic of foraging wasps to and f rom a nest is <'.SS<:nti ai to normal developn,ellt o f a COlony. Yet a colony of VesPlt/a "if /pili/rea at Cupertino, California, lVas <:stahli.lIed in the earth behind a twn-iuch concrete retaining ":ill in a place which was accessible only thro ll 'II 3 uarrow and tO rtuous passage in the c<lllcrete, " flaw resulting from the imprisonment of ai r in the concrete mix wben lhe wall was built. T here wa barely rOOm enough in the passage for three workers a, a time. \ orkers at intervals attempted to enlarge Ihe pa sage,voy, but w'tllout success . The nest \,; a' dug our on October 5, 1930. Tt was quite >lm U, in spite of the lateness of the season. The total POI)uiatioll was one hundred and seventy-three wasps, of wbich one was tbe '1u~'CJl mother who bad f a unded the u<:st, six were new queens, and one hundred and thirty-fou r were workers. The nest contained a few male pupae, but ap­pareurly no males had yet attnined maturity_

An ideal n' t site hauld afford protection from marauding rruunmaIs. Yet aerial nests, which would afford complete protection if btl;lt high enough from the gTound, are olten placed so low as to be within easy reach of any mammal ro wh 111 wa p brood is attracti ve. The nest of lJu/ichQlIGs/>u/11 ""B'IlJrn., twice referred to (pp. 126 and 128) as having b<:cl1 buil t at the surface of talu rock, had been tom open when found, an" a large part of the contained brood bad been eaten. Even Dolichuvu­pulu lIlac.lata, which perhaps more often than most species nests high in Irers, occa, ionaJl y errs in Lhe choice of a nest site. Rau (107) records nne L1w was placed bu t thr~ feet from t e gTound in an elm shrub. Sub­terranean nests. which would be we.1I protected i f loc:lted a foot or more I"'neath Ihe surface. are frequ""tly so placed as hardly to "" covered at all. I lnc o f these found on the Stanford University campus had been ripped Ol>en and tho I' ugh ly ra wged by a skunk.

J ust what it is, then, that caus a queen wasp LO select a particular , ile is not clear. The only actual ~uirement seems to be that the site

[ t29 I

130 NORTH Ali4£RICAS V -rf~'E WASPS

provide" suitable surface {rom which the nest may be SU5~ded. Such a sudacc may be anything whicb , ill give overhead support. The nature of such support in the case of Dolrel. " rspula nestS is well known . In the case of V up .. la Ilests built below groUD I, a root or pebble projecting into the eavity cho CD the site usu lIy prm ides the sudace ior attach­

ment of the ncst ( 'igs. 199 and 2(0). TIo_ .. 1] ...... ''''sl.''~n a site bas been selected the queen gathers

paper pulp and begins work. The first rtructure built i generally a p"diC('1 or suspensorium for tbe support of a small comb f cells and irom Olle to four envelopes. \Vhether the e><1lct spot chosen [or the at­tachment of the suspensorium always receives prcliminary t"eannont or not is unknown to th ' writer, though some such trealment would seem neC(' 'sary to iosure tIkIt the paper pulp wi ll adhere. Janet (7" ), in de­scribing the rt!building, by the European Dolirllq; ·csp .. la IIIcd.a, of a nest that bad be"n destroyed, says the wa. ps cooted so.ne five centimeters of a twi ' wilh a glutinous substance. The fi rst few loads 01 paper PuJI' may all be applied at one . pot. in wbich case the pedicel pid!y takes shape. n the other hand, in Do/ichovespu/a nests, at least, the hr.t few loads may be applied to a number of distinct spOt scatiered about within a few centimeters of one another, as if the wasp were unable to choose between tI,em. Eventually one of these is enlarged to become the suspen­sorium and the others receive no \urtbt!r attention.

The base of the uspcnsorium in all the nests examined by the writer (Figs. 199. 200, 202, 214) is a thin. hlade-like structure iastened by one edge to the ,upport. A horl distance below the point o f attachment tbe su pensorium narrows and becomes more or les ' cord-likt!. It I.h"n expands gradually into the first cell of the nest. Thi is true not only of the initial uspensorium of the n~t wh ich i buil t by lhe queen and which upport the first comb but a lso of those of all the combs built subse­

quently by the , orker.. The fi rst comb. then, is tied se lI rely to the nest support by means of the suspensorium. and not until this has been accomplished a re paper envt!lope built around tile omb.

Variations in the mode of beginning a nest may exist, though more observation i needed to establish r disprove this point. I n mid-Aull"'t, 1929, a t Cupertino, California. the wriler collected a nest of Do/icioovesr·lla

art .. Gru., the structurt! of which suggested that one or more of the en­, elopes may have been started first and that a portion of ne of tI,ese may have been continued into the sus~sorium supporting tht! fir t comb.

ouper (34) holds that Do/ichOf)espula ,noculala constructs first an ~n­velope and say, "After it is perfected. she attaches a single tier oi cells ,. Saus.ure (119) makes" similar statement. referring especially to nest, of Do/iclllw.srllio ", .. dia, but implying that other wasps build in a like

[ 1.10 J

BIOLOGY OF TIrE VESPI NAE 131

nmnne r. pecifically be stat.,. thal a disk or skullcap of a cellular nature i lIIade fin;!. The initial comb is tben attached beneatll this at a number o i points. Janet (75), on the other band. 6gures n.... of D. ",.dia that w ' re begun in exactly the mann"r described in the I'recL'<ling paragraph. Couper, judging from other data, maniJestly was in the habn of incorpo­rat in T h is preconceptions in reports of his observations. Ilence ont! may wel l 'I"estion if he ever enrdully scru tinized y ung nests to determine tlu ir ctual structure. And Saussure, though generally highly accurate, "PI~'~ rs te) bave based hi slateJ11C11t.!, on obser\'auoll.S of large n ts only. It •. doubtful, therefore, if ti,e mode of nest origination reported by tl,ese

111t'T1 . actually occurs. The lirst cell is circular in ero section . and is longer, straighter, and

l11(1re gradually tapered lhan any of the later ones ( i-igs. 89, 90). It is huilt tu " consideraL\c length be.fore any others are tat ted. The base of till -l' lid cell i laid down i11 tl,e form of a thin crescent of paper on thr ide o f the first and SOme distance from tbe base of the laner (Fig. ~" ). A dditions are made to the edge of tbe crescent, the n~wly added >tnps of paper being arched SO as to enclose a lumen which is at first " :llli,-ircuJar in cross cti n but which , as i l is built up and surrounded f, ,, IIth" r cells, gradually assumes the typical bexagonal form. The lbird cdl i tarted in much the same manuer. II may be begun closer to the • a.-e .. f .he first cell tb.m was the second . or farther away, though the latter is probably u ualJy the case. In any event, the third comes to occupy 011 ' oi lbe angles betw~ the first and the second. sa l" ult the ross ""ction 0; its base is s ruewb.at less than a ·emicirele. T his i true of all the cdl ' subseql1ently built in the nonnal tn.'UlIler. Each is added in the am;lc I", W!!en the walls of two already exi ling cells. T he queen unaided iluild. but a ;mall Dumber of cells : Buckhurst. StaniJand. "nd Wa[<nn (21 " I >"Y about lwelv"; Ra u (107) r cords ten ; and the author has a 11,· .. t . presumed to b · that of Do/ic/ouvcsllllia macula/a, containing eighteen (h;[. 9 1) . a probably incomplete nest of Vrspula P" tlsy/t"'/tica. \\~th t\\ d yc. and another of the S;ln'e pecic - wi th at least nineteen .

~hortly " fter the initial uspensorium is completed and the first cclls ar,· 10< '~n, the queen begins to huild a sph~rical o r oval envelope, the first '·' "DIl<". ng the nest wall. As therc seems to be a difference in this praced"re I ,·tw, :n wasps of the two genern DolichOV~Sf'ula and ~ tS/,uln. the method '·.npl"~l'd by Do id,owspllia spp. , ill be described first and that followed. loy I ,sl'lI/a spp. prt!sented afterward. T he envelope is beglm at ar fl ear the l>;Ie f the suspensoriwn. Th paper composing it is Wd down as a con­cave di'k with lhe IIs~sorium pas ing through its center. Additions are n ,,, ~ to the disk unbl it i, tmnsformed into all inverted. bowl-like stTUC­tllr . During further additions the em-elope is narrOw d unti l it becomes

I 1Jl1

r ughly sph"ricnl or ~gg-!hapcd and c mplctcly cndo es the comb of cells, except [or a hole left 3t the bottom. T hrough this hole the qu~ wters and lea • Space is left between tbe comb and the em·elope so that tbe queen may circulate f reely inside e nest.

J\ second ,,"velope; now Sill ned a short distance above the Ilase of the first Or a lillie farthe r from the b~ o f the suspensorium. depending on whether the suspensorium is attached to a .I,,"der object such as a twig or to a broad urface. This envelope i con tructed ill the >arne nw mer as the firs t and is brought downward at 3. relntively constant distance out­side the first. The same i tr ue of the third and th fou rth, whenever these are built. It is doubtful i i a queen wasp ever builds more than four en­velopes : oiten she does not complete three. TIle distance between envelopes, as well as that between c.onsecutive comb, and bet\V~ the: innennost en­velope and the « IIllbs in Ider n""ts, is roughly correlated vith the .izc of the \ p., body. It , therefore, "an solllewhat with the pt.'C.ies.

In nl least one species, Dolid,of"Bsf'ula maclliala, the qu""" frequcntly builds ne addi ti orol structure. ne o i the lir t two or three envelopes i. ""tended below to form a tubular beak , bout one-half inch in diameter and f rom two and Olle-h •• 1[ inches to fOllr and one-hall incbes in lengt.h. This beak has been noted by vanolls wri ters. It bas been figured by Rau (107) . by mith ( 124 ) , and io the issues of the C uidr tlf NaJurl ior June 19 17, May 1918, nod September 1918. T he ne>t figu red in the September 1918 issue of the Guide of Na/tl re had a body two inch in length and a beak slightly over four and one-half inches. The nest came from Ohio. P rcswuably no addition are made to t.he beak by the \\" rkers when they assume the duties of nest construction, and as the oest is enlarged a Ii lt.1e later, Ihe beak is tom down.

Quem W35PS i the genus V ts/",Ia em to begin the nesl envelopes exactly as do those of Dolicl.l1IIupula, but di fferences soon becom. vident. T he second cnvel pe Illay be staned before the fi r-t. is completed, and simi larl), ior Ia t r envelopes. T he writer collected a small nest of Vespul(J pensylv"";c,, (Fi '5. 200-2(2 ) ill the process of being conSlructed some four ind,cs below ground in an old gopher burrow in Lake ounty, California, which possesses IwO envelopes. The fi rst extend. downwa rd but three­eighths of an inch on one ide ano not ver 6v,,·eighths of an inch 00 the. other. The econd is much larger, being a (ull inch in extent along ils curved sur face f rom the suspensorium to its iree. margin . The two en· velopes are considerably closer together, also. than the corre ponrung envelopes of Dolichovesp .. la queen nests. Rau 104) records simi lar fea­tures in a queen nest of "Vupa gcnnanica" (V,splfla lHan.lifrt>,LSf) . say­ing. "The nest was small and had four layers covering ovor the top and p;!rtly overlappinl( the sides." Nixon (95) , in England (1), describes a

[ 132 J

BiOLOGY OF TUE \ "ESPl SAE I I I

queen nest of "l 'c~pa (Vupulo) ~'ulgarir' collected fr m n "wouse dwell­ing " in n ra ilway bank as having nearly the size of n golf W I and "one complete. outer covering and one which was hal f completed." ' The 3uUlOr has a qu~en nesl of Vespu'" fJ1Jgaris (? found within n bollow stump in ' equoia Natiorol Pa rk in 1933, which has a sowl paper rusk above the comL and, in addition, two envelopes OIl one ide and three opposite (Figs. 2 13. 21 4) . A · the nest had been abandoned when found, identification was ha&ed solely on Ihe character of the nest paper. Another nest oi V. pc'~'I­t"(lII ica (Pigs. 197- 199) rrom Lake aUDry, ifomia, bas three envelopes, the fi rst two oi which are complete.

onsiderable variation i shown in the mode of attachment of the second aud In.ter nvelopes in Vup .. la queen nes . In the nest of V. pcn· sytmnica just mentioned, the thi rd envelope is attached, not to the suspen­, unum, but to the second envelope in a circle a full half-incl' outward from the suspensori"m. Thus it does not extend over the t.op f the nest at nil. Mor vcr, it does not completely enci rcle tbe nest . thougb most of it is buil t outward a qnarte.t of an inch f rom the line of . raning.

W; ps or the genus Vespa, in a far as generalizations can validly be ba<ed on puhlished descriptions ( the a tivities of V tspa crabro, vary their ne. t-lmilding behavior according to the nature. of the nest si te. \ Vben nest.s are I.l.1 il1 in the open, or in ,~ti"" in soft earth. envelopes are built a.round the combs es.'eotially as wi th nests o f Do/ichov"spula. Nests built in hollow tree. often ar" provided with DO envelop at all, exc~Pt perhap a few lu,li ments above tbe fi rst omb aud one or more sheets of paper built SO a panl ' to clo tbe hole in the tree which provides access to the hollow interi or. ubterranean nests lack envelopes only when built in very hard awl , touy ground in which Ih stones prevent normal enlargement oi the

From the time the care oi th,' fir brood begin,; to make riou de­Ill' <Is [I the ti me of the quecu until Ihe brood matur and the resultant wurkers take up thd r duties. the nest struelure remains essentially static, al lea. t in outward appearancc. Some work is done all lhe comb, for the cclls are not built to their fell l length at first . Eggs are laid in them wben thl'Y are quite small, and the ceUs are then extended pa ri passu with the growth of the larvae. Hut from Ule. time the fi rst workers appear until the colony declines at thc end of the season the nest undergoes a continuous ,ue -sia n o f changes_

.\11 old belie f, e..'<pressed by L ch. according to W .. twood ( 157). and loy Couper 34), held thaI when e fi t rood f workers emerged they ,icled the queen in building an entirely Dew and much larger nest. In this a <-"<:ond no r! mo re numerous brood was reared, after which a third and . till la rger nest was built. Presumably the same sequence of e,'enls COD-

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134 SO'R1'1t .O\lf£RJCAN VESPL'lS WASPS

tinued throughout the season , each nest being conceived as complete aod ioc:apable of being modified. T he belief i eDtirdy erroneoll.'! , there bei ng no e"ideate whatsOever that sucb a performance ever occurs.

Modificatiotl of Ihe nesl'ng sile.-As a general rule a nest site requires more or less modification to adapt it to the needs of a growing colony. Vespa uabl D, according to Saussure (119), Cory (33), Janet (is), West­wood (157) , and others. when nesting in bollow tr. or timbers, go"w. the wa lls of the cavities to smooth them and to enlarge them. The wood thu­gnawed off either is used in the. enlargement o( tbe combs or is dropped to

lhe bottoms 01 the cavities. The species of Dolicho".s/lula, according to the author's obse rvations,

do very Iitde work upon the nest site. ery often nOlle is necessary; bur when nests are built in dense foliage Or close t the ground, the clearing away of leaves and twigs. and f grass and weed stems, respectivdy, would be o( decided advantage. Yet this seems never to be appreciated. n,e wasps, instead. incorporate all obstructions in the nest wall (Figs. 215, 216. 219. 221) and then later. as the nest i· enlarged, chew them away from the ill.<ide, bit by hit The author', observations b.lve concerned colonies of D. aunaria and D. matula/a, chic:By lhe former. Davis 43) shows a photo­graph of a D. ",a"u/ala nest which has lea,'es and a mall branch included in its nest wall. Janet 75), in a drawing, illustrates the same thing in a ","Sl of the European D. ,,,.diG.

Spec.ies f V slm/a, as would be expeeted in view of their SlIuterranean habits, do a great deal o f work on their nest ~i t , They regularly clear away leav ,grass, and olher litter over a small ure.~ surroundi"g the co" traDe' to their hurrolV (Fig. 235) . 1\ colony of Vtsp"la vlIlgaris, found durmg the summer of 1930 at Stanford University, prcvided a .lliking example. The nest was located in a grassy field. During the preceding spring Lhe grass had grown luxuriantly. and with the advance of SW1101er had become matted down into a dense layer about eight inches thick. Around the nest entrance the wasps had CUt the gr= away as to (om, a completdy cleared wcll-like approach aboul four inches in diameter through the mat a f grass.

V <spu/a spp. also d a greal deal of excavating to enlarge the cavity in which the nest is buill This is done ordinarily at a rate approximating that of the gro\\1h of the nest itsel£, so that there is alway n space between the wall of the nest and that of the cavity (Figs. 228. 229) . The clearance ranges from about tbsee-e.ighths of an inch to a litue over an inch, and averages slightly over a hall inch. Since r' os/,,,Ia nests are apt to be quite large, often attaining a diameter exceeding a foot, the amount of excavating done is considerable.

Because l obvious difficulties, the actual .. '<cavntion of earth bas ool

[ 1341

810LU CY "'" TUP. VESpL:' A£ 135

yet been observed by the writer. nor have direct observations been recorded loy ou,ers so far as he i. aware. Nevertheless, the l1L~nner in which the wurk i done can be in ferred with fair accuracy £rom numerous indirect indications, some of wh.ich have loog been known, and all of which may easily be observed. The ioterior of a nest cavity has the appearance o{ a tiri<!<l mud surface. The same is true of the passageway leading t the ca"it)'. Wasp may be captured as they leave the nest carrying r>cllelS of earth that either are moist or give evidence. by their form of having been >0 re"...uy, These things are true whether the ne t i located in " humid or an arid region, in naturally moist or in dry soil. In the active season, also. worker was s are abundant at sources of water. That they are not always .,impl)' drinking to satisfy their own needs may be easily determined. F~r xample, on July? , 193O. at Stanford University. the writer observed n

r".'tura s1llpJlUrra worker which returrled al intervals of abou t five minutes In a p.~rt.i cular spol at the edge of :' small lily pond. Each time it drank deeply before it fl~ away. It was stiU coming for watef when the obser­vation was ended. Wasps of other species were also observed at the pond. ~Inn}' of them drank so much water ~, to "'-perience difficulty in taking fli/:ht. From such observations. therefore, it appears certain that water is

r ri ed to the Dest CIlvity and regurgitated upon the wall o f the cavity. The fI"ult-.J11t mud is then scraped off in rcllOdish peUets, eitber by the same or by a different wasp, and then arried away.

The earth reruO\'ed in excavating is variously disposed of. Much of it i, carried utside the hurrow and dropped '1 a dislaJ lce. A worker [ , ·t'sp.tia P"!lsyiv(l],ica ob 'en'ed by the writer carried a pellet at least seventy ice I [rom the nest ilefore dsopping it. he may have carried it much [an Iter, but as he had attained an levation of about twenty-five feet in lhat dista nce it became impossible to determine the final disposition of the I'"llet. The same mode o( d isposal of excavated material, was long ago ohserved by Westwood (157) .

SOt all of the mud raped off in excavation is always UlllS carried """R.,. ltal1tities of ule mud pellets ,Ire often uoed to fill cavities leading fJUl through tbe soil {rom the nest chamber (Fig. 232). Such cavities are (lmmonly branches of the rodent burro\\' or c.revice in which the n""t was

,tarted. ttasionally they arc cavities left by the decay of a large root . In un. nest i / 'espuia phlJ"ylvall;ca built between two large rools o f a dead pine, the author took three quarts of dried pellets from a cavity left by a decayed root.

The rrcatll1col accorded large ob taeles encountered in the soil varies with the natu re of the obstacle and lVith its ,i.I!. Pebbles too large to be earned to a d.istance, but ~mall enough to be moved, are dragged to the mouth of the burrow and left ncar by. 'Vben the nest entran<e is located

[ IJS I

136 N ORTH A &'1ER..lc.AN VE.5PISE WASt"$

in sloping ground. pebbles are sometimes brought just to the entrance. wbere they are dropped and al lowed to roll down the slope below (pig. 22J) . Pebbles and stones too large io be mov~1 :If. left in place or are dropped t the boltom of the nest cavity by the removal of the earth beneath them. mal l roots are cut off bit by bit as the cavity is enlarged and thei r substance is either carried away or converted into paper pulp. When large roots or stones are encountered, the nest is simply buil t around them. the paper of the nest waIJ being- attached to them so as to incorporate them into the nest s ructure (Fig. 226). T he combs "~lh in , on the other hand . are but I'lI.J'l!ly attached to roots or stones. A spacious passageway is usu ­ally ldt around such included Objects, thus giving the wasps f ree acees ' to all parts of the oomb.

lr. occasionally happons thal the passagewa), leading to a nest "'I\'it)' is too large to be sati'(""tory. as when the deserted burrow of a ground squirrel has been selected as the nest sit,,- I n such case ', mud removed f rom U,e nf t cavily may be packed into the sides of the passageway so as to reduce i L~ lumen . uch packing is PUt in raUler loosely, nwnerous spaces being left between the conlIlonent pelle (Fig. 23+) . 1n one case hserved Gy lIle writer, a gigantic n .[ of V u/,ula 'II'Illgaris had been buill in a rtteS beneath some timbe supporting the edge of " roadway (Fig. 255) . The r/!Cess opened out widely beneath the road , and along this open ing the wasps had built a large earthen buiwark two or three inches thick and several inches high to encl ose the n l (Fig. 233). The fragment of p.~cking shown in Figure 23+ was taken from this bulwark. The expo!>ed urfaces of such mud packings. wlmtcver their (unctioD, are covered over with mud

[ " li ttle softer consistency so that no dll [ or loose granular material remains. T he lIItim.~te sur face suggests that of n rougb plaster_

Ordinary runwa)'s, too. even though no chMge in dimensions is neces­sary. have theil' suriaces plastered in the ame manner (Figs. 226. 228-230) . T hat tile nlud used in me plastering u brought from the nest cavity.

r may be so brought, and is nol merely me .. worked surlace of the pas­sageway itsel i, was clearly illustrated by a nest of V"pwa vulgaris col­lected at Cupertino, CaJifornin . in October 1928 (Fig. 226). The surlace soil, through which tile passageway to the nest ca"ity ran . was a daTk gray friable loam. T he soil beneath . in which mo t of the nest ca"ily had been excavated. was a ligbt brown adob,,- T he adobe had supplied the material for plastering the passageway.

lle other use i~ made f the .,,'lttavaled mud. A &mall mound is often constm cled at the entmnce 10 the nesl burrow. In the making of th.,,;. mounds, and perhaps also in the plastering of the runways ,,;thin, til. mud is applied in little vermiform masses (Figs. 223-225) . A few refer­ences to such mounds iD the Iitemture interpret them as being designed to

[ 136 J

ntOLOGY O F TIlJ::: VJ!.SPLS/l.J! 137

keep rain out of the nests, bUl thi $eems unlikely. T hey are rarely high eDough to be o f much value, many nests have no trnce of a mouDd. and nests wi th compa,rutively large mounds are often buil t by wasps of reL, ­tively arid regions in which the probability of a nest being flooded i extremely remote.

An occasional underground n tis prov ided with two openings leading to the nest chamber, though not, 0.$ some have. supposed, with one for enrrnnce and one for ,,-"'t. hcn two entrances exi t . wasps tnter and lea " 0 indiseri minately through both.

E,rlargtmumt of t hD cotllbs.-Throughout Dearly lhe entire active period of istence of a wasp colony the comb are enlarged by the addition of ceJls to thei r margins. and new combs arc bui lt below U,e ones alrea.dy. in exi ence. Not until the worker population dwindles ~th the onset of U,e col uny' decline does this work of enlargemenl cease, except in the upper-0Iru.'t combs. These apparently become too far removed from lhe nest en­tr:lllce (or addition to mem to be practical. Moreover, Uley eventually b" conlC unsuitable fo r brood rearing-. Addition of cells to these combs, tIl.rdore, ems some week, before additions to (be lower combs cease hei ng made .

• ,,\I' cells, except the Iinrt three or fou r that are begull on the periphery of the initial cell of a comb, are always started in the angle between lWO

pre-existi ng cells (Figs. 90, 91) . The paper strips of which t.heir liases are COlli posed are laid down at rigbt angles to the radii or growth axe of tbe comb (Fig. 248) . T he wall o( the new cell is crescentic in omline. the he..xagollal ,hape of the cell nol being attaineti until the cell acquires some heigbt and newell. are built beyond it. Marginal cells. in [act, rarely ha 'e '" compl tely hexagonal shape (Fig 199. 201) .

New cells are not built to thei r fun height immediately, nor is their '''~ (Id "yed unti l they have been completed. n the contrary. me queen lays an egg in each new cell shortly after i inception. often when it is lit lIe mOre than a crescent of paper on the edge of a comb. The height of tho, cell is then increased pari p u with the growth of tile larva within.

When paper i' .dded to a cell to i ncr~ its height. it i put on in little .trips which run around the cell transversely (Fig. 212) . W illie the cells are <Juite small each trip is limited to the free portion of the wall of the new cell. u.ter. a fter the cells iD " ne\ cell row ha\' e grown until tbeir ase. touch, a single paper strip may enler into me conlpositioD of tbe

wullll of two or more cells. nd in a group of cell. of the same height whi ch are being enlarged simultaneously, Dew paper is added to the walls wil l!J'e"er needed, with no attempt to maintain the integrity of individual cclJ walls. Thus it is that the walls. rather than the cells per St., are bui lt lip. The rull is an interlacing of the ronlponetlt paper strip. which

[ 13' 1

138 SORTD AME:IUCAS VESPlNE. WASPS

greatly increases the strength of the combs, as has heen pointed ut by Janet (75) .

W ith hexagonal cells as the units of comb construction it might be sup­poliW that the combs as a whole would assume a bexagonal shape. This shape is sometimes approximated (Fig. 210), but more often the combs an nearly circular (Fig . 252), for the fOllowing reason: Tn nests toot are developing rapidly and symmetrically, new cells are commonly .dded along the sides of the hexagon formed by the central ponion of a comb before they are added at the angles (Fig. 2\0). Th i. [acilitates the economic utilization of pace and materials in nest building. It would not be justi­fied , however. to assume that tbe wasps sense the economy which rtSults from their mode of enlarging the combs. A simpler explanation is that thei r comb building is regulnLed, more or less, by the distance between the comb margio and the n""t wall. In growing nests of any size a distance is maintained between the combs and the ,\'311 which is just a linle larger than the minimum requirerl [or the easy passagc of the wasps . When this distance is reduced to a miuimum all around :l comb the wasp. lear alVay a layer of paper from th~ nest wnll They then build the comb outward until the wall is again approacbed, whereupon the proeess is ~peated.

W hen a oew comb is built, the main suspensorium of the Dew comb is attached directly ben",th tbe main SU5pensorium of the comb above, or nearly so. The effect of thi practi.;c, is to center the loo.d bome hy he suspensoria of the variOl!' combs and by the nest a whole, and to give the nest greater effective st=gth than would otherwise be true. But again it i. quite improbable that this [act i, appreciated by the wasps . The center of the lowest c:tlmb is doubtless chosen as the Incus {or building the sus­pensorium of a new comb because it i the place where Uler. is the greatest distance b«weeo the comb and the nest wall. This supposition is borne out y the fact tl13t io underground nests built in stony ground the initial suspensoria of new combs ar~ oftw eCCCltric in position, their location apparently being determined by the opporhmity for growth, And in .. "­trenely large nests, such, th.,t shown in Figure 255, in which the usual oval or spherical form has ~ lost and a relatively Oat wlder sudace has developed, new combs may be started simultaneously ill two Or three places On the faee of the last pre-existing comb,

The initial suspensorium of a nell' comb is usually begun at the point of meeting o f the wnlls oi three adjacent ccl!. in the . mu above. r",v 1"'1'<" is added to the illtorsectiog walls thickening and 5lrellgthening them. The new addit;ons are carried downward and sl",ped into the su pellsorium of the new comb. The new suspensorium at first is not very rigid but is capable o f conside.rable bending and twisting, as the following notes made while ous<rviog a srnnll nest of Dolic/lo" •• pu/a w6nana will .how.

[ 138]

I:IOLOGY "1'- THE VES1'lNA£ 139

"T he nest now contain. two combs of cells. The {ace 01 the lower comb 10 one and three-quarters inches from d,C base of the nest. It contains lar".e at least one-third grown in several of its central cclJs. This comb is much less ,<""rely iastened t the first than might be supposed. As tbe w-olSPS \'."lk back and iorth on the face of it, or move between the two c mh . . it rocks back and forth as much as one-eighth 01 an incll:'

Once the suspen.wriurn has been fashioned, the developmental history of ""ch new comb is a repetition 01 that of the preceding comb.

As the end of the season 01 activity approaches. the workers of a colony c ...... to build cells 01 the size previously constructed and thereafter build cell5 of a larger size (Figs. 248, 250. 251) . The large cells nre designed primarily [or tile rear ing of ne w queens, but males also are reared in them. T he smaller cells built pre '~ously serve principally for the rearing oi workers, though SOme of the latu-built ones are used for the rearing o(

males also. T hus worker wasps are always reared in small cells and qUl!ens in I"rg~ cell~, but males may be reared in either small cel ls or large Ooes.

In colonies o[ Dot;'''01Jcspu/a the change in the type of cell constructed apfll"'f> to take place somewhat gnadually. At any rate tbe boundaries bet"'l!en areas of smnll and large cells. respectively, are not always distinct in nests 01 DolicIlOfJ"puia. In colonies of / ' '/'l<la, on the other hand, the change tak .. place ;1bruptly, and the boundaries between comb areas of large and small cells. respectively, may readily be r~ogn~, even on the upper surface of the combs (Figs. 248, 250. 2~1). The bases of the large edl, are more convex than those of the small cells, and are of greater dimensions.

'orne developmental cha,nge within the colon)' doubtless provides tile , tlOUllus to which tbe cbange in cell building behavior is due, but whnt the , lUHUlllS is remains unknown. It cannot be the mere incr= in abundance of inSttt food, as is implied in the hypothesis to explain the origin 01 the weICk.r caste which \Vheeler presents in hi Soci4l Life among lite illsec/s ( I h i) , The abruptness with which the change in cell type is made and the fact that the queen reUs are larger from the moment their hases are laid clown invnlidate this suggestion. Moreover, j f the e.~lanariol1 lay in an illcrctlse of available food . all species of wasps in anyone locality would ""l:in qu -rearing at the same time, which they manifestly do not do. j)olic/ICn:rspul. ar •• aria, in c=1 California, may rfar new queen. as , arl )' as late June, whereas none of lh" species of Vespu/a inhabiting the > territory normally produce queens earlier thau September. In some n"" of V cspu/a spp. queens arc not produced unti l late October.

The wasp may attach the large queen cells 10 the edges of combs aI­readl' built, thus fonniog marginal zones of queen cells, or they may con ­struCt new combs consisting entirely of queen cells. I n most nests bolh

1 139 J

140 SORTU AMEII.ICA..~ VESP1~E WASPS

procedures are foU wed sO tImt at the end of the _son a nest will contain , in iu uppu portion, combs ronsisting of small ells only : then will follow from one to three Or four comb onlaining both small and large ceUs : below thtSe there will be olle to three combs o[ large cells only.

DeparturtS r rom the u,,-uaJ procedure occur in rare instances. Baldu f ( 7) reports a nest of "Vup"lll" (Dol~"ovespu/o) IHOL11/olo taken at · r­bana, Illinois, in 1931. in which the queen cells occurred "principally in the distal half of the s~ond comb [there were [our combs) whereas none o[ these were present on the firs t and fourth. and relatively few on the third ." Rau ( 107) repon.s a nest of the samopecies as having queen c Us scattered '·throughout two central combs." No explanation f tbis anomalous dis .. tribution of large cells e ... cist in the normal behavior of the wasps . 0 far os the present writer is aware.

T he only nest that the wriLer has actu.Uly seen in which queen cells Wert surrounded b" work r cells was the enormous n t of V rspuJa t lll/llaris shown in Fi~re 25 - f Plate LlY (see p. 159). This nest was the product of a colony which SU/'\'ived the winter of 1932-33. ughter queens to the number of twenty-two or more-all progeny of the queen mOLher which founded the nest in 1932-\\,er present and active in the nest. With the beginning of acti vity in 1933, snail cells were added by new workers to the edges of the old comb made in 1932. T hus, in ..,veral combs, worker cells formed zones surrounding queen cells which in Lurn surrotmded the worker cells that formed the centers of the combs. It i ' highly improbable tbat the nest "poned by Baldu! (7) had any sucb history as this V rsl'«io ""igan., nest, for lbe prObability o f an)' DolicirovuplI/(I colony surviving into second s~on seems very remote for an)' locality and practically nonexistl:Jll for a colony subjected to the rigors of an Tllinois winter.

When quee" cells are added as marginal zones surrounding worker cel ls the zones often are not continuous but take the [arm of a series of lob~ (Fig. 196,24 , 250). T hese lobes appear to result from the impossi­bility oi joining large-celletl comb to snall-<:dletl como without destroying the uniformity of comb pattern. Where"er the two size, of cells arc in canlllct, cells of abnormal shape are built. T hese may readily be seen in the "gur . uell cells must be built, for there is no method possible of increasing Or decreasing the size of the units of any geometric pattern with­aLit distoning them. T he abnormal cells are often quite unsui ted for u..,. The lobed combs, therefore, •• eve as a dc,~ce fo r reducing U,e unavoidable abnormalities in comb construction to a minimum.

The ratio of the number of large cdb to Lhe number of o;maIl cells pro­duced in a nest varies according to locality and seasonal condition . Beyond thi available data are insufficient at preseot for any extensive geoern.liza· tiod. To be significant, comparisollS must be made between nests at the

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BIOLOG Y OF TUB VF-SP INAf; 141

sallie 'llIgt of developmenL r at the close of the .season "fter all activity 11'llI ctaSed. Most of the. published data on wasp nest· are not reported in stich a manner as LO make this possible..

• \ t least ne condusion. ho\\' vcr, Sern\! safe, n;l111ely, thaI colonies of /)oli Irovtspula spp. build a larger proportion of Ilueeo cells, though 3

sma.llcr total number of cells, than d colonies of V I!SpuJa spp. For ex­aml'le. a. nest.of D a/ieirbIJe.rpuJa Dr.nana <ollected by the wriler at Cuper­till'l, CalIfornia, on ugust 18, 1929, after building operation. in the colony had ceased, conmmed 1,199 small cells and 1.290 large celli, distributed amOIl fi ,'c comb . A much larger nest of V.spula p.nsylvall ;ca collected at Stan ford University, California, on August 19. 1922, contained seven "OIlIb, and a total o f 10,456 cells, all of which were =11 cell •. The < lOllY \\ a3 at the height of its activity and bad at least a OIo",h longer in which 10

,1e\·c1op. :A. ill larger nest of Vupulo pert.rj'lwnica coli cted in July 1924, wht'n 1Il I ts second season, cQntain~d twelve combs. only four of which ",,"tained any large cells. These four comb, though" third of the total number. contained scarcely a fou rth of the cell . in the nest. The fact that thi_, n st was in its second season does not invalidate the present compari-5< '11: for the nest bad experienced only one period of male and queen prl)­duel ion, and no Sm.'11J cells had bettt huilt in it subsequent to the building of the queen cells. Furthermo ... its second se:u;on had been a weak sea 0 11 and tl:t· c,,1 ny had almost died out at the Lime the nest WlIS collected. This co! my had bc~ under observation since JaJllLlry 1924.

Tn tit" history of every comb, no matter whether it be made of smao I)r of I .... rge cells, a time is reached aftcr which no 1urther additions are made to it. Two distinct and unrelated mani iestatioD.'l o f this phenomenon OCcur. n Lhe One hand. after a certain size is attained by a nest , new cells ar . added only occasionally to the upper and older combs. Eventually the addlt.rons cease completely ann U,ereafter these combs remain constant in siL<o !ven thou b the colony is still ,oigorous and illcrca5ing in worker p"pu latr n. n the other band. additioru to Lhe lower, Or to all , combs in a 11<'t ce e as the colony declines toward the en of the season.

. The explanation of the latter case is simple. T he worker population ,hnlll1lsh as fewer new ones matur' to replace those that die and the needs 'II the emerging young queens and maJes re(luire an inc.reasing e.'<penditure uf energy On the pan of the remaining workers. There i~ no time left for bUilding. The fo n ner case i not easy of explaoatiofl and i probably au expr · "on of t.he interaction of severol factors. For", ample, old combs ;!'P r to become unsuited for brood rearing a.nd are therefore abandoned. . he old combs are farther and ia rther removed from the entronee, and "'coming wasps carrying paper pulp apply their burdens to more accessible <onlb . The increase in Dest weight, and of the load on Lhe upper comb .

[ 141 J

142 • ORTtt AWERJ N V.liSPIN~ WASPS

causes thu~ comb. to assume more and more the function of suspensoria. This in tum requires that ch ug .. be eliminated and a condition of stability attained .

pedfic if not generic differences apparently ist in the e.'CIwt to which th~ statements hold. In nests of several speci .. o( V upula the combs, from above do"nward, show a gradual increase in the si~e attained before a. tnlic condition develops, the upperrnost c mb being small, the next 13 cr, and so On till the middle or slightly below the middle of the nest is reached (Figs. 127. 229, 230). From here on downward the combs d=ease in actual size but are still being enlarged. In Dolk/'O'IJul'u/a nests the largest comb is generaJl ~' at, or above, the middle (Fig. 216). In nests of Dol;chov6sflu/a ",ocula/a the toplIlQl;1 comb may be the largest.

The combs of wasp nests "r. commonly described as being horizontal. One should not infer from tllis that they are also plane, tllough they ap­proximate planes. In realil)' each comb, as viewed f rom above, is begun 3< a cone (Fig. 90) , and in some cases the cone shap<! is p<!rmanently ad­hered 10 (Fig. 230). The cone shape results from the fact tbat the initial cd l de, doped from the lower end of the suspensorium is long and conical, and the b,"l5es of th~ next few cell ar begun a lillie way down tlte sides of the firs t cell. The combined c 11 bas~ then [orm a cone-shaped surface. ~ the comb i. fu rther mJarged the cone ~nerated by the bases of the cdls becomes 8atter and flatter and approaches n plan,,-

In nests of V6spula spp. a plane haracter is often, though by no means ruways, actua/ly attained by the combs. This results from the fact that, once the comb is wdl founded, the ceJJs are constructed of nearly or quite uni­form diamel"" throughout their length and their bases ar~ not overlapped. Moreover, the cdls tend tn become straight instead of curved (Fil:". 212, 227, 229, 230, 246, 248. 249, 255). This is especially true of V upul4 'fIl/' gans nests, in which the comb. are often completely plane.

In nolichov~s{l"la neStli. on the contrary, a plane f rm is ne\'~r main­tained by the combs. Once attained it is quickly abandoned, only a narrow circular zone of comb lying in the sam plane. Beyond this circle the comb surface i recurved so that the c mb as a whole becomes conca"e or saucer­shaped (Fig '. 216, 240, 241). Two factor. are responsiule for this change : Fi r t. the cells continue to be built with bases that are narrower tnan the cell mouths. Second, the cell bases continue to overlap. In fact the concave form of the upper comb surlae would be much 1110~ pronounced than it is were it not for one circum tance-the cdls are built in a curve 0 the mouths will open downward or nearly so.

lt happens that tlle curvature de\'doped in the cdls is admirably adapted to Ihe curved body of the larvae which are reared in them. The plane de­t rrnined by the cell curvature, in perfectly symmetrical combs, occupies a

[ 142 J

nIOLO(; \' 01' TUE VESPINAE. 1-13

radial po. ilion with respect to the omh a a whol • Whw the combs are unsymmetrical the planes of .I I curvatun: f succ~ive cells describe a C) indrical surface, the curvature of which follows tbe axis of comb growth along which the cells in questiOD lie. (See pp. 155, 156, and Figs. 195, 196, 23 1,2 .249).

Cilallgu ill comb and " csl susp.,..rit»I.-TIti. topic will be discussed und three divisions, as there are tltrce categories into which suspensoria in wasp n ts may be segregated. These are as CoIlow. : first. the comb ''''pe1lsoria. Th~ are nlore or I .. verticil in character. They suspend one comb fro l11 another, except in the case of the topmost comb, which is ''''I>ended from the roof of the nest wall or from the support to which tl,c nest as a whole is attached. Comb suspensoria are of two sublypes: initial, or taltrai l or main suspensoria; and lateral suspensoria.

~econd . more or less horizontal bracing suspensoria, which may be built bel ween the comb edges and the nest wall.

Third. in the case of Vrspula nesl~, secondary nest suspensoria placed at "anous angles betw""" the nest wall and the wall of the nest chamber, and in Dolitl.avcsplIla n ts, add itional layers of wall paper which intersect the nest support and therd ore contribute to the support of toe nest as a whole. The three classes of suspensoria will be cotlSidered in the order m which the ' are gi ven here.

. \ s wasp nests are enlarged, and their weight, in consequence, is in­eren< ..J , additiotts arc nude to the in itial suspensoria of the combs and la«'ral suspensoria are buil!. ddilioDS to existing suspensoria of all types are n",d~ by plastering pap<!r pulp on them and On tbe adjacent surfaces of the ombs. 'This thickens the suspensoria and broadws their bases. The ,l r' llgth of the suspensoria is still further increased by applieations of So; . \'ary st!Cretion.

The I"t"ral suspensoria in DOlichavcsl",kJ ne.'!!s consist of comparatively Ihin. irregular, vertical walls of paper. orne of which are built out as ),uttrt.SSl!S from the main su pensoria and me of which are built inde­I'endently (Figs. 216, 240, 24 l. 247) . This fe"ture was observed in nests <I f IJ. media, though not correctly interpreted. by 5aussure ( 119) and it has l.ee.n figured by Rau (107) for .0. ,rwculala. 1n nests of .0. arellana the lat ral suspensoria are often composed of an ~en-textured paper similas to Ihnt "ompo iog the edJs in the comb. and thick r than that in the nest wall. In twu nests of .0. macula/a obsen;ed by the writ"" most of the 5<'ConeJary sU>'Jl< D..oria consisted of thin paper exactly like that composing the wall.

In building these u pensori. the paper is added in strips Uke those in thr. ne>t wall but running up and down. The inside of the arcs of paper fare Clutwnrd in some ~S, inward in others, indicating that the wasps when building th.-m faced inward at times and on other occasions outward.

r 1~3 I

144 !'lOflTH A.MF-RICAN VESPlN£ WASI'

In only a .ingle nes of V.sp,ua lip. , a nest oj V. Vl4lgaris, have buttn:s -like su pensoria been found by tit· author. I n this nest, a mmll number of the buttreS!-lik. suspensoria were grouped around the main suspensorium of the topmost comb.

The upper edge. of latern! suspensoria in DO/ichtn!tspllla nests, except those above the first amb, u ually are joined to the edges of the cdl walls of U1 ' comb fr m which they depend. Th y thus. perforce, have a zigtag form above (Fig. 240). ""ondary suspensoria built above the first comb nre atlnched to the roof of the nest wall, and are inclined to be more regular (Fig. 247).

The structure of all comb suspensoria in Do/ic/'/nJcspu/o nests tends to enter the I ad 0 1'1 the main suspenso ria , which are central in location in tbe combs, and arc placed one below another. They lhu constilute a verti ­cal axis around which tbe weight is evenly di ·tribllled. This is Ihe best possible structural arranglmcnt for an aerial ne t hung r rom a swaying support suc h as the branch of " tree. In a nc I hung from a rigid support such as the timbers of bridges or buildings cenlrn! suspension has not the sam~ value.

All typical comb suspensoria in nests of V S/,II/O SPI). (Fi~ . 227. 229, 230,246, 249), both initial and I:lternl, are roundish ill section and more or less cordlike. They expand below into much-flattened can ,forming broad surfaces for attachment. It is worth notiog h...-. that, ulOugh laternl and initial suspensoria in F"/,,,Ia nests are similar superficially, they differ in aile important respect: Like the suspensoria in Dol,c/luvlSpula nes , lhe initial suspensoria are intcgrnl parts of the comb wh icb they suppon , for each .xpands belo\ into the first cell of the comb supported, whereas I:lteral

lIspensori. nre purdy applied structures. The upper ends of all comb su pensoria in V"spII/a nests, except the

initial suspensorium of the topmost comb and therefore of th nest as a whole, are attached to the face of another comb, usmtlly at the meeting place of three \\'al.ls betw as many adjaceJll cc:llil. The bases (upper mds) of the suspensoria are built oll tward in little fanlike buttresses along the thr~ cell walls concerned. ome 0 f the component fiber are extended deep into the supporting cells in lhe angle betv.·et!f1 the walls. Often the fanlik buttresses extend outwa.rd along the wall. betwero cells fo r a di.­tance of three or fOllr cells [rom the centrn! point of attachment.

Lateral suspensoria in I upula neslS are "13de of paper pulp that ap­pears to h \'e ueen chewed at least twice. for they generally are quit homo­geneous in Ie.xture, as the figur how. T he source of this twice hewl-d pulp is the nest itself. Paper lorn from the inner laytrs of lhe ncst wall t make room for lhe nlargcment of the comhs (Figs. 244, 245). and that obtained from old abandoned comb by cutting the ells down to lhei r base

[ 1441

BlOLO .'l OF THE Vr:srt.NA £ 145

(Fig. 243). appears to be U,,: material of which lateral suspensoria <tI1d additions to initial suspensoria nre made.

• ' ew latern! suspensoria in V ~spu/o nests appear to be started most frequenuy on the face of the coolh abo\'e the one to be supported and then bu ilt downward. Th i statement i based on the fact that the lateral sus­pensoria in nearly 011 cases extend directly downward from the edges of the cells in the combs above. I n other cases, however, new suspensoria are begun on the back of the comb I be supported anll arc built upward until the comb above is reached. This is known to be true , the bases (lower ends) 0111)' of suspensoria may frequently be found on the backs of the combs . Another fact upporting this statement is that the upper ends of orne la[(~ra l suspensoria pass directly across the mouths of the cells in the

comb above in lead of being buil t along the walls of the cells. I n such cases the cells above, if empty, are 6tted with speeial paper caps to r""dve the suspensoria. If the cells above contain cocoons the new suspensoria are attached to the cap' of the cocoons. Sometimes tile upper end. of suspen­soria block off cells in the comb abo\'e and prevenl their being used again.

The arrangement and character of !atern! suspensoria in VespII/a nests, in contrast to those of Do/ic/Jw,s/>,,/a, tend to distribute the load raUler than to ceJltralize it (Fig. 246). On any comb the suspensor' are fa irly evenly distributed though there are great individual differences. For ex­ample, there is 110 massing of any large proportion of the suspensoria on any Oue side f the comb, yet two individual uspensoria may be five limes as close togetl1er as either is to its nearest neighbor. rn the upper part of the nest, also, where the comus are progressiv ly lllallH upward. the sus­peasoria on any given comb may be closer logether along a circula r line immediately below the edge of tbe pr.ceding comb (Fig. 249) than they are elsewhere.

The number of latern! suspensoria in nests of V.spula spp. is correlated to SOme extent with tm weight of the combs to be upported. This corre­lation is not nearly as high as might be expected, however. nest of V. v.Jgaris, collected at Cuperrino, California, in 1929 bowed the follow­ing: omb I (the first one built) had 19.5 cells per suspensorium; comb 2 had 28.04 ceUs per u pcosori urn: comb 3 had 36 . 4; comb 4 had 26 . 74; mb 5 had 40.4 1: comb 6 had 55 .54; comb 7 had 49 .65; omb 8 had 35.00; comb 9 had 4 .37; and comb 10 had 41. 00. fn interpreting such data it is necessary to reme01ber that the su pensori" of comb 1 upport all ten ombs. those of comb 2 support combs 2 to 10, tl10se of

comb 3 support combs 3 to 10, etc. T hus we see thaI while in upper combs, if each bo considered by itself, there are fewer cells per suspensorium tha n in the lower ones, this featllre is not nearly as marked as it would be if the weight of the 11 I wcre distributed evenly among all the suspensoria.

I 145 )

146 ~ORTH AMERICAN \I E. P INS WASPS

Taking the whok nest ill to consideration, " calculation of the loads borne by th individ llal suspensoria in thi n .... t showed that the SUSJ><'ll50ria of comb I supported 5 2.93 cells eacb, those of omb 2 supported 419 .82 cells each, those 01 comb 3 upported 359 . 13 cells each, Ihose of comb 4 supported 154 .99 cells each, those of comb 5 supported 142.50 cclls each, tho e of comb 6 supported 139.21 cclls each. those of comb 7 supported 95 .21 cells each, those f comb 8 supported 62 .92 ccll each. those of comb 9 upported 66. 26 cells each. and those of comb 10 supported 41 .00 cclIs tach.

This obvious di parity in the loads put upon di Ire rent uspensoria is partly compensated for by several iactors. F irst, as bas been noted, the s lIspensoriain the upper part of the Dest are considerably thickened as the nest ~"'ow •. and their indiv idual strength is thereby greatly increased. S~ondly . in typical o\'al Dests each of the uppermo t three or four combs is smaller than the combs immediately below it. T he suspensoria on the projecting ·ections of the I wcr cornu. are often attached to the Dest wall rather than the comb above and hence a part of tbe weight of the lower half of the nest is not borne by the uppenno t combs. Third, the abandon­ment in large measure, so far as brood rearing is concerned, of the cells in tl,. uppermo>t combs lightens them and thus lessens the total weight to be borne by thei r su.'PC"soria.

Tn general the combs of "espine wasps are free from the nest wall (Fig. 216) . Th~y also, in nests of VU/lulo. spp . at any rate, a.re kept frtt i rom foreigTl ohjects. such as large rOOlS which may pass through the nesL Tn fact, this free c.hnrncrer of c mbs and wall in the nestS of certain genera of wasps, md the lack of it in oth~r genera. forms the basis for the pri­mary classification of vespid wasp nests and of vespid wasp into pbrag­mocytlaroUll and stelocyttnrous types that was propounded by Saus;ure ( 11 ) orne seventy-five years ago. Vespine wasps and thcir nests belong typically to the stelocyttarous type.

In large nests of vespine wasps the freedom f the combs f rom the nest wall is partially lost, fo r tlle wasps huild mOre or less horizontal, bracing suspensoria between the edge of the combs and the adjacent nest wall . Such horizontal suspensoria o<;cur only in the stntic upper part of the nest where the combs are no longer being enlarged. T hey appear to be com­moner in the underground nests of V .. ""la than in the aerial nests of DolkhoolS/,.lla. Still, nests of D. arellDria ( f ig. 2l6) and of D. IHIJ&IItaJa sometimes sh w numerous onnections between the upper combs and the nest \\'all. Zabrislcie ( 164) cites a nest of D. maw/ala in the collection of the meriean ~ruseum of :-<atmal History with "some of the combs at­tached solidly and vertically to the inner surface of the protective covering, which seem 'to be abnormal instane ." In very lArge nests the horizontal

( 1461

BIOLOGY O F THE £SP1~'A£ 147

susp 'nsoria may be so tensive tha t the upper combs and nest wall are consolidated into a rigidly continuous structure. Such developments hinder the ci rculation of the wasp about the combs, but as the combs involved arl! usually old combs which have been largely abandoned fo r brood reariDg. the hindrance is not a g reat handicap. Still, iD one nest of V"""la pensyl-­",mica collc"Cted at Stanford 'nh·e.rsity the fi fth comb was attached fimlly 10 tbe Dest waJl f r mOre than half its circumference, a very unusual oc­currence.

As wi th the vertical suspensoria . the horizontal uspeosoria of V rspllia neBts are thicker and stronger than those in Dolit/u1IIcspula nests. The lat­ter consist, as usual, of thin paper essenlia lly Ijke that in the nest wall. (See also p. 143.)

In Vuputanests additional susperuoria for the ncst as a whole are built betwe"n the outer surface o f the nest waU and the inner surface of the chamber in the soil in which the Dest is constructed. These toO are built only over the upper part of the nest which has become static (Figs. 228-230) . T hey may b venical, diagonal, or horizontal . according to their position.

In large nests of species of Dolichovupllta there is obviously , in most case" no possibility of building Dew suspensoria between the n t wall and the nest support. In such cases additional support is gained in an entirely diflerenl manner and is a nonnal concomitant of nest growth ; lor the nest wall never attains the static charncter that is to be observed iD V cs~"ta nests. Throughout the de \"elopnlCDt of Dolichooespu/a colonie new I;,yers of paper a re arlded, though less frequenuy. to the lop of the nest as well as to the sides. As Ule nest enlrlrges it encroaches 011 neighboring objects sucb a [\';gs and branches (Figs. 21 5, 21 7- 219,221 , 247) . Thus new up port is gained.

\Vh n Do{icilovespu{a nests are suspended from plane surfac' such as the surfaces of <imbers, only a fel\' additions can be ml)de to the root of the nest before the timber is encountered. Thereafter each Dew wall layer i_ atmcl,ed to Ule supporting timber in a circle surrounding the last layer buil t. T he layers of paper in the wall thus intersect the support in a series o f ncentric circles.

In very large nests f Doiichoors/lllia the additions to the n .... t roof ma .. not all be iocident31 to normal growth. In nests of n. 1II1Ulltaia and of Ih European D. "",diD. pecially. sheets of paper of small e.uent are laid down in an overlnpping fa hion across the top of the nest, prod\.cing the well-known ccllular cap 10 these nests (Figs. 219. 247). It is quite (lOS'

5ible. that these additions have the specific fUDction of extending the sup­porting surfaces of the nests.

In addition to building Domm! suspensoria, which come as near e~pr..,s·

[ 147 I

148 SOJlTU A)fE FU :~ VESPINE WASPS

ing pUrpose as logically may be grunted for structures buil t by insects. wasps do quit~ a I t of rratic pa~r building Lhat contribute> slightly, thougb purcly incidentally, to the support of the combs in 3 n<:;1 . Such rratic pa~r-making is especially characteristic of Dolicil0fJ6Sp"la spp . In

large n ts of D. orenoria • nd D. onoc"lala, for example, vertical .heets of pa~ ase commonly built between the edges of consecutive combs in the upper part of the ncst (Fi . ~41) . Sometimes they are attached to the edge of a comb Or to the ne t wall and are carri.d inward and attached to the face of the comb some distance £rom the comb edge (F ig. 242) . In the lauer case the paper beets cover over marginal groups of cclls aod prevent thei r being longer used (or brood rearing. In li e IS of D. lIIo",/ala , also, one or more combs sometimes arC not built to full size, but extend only part way across the n > . I n sucb cases a very irregular <cllular mass of paper is built, filling in the space that would normaU be occupied by an extension of the combs (Fig. 220) . aL\SSUre (1 19 describc5 a similar cel­lular mass of pa~r "" occurring in nests of "VlSpa" (Dolichovcspula) ",~diJJ beneath the second comb. He describes it .5 nonn: I in n Is of this species ,md says it is built. because " In l1alu~6 slmlble (J'I.'()ir horrlJUr d .. vide," in ti,e space in which a t.hird comb is destined to be built. ""hOJ the third omb i built the ir...,gu la r pa~ is tom away and used in building the comb. Saussure most lik.,)y based his mteml!Jll on observations oi allllormal nests. Jallet (75) shows no such anomalou.s strucllorc in n~sts of D. lIl11dia.

In V ~.r"ullJ nests sucb upcrftuous paper is less commonly met with. When found it may take the form of a vertical sh<':t!t connecting the edges of two or more combs (Fig. 227). Such sheets in V~sp"la nests may pos­sess a real function and not be erratic in origin, for they oft"" occur just inside the entrance hole through the ncs! wall, thus having the position of a screen within a doorway. In other cases "crratit:illy" built paper in V .spula nests is found 00 ti,e upper or back surfaces of composite combs. To elucidat> : In large Vespula nestS a comb is sometimes begun in two separated places. II'bell the two sections e ... entually are brought together tbey often do not occupy the same 1.,'e1 and " jog is formed on the com­posit. comb which their union produces. This jog wiU be covered ove r with a . heet of min paper similar to that in the nest wall.

Till fait of old combs.-As has already been noted (pp. 144, 145), wben a cornu has produced several successive 10 of brood U,. cells in it are lasgelyabandoned for brood productioo. Thus the upper com~ in a ncst arc often used very little after mid-sea>on. \Vhen this stage is reached, U,e walls of the cells are frequently cut away considerably and the depth of tile cells thereby reduced. Occasi'>nall)' the walls may he Cllt down almost or quite to the fecal pellet.; in the cell bases Fig. ~.j3) . The paper thus ren,o"cd

1 148 J

otOLOCV OP THE VESPJN AE 149

appears to be used f r building new suspensoria or for the construction of neW comb. Cornb with cut-down cells bas been observed by the writer in nests of V cspula spp, only.

Anotllcr phenomwon observO!(I fre<luenUy in the nests of V8spula spp, i; the pa~ing over o( the race of the old l' combs of the nest. No men­tion Qf this phenomenon in tlu: literature on wasps bas COme to the atten­tion of tile "riter. The extent to whicb a comb is pa~~ over "aries grcaLly. O ften but. few 0..,)1. are papered over and those are usually. th uj(h not always. marginal in position . At other times large areas of comb ur even wbole combs are covered."

The manner in which the papering over is done also varies a great deal . ~I ~ t ommonJy each covered cell is provided with a 8at, close-fi tting lid ",ade f pa~r of uni fonn texture and color but of little strength. The pilIp for making this pa~r i. probably secu~ from old comb or wall parler which is tom down. Each sucl, lid is usually built Husb with the tOP o j its c U Or even a trille elow it. More rarely the lids are made by the wa ps adding little strips of pa~r aroWld the rims of the cells and grudu­ally working inward to the center. The lids thus formed are rough on the inner surface but smooth n the outer sll r(ace.

Occasionally the papering over i done in the manner described on I'" 148. Several to many ell. may be covered by a single coovex piece ot paper constructed of fr bly gatbe~ pulp and similar in every respect to the paper composing the wall of the nest (Fig. 242) . Such pieces of paper are sometimes built over cells whicll have alfeldy bce.n pa~red over in tht manner described in the preceding paragraph.

A third ~ oi pap<.ring over of the old combs occurs in nests 01 Ves­(, ula nJgoris. In this case the material used consis of finely comminuted rotten , ood applied so smoothly that there is no suggestion of strIP oi paper (Fig. 239). The whole covered surface is 5t1Iootbly plastered. The S11t1e material is applied also, and in the same wa)', to the inner surface of the ne,;t " all in the upper pan of the nest.. In on nest from Santa Cruz, California the wall surface from the filth comb upward was thus plastered. A the le;.l of the first comb the pl.stering of the nest wall, instead of bli llg continued on upward and across the roof of the wall above the first e',mL, was curved inward and joined to the edge of the first comb. The 'p'te wbich originally existed betw~n the edge of the first comb and the tle.t wall waS thtls completely bridged with the plastering materi'" and tile Ii"t cornb thus made a part of the nest wall.

The purpose of this papering over of old comb is nol clear. It obviously impr ves the s:mitary condition of the nest by covering a portion of tile older cells which contain the largest amount o( fecal m'lterial and bwce constitute the greatest potential source of infection within the nest, It is

[ 149]

EO NORTH AM:£JUCAN \ '"ESPINE WASPS

very doubtful. however, if improvmtent in nest sanitation has been the fac­tor which h.~s devdoped the instinct which leads to the papering over o[ old comb. I£ such \ ere the case we sh uld Xpccl the cd ls to be papered Over in the order of their ages beginning always with the top comb. The paper­iog would also begin at the center of each comb and proceed outward. In actuality this order of papering is not followed.

Tbeft' are two otber possible exp\ar"' tions. Papering over of old comb may be a partial expression of the in.tinct to increase the trength 01 the growing nest 5 tructu~ (see pp . 141-147) . This would seem to be true in the Santa ruz nest of V upl/la ~'U19ari.r already referred to. In general. how vcr. the paper used in covering old comb is very weak. It is not can­neCled with the suspensoria, and it adds weigbt but nol trength. The third possible explanation is that papering over of old comb is merely a part of the activity which wasp mani fest in moothing over th illlerior of the older and mare static portion of the nest wall (see p. 151) . Aft" a nest has attained considerable size, the combs in the upper part are 110 longer increased in size. There is, then, no further need for removing the inner layers of paper from tbe wall to make room for comb growth . and the wall thereafter remains constant in thickness. uch older portions of lhe ne,t wall are carefully worked over by the wasps, and a <datively smooth thougb irregular inner surface is produced. P rojecting tags of paper, leit when the last inner layer was torn down, are removed. and holes are Jill in . It seems possible tbat this " interior finishing" may be extended to include more or less of the uppermost combs. This e,'<planation. if correct, accounts also for the fact that tbe marginal cells of a comb are the ones which are most frequently papered over.

C/oonges it! l/oe l1es/ 'RtJJ/.-In the oests built by queens f 'JX.'Ci of DO/"/oUl.·esp"la. and in lhe same nests for some time after their enlarge­Illent by workers bas been in progress, lhe nest wall consists of a series oC spherical or val envelopes that are everywhere di. tinct and epara!. from each other (Figs. 203, 205-2(7) . This fact has long been known, as man)'

publ.i.'lhed d riptions and figures attest But by the time a nest has at­tained any considerable size tbis perCection of pattern begin., to break down

Figs. 217, 218). E ventually, the heets of pape r comllosing the wall Figs. 21 -, 219, 221 ) . though still very e. .. ,tensive as compared with the

"shclls" of paper in lhe walls of VNpula oests (Figs. 226, 228. 236, 237. 244.24- ). are much smaller proportionately, n one covering more than a i raction f lhe nest surface. No I nger are they always started at the tOP of the nest and brougbt downward all around the ne.t. They may be be­!run. and completed, at any level. On the roof 01 nests 01 D. ",oclI/ala. as has alreadv heen noted ( p. 147), the paper addition. may be oi quite small exte,; t (Figs. 219, 247) . ~e,ertheless. the wal l of " Dolicl"w,sp .. /a

1150 J

BIOLOGY OF TUE VESPJNAE 151

n t remains a wall composed essentially of ""P3rate layers £ paper (Figs. 216. 220) .

J n queen nests of V CSPIlIa spp. the wall envelopes occasionally nre scp­arate from each other (Fig. 2(2). More often, ho\ . vcr, the second or third layer is begun sOllle dis tance from the nest pedicel on the surface of the /irst

r econd b yer (Figs. 199.213.214 . Later layers may be started either I rom the pedicel above the nest or on the urface of one of the existing layers. Thus. though the queen nests of V UPllia spp. strongly resemble those of Dolicltotlupula spp .. they quickly lose the feature of concentricity of waUlayer. So characteristic of the latter. And within a short time after the V Spllla workers take up nest building all suggestion of the concl!ntric pattern of the queen nest disappears (Figs. 208.209, 211) . The nest wall acqu ires the well-known "shcll" Or "scale" pattern which it retains through­out the balance oi its duration (Figs. 226, 228, 236. 237, 244, 245).

\\ hen lhe workers begin adding new layers to the nest wall lhey also hegin tearing away the inner layers to make room f r the enlargement of th comb. Thereafter, until the decline of the colony, ule wall undergoes caminual change. The de. truction of the fi rst envelope inside the nest, Or the fi r.t few, nlay be begun at the bottom Fig. 2(7). though with later elll'elopes the middle portions closest to the comb edges are removed first. The ill ncr layers are chewed away bit by bit (Fig. 244) until only a disk­like remruuu rernaills above the topmo t comb (Fig. 2I t ). The edge of eacb disk is then fastened by the wasps to the next disk outside it. 'The final r~· ",I t is the fonnation of a sort of Ceiling to the nest (Fig. 21 6) .

In Id nests, when en largement of the upper combs ceases and the upper part of the ncst acq ui res a permanent r relatively permanent struc­ture. the removal of wall mate.rial takes place only in the lower part 01 the nest The upper part, of the side walls then, as well as the .. eiling," are smoothed o".r, all rents are mended, and projections are removed. This ha. already be<:n discussed lp. ISO) .

The paper of t e tortt-down e.nvelopes is recllewed and used in the manufacture of the new comb and of suspensoria. or it is brought outside ,,,,,I ill orporated into the nelV layers of paper that are being added to the watl. The last use of it can he obscrv~d readily n allY ncst of Dolicho­t','spula ,0 situated that it may be apilroached e1osely. 10 such nests also one Ill"}, distinctly hear the wasps tearing away the paper inside.

Tn large nests of V tspula spp. after the addition of new cells to the upper few combs has ceased the addition of new layers to the upper part of the wall also ce:tSes in large measure. As a r ult the upper part of the wall omes to he constant in foml and in thickness. whi e the lower part continues to grow through the additions to it. This lacruizing of growth in the apical half of the nest occurs to some extent also in Doli,ilovupula

[ 151 I

152 NOIrn AlJE.RlCAN' £SPIl'-'"E WASPS

nbts, but it is not so obviou a (eature of tbem. At any rate the wall in nests 01 Dolkillwespuia spp. often differs much less in thickness in the upper and lower ponions d,an the ",nil at Vcspula nests. In One n t of Vespula p8nsyl'llanka Sauss. at tan ford Universi ty , California, the lower part 0 I the wall was close to six inches thick and formed 8 large hemi­spherical ee1lulas mass. On the top 0 f lhis same nest the wall was but

nl!-ha]£ inch thick, and on the sides it was from one to two inches thick. The giant n t of V. vulgaris shown in F igure 255 had such 8 ce1lulas mass (Fig. 245) that was elev n inches thick, forming the lower part of the Dest wall. Such masses of ceUula r construction suggest those described by Saussure (see p. 148) as occurring in Do/;e"rrJtsp .. la m.dia nests. but whether they are to be thought of as abn rnm! in any sense cannot be de­cided at present.

T he number 01 wallinycr- present at anyone time in the lateral portions of the wall of large-sized nests of Dolic/Ilwupula artnaNa var; [rom nine Or t n to ahaut seventeen (Pig. 216) . By the time a nest 01 this species has attained a diameter of six to eight inches its builders will have con­strm:ted (rom twenty to thirty wall layer., o f which ten to fifteen will bnvr b en tom away to make room for addition to tbe combs. In nests twelve inches or more in diameter, from forty-eight to fifty-five wall layers will bave been built, and from thirty-five to forty of these will bave been torn down. In Vespa crabro nes that are built within hollow trees, or in other restricted cavities, according to ,,,,riOliS published accounts, the n t wal l may be completely eliminated when lhe nest attains lasge size and requires all the available space for comb building. The writer has observed a comparable partinl elimination of lhe wall in nests of Vsspula flll igaris, V . sulpl,ur.a, and F. pensy/wllica.

L ocation of Iht tIIlran," '101 • . - The entrance hole through the nest wall into the Tl • t interior is at 6rst ventral in po ition in all kinds of vespine wasp nests, and may rt'I11ain so eyen after the nest bas attained considerable size (Figs. 197- 203, 20:'-207, 2(9) . In most cases, however. the entrance comes later to be lateral in position, especially in nes o i species of Ves­pIt/a. In old Ilests of the species of Vrsputa known to the writer, the entrance hole, though sometimes 0 11 the hattom f the nest a figured by Janet (75) for a Vesf'uta nata n t, is us Ially high up on the nest wall, generall}' above the middle of the nest and occasionally even on the upper surface ( Figs. 226, 228, 230) . This position is undoubtecUy correlated with the fact lhat since Vtspulll lie as ' umlerground they are approached from aha,'. Or rrom the side and not from the bottom. In old nest of Dolic/rotJes/,lIla spp. the entrance hole is never very fas up on the 'ide of the nest (Figs. 215, 217-219, 221).

J t is thus obvious that as a nest is enlarged Ule entrancc is shifted r rom

[ 15!)

DIOLOGY OF TIlE VE;SPINAR 153

a "entra! position to a position in tbe direction which afford the easiest ace 5 and departure to foraging workers. aUSSllre (1 19) says that in nests of DolicltO'llcspula ",.d;" the entrance is commonly on the north side, th side (rom which the rai n does not come. Th · shift in the entrance slIC;gests that the wasp. are aware of th. ad"antages f the change. The suggestion, bowever, is not borne out b . the facts; for another and more probable explanation is available.

As the workers which are building wall layers bring the edges of the n 'w layers 10 the vicinity of the Dest entrance, their work is interfered with more or less by the incoming and outgoing workers. This interference is gn,a~eSI on the side of the entrance from which tbe lasgest number 01 foragi.ng wru.'Ps lea"e and to which they rerum. The workers adding to the wall layers about the entrance are prevented by tbis interference from building the new layers as near to the apex of the Dest as the pre-existing la' ers w re bu ilt when there was I interference. Conversely on the side 0; the ntrance where the entering and departing foragers ase fewest, the I 'cr interferen e in building pennits the new envelopes to be huilt a little fasther than ,wy preced1ng envelopes. Gradually til"", but quite "ullintentionally," the position of tbe nest entrance is changed to the most at vantageous position. One" this positi on i reached . there is no further cLallge, herause there i then thc same amOlUlt of interference in the build· in.: of ,,,,u layers on all sides of the Ilest entrance.

The probable corrrctness of the foregoing ""planation is Lome out I,y tht behavior of wasps in repairing damaged nests. The writer has 01, erved til repair r two such ck'lmaged nests of Do/ie/r()fJcS/>1Ala arellana. Un~ of tbese. a nest attached to a branch of wild gooseberry at Cupertino, Cnlifor ·a. bad the bortonl of II c nest wall torrl off by some mammal. Th. "ther. a nest built in a sheu at Stanford niversit)'. had the bottom broken Ollt as a result of the combs millg loosened and falling through the n,' t \ ·" U. I n both case , had the IV, flS been really aware of UIC advantages ui the lateral position for the nest enlrance. the waU could have been rl!­p ... irtxl so as to ulclude an even more lateral entrance than the original one . ,\ , Ii IlllUter of fact when the: repairs were completed the neIV entrance in )'''th nes was ventral in po ition . Janet (75) records an essentiaUy similar .i tltatiOD with a colony f D. fflI!d,'a.. I n thi. cale also, the Inrge nest that ""'" destroyed bad a lateral entrance and the small Dest that was built to replace it had a ventral entrance. \Vhcn the latter nest was destroyed, and a .hin! was built to take its place, the first envelope on the third nest had a ~lightly lateral entrance bole. This, however, was subsequently lased with paper and a Dew ent:raru:l!, ventral ill position, was opened.

I t will be suggested at this point that the ;arne factor ( interference by foraging work"'s) which caused the rigina! shift in position of the

[ 15J )

154 NOaTH tE-1UCAN VESPINE WASPS

entrance might"" exp«ted to operate during the period of repair and tbus cause a lateral entrance to be constructed. T hat the interference factor doe. not materinlly influence the course f repair work appears to "" due to two thing~. Fim, e urgent need for repair of the nest w:ill r ult! in SO large a num""r 01 wasp. working on it that it is rebuilt too rapidly lor the inter­fer-once by foragers to "" effective. econd, the diversion of Ihe wasps f rom the foraging grollp to the building group greatly reduces the amount of interference. As a result of these two faClOrs tll< repair work is gov­' rned solely by the original building instincts of the wasps, and the recon­structed entrance is ventral .

The entrances to the nests of American vespin wasps exhibit two dis· tinct types of con truction correlaled with the taXonomic group to which the makers 01 the nes . belong. I n nests of Dolich wespuJa spp. the en­trances are usually simple passagewnys througb the several layers of paper composing the oest w:ill (Figs. 216, 222). In these nests no attempt is usually ronde to finish the entrance into a definite lined IU"" traver ing the wall . the edges of the wall layeH ~.nding freely at the passageway. An exception, however, e.~ists in the so-called beaked nest. of Doiic/.ut'csp"la .nllCulaia whicl1 have been recorded by various writers (Rau . 107 ; Da\'is, 43 ), In this species, when nests are quite sm:ill ne of the first two or three w:illiayers is produced at the nest entrance into a tubular passageway which extends as much as two inches beyond the usual position o[ lhe Dest entrance. Through .is the queen ent"s and leaves. The additional wall layers arc simply bu ilt down to this tube. Eventually, however, when the inner wal l layer. are tom down as the nest increases in size, thi tubular passageway is destroyed and is not renewed. 'milar th ugh much less dev~op<!d passageway through the nest wall is figured by Janet (75) [or "V,splt' (Doliclowupula) syllleslris.

[n nrrast to the simple condition o{ the tl1'ical entrance to mo>t DolichQ1.:fSptda nests, we flnd that the entrances to large V.slmla n t.s are nicely " finished," 0 i ree edges of paper Ill"e to be found in the entrance to a V;spula nest, but there is one continuous rounded and smoothed surface from the interior of the nest wall to the e.'<t"rior. This "finished" condition does nOt exist in very small nests of Vespula. iL'i i. well shown by Figures 197-202,213, and _14. In fact. to judge from the nests shown iu Figures 200-202 and 209. it would appear tllat VeJpula nets do not always have

ntranCes in the proper sense of the terro until they increase considerably

in size. CloolIges i ll /h' gellCraJ sflap. of tloasps' nos/s.-T he nests built by queen ps are nearly always appro"imat~y sphericnl in form (Figs. 197-203,

205-207, 213, 214). At most they arc only sUgbtJy elongated. As long as optimum conditions e."i.t and the nest i. still relatively small, thi~ fom. IS

[I I

OJOt..OGY O F TUE V£SP1 N"J\.E ISS

ma intained (Figs. 208, :2(9) . New layers of paper are addl'd to the wall at fairly regular intervals outside those already existing. Yet it is not long ~fore the nests begin to deviate from the spherical form. They tend to 1x.'COme ovoid or elongate, wi th tJle longer axis tbe v rtical one (Figs. 215, 217, 219, 226, 228). Usually also they ~me more or less asymmetrical.

T il change in nut shape appears to "" the resultant of several inter­act ing factor ', more of which reside in the mechanicnl interrelations be­tween nest and environment than in the instinct-equipment of tJ,e wasps. T he permanence of the main point of suspension is probably the most iUl­I",rt""t of the purely mechanical facto rs. I ts fixity prevents expansion ttl'" ard and forces growth to take place downward or lat rally. The neces­sity, in old nest , of strengthening and making more rigid the upper pan, i- anolher mechanical factor which inhibits growth in the upper pan of lhe nesl. In the case of Vespulo nests gra\.itation nlay be a signi6cant factor, >i llce it is undoubtedly easier to excavate the floor of the nest cavity than the \ lis or roo£.

ne b ctor responsible for the development of an asymmetrical shape has already been discussed in conectiou with th change in position of the nesl entrance. This faclor, nrunely, that environmental condition. deter­mine d,e di rection from which a nest may be nlost easily approached, in­fluen es ualIy b th tbe subterranean nests of V.spulo spp. and the aerial

tI of DoiichovlSpula. ""cood (actor oltsi ts f the actual physical hindrances to work encountered by wasps. This generally h, a m re pro­iound influence on nests of VIS~ula than 0 0 those of Do/ichavespula. H ind nces to exC3Yation range from simple variations in texture and hardness of the soil, producing such slight asymmetry as is exhibited by the nest of V . PClIsyl'IIGllica shoWl. in Figure 228, to actual obstructions in

~ foan of roots or slones, leading to the development f extremely ir­regula. nests, like that hoWD in Figure 23 1, This nest also was a V. i'.I1-sylvallia, n . t.. In th caSe of Dolicflwesp"/o nests ob truetions may cunsi t of neighboring tree limbs or branches, roof timber.;, or the confining wall. of cal'ities in which nests may be built, T he nest of Dolichrnlespllia or, 'Iorio shown in Figures 221 and 222 became, ~use of CODt.~ct with the ground, an irregular hemisphere.

In considering asymmetrical fornts resulting front hindrances t nest CI)O~truction an interesting reaction of the wasps merits cxanlination. In enlarging a nest wasps follow the tine of least resistance. ew cells nre added all Ill"ound the margins of the g rowing combs with uch consistency that the growth of a comb may be compared 10 the spreading of a quantity 0; liquid over a smooth level sudace. The result generally i . that the comb (l ike the liquid) maintains a circular form (Figs. 246, 252) . If obstacles are encountered . advance is confined to the channels ""tween the obstacles,

[ I 51

NORTH AME Q. tCAN VUPtNE WASPS

and the advancing portions take the (ann of ann-Uke projections. When the obstacles have bem passed, each arm of the comb spreads (an-wi,., beyond. Eventually two or mo re o( the advancing arms may join and again spread as a single mass. This mode 0 f expansi n i mani (ested almost perfeetly by V"pula nests, like Um! of V . pcruylfJanita shown ill Fig­ure 231.

That the growth of the rombs of such nests occurs essentially as stated can be determined by examination of the celI bases on the back sides at the combs. Since, as has been pointed out earlier in thi paper (p. 137), U,. paper strips composing the cell bases are laid down at right angles to the axes of growth of the comb, the course of these axes can readily be determined (Figs. 246, 248, 249) . Figure 195. hawing in diagram the ouUine and the axes of growth of one of the combs in the asymmetrical V. p.nsylvallica nest hown in Figure 231 . was made on the basis of this fact. Asymmetries of the sort just noted should not be confused with the bwlding of lobed combs composed of queen celIs such as are discussed on page 140.

Some instiIlctive. or at least biological, iactors thllt influence changes in nest shape are the (allowing: First, cells are built to Ope!l downward_ They mu t, in consequroce, hove suspension from above. This fact, there­fore, IlrevenlS growth upward in a wasp nest. Second, wasp. exhibit the general indolence lhat is manifested throughout the animal world and i. expressed in the tendeney to take Ule easiest conr,., that is open . As nests are increased in si.e the uppermost cODlhs become farther and farther re­moved from the entrance. Comb-building workers, therefore, are increas­ingly impcUed to apply their loads of paper pulp to the lawer combs and neglect the upper ones. Third, .. ,ch grown larva, just before pupation, deposits in the celI it occupies the indige bllles and wasres accumulated in its stomach during its entire larval life. The cclls in the upper comb, . then . each of which bas produ ed two or more young, become increasingly unsatisfactory for brood-rearing. a.re neglected, and i'nallyabandoned.

Tbe various factors which lead to change in nest shape produce the folio "ing general final results. In aerial n!!S1S (i.e. , nest. of VoUcha­vespufa) growth in length always exceeds growth in diameter e"cept where obstruction are encountered (see p. ISS and Figs. 221, 222 ) . whereas ubterran.an nests (Vesp1l/a RestS) may grow as much (Fig. 228) in di.tIl­ter as in length (height) or more (Fig. 255) in diameter than in length

(height . DoiicilOflCS/lulo ne ts tend to become slighUy asymmetrical ovoid bodi with their greatest diameter above the middle Figs. 215, 21 9) . V upula nests tend also to become ovoid in form but to develop their gre. t-est diameter below the middle (Fig •. 216, 228). That factors additioD31 to those presentod above may inRuence change in nest shape is suggested by

[ 156)

BIOLOCV O t" TUF. VESllJ='A C 157

the n La of V uta crobro. T he columnar fonn of thei r nests when built in hollow tree. would seem to support the explanation given, but Buysson (2·' ) fi/,'llre5 a nest of r·. crabro which . though aerial, was n ertheless coiumn.a.r in fonn.

Irregularities and oimorllloli/ics in n.n CI>nSI,."c/iotl .-vVhctl a comb is started from two or lIlOre centers, as occasionally happen in V ... "ula nests, Olle o f the pieces may he deflected downward wben it meets the other a"d be extended at a lower level across the face of the otber so Umt tbe uU,er is interpolated belween the deflected section and tbe pre<:eding comb Fi . ~-- .

Whenever ,he separate sections of a comb started in two or more n'l1tcrs meet and are joined together. various irregularities may arise. For example, the separate secti ns frequently differ in level . owing to differ­ences in the lengths of the suspensoria supporting them, and an uneven compound comb re ults. The direction of the rows of cells in one secti "f comb is rarely the same as in the other sections. Ther e seems to be no (actdr other than chance, as expressed in the position of the S<!CDnd cell oi a comb 011 the side of the first, which determines the dire<:tion of cell rows in a comb. ~ fusion o f two sections, thererore, re5l,1 lts in a "<llllpound comb which is lacking in unity of pattern. Then, in the case oi any cumpound omb. irregularities in celI size, the omission of cell logi­,. Iy to be e:"pected. and the interpOlation of accessory cells along the line of j uncti(ln and to some extent also around the periphery o f the compound ctlln b al ways follow the fusion o( the separate ,.,.tions.

When nests are damaged th rough any cause . the wasp repai r the damage mOre or less success fully except when the nest is totally destroyed ur \I hen the d:unage occurs at the ery close of tl,e nest' teIllU1C)' . In the ca," uf nests wbich art! t mpletely destroyed, any wo rkers which remain afterward usually artcropt to rebuild on the site occupied by the original 1I0st. This fact has been noted by Janet ( 7 ) . Rau \07). and othe ...

In all Or nearly all cases, repaired Dests show irregularities in COD­

,truction which are not normally pre5f!nt. n,e wasps seem unnble to rebuild a " <Ir"'al type I nest. The repair of two nests of Do/j,lwves~uia arenaritJ i rolll whic.h lb. ape.~ oi the .,all had been torn has already hecn discu ed hridly (p. 153). In this case the repaired nests poSS<!ssed strictly ventml ent rance holes, whereas the undamaged nests had had laterally directed (':0\ rn nces.

. One of these nests was built in a woodshed near Stanford Uuiversity In the spriu oi 1928. This nest had by May 20 nttained a diameter of . hout five inches and a length of ix inch .... when for some reason. possibly a weak SUSpensoriulll. the comb$ fell out, tearing out the bottom oi the nc, t wall as they wwt. The damage was probably not Ibe act of a marnud-

[ 157]

1" :sORTH AlIIERICA!': VESPISE WASP'S

iog mammal, unless the latter was routed by the stings [ the workers, for the combs w re found on the shed Boor the next morning with the brood undisturbed except ior the injury resulting from the fall , A few dozen irritable workers were clinging to the comb, but mo 't of them were clinging to the old nest wall attached to the shed roof. From one edge oC the hole in the bottcm oC the old nest wall there dangled a large piece of one of the outer wall layers which had 110 t been torn completely loose when the tatastrop~ occurred.

Til two or three days the workers had rebuilt two or th r pieces of comb, which could be seen through the hole in the bottom oC the nest, and had begun to repair the wall. By May 24 the h Ie in the nest wall had been closed. except for a small apertu re left for an entrance. by a sheet of paper attached a little within the edges of the Oll to< layers of the original wall. The queen apparently had been ki lled , Cor the nest was enlarged no further nod the colony ceased to exist a few weeks later.

\ hell the nest was tak'n down. ten di tinct sections 0 r omb were found within it. These were all , nuill, suggestive of medium-sized Polisles n IS, and were suspended in haphazand fashion, partly irom the nest wall and partly [rom one another. One piece had 11 very peculiar location. In the repair of the nest wall the wasps had leCt a large ri ft between tile old side wall and the new layers of paper cl ing the gap. In this ri Ct on of the com b. , located.

Many of the cells of these combs appeared to have produced adult wasps. These no doubt were males, as they must have been produced irom eggs laid by worker . J n all the repair work on the nest, the wasps had fai led to make use of the dangling remnant of the old wall paper hanging f rom the bottom of the nest. T he remnant was useless in its position and contained cQn iderabJe paper . yet the wasps took no cognizance of these facts.

miscellany of small irregularities occurs in nests. In nests of Dolkllo­ves/,ula spp. single cells or som times two or thr .. cells will be built in a horizontal position across the back of a comu. L1 one large nest of D. ort­flarU> a peculiar piece of comb Fig. 2~) was found attached to the side wall. It consisted o [ a number oC cells bui lt in such a fashion as to sugge I

a wi nding staircase. Each cell in it was boilt at an angle instead of verti­cally.

SrzE ATTAINABLE BY \VAS P CoLONIES

T he largest nest of any species of Do/ichOt'cspula which has come to tile attention of the author. a ne t oC D. or."oria, measured a litlle over" foot in d iameter alld fourteen inches in length. L:!rger nests hllve been recorded . but it is probable that most nests do not • ttain evell this size. D. "wclliala

llS8 I

IIIO l.OCY 0,. nu! VF.srrNAE 159

nests certainly attain gemler dimensions and may av rage larger, but few nc.ls of D. are1wriQ in the vicinity C tan ford Uoh'enit}' exceed ten in ches in diameter or twelve ioches ill length.

:-' ests of Vcs/JulQ $Pp. in the vicinity of tanford University attain" milch la rger SIZe. One of the largest typical nests taken by the writer meas­ured seventeen ancl a quart"r inches in length, sL'<teen inches across oe way and ten illch across On a line at rigbt angles to the first. This was th~ ne, 1 built by the colony referred to elsewhere in this paper (p. 141) as ha" ing endured for part of a second season . It was the work of V upula I'I'/I .\·y/~·allica and was somewhat larger than is usual for n ts of this ped .

The only typical nest of large sile of which a complete count of the ('clls is a, alIable is the irregular n t of V . pensyiVllJliCQ the combs o[ which arc sh wn in Figll re 231 and refured 10 on page 15 . It was dug up before the. cason for this speci was more than two-thirds over, yet it contained 10.4;6 cell ", hicl l had produced an average of two adult wasps each or a tOlal of at I t 20,000 wasps for the season up to August 19. None of the c~lI s in this nest were large cell for queen production. This colony con med 4,768 Itve workers and a queen al the time it was taken. Had it ueclI allowed to continue until the end of the season it would undoubtedly have ueen increased by 2,500 to 3,500 additional ceUs. and m ny more wa,ps would certainly have been produced. Its population at anyone time, ho,wvc'l', rOJ o>'ht not bave lDereased materially, owing to the dying If of old workers as new on~ emerged. A nest oC Vuspula vulgaris dug up by l'm J. pru)'1 al anta Cruz, Cali[ornia, in the fall [ 1926, contained the queen moth .. , 5 males, and 2,9 - 1 workers.

( )nly typical Hests have been discussed thus fa.r in connection with size. :\' 0lHypicuJ nests in a second year of development. and mothered by '''' eral Instead of but one queen. develop in CX I remely rare instances and attaill enormous s.ite. uch was the nest shown in F igure 255 whicl. is 10 be t he subject or a specml paper. I t was " nest of V cspula ;ulgo.ris billit 1Il a. re.:ess beneath sonle timb ring at the edge of a mountain road in Big Ila'ln ate Redwood Pork, anta rul unt:y, California. The structure (,f the nest and the nature f its OCcupants ~Iearly indicated that it IV"S

the work of a c lony ill its econd year. The 11 t measured forty-six inches ~rom lef t to right, forty inches from front to back, and thirty inches in "!:h . Only about hal[ the heigbt was occupied by combs, but lhere were

SOllIe twenty-one c.omb levels. Twenty-four combs were counted' but there were !lot twen t),-iour comb levels, for on some [ tlte comb levels n~w combs had been developed from separate center. No count was made Ot the wasps secured, but lhere were approximately four gallons of them. T hese were mostly worker •. but induded newly transformed males and

[ 159)

160 HORTU AMERlCAN ESPINIl \'ASP"S

qUe<!M and some twenty-two I) fune ional qu«n mothers. The latter were recognizable because of their worn condition. Their wings were frayed at tM tips and thei r back were worn fr« of setae.

LIFE HISTORY A 0 METAMORPHO IS

T HE EGG

The eggs of vespine wasps (Figs. l73, 212) are very delicate, so much SO that it is difficult to detach them from the walls o f the cells to which they arc glued without destroying them. Each egg is eloo!:"te, about four time as long as broad, sJighl1y eurved. sli ghtly narrowed aDd rounded at the anterior md, and more narrowed at the posterior end, which is also less blunt. Color m ilk white. The ~ is glued by its posterior end to the wall of the cell, being usually attached in one of the angles toward the enter of the comb. I t projects out into the lumen of the cell, and toward the mouth o f the l:uter, at an aogle of about +5 degrees.

BROOD PRODUCTION

Egg-laying begins shortly after tbe first cells Qre laid d wo. s in the cnse of cells newly constructed later on by worker, eggs are laid in th"'" long before they have b«n built to full size. Thi fact may be due to the urgency o f the instinct to oviposit : but whatever the origin of the habit, it facilitates feeding the larvae. The egg is placed near the bottom of the ceU and the young larva adheres for a fe w days to th< wall of the cell. The cells are increased in depth as the larvae grow, so that after the first few days, their heads are alway- at about the :une level with respect to the r ims of the cells. In the case ot an old cell. which i being used a econd or third time. the egg is laid deep in the cell. ft is obvious, there-

fo r , lhat the ea5<' in feeding larvae, which obtains in tbe case of new ceUs, i (II) incidental r ult of eggs being laid in th"'" while ey are yet small, and is not the result of adaptation.

Durtng the COUT5C of her lifetime the queen of vigorous colony may lay as many as twenty-five thOUS:llld aT thirty thousand egg.. This means that he must insen her body into the paper cells of lbe nest twenty-I",. to thirty thousand times, as the case may be. The result of the friction involved in SO many layings of eggs is as would be el<pected. The wings become frayed "t the tips and worn shorter aod .h rter. One such queen in the author's collection has wings which are reduced to mere stubs less than one-third of theiT original length. T he setae, which at first clothe her completely, become broken off "lmost entirely, none remaining except on small protected areas such as the posl rior face a f the propodeum and the

[ 160 I

OlOLOCV OF THe Esr l NAE l61

.nterior face of the first abdominal segment. The integument b«omes rfectly smooth and poli More or Ie .. e1iscolomtion of the body,

especially of the dorsum of the second and third abdomInal segments, (J<.-cur; . Th is dis olomtion con ists of th suffusing into the yellow or white maculation of a brownish r blacki h c lor. Thus. by the end of the seaSOIl, the queen mother of a colooy becomes litemlly, as well as figura­It\"ely. worn out.

ECG-LAy , :> BY \VOIlKI:1lS

111i. phenomenon. which remains yel to be adequately inlerpreted in terms of in tincts and reflexes, i a fairly regular occurrence toward the <III'" of the life of a colony. Egg-laying workers are less sys!em.,tic than the queen. The queen lays but olle egg to a cell, whereas workers may lay ;","eral (Fig. 212) . The author has found as many as nine eggs laid by ,,", lrkcrs J.Jl one cell. Tlte queen also, as already stated, generally lays eggs ill the angles 00 the ides of a cell ne:u-esl the cmler of Ihe comb Or in a lille witb a growth a.'<is of the comb. W rken; lay theirs in any angle or Ull an ide of a cell. The general belief is th."lt workers do nOI begill egg­laying until the death of tlle queen; but if the e."istence of supernumerary <g"gs in a nest is evidence of egg.laying by workers, then lhere is proof to t .e contrary. A nest of Vcspilia /,ells)'h'Gllica dug up at - tanford Univer­,ity un :.lovember 9, 1929, contained many. upernumemry eggs, yet the 'I"cen was till present. She was, however, extremely dilapidated .

TUE LARVA

T he newly emerged La",,, is very little different in appearance from the . :;g. It is a triB thicker and shorter immediately after emergence. hilt that i. all. In fact it is sometime difficult 10 determine whether the ohj ect deep in an old cell is an egg or a newly emerged larva. The egg ml!Dlbrane is very th ill and delicate and i simply wOt"ked down over the l:"Crging larva till it becomes a little wadded mass surrounding the POSIe­rlOr cnd f the arva. The egg membrane o f a wasp is therefore very <ltffcrcnt Ir m those of many insects in that it is not shell-like and does n"t maintain its fo rm after the larva em rges. T his is true o[ the eggs of DIan )" Hymenoptera.

The newly emerged larva clings to the spot n the tel! wall t which the :g had been lued. 5 was first pointtod oul by Du Bu. on (22), for about

th~ first twenty-four bour the posterior part of the body of the young lar a is surrounded by the egg mern rant. rt thereafter ' adheres to the di ,carded egg membrane and to the cell wnll by virtue of a viscid mucus which it secrfles. This may readily be ~en in the case DC a larva which is

[ 161 J

162 NORTll Al.lER1CAS v -rISe: WASPS

two or thr~e days old . The muCUS is 50 tenacious that if an au .. mpt is made to pull a young larva loo.se from th~ eel! wall. the larva will often be pulled in two ~fore the mucus yields.

The larvae gmw very rapidly and . ae ording to Du Buy son (22), un· dergo three molLS before the final tranSformation to the pupal stage. The exuviae at each molt are worked backward over the body and massed against those which precede them, the entire lot being glued together by the mucoUS secretion from the larva. When the lan'a gets large enough to fill the lumen of the cell it frequently transfers its point of attachment from the side wall to the bottom 01 the cell. All the while. how" ... in the great majorit)" of cases, it retains the same p<lsition .11 the cell . i:e., ."ith .'ts ventral surface dirrcted toward the center of the comb or held In line WIth a growt h axis of the comL. The large larvae have paired plAl/ral lobes (Figs . 170, 174: LPL) on each of abdomin"l segments twO to SIX. The lobes on segments three to 5ve are more pron~ced than. tbe others. These 10~5 doubtless aid th~ larva~ nsiderably 10 Illatntamlllg th=­selves in Lhe cells. As the larvae become larger they depend less and less on the mucous secretion and eventually depend entirely on lateral (riction t.o bold them in. uch old larvae may be readily pulled from their cells without injury. Often wh I a colony is asphyx iated some of the large

larvae fall out of their cells. The position maintained by th larva. that is, with the venter directed

toward the cenl~r of the comb. or in a l 'n~ with a growth axis. is correlated witll the usually curved shap of the cells composing the comb. The curve of a cell and that of the body of the larva coincide. The diameter of lhe body of the larva and that of th~ ceU which it occupies also show n high degree of correlation. The cell is widest at its mouth a.nd tapers .gradually to its base. Th~ lan'll is \~idest just back of O,e thoraclco-abdorruual Junc­tiOD, whkh is commonly held at lh ' mouth o f the cell or just belo\\ il. and taper. posteriorly at about the same rate that th · cell cl<lC'.

Cbarac1erUtics of Ih~ maiurl larva ( igs. 170, 171. 174} .-The mature larva is elongate ami fl attened-cylindrical, the flattening beil~g dor o-ven­tra\. It tapers rather abruptly in (ront of Ihe first abdommal 5e\:ment. wbid l is the broadest part of the body, and more grndually backward. The body is arched slightly ventrad. h s form is thus seen to suggest that of the adult. but there arc several notable ,liIl"erenc... The lar"ttl head IS only aLout one-third to one-half a. \Vide proportionall)' as is the head o( the adult. The head and thorax of the larva make up only about one-filth Its total length , whereas ill the adult they constitute well over a third of the

~Ien~. . . The sid f the n"'lUre larva are nlarked with longnurlinal rOlln ,d

rioges which are produced. on aLdominal 'egment. twO to six . into the

r 162 I

PI LOGY OF THE VESPINAE 163

pleural lobes (LPL) befo re mentioned 115 aiding the larvae in retainiog thcir po ition ;n the cells. The intersegmental grooves are indisunct where they cross Ihese ridges. The 6rst twO pairs of spiracles (Sp) are located just in frOllt of the intersegmental grooves, the third pair in tlte inter­segmental groove; the last seven paira are locatffi On segments five to eleven just caUdad of the intersegmemal grooves. The thineenth seglU~nt bears laterally. at the level of the laleral ridges on Ihe other segments, a pair of elongale. blunt calldal papillae (CP) .

The segmentation of the body of th~ lar\"3 exhibits som" inleresting f ""tur . These are well shoIVn in the figures. Wh~her or not th~y ."ist in the same degree in other specie' remains yet to be delermined. The first i ur bod)" segmcnts (those giving rise to the thorax of the adult- really the thr~ thoracic and the Ii t abdominal) are much shonened venlrall),. the result being to produc~ the charact~stic ventral cu n 'at ll re of the body. -n,. iotersegmenral groov between the anlerior segroenls are strongly iJent and their low« portions shifted fo rward so that lhe lower ponions he helow O,e middle of the terg-al portions of the segnlents. The sixth inler­S<'gmcntal groove (Ihat between abdominal segments three and four) is the first to run in an unbroken cun'e around the body.

The !.~rval head (Fig. 171),· in cephalic aspect, is sl\ggestiv~ of that o( Ihe adult . [t is rounder, how ·ver. than the head of the adult. The d ."PlUS (Clp) i faintly mark l out above, but is distinctly set off from the shon. t ransve .... hilobed labrum (Lm) ~lolV. T he labrum is about four times os wide as it is long and is ~J11arginate medially, and the lateral lobes arc /:ently rounded anteriorly. The mandibles CUd) are of moderate length. and :JJ"C truncated and nalTowffi apically where they end in three prominent teeth. They are quite suggestive of t.he mandibles of the adult except that th" teeth take up the !:ntire apical margin . whe.reas in tbe adult lbcre is a knife-like utting edge on the apical ma rgin anterior t the teeth as seen irom the side hut mesad and dorsad when viewed from the front. The opicallhird of the mandible i. strongly sclerotized. the basal two-thirds are

Diy weakly sclerotized. The maxillae M x) are in tbe lorm of low, eloIll,'lIle. rounded prominen . Each bear two minu te, brownish papiUi­iorm tu~rcles. One of which may be the ma.>cillary polpus. ince. however. lhi point has Ol ~ determined. and there is an equal chance that the tuhe.r les. may be. the galea and !acinia. tlley nre here referred ttJ simply as lhe ma.rlllo,)' lob" (LM x ). The labium pr~sent5 two division : a broad basal one, th~ labiobau (LM). "hich bears tlle other ; and a projecting

.. The: nomenclature or larval head structures u~ here il patterned aitt!r that u=l by Parker and Thomp.on (98), "ul.ini/llr to larval ch.alddoid flymenopl<ra, ond ,bat u><d by Va""e and Smith (1~9) . in whieb ""rasi'i. Hymmapt<ra g<nl!rally art! considered.

[ 16J J

NOIlT H A H~IU N £.sP INE WASt'S

median I be, the labiGJ prolll ;II,nu (LbPr) . The labial prominence bears lal~rnUy a tydir of minute, brown-ringed tuberd " the rudim<cnts of Ibe 10biGJ ~ol/,s (LbP , and bet",,,,,n thC!rl1, showing fainUy, the arC1l:lte orifice of U,e labial gland (SO). T he antennae (A) are represented by a pair of small tubercles borne on stightly elevated disks, They are located a short distanQ latero-dorsad of the clypeus. Vance and mith 149) Iabcl the circular areas forming similar disk. on the head capsules of parasitic Hymenoptera, all/e.",ol foro.nilla (0/0). The same teml is applied here, tentatively, to the disks on larval Vespidae. The coronal suture (fs ' ) is well de\"~loped, and from its anterior end, on the vertex, the frontal sutures Us" ) diverge to the upper laternl angles of the dypeus. On the head capsule above the compound eyes a pair of prominent, d ivergent brown lines . tlle /'Illporal fos.ro~ tf), diverge. They are forerunners of the compound yes.

The general c lor of the fully grown larva i ' a creamy white. The more posterior parts of u,e body may, however, appear somewhat blacki h. the color bei ng du" to th" dark contents of the gut. Th",,, con ist of the indigestibl" portions of the food . with the addition perhaps of certain excretions, which are retained within th" gut until the timc of transform:t­tion to the pupa.

Spin"illU and lronsfarma/;Qn.-\Vhcn the lan'at reach maturity, each spins a cocoon. The ",,,,II of the cocoon varie in thickness in the different portion, being thickest in the exposed end which fomlS a cap to the cell . Wilhin the coU the cocoon thins basally to the frailest oi membranes, In Ule case of some species the <;QC()()n is incomplete at the basc and U,e anal segmcnt of the Iarv~ i ' in contact with the pCpI!r of the cell or with thc exuviae of the preceding occupant of tbe <cU, as the case may be. T he cap and nt I 'tlSt ~ portion of the walls of the cocoon consi t of twO distinct heeis at silk whicb may readily be separated.

At the completion of spinning the larva within the coco n resUllIes the posilion in ule cell which it formerly held, i. " with the curve of its body coinCiding with that Df tho. cell, and \\;th its venter dirttted toward the center of the comb or to,,,,, rd its growth axis. This fact WlIS readily deter­mined by cutting off the lOp of cocoons and noting the position of tbe larvae il ide. The same position is maiotained du ring trolll formation to tl,e pupa and througbout th" pupal stage.

ery SOOIl after the completion of the cocoon the larva \" ids from its int~ine th~ blackish mass of undigested food remnants and excr tions, the f116COII;lIf11 . which have been accumulating throughout its larval life. Th is mass becomes packed tightly into the bottom f the cocoon, Or the bottom of thc cell in the case of those spI!cics in which the cocoon is incomplete basally. Eventually the meconium dries Out more or less com­pletely and becomes a brownish pellet of granular m~terial. The number of

I t64 J

S f LOGY OF' TUE VESPlN AE 165

til e pellets in 'l given cell indicates ho v many wasps have bI!en rcared sucCl!Ssively to matu ri ty in that cell.

Du Buysson (22) gives from twelve to eighteco days for the develop­ment of a queen larva, about fi fteen days for that o f a male lan'a, and, in the summer season, consider.lbly less ("braucotl~ m<JiM' ) for the worke.r. The length of lb. larval stage must certainly \"ary, bowever, with the species and witll the. ecological aDditions undt.r which the individual colonies develop.

TilE PUPA

The pupa (Fig • . 169, 176 i of the. examte type. baving legs and wings frc from the body and, except for its whiteness and delicacy, is very similar to the adult. An extended description, therefore, is un necessary. A few facts of espl!cial interest will suffice. T he derites are less distincUy marked out than in the adult. The diffe.rent castes may be easily distin­gll ished, bowever, the males by the longer amennae . and the qu",,"s and w rke.r · by tbeir discrepancy in size,

In the pupa the mouthpartS are extended and are. bent backward over the fo re cOxae_ Tbe antennae are laced with the scapes (Sc appressed tigbtly aga inst th face and turned obliquely outward and downward, cross­ing the compound eyes about midway between the eye sinus and lhe lower margin of the eye. 11,e anteonal ftage11a (FJ) are e.'<iI!Dded caudad over the sides of the head, crossil)g over the bases of the mandibles (Mil) . and <'" tending diagonally a10ngsid the extended mouUJparts to meet in the midv""tral lille VCr the thoracic temum. I'll the queens and workers The tips of d.e flagell a nd just over the base of the middle coxae (Cx). [n the males the tips of the flagella attain the base of the abdomen. Th legs of the pupa are folded tightly against the body. The f ront femur (Fv ) ex­tend!; upward and fo rward along the anterior lateral face of the meso­thorax to about the level of lh .. sinus of the c mpound eye (CE) . The tibia (Ti) i ' flexed on the former and lies against the gena (C~) . The tarsus (Ta) is turned backward and parallels the antennal flagellum. It reaches to the middle coxae. The middle fenlUr, flexed against th meso­" ICUrOD, extends directly toward the posterior lobe (LTl ) oi the pronotuD1, the middle tibia is flexed against the back of the femur, and the tarsllS is carried diagonally bacl.:ward along the oultt margin of the hind coxa to meet i fellow of the opposi te side on Il,e midline, uear the posterior margin of the second abdominal egment. The hind femur is Rexed against the metaplell ron and extends toward the bru;e of the hind wing. Th. hind tibia is Rexe" IIpon its femur and extends obliquely backward and mesad parnllel to the middle tibia. T he hind tarsu. continues backward in the same course, gently curving toward the midline to meel its iellow

1165 I

166 NORTU AM..U.rCAN V£'sPJNE WASPS

over the fifth abdomi rtal sternum. The wing pads (WPZ and WP3) are folded against lh" sides of the thorax, t.xtending obliquely downward and backward in til<! spaces bt>twcen the middle and bind libiae. Th" a))"X of the front wing pad ends opposite the apex of th" hind tibia. The hind wing pad lies bt>neath that of the fore wings, only it5 extreme base being ,·isible bdlind that of the fore wing.

When first formed the pupa i. entirely cream white. Pigmentation develops gradually. The eyes are the first parts t develop pigment, and they are frequently quite dark before any other parts become colored. Tbe thora.'C deVelops pigm<!IJtation next, the color ~howing fi r t in the meso­scutum (Sci) and the mesopleura (PI2). Later the posterior lobes of the pronotum (LTJ) darken . Followi.ng this. the other parts of the head and thorax, the legs, and the abdomen begin to develop color. The antennae and the abdomen are tbe last parts to be<:Ome hea"ily pigmented.

E,,,ntually the pupa com..s to look much like the adult except for the mall size of the wing pads and the generally paler coloration. The pupal

integument i recognizable as a thin, delicate, transparent membrane cover­ing all the parts.

Up to this time the pupa bas remained in tire same po itioo as the lan'a and absOlutely quiescent the end of the pupal stage approaches, muscular twitchings occur. EvenruaJly lhe pupal ~"ticle ruptures and is worked back ward over the body to the tip of the abdomen, Where it is cast off.

Du Buysson (22) states thn.! . although th.e length of the larval stage differs, as noted (p. 165) . in tbe three castes. the pupal period is about U,e same for all castes. At the height of the season this is aboUI t-wel"e days.

T SE ' EW ADU LT

The young adult is at fi rst , olllewhaL pale. The pubeocence especially is light colored, being ommonly grayi h. even though it is destined to be black later. The new adult is feeble at first also. bUI it begins to move around in the cell. It 00 longer maintain the position with the venter toward the center of the comb but rotates at1 iU longitudinal axis within the cell. That this rotation occurs after the final Itlolt and not before is proved by ti,e fact Ihat pupae, however strongly pigmented . are found almost withouL exception in the u ual position, whereas newly transformed adults may be found facing in any direction witlnn their cocoons.

When the new adult is ~trong enough it cuts it way out of its cocoon. Sometimes it cuts a ragged hole in the center or toward One ide of the ""d of the COCOOn. At other times through rotalion within the cell it cuts a neaUy circular piece from the end 01 the cocoon .nd emerges.

The newly emerged adults do not trim the ragged edges of the cells

[ 166 J

BIOLOGY OP THE VESI'INAE. 167

which tbey have just left but lea\'e tbis task for the older and more t.xpe­Menced workers. The latter apparently u"., the trimmings, in some cases at least. to incorporate with vegetable fiber into new paper. The newly . erged. or teneraJ individuals as they are called . according to Janet (75 ) . immediately seek out the larvae in the comb and bite their heads to induce the secretion of salivary liquid. This secretion. then, is the first food r 'ved by the newly emerged wasps. Shonly aitenvard they a re fed by the older work... in the nest on regurgitated wee ... , or they may be given tll r .. Is £ malaxated insects fr m which they suck tit. liquid cont.nts. The newly emerged individuals remain within tbe nest for the first few days, ,mly gradually, il workers, assuming the duties of nest-building and urood carc.

Teneral individuals nre usually mild-tempered, and if the nest is dis­turbed many will attempt to escape by crawling head first into empty cells . OccasionalIy. however, as observed in the case of a nest of Vtspllla pm­u /t·Qllica. which the aLUhor dug ollt in \ .shingron. teaoral work.,.-,; wiU j;,m in the defense of the colony and bite and sting the trouble-naker.

CASTE DIFFERENCES

previously noted ( p. 139) , queen wasp are produced only in large eel , workers lypically in small celt.. and males in cith · r large or mall c . With lew ""ceptions one may easily distinguish betwe"" large ceUs c ntainiog queens and those containing males in the pupal stage, for queen Uu-me , pin long cocoons tbat project high out of the cclIs, whereas mnles "Ili n cocoons that end flush with the cell margin. Or evert slighlly below the lIIargins. The only way to dist inguish between small ceU containing worker pupae and those containing male pupae is to cuI open the cocoons.

P ROPORTrON OF MA1.ES, :-lEW Q UEE-,( , "'''0 WORKERS

~Iales and new queens appear to be produced in about equal numbers, but the actual numbers \'3r)'. From two bu ndred t St." raJ hundred of each are produetd in a vigorous colony. The percentage relationship which the number of males and young f.,ma]es hears [0 the total yearly population of a colony also varies considerably. The percentage of males and fenhlles is rlistinctl. less in large col nies than in small ones. the reason being that in lar~e colooies • much greater number 01 worker cclIs are built and a larger number of workers are reared befo re the season f male and queen pro­duction approaches. The percentage of males and females is therelore much less in colonies of V cS/>II/a than in colonies of Do/i" l(r~,spuia. V (S -

1'"la colonies endure from one to four months loDger than colonies of DO/j"'utJtJ/,lIla. In conse<luence, th y have an opportunity to de"c1op a

I t671

168 NOaTH A¥ERICAN 'ESI~,sa WASPS

vastly g-r ater number of workers, actually and prop<m ionally, than ;s true of Dolichovupulo colon; ....

TDE A UTUMN CRO P OP ' E\V QUEENS AND MAL£S

s the deDlise of lbe colony approaches. the eggs laid by the queen e to produce workers but begin to yield males and other fertile fenl.,,1 like herself. These last are to oecome the founders f Dew colonies in the year following.

The males arc commonly said to be developed from unfertili .. d eggs (Lubbock, 85; Wheeler, 161; and others), but lbe mechanism which d.­termines ulac some eggs shall be fertilized and olbers not remains yet to be disco,'ered . The suggestion has been made (sec Wheeler, 161) that !.he queen may be able to control !.he litteration of sperm fr m her permatbecae and lbus to lay fertil ized or uDfertilized eggs at will , the fonner producing the new crop of queen., the latter lbe mal '. Whether or not !.his is true remains to be proved. The suggestion has also been made that mechanical factors, such for example :IS the differi ng diameters of !.he cdls in which the eggs are laid. might influence the escape of sperms from the sperma!.hc­cae and thus result ill the laying of some fertilized eggs and some unf«­tilized ones. This uggestion, howe .. er, is not t"nable as regards wasps, for the worker and queens, bo!.h of which are conceded to be developed from fertilized eggs, are produced in small and large celis, respecti vely, whereas the males may be produced in either small or large ceils. Yet another possibility (Ormerod, 97) is that males might be produced only after the queen'. supply of perms is exhausted. T his sugge>tion, also, i readily shown to be untenable. Both males and females are produ ed in large numbers at the same time, a circutnSL,nce which obvioll Iy could not

xi5t if the male were developed from ggs laid after !.hose which go.ve rise to females, unless !.he males had a milch shaner de .. e1opmental period umn the females . That the lauer is not true is evident from ihe fact that the developin' males and females, presumably developed f m eggs laid at about th same time, may be intermingled in the ceil of a comb. yet tbe whol" lot will spin cocoons at the arne time. If lile deveiopmClltal period for male'S were much .horter tban that for females, the males of such a lot would transform befor lhe fCOlales and there would be formed a sprin­kling of cocoons ontaining male pupae intermingled with uncapped cells containing female larvae; and ,uch i. not the case. Furthermore, surrow"l­ing !.he zone of cells containing male and female pupa in any comb there is always to be found a fairly wide zone containing eggs and larvae in all stages of development. This zone f brood, SO fnr as present e"idence indicates, produc"li both males and females. T Il"re nrc thus female-pro­ducing egg laid after m:my males have completed tlleir development an

f 168)

Ilt01..0G¥ OP Tltlt VESPINAE 169

emerged. This fact alone precludes the possibility that the infertility of male-producing eggs is due to the exhaustion of the sperm upply of the queen.

fu.i.EItCENCE DATES OJ!' 1 ALES AN 0 QUEE!'IS

According to the writer's obsen'ations, new queens and males emerge at the same time. P ublished data, however, are contradictory. Rau (107 and Balduf (7) report On nests of Dolic"()'IJe .. ~"la macuJaJa in which most of !.he queens emerged befor" most of the males, and F. M. Duncan (51 reportS the :;;ime thing for Vespllla t..ugaris. Many more observations need to be made before reliable conclusions can be drawn. ollection data give lil tle or no aid, , ince there is enough casanal variation to obscure lbe in­formation desi red. Furthermore, the radical differcoce in behavior of queens and males militates against the securi ng o( reliable data from col­lections. The males fly about a good deal a ft r leaving the nest, visiting flowers and watering-piaces, wh«eas the queens go into hibernation almost immediately a fter mating. The males therefore are much more likely to be secured by tbe general collector than the queens. Hence collections con­tai n many fall males but too lew queens (or the collection dates to have any significance.

On the oU,er hand, one thing is certain, namdy, tha t new queens and male. usuall v are produced much earlier in colonies of Dolichovcs~ula than in colonies o( V csp .. la, at least on the Pacific CooSL Fo r the substan­tiation of !.his statement, collection data pertaining to males are as valuable :IS anv other data ; for if mal are being produced in a c lony, so also arc qu~s. Tbe following colleclion data show clearly the differeDce between

sspllia and DolichO'llcspu/a ill respect to ,nate and que.en production. Mal . Imve been found in fair nunlbers in a n.s! [D . or"wria as early

as Mav 30 in the Santa lara Valley. Male production . and h"Dce also queen production, in tbis species is in full swing in !.he Santa Oara aUey by lbe end of June or early J uly and is largely over by mid-August. Males may be taken, however, for several weeks after this date. The latest coi­I nioo date for a male of this species seen by the aulbor. lrom any locali ty, i October 1, 1923. The , pecimen in question is from East La nsing, Michigan. There is 3 male of D. 'Ho~/ata from ranbrook, Bri.tish Co­lumbia, in the Stanford collectioo dated Novcotber 1, 1919. Two males o(

. "Jocl/lata in he author's colltclion wer" c Iieeled by T. H. Frison at Mahomet, tIlinois, on clober 29 . 1911 . Such late dates. howe".r. which are dates 01 capture, may r 'present ind ividuals wl'ich have lived for somt time after emergence and are not, !.herefore, indicative of dates of produc­tion. On the o!.her hand. it seems probable, f rOlll published records, that colonies f D. moculaJB endure longer tball lbos<: of D. or'fluria, and there­for. produce mal and queens later in the ye.~r. Or the .xplanation may

( 169 J

170 NOltTn AMERICAN VESPINE WASPS

be as follows: The establishment of n w colonies is ddayffi in eastern oiled tates until May. (In cwtrnl Califomia they may be establish.d

as ... rly as late March. T he season of male and queen production is cor­respondingly advanced from midsummer to fall . Hence in colder regions the difference, in respect to male and queen production, between V .. pula and DoliehovC$pula i coDc ... lffi by the effect of ecological influences.

The earliest date for the production of males of Vesl",la of which the author has any record, except for a group to be discussed presently, is one of Juoe 16, 1923, for a male of V. rufa Br. ill/mn.dia, taken by J. A. Kusche at White Horse, Yukon Territory. Canada. This date seems to b. quite exceptional. The next earliest date is August 2, 1891, for a male of V. , ufa var. AmerieQ'14 (= V. rufa var. Qeadieo . according to Bequaen. 14), bearing a Palo Alto, California, label . Aside from Lhese there are no specimros bearing dates earlier than September 16 in the material accessible LO the writer. Most of the males of species of Vupuia bear dates in OcLo­ber. The latest date ior the collection of normally producffi m.,les in the author's collectioll is December 22. 1929. On that day, it being warnl and sunny. a number of males w re set!n Hying about and alighting on

egetation about a seepage area. few of these which were captured proved to be V lSl'ula pl1ts,),lvalliea.

Ther are several males of the saule pecies in the collection.'! available to the author \ hich bear htbels dated in the early spring from March 27 to May II. Mo t of these are specimens which were takeo in the spring f 1924. II seems probable that all the e spring males we.re produced in over­winterffi nests. ne such nest was b erved by the author from January to J line 1924. It \'13$ de6nitely producing males in April of tlmt yea r. The author also has twenty-three !TUlles of V . moeuiifrolls taken by F . W. Walker at Rock Springs, Florida, on April 7. 1923. These also pr bably represent males produced in an overwinter I nest.

MISCELLANE US

;\Lu."TL'IANCE OF' O.IB FOR BROOD PaODUCT!O';

Among the social acti vi ties of the workers are those 0 f building new cells to receive eggs as rapidly as the increase in the numerical trr.!lgth In

the colony permits and of renovating old cells rendered available again .by the emerg Ce of adults or by the dealh of existing brood. Eggs which fail to halch and larvae which die Cor any cause are probably in most cases removed promptly by the workers. That dead larvae are disposed of in some manner is ev ident from the fact that, though nearly every nest of any size contains scatterffi cells in which are 10 be found brood of a rage differ­ent from that characteristic of the portion of the comb in which they occur, it is extremely rare to find a cell containing a dead larva. Ormerod

I 170 J

BIOLOGY 01' Ttn: VESP1..NA£ 171

(97) states that they are auTiffi OUt of the nest and dropped some distance away. Pupae and unLratlJlformffi larvae which die within their cocoons appear not 10 be detecttd by the workers and hence arl! not removed. rna nest oj Dolichuvc.tpula arc"ar;a colJectffi at Cupertino, li foroia. on Jlme 28, 1928, two cocoons containing dead larvae which had been killed by a fungus werl! found . Their dry and shriveled condition proved that they had been dead fo r some time, et the cocoons containing them re­mained in . ct. Ordillar~y cells with uch cocoons are nOt u ed again. That sitch cdls may sometimes be used 'IglIin is hown by the following: In the o\·erwinter.d nCSt of V. pensylumica discussed in the preceding section t. p. 170), there were found two cells contailling d .. "p within them male which had failed to emerge. Ahove the cocoons of the dead ma les, in tbe >Ame eells. were typical meconia left by transforming wasps. This indi­cates clearly that after the death of the aforesa.id males these two cells had Ill,en lengthenffi, eggs had been laid in them, and an additional generation ol f young had been brought to maturi ty in them.

\\ 'bene,er young wasps emerge from cocoons and leave, as they do. ragged rims of cocoon projecting slightly beyond the mouths of the cells. the worker already pr sent trim off these rugged edges and remo"e from the celli any I e uebri5 which has been le(t bellind by the emerging wasps.

0 111 times the workers completely cut off all the projecting portion of the e \ hich caps the cell . At other times the longitudinal part is Iel t to iorm a part of the cell wall. When such cells nre used again for the rearing of ncw larv .... the workers may add paper to the rim of the cell beyond the si rip of cocoon material. A portioo of such a cd.I wall Ihen COmes to coo­,i,t of alternate bands of paper and cocoon silk.

t.JCCESS10N OF BROOD J N THE C OMBS

. \ 5 previou Iy notffi, each comb io a wasp nest is begun as a nucleus of one cell attached to " cord-like pedicel. Around this nucleus new cdls are addffi more or less continually throughout lhe season of growth of the comb. Other things being equal, the DC'V ells are added in equal numbus Gn all sides of a comb. the comb thcrdore steadily increasi ng in diameter and ttlaintaining a circul .. r fo rm. Th. first cell. or first few cells, are pro­vided with eggs shortly after being started, and an the later cell have eggs laid in them as soon as tlley ha\'e been buill large enough to hold eggs. A. Ion as a comb is being enlarg ·d. lhi method of building and l-gg-Iaying r . ults in the outer cells of a comb always containing eggs, the cel l just back irom the edge containing lanae, and those farther from tbe edge containing pupae in cocoons. Xow as soon as a pupa transforms and the n . w adult emerges from its cocoon. the ceU which it occupied is provided '''th another egg by the queen. This means that if a comb be large enough

[In J

172 NORTli A1t,laRlCAN VESP]N"E WASt'S

it wi ll have. interior to the c.us with cocoons mentioned above, a second zone of cells with eggs. Inte.rio r to these will be a second zone of larvae, etc. In other words, as a comb is built larger and larger it comes to exhibit a senes of concentri .Ones of cells containing brood of varying oge, but in cach complete zone the outermost cells will contain eggs and the inner­mo t will contain cocoons (Figs. 252, 253) . This zoning of broO<J has been noted by other workers (Janet. 7S) . Concomitantly with the increase in diameter of the combs and the maluring of the brood. these zones of brood move outward in a wave-like manner as ripples move concentrically OUI­

·, .... Td from an object dropped iltto a pond. 10 any comb with more Ihan one zone. and all combs except the newest

ha"e more than Ol1e, the wi, from the edge of the comb inward to the nc"t zone of eggs are producing their first generation of wasps. those from the ouler margin of tile second zone of eggs inward to tbe third zone of eggs are producing their second gmeration f wasps, and SO on. I t is thus possible, within the li mit,; noted in the ub. "<Iuen! paragrapb. by noting the zone in which a given cell is locnted. to de termine how many times il has been used, and <:onscquently how many adult wasps it has produced. Using tile dam thus ob tained. the Iota I number of wasps produced in a comb or in the entire nest =y be estimated witll a high d~",e of accumcy.

T he regularity in growth of the combs and the wav..-like succession of brood generations in the c.us is modified greatly during the latter part of the lile o f a colony. H ence its use in estimating the IO t:J.Inumber of wasps produced in a lony is reliable onJy until tbe irregularities begin to be sig­nificant. Thereafter, estimates of tOlnl wasp prodUction based on the zoning of brood become les. and less accurate and ultimately Quite unreliable.

The irregularities in comb development and in th~ zoning of brood re­sult from sevenl causes. First, if tile nest be :ill undergruund nest which is started ill stony soil of uneven texture, symmetrical growth will be im­possible and great irregulari ties may occur ( Figs. 195, 231) . Such irregu­larities commonly have little direct influence 011 .uccession 01 brood except to inlert1lpt the zones wherever a barrier to growth exists. Occa.ionally, however, where the progressing arms of a comb become joined together again beyond a barrier, it happens that the cells 01 the joined edges do nDt belong to the same tone, and hence i rn:gularities which are difficult of in­terpretation develop. Second, after a certain sl7.e is attained by a given nest. new ells are added only occasionally to lbe older combs. Eventually the addition of new cells in the old combs ceases altogether and the combs there­after remain constant in size. Third, eggs, larvae, and pupae occasionally die in snuller or larger numbers. As noted above, eggs and tarvae a~e com­monly taken out of lhe cells by tile workers and the cells renoYated. The dead pupae are left in their cocoon . Cells in which brood bave died thus

[ 1721

!HOLOOY OF TH"E VESPI"N AE 173

come to contain ne" brood of a J ifferent age from that normally present ill cl leir xcne, Or tbey become inactive as brood-pro<lucing cells. Fourth, in Ihe uld"r combs, also, the cells are abandoned in I"rgo number. Fifth. as Ih" weight of lbe lower brood-filled combs incre:.ses. additional suspensoria arc buiLt in the upper part of the nest. Many of these suspensoria block the ceil .. to wbich they are attached and thlls r","ove these cells from usc.

Tn very old nests the irregularities due to these factors may i>ecome so g rcat as to render the Ixlllndaries between zones indistinct or even to ob­literate all appearance of zoning.

In the older combs of a nest wbere the zoning has been broken up and uhliterated , an accurate determination oi the total number 0 f wasps pro­dll ed can be made only by a tedious ""amin.tion of each cell to determine the number of wa.ps produced by it . This, however, may be done with .. ase and certainty. The meconi:li pellets ejected by the larvae at the time DI tran.formation to the pupae aCOlmuiate in the bottom of the cell. These may 1lC! counted. Another method is to count the number of silken linings. i.< .. the number of cQC()on , in the c.us. This will be found to agree with the number 01 fecal peUets present.

OLOR PATTE:R..~ IN VESP (NIJ V ASPS

The color pattern. presented by North American vespine wasps consi<t uf two distinct element,; (Figs. 177- 194). Une of these is a dark color, allll" , t alway black ; the other i light and is most commonly yellow, though oll<'ll white. These pattern .Jemeot,; occupy all the pigmented portions oi the integument " '''cpt certain cry .mall marking of minor importance wh ich will not be considered here.

T he black element appears to constitute the ground color of the integu­melll . The light element is present .. a eries of maculation. which \"Ory i exlent with the. indi"idual. II the maculation. are xtended, the black is c rre pondingly reduced . and vicc versa. Certain centers of black pigmen­tation are rarely i 1 ever encroached upon by tbe yellow pigment. The ('c,mvenie is ruso true.

For each species of wasp there .,xi ts" definite and distinct pattern plan, Ilctcrmincd, first, by the relative position of the permanent enters of black and yellow pigmentation. and, second, by the manner in which tbe exten­. ivlI · of black ;LI1d yellow pigment from the .. permanent centers mutually encroach on the intervening ""'''''_ The individuals of a species. however • • lisplay " great number of indi idllal or definitive patterns according to lIle Iker.., of extension of Olle of tbe pattern elements at the expense of the other.

T he distinctive features of pattern plan are most strikingly exhibited in the abdominal patterns, hence these are used here .. ill ustrations.

[ 173 I

174 N'ORTll A.1U:::aICAN VJ:;SPII'tiE WASPS

In ~ up,,/a pmrylvallica (Fig •. 177-179) each t rgum of the abdomen except tbe last is provided along its apical border wilh a yellow wcia which is deeply indented medially by n triangular extension of the basal black area, and is notched once on the sides by squarish tensions of black At the ides of the tergum the fascia narrows and turns forward along the lateral margin of the lergum. T he fascia thus .xhibit. " continuous apical portion and four forwardly directed lobes. In many individuals, most of wh ich are Queens, the two lobes f ea h side extend forward and encroru:h on tbe lateral hlack notche' in such a manner as nearly or completely to cut them off as black spots surrounded by yellow. The e.'Ctreme definitive pat­tern thus produced consists of a broad yellow lascia deeply indented medi­alJy and provided with a pair of black dots lalerally (Fig. 179) . The median indentation is ne,'er cut off from the basal black portion of the tergum, though it may be narrowed somewhat by encroachments of the yellow. In some individual. the lateral black notch in the fascia may be converted into an isolated spot on only mle side of Ihe tergum, or the conversion n!.~y OCCur on some lergll and not on thers . . \5 a gene,.,,1 rule Ihe m.' cuiations tend to be more extended on the mor posterior segment fi nd to be progressively reduced as tlle base of the abdomen is approached (Fig. 178) . Occasionally th reverse is true, and in me specimens the maculations are "xlensive on both anlerior and po terior sej,'l11ents and are reduced on the middle seg­ments.

The pattern plans oi different species commonly differ much that in-di vidual wasps in most cases may be readily identified by pattern alone. Such idOltificaliou can be sa fely made only ill the light o( kno wledge of the significant features of (mttcrn plan, reoally Il,e manner in which "''<­lension or reduction of the maculauons occurs. Identi5cations cannot safely be made on the basis of the defini tive pattern of single specimens, for the reason Ihnt very nearly id · , tical del;ui uve patterns may be developed from quite different pattern plans. por example, Vespula pensylvanica (Figs. 177- 179 nnd V spula rr1" var. alropilo." (Figs. I l82) belonging to different species groups, each of which has " different pattern plnn, and Dolic"CItI#sfula arMaria (Figs . 186-188), all pr=t, in their most highly maculated indi,-iduals, sentially the me definitive pattern. This point is illustrated by the figures, which show for each of the.e three species the patterns of individuals \\ il1, reduced pattern in which tbe pattern plan is evident, and those of individuals with extended maculations which obscure the pattern piau.

Thus the contentions of leading systematists that color pattern alone is often quile unreliable as a bn is for making spe<:ifie determinalions, and tl1at new speci"! cannot be dcs<:rib<od adequately from the color pallcrn of individual specimens. are amply SUllported by iaet. At U,e same time it i.

( 1741

SlOLOCV 0 ... TlIp. VP-Sl'lNAE 175

d ear that if sufficiently extensive series of specimens arc critically studied and the pattern plan dd.ermined, color pattern will take its prop« place along with other characters of diagnostic value.

COllW ~all.", .IId phylog,"y,-There i a definite correlation betw"",, panern plnn and phylogeny in the vesploe wasps. This is so clearly marked thaI several species may be referred to the proper species group on sight. T'rspula rut" (Linn.) (Fig . 193. 194), and its varieties ill/cnn.dilJ (R. du Buysson). aJropiloStJ (Sladen) (Fig . 180-182 . vidUD ( aus.) (Figs 183-IS" ), a,adica (Sladen), sladelli (Bequaert), and consobrilla (Sauss.), to-

ther wi U, V. all,rlrWi:" (Panzer), V. sl""lor (F. mith). V. sq'",IIIosa (Dr.) . and V. sulphurea ( auss,) . form a well-marked group of species, which Be'lllaert (14) has called the rofa subgroup (o f Vcspula s. I. ) . Ves­tula tNigaris (Linn .) , V. gDNII""i," (Fabr.), V. IIIaculitro/lS (R. du Duys­. n , F . p~nsylt'OIIica Sauss.) (Figs. 177-179), V. Ift<.i.sii (P. Cameron), and T·. koreemis (Rndoskowsky) c; nstitule anoth .. r well-roarked group, It'rrn I by Bequaert the 1"Hlgari.s subgroup, The , ufa group and the fJulgaris !!"ronp are the only two groul" of the genus Vasp .. Ja. (5. str.) which arc rep­re;tnted in the United States .

1)n1y the V . nIta group will be discussed in detail This group is char­a~lcri"ed by the following pallern plan. In pecimens with reduced macu­lat iun.o; the aildomioal lerga usually present apical fasciae with three llllargin:uions and four nnteriorly directed lobes, comparable t those de" M ribed for VCS1lu/" f'tlU)'/t'tlflu:a, though not always of the same shape or lxtent. Til .. '<ten sion of the pattern in the T'. TIl/a group, however, is Iirought about in a different manner [rom Ihat in lhe V .• It41garis group. Tile lobes of the fasciae advance bUI little. Inslead, tausverse, oval, or ellip­tic yellow spots develop just in front of the laternl notch in Ih fasciae ,rigs. 180, 189) . These spaH enlarge laterally and 'entually c nneet witll the I be. of the fasciae, cutting off the black notches as dots. Con-1I<'Ction with the outer lobes of Ihe fasciae is usually made first (Figs. 181. I'll, 192) . the effect being to produce a pattern in wtllch the lateral notches appear to have developed from the median triangular marginations instead

i i rom Ihe anterior margin.~ f the fasciae. The llIembers of the mfll ;:ruup agree generally in structural ch."lr.lcters as well a in pattern plan ; SO

llilT i. no doubt th:.t the agreern It in coloration i. indicalive of lose rdntionship.

Within the group, degrees o( relationship appea.r to be indicated some­what by vnriation. in the pattern plan. V Ds!,ula squall.o!a and I' . sllll'lI""11 hotb have two longi tudinal yellow lines on the mesonolum, whereas I' . ruia, '·ar. vi/11I11 and. '. ruta. lack lhese lines and have reduced patterns.

Cnlor !,allCrn alld caSlt diffrrtlliialion.-Tbe llifferences bctwem worker alld queen wasps. aside from ,i.e difference., and the faikre of tlte repro-

( 175]

176 Noan[ AlIER.lCA!"" VESPINB WAS.~

ductiv~ organs to develop in the 1V0rker, are manifested in color ttero and behavior. The difference in color paLtern i5 geoerally slight. though definite in charnct.,... It has bew ov rlooked by most wrilers, and is not constant in any species known 10 the writer except V espur. sq.Ul."'OS~ (Figs. 189. 190) . The basic plan of the color pattern appears to be lb. same in the workers and queens of all species. The black element of the patlern is mor~ extwded in the workers than in the queens and the ycllow r white element is correspondingly reduced . Thus in VespukJ PrnsY/fJlJrt;ca the yel­low fasciae of the abdominal segmen , as exhibited by typical Pacific Co •• 1 specimens, are commonly narrow and Iri-emarginate in the worker, as in Figure 117. but are broader in the queen. and the lateral cma.rginations .re replaced by isolated black spots, '5 in Figure 179. T his difference is not (Oostont. since indiv idual workers often have One or both of the late.-..J emarginalions replaced by a black spol as in the queen, and occasional queens exhibit the reduced pattern typical of the worker. Such variations in color pattern are corrcl,\ted to some extent with size. ina,m uell (IS the more highly colored workers are apt to be the larger ones and the less highly colored queens are most often the maller ones.

In Vesp"/I> squamo.fO the difference in color pattern between workers and queen i- the greatest in all merican wasps. The worker and qu"""-S were at fir-·t described as distinct species. and nOl for many years did their cospecifi identity become known- 1n this speci Ihe patt"rn of the queen (Fig. 190) i almost entirely rich orange yellow. [he black being restricted to a na rrow ba aI band and three mall indisti llct sPOLS conoected by a faint arcuate line. T be nbdom of the worker on the othe r hand (Fig. 189 ) presents a series of narrow, a1tema/.ing. strongly contrasted black and yel­low bars. There is no intergradation in color pattern between queens ami workers in this speci' .

TERATOLOGY

T wo specimens o ( VespuJa pensylwll ica, one a male, tbe other a worker, in lhe writer's collection. present a comparati ely rar ' type of defonnit)' "'''pressed in imperf t segmentntion o f the abdomen. Janet (75 ) records a similar C!ISe in a rnale of V. !'Vfa, and Chapman (26) records one in a worker V. gO">lollico . Ln the male in the author's collection Fig. 130) lh, right-hand portion of the fourth tergum is partially se[,.rated from the rest of the teq,'Utn . The left-hand portion i. apically in rlircct continuit), across the middle with the right-hand portion of the Ii!th tergite_ The left-hand portion of the fift h tergite is completely separated as a distinct pil'CC. Th~ worker specimen (Fig. 129) displays II similar deformity affecting the third and fou rth terga. The sterna in both these specimens are normal. The cause of uch irregolaritil!> in segment.tion is lIS yet undiscovered .

[ 176 I

LITERATURE CI I EO

1. MOM1£AD, \ VILLlAt.t H . 111\ Proposed ".tural AfTangemmt of the H)'mmop& Lerou.s F-amilics," PrMltdings oj 'M EnJOD.olog;cGl Soci~'Y of rYashinglOJl, I : ~99. 1888.

2. - -. .IClaui6catian of the Fossorial, Preda~w, and PArasitic. \Vu ps, or the Superrornily \ '''poidca,'' T", C.IKId;mt EOI.meloO"', Vol. XXXlI, 1900 pp. 145-55, 18>-88, 295-'16; Vol. XXXIV. 1902, pp. 711-88. 131-37. 16~ 203-10, 219-31 , 268-73, 287-91 ; Vol. XXXV, 1903, pp. 3-3, 39-14. 95-107, 15.S-58, 199-205, JOJ-IO. m-JZ; oL XXXVI, 191M, PI'- 5-9.

J. - - . "P~pcn from the Rarrinlan A1 .. ka Expe<!iuoll. XXVIH. Hym",oplC!1l," P'.cwJ'"flS a' I~. WQ.lhingl"" (D.C.) Acad ... , of S,i4.us. 4: 117-274; pl • . 9-11. 1902.

4. - - . Remarks on Vu~ doryloidu and fl. munaarittiD. Procudings of Ihl £1f­," .. ol.git.1 Son.ty of WasJri"flIOIt. 5 (4) : 2801-85. 1903. (In ~{inuI",)

-. - - , "Pro\·espa., til N~w Gmw in w VespidAe." fintDmofotlkaJ Nt"fDI, 14 : 182. 1903.

6. BAllRCo . W . ) . ·'Not .. 00 tbe Nett o.nd Ibe Popul.lion or a 10DY! f/. Sf!tJ diubolica," ; .. ",.1 of E,o .. ""ic Enlomolorn. 14 : 509-10. 1921.

7. nALDUF, W_ V. "O"'erntjons on Podtll""i. '-;olo"i,,,,,.,, (Lcp.) (Sphocidae) illtd Ytlpw/a Rule.lata (Linn.) (Ve!pidae) ," Tit, Ca1Hl.ditlll Enlotna/ogi.st, 68: 137-39. 1936.

8. BA t-t n&. ASD WAL"fI!Jl.. f.lNOt6 01] a Ne!l of Vupa gtr'lflall;cfJ/ ' Proctf1Uinf)1 (Jf I~, e.'omological Soci." 0' W.uJri • . gl.R, 6(2): 70-71. 1904. (l.o minutes of .ociely.)

O. Bl-guAnT, J, OIO n the Occurret'lCC" of Vupa QustrUJca Panzer in the Northeastern States," BoII,II" "f th. B .. okly" Enl_.lalJical Soo.' )I, 11 : 101-i. 1916.

IU. --. "A Ren,;on of tbe Veopidae of the Belgian \Aogo Based on th< CoI­lect i.,.!! of th~American Mll!CUlD Congo Expedition, with a List Df Etbiopi-.o Diploplc:rou.$ \V~5P1, " Bullltin oj Ih~ A"un~cu" MK.lIHIm of NnJIln.d Hulo'7/. Vol. 39, An .. I, pp. 1-384; 267 Itxt fig., Pis. I-VI.

II. --, .lHym-eooptcn. Collected Near Botton., M;us .. with D~ript iOD of i'l

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14. - - . "A T<ntative ,nopsi. of lb. H0<nct5 and Yellow-J.ckets lVesplnae; flymen"",.r.) or America,' e.lo"",l""i,o A~ca"o. 12( .... Z) : 71-138. 1931.

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[ 1771 [In

178 NOH.Tn A'MERIC'AN VI:::.SPrN£ WASPS

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text fill.· 192-1. 20. BUDL.EY. JANU CHL5TKL "Tb~ Taxonomy of the ~hu .. rid \V .. ~ Inc1udinc a

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26. CUAf'KA..'i, DR. "Cuncemilll a Teratologic \Vorker o[ fl. PU'1fUJ"i4{).~ PrOtlM­I/Igs of I~t Enl ... "logiral SOt;'ly of L •• dun. p. xl;", 1917.

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[ J8J 1

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11 3. Rrrcnaa. P AUL O. "Tho Ext«nal ~{orpbology of IArvlll Bremidae and K<y to Certllin Specie! (Hymenoptua)," A,ul4/, of Ihl En/O'tt{ologicol Socii., 0/ Amrriea. 26 ( I : 5J-6J. 193.1.

l ila. ROllW';a.* S. A. ''Vespoidea and Sphrcoidea" in ··Guid~ to the lns«ts of Con­"OCUcut, Part ilL Th. Hym<l1optera. ur Wup-I'k. Inl«" of CoDD<CtiCll~" C..,nrClkHl G •• logie.land Naroral Hislory SUM. .. ,. Bull. 22. pp. 1iOt>-97. 1916.

114. ROIIWU, S. A., AND G AlIAlf, A. B. "Hori!mo10&)' of the Hymt'tlOptuotlS \VinB," P-.u.di.g, 0/ I •• Ent<>mological Socicl, ./ WllShiJtglon . 18: 2(1..76 ; plJ. I-J. 1916.

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116. ROTUn, l>{AX. "f/upo cr.~'D L. in Pennsylvonia (Hrmen. ; V<spidae)." E.,.· H,ol.gical N<ws. 48 8) : 215. 1932-

117. ROUIJAVD, E. '"evolution de I'ulitinn chez Itt Vespides ; il~~ bjoloc;iques ~tJr lei Guepe. aociaJ9 d'Afrique du ge::IlTe B,loJfga.s1,,. ~u.!il.," Compttl Rlrtd~1 AcGd,mir d. Seima. lSI : 553-56. 1910.

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11 9. DIS. !;Aussua.r. HL"Ol l. trfldts SNT la !ONlilt, du VU/"1'Jn. 2; JflMcgrathic des C .. ilu SoC'ilJlu, uw d, In .rib" dll VnpLtns. 256 pp.; J7 pls. ' Mauon. Paris: J. Ke.>m3n, ~neve. 1853-1858.

[ 1821

UTER..ATUll.£ CITED 183

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[ 183 )

184 NOR.TU AJdt:, IUCAN VESPll'lo'"E WASPS

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147. rUllN.R. C. 1-1. ' 'Tbe Worker. of V",., ,,,,.liM Dr. R.....ml. in Coloration tb. Male .. A Correction," PI)·,h., 15: J-J ; pI. 1908.

148. LRJcn. WE.INER .. IDie .Mundwerkzc:ugc: der Sphec:idc:u," Z~ils,hnfl fiJ,. Mor· pMlogl. IIJId Okologi. (/-. Ti.", 1 (3) : 539-636. 1 92~ .

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151. W.,,,u,,,,,,, F. L. '''Th<; H"ymenQP'o", of /Jinn<><> ..... in Thl 171h R,porl of Ih, Slo', Elllo",olqgisl of Min""ola, 1918. pp. 14>-237, pis. 1-3, "i'- 26-125.

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156. WX~r --. "Note on Vil/'O ndgw/' BUl/~tjK 0/ 1111 B,.ook/Y1f &tomo­IO~QI Sod<ly, 10: 41. 1915.

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1.59. --. '"11Jc Pa.l1l5itic Aculeata. A Study in 1!,vo1ution." Procttdillg.l of til t "hn .. ricClII PhiJo.l'(}/,hicoJ Socidy, 58: 1--40. 1919.

160. WH&£l..U:t W. M., AJ4D TAY'LOa, L. H . ·'-Vupo. grcl jca Rohwu. a Parasite or v,.t/'O Jiaboli«> d. Sa.55.c.," PlY''''', 28 : 135-44; pI<. 6 arul i. 1921.

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163, WOOD, J. G. 1",,,,. al Hom • . 670 pp" 700 ilIo •. a .... Scribner &: Co., Now ark, 18i2.

164, Z-Ulll.!UIt., J. L WNotes on Some Parasites of Vt.jHJ/~ Jounl41 of tI.t Nrw Yor. E,.tottSDloglcol Socitt, . 2: 91-.86. 189-1,

[1 84 J

UST OP ILLUSTRA nONS

PLAT" 1. Rea o[ I'fs/'Ula prnsylvl111;ca, queen Pig. 1, Lateral upect; Fi!). 2, Cephalic .. p<:ct; Pig. 3, Antenna; Fig. 4. CAudal aspect with mouthparU rrmov<d; Pig. 5, Dor",1 .. peet; Fig, 6, Ventnl azpcct of head with mouthpa.rtJ removed

I'LATE rJ . Read shlcron o[ flupulD I'.nsyin,"ica, queen Fig, 1, Inner as~t of tatenJ halr; Fig. 8. lnm:r ~~t or ~5lerior half; Fig. 9 S..:tion tbrougb .. · •• 11 of antonnal socket; Fig. 10, Lateral "p<Ct with s~d<: tut aW3)'; Fig. 11, lnnu aJpect or front haH

f' !.An In. v up"/o {>clIsylvanica, head structures and mouthparu of quem ,

Fig. 11. Caudil l upec.t oi phar'}ox and adjoining :ll.rUCture!l ; Fig. LJ, BlUe or n!.dUm. with mUICIc:a; Fig, U , lnnu aspect or tgtr or bad aktlc=ton, and ce.rta.in muscles: Pig. i5, DetaiJ of sensoria and plC3I pecten; Fig. 16, Cephalic Iltpttl of labru!U and p~tynx; Fig. 17 r MeW a!~ ?( ~jlla and its mU.5Cwature; Fig, 181 Head skeleton and mouUtparlJ LD sa,gltlal Jle.ctiQn

PLA1" IV. resp"la p~nsylvanica, head structures of queen Fig. 19, ltmcr right balf of head IkclctO!1. Y;'ith certa~ musctQ; Fig. ZO, Musculature of Olntt'Jlnae otnd rnoutbparn m 'ltt!, front View ; FlU. 21. Samt:, from bchlnd; Fig. 22, Right ,"""ilia ; Fig. Z3, MDXUlaturc of ant<nnae, phar)'llX, ~ c:emin mouth~rt.s in side: 'Yicw

P LATE V. V~sp,,/11 p""syivanic.a, cepbalic stomodaeum, mouthpa rts except ma1ldibles. and associated muscles of queen

FiR. U, I:..,e,.," aspect; Fig. Z5, Sagi'ru section; Fig. Z1i, Lateral a5pret, without maxilla

P LATE \ "1. V.spula pfflsylvallica, derails of mouthparts and muscula-ture of queen .

FI'g. 17, !.bxilla, inner 2.,pect; Fig. 28" t..abio-ma:dlb£r compt~ audaS ..poet ; Fip. 29, Detail! of maxillary ll5<uatur< ; F.p. 10, O<toil5 o[ labium: F.g . 31. S;une i Fig. 3Z, Cq>balic .. poet or 1ab, .. maxilla.ry <00l­p1ex (except rigbl mulll.)

P LATE Vli. Vupllia p.",,)'/vallica, moutbparts and musculature of qu...n

Fig, JJ1 Ce.ph.uic ~l«t or left mandible ; Fig, .H, Caudal aspttt or !Qtut; Fig. 35, Detail. of labium : Fig. J6 , Lateral aspect of lef, nwtdibl.; Fig. 11. Derul. of labium

PLATE Vill . Vrs(tllia I'tnsylooll;ca, thorax of que"" Fig. 38. Iullic "PC:C:' ; Fig. 39, Calldal .. "",t ; Fig. 40. Vett<n.1 .. prel: Fig. 41, DorRaJ "'p<Ct ; Fig. 42, 1:...=1 aJp<C1

PLATE IX, V sf"'!a {>c'sylt'rJnica, thoracic struclur<>s of queen Fig. 43, Sagittal 5lction, right hal!; Fill, «, Some, with cenain parts .... moved

1185 J

193

194

195

1%

197

198

199

201

202

( ISS

186 ~ KTI:J AlI£RtCAN VESPIN£ WASPS

,,,g .. P LATE X. VrspuJa pCIIsylvaniCII, thoracic structureS of queen . 203

Fig. 45, [nner "-'Jl<d of right half with prostunwn and base of m ..... 'utW apophysil rClDO"Jed; Fig. 46. Same. with ptOpectU5 and scutum r-mlOYcd

PU.TE XI. Vespula pensylVIJnica, thor.cic structure' of queen 205 Fig. 47. Detaii.J 01 right .id • . inner upect; Fig. 48" Section of right tho-racic. wall anterior to propodtal spiracle; Fig. 49, =>cction of left thoracit wall between pasuqul;a and propodcum; Fig. !J(J. Scutellum, m<.opo"­""- and r<!lion of w"'" b;ue. viewed from left

UTE XII. f' upula pellrylfJanica, thoracic structures of quem . 206 Fig. SJ, Octaib of right w!ng reg~ inn~ "'pert), with 3.i~ociated pam of body w;a\I .nd nluscJe.; FIg. 52. S.me; F,g. 5J, ~e; F,g. 54, Some

PI. ATE Xl fl . V.spuia f'tllSy/'KmiclI, thornci. structures of queen 207 FiJI. 5;. Oute:r vicw or artiallar rt:giOll of leCt fore wing; Fig. 56# Ventral Yi~w of tegula in Si lO; Fig. S1i Left third uiUary or the frorn wing. with its intrinsic mUKle : Fig. 58, Internal view of articular rqrion of iore "';0 ' ; Fig. 59. La',ral upeet 01 mo"!'O"'p/tr.lama in .iru ; Fig,6/}, I.tm<r view of articular rqion of left fore ''''''In&

P LATE XIV. V~<p"la pellsylvaniea. prothoracic structures of queen . 209 Fig. 61, PropedU5 and leg base, in postCro-dorul view: Fig. 62, S;une. in pOOtero-.. "tnl view; Fig. 63. Inner "'J)eCt, righl ~I f of propect ... <UId a.rtkulatiou with bead

P UTE },,:V. Vrsf>llla p.rosyivllmca. thoracic structures of queen 210 Fig . 64, Rish' prop\euron, righ' CO"". ond musel .. ; Fig. 65. Delail. of first thoracic: spu-ade; Fig. 06, Left IISpec.t o( proslc:rnwn 3d. its muscles; Fig. 67. Postcro-dorlal ~t of propeclu.l, wit.h certai,D muscles remo,'ed : Fig. 68, Det:lil. of "",ond thoraae .piracle

PLATE _ 'VI. V,sp"la ptllS}'ivaniea, thoracic structures of queen . 211 Fig. 10, nterior pen or thorax with haH of proootum removed: Fig. 70. Anterior pan of me!lOtho:ru, sbowing indirtc.t wing muscles: Fig. 71 , Ria:ht tort' coxaj showing mlUCie insertions . Fig. 12. Riehl rore coxa ;U'ld mu 01 .. 01 tbe leg b;ue ; Fig. ; J, RiJ:ht ",pee' of propee'WI and usocia .... nuud ..

PLATE XVI l. VcsP"Ja p.nsyh'allica, thoracic stntctllr of queen . 2iJ Fig. U . Ce\>halic upect of thQrax; Fig. 7;. Oe",i" of propode.1 .apincle ; Fig. 76, Ve:nt:ral view of prostunum: Fig. 77. uud.aJ aspect of IIlClOlhorax; Fig. 78. Propodeal Ipiraele and ill muscle>

PLATE XVllI. V.spilia p"lSyifJllnica, morphological details of quttn 21 __ Fig. 79, lnner aspect of metatl10rax and abdominaJ ba..~ as seen from in fron'; Fig, gO, C.ud.:U :upect of ventral hall 01 melllloorax and prop<>­d.um; Fig. 81. Detaili 01 .rtieulll,ion between m<:bltboru and abdomen; Fig. 82, Spatub.te &e:tae on glouae

PLATE XIX. Detail o[ morphology and nest structure Pigs. 8J-86J Diawms or paper "shell .. " from wall o( V t'J/,KItJ ~n.ly/­w"ica bell; Fig. Hi, V u/)lIid p~n.s')lltr(:tp.i.ca, hnmu h on hind wing of queen FiO. 88. Detail, of l1\60thUru. and its mUJCubturt. in caudal ~ Fig. 89, Do1i~hotln" .. la art'Naria, iuitiOll thr~ cell, of :a. new comb

[ t86 J

215

L IST P lLLUST1lATfONS

F'~. 90, Dolidu'Ir*rJ"Ja mOnda/G •. ,ide view of comb from qUCt:fl f'ldt: Fig. 91, Dorsal aspect.o[ sante; Fig. 92, V,sfivla Pf!71$Jlt.v7Iica. dc.taib o( ~ot.bora:.c: of queen to antero-dorsaJ epea

187

PLATE x....X. V"fJ,,/a pfflsyifJanica, leg and wing tructures of queen 217 Ftg . .93, Left hind c:axa .Iod its mu.fjcl~s : Fig. 94, Sue or right hind wing .~ Ill, muscles; flv- 95, [..eft alIddl, cox. .and ill Dluscl .. ; Fig. ~. Ri&"h' hUM ~mg base, direct Wing musclea :and axillary Jderites i Fig. 97, Base of lef, hInd 1<Jr and mIBC\es

PLATr.)(XL VesPU/1I I'rnsyiw"'ca . structural details . 218 Fig. 98, 4ft 10« 1<, 01 qu<en in,oolli.1 aspect ; Fig. 9!', Same. in ccpbalic .. peet ; F,p. 1{)(). Hood of m..le In cep~hc "'pee'; F,U_ 10/ Oi, .. 1 'wo­thirds of ng,h, hind Iq of queen; Fi~ .. J!!l, Righ, .n"""'" c\";"" of !lime; FIQ. 103. R~ltl an'<nn;a of male; " Ig. 1~, Rlg"h, middl, leg of queen' FIg. lOS. Ba.. of Ief, lruddle leg of qU<dl; Fig. 106. Rillht hind 1"11 01 queen

P un: x...XU. Wing venation and demil o f abdomen 219 Fig. ]m, Vtsta erabro, fore wing with Jurioun nomenc.lature· Fig.r.l08-109:.. VupwfJ ~tll.SjhvltkC). queen, fore and hi!1d wings with n~ature of K~; Fill. LlO. Some, b;ue ,!I abd""","; FI!J.111, Same. detail ell huIIa on cubltl.l5 16 of (ore Wlng; Fig. 112. V U,tt/4 svlplturea fore wing with n~claturc of TiU,. .... d; Fig. 111. D,4/khO't'n~ au:Wria. fore win, W1th nomenclature of the Comstock-N«dham s)'$ttm as intupuled by Br.adb,y

Pun XXliI. Vespula pelU)'ifJtJtliea, abdominal structures of queen . 220 Fig. 114, loner ""pee' 01 abdominol n."",; Fig, 115. VCll,llLI aspect of :lbdomm; F,g. 116, Latoral .. peet uf abdomen; Fill. 117, Inner upec t of left IWf of .bd<>m<:n

P' . ' TK· XIV. f/ss/,ula p.II,y/wniea, sling mechanism of tbe queen . 223 Fig. 118, Later.1 aspect; Fig . 119. Dor .. l ... """; Fig. !20. 0,\3il. o f do".1 aspeel; F'g.I21. Ven,ral upectj F,u. 12l. Lef, ""pee, f poison duct ~ sting IJulb in I.itu i Fig. W, Lateral aspect of ~ developed froOl IlJD~ ~en!: Fill .. 124, u.ten.! aspect ;after ,removal of ei"lnh I<~; FIf/. US. TIp of .una lancet ; F,g. U6. E.xu-em,ty 01 same. a-reotly <J1I.rg<d

PLAn XXV. ?Ii ceJlaneous demils of morphology 225 Fill. lZ7. V uplll" /¥n.rylt'Olfico: abdomen or male; Fig. 1~, D()Ii.r/ro'(-'opula ".QN4Itlt antenna. ol.m:l~~i Flg.1Z9. J'up*/o ~h'Onll(J. abnorn:tal 0I0i>-­dom<n 0 worker.; Fill. LiIl, Same, ahmm.W abdnmen of male; Fig. 131. S.Ime queen. detad." of abdomi.nal ce.rga; Fig. 132, Doiic.nutouiu/IJ. D,~n.aria -, left a.ntenna of tmlt:

PLATe XXVI. lIespw/a PCIIsj'lvQI,;£a, abdominal Structures of male 227 Fig: )1.J. ,POS!'=nor .end of abdomen lrom b<low; Fi(/.lU. Do .... 1 ~t 01 S<"OJtaha m 'Silu : Hg. 135, P~1eri ,. end of abdomen from above' Fig. 1.16 Latef!J aspect of gel!italia in situ; Fig. 137, Vmtral upect of fuSed eighth and runtb alC.ma: F.p. 1.18, Ventral upcct of fItnlmlia in situ' Fip 1.]9 Dorsal aspect of gerutaJia.: Fig. 140, Ventral aJpt:ct of g'eltitaJia" Fig 14/ yt~ta! aspect Of ~enit.Ii;t . partially cXIKJ5cd; Fig. 1-12. Late:r;aJ' as~t of' gen ll~ha, tully upos~

11871

188 NORTU AMERJCAN ESP':-:-!! WASPS

1'A.0t.

PLATE xxvn. Th!tails of mal~ genitalia 229 Fig. 1#J. V"p. lo p.nrylvGni.<0' r1,b. inocr aspoct 01 cooocardQ '!J'd 10~<>­otipcs: Fill· J44 Sa..." 1.1. 'Mer upeet 01 I"""""rdo and gonosb~, ,,~'h ~ in lito; F:g. 145, Sa.m~ • .innu .a.sptCt or sarntA and \'olsellA; Fig. It6. Same. ventral aspect or pc:nlSj Fig- 147. ~ d~l iUp«1 ?' penn ; Fip. 148. Sam., right inner upoct 01 ,,!,,~PCS .wlth "'I""",! IR P""": F'll. U9, Same, proximol uP"C. 01 ,,,mtilia. : FIg. 1~, Doll(h"",~pola a,.~IIDIi.a, riaht 1JlIlU aspect or I000ca.rdQ ,and g~LJpes, WIth peDJ..5 10 situ; Fig. 151, Same~ ~onat aspect or gegnaha; F.g. 15Z, Same. '-tent UIl«t 01 goniWia; Ftg. JjJ, Same, loft Inner .. pect 01 g?""""rdo ~ gOllO:!l ti pes ; Fig. 154, Same. ldl inner peel.of ~ono.tfPC With I(luama 10 place; Fill. J55. S __ <eotnl upoc. 01 gerutllIA

PLATE X),,'VllI. Morphology £ Dolil:llo'Utsplll4 and o f Vespa 230

Fig. J56, D. a' ..... ria, tho,..,. 01 quem' Fig. 157. Same, cepllAlic HPOC' of lKad; Fiq. 158, Same, lKad 01 male; (:;g. 159. V. ,~ob~o. dorsal "poc. 01 male a<rUtalia; Fig. 16Q. Sam •• -.en"",! .. poet 01 g<llltallA; F,g. 161, Sam<:, latc.nl1 aspect of ~nit11ia; Fig. 162. Same. Idt lnDl!r aspect of gonocardo and so_tipcs : Fi~. 163, Same, right inner upoet 01 80n0cardo aM gooo­.tipcs. with penis in litu

V _L_ '_~31 PLATE; XXlX. Morphology of espa cr"",o Fill. 1M, Tho=< 01 queen; Fig. 165. Antcnllal llIIa;ellum ol male : Fig. 166, Qu<cn, cepbaJlc .. pect 01 hoad; Fill. 161. Same, doruJ .. poet 01 head; Fill. 168, Male. copllAl;c "p«:t 01 head

PLAn: XXX. Vespula pcns.I'lvullica, inunatur~ stages and miscellaneous 232 Fig. It/Y , V<nlral .. peet o( qU«Il pupa; Fig. 170, Vcnt~ ... peet 0.1 queen larva : Fig. Ill, Samr. cq>bahc aspoc. ollarnl bead · FII/. 17Z, Dlp~ (1) egg Itt2ebed '0 nest; Fip. 17], La.era! .. poet oIVlIpo/a egg ~OWUI8' att2chmcnt.o wall 01 cdl; Ftg.174, Lateral U\l<Ct 01 queen Jan. .. ; FIII.IlS, SI!Ction 01 "" shown in F'1g. In; Fig. 176, Latenl "'poct 01 quem pupa

PLATE XXXI. Abdominal color patterns 233 Figs. 177-1,9, V"polo #rUY/.VllkG .. FillS. 18(}.-1$1, V<S~~/o ",/a (lJ'!'Ll , var. alrotiJo .. (Slad",,) ; FillS. J.8J-18S, VfS~/. "'f. (Lann.), var. ",dNa (Sau ... )

PLATE X,' , 1. Abdominal color patterns . 234 Figs. 186-188, DolicAowspoJ. ",,,,aria .. Fig •. 189-190. ~ up.1D sq."moso (= V. coro/i ... ) ; Figs. 1P1-1PZ. Vupul. sulph"r", .. FIgs. J9~194, V,,-polo "'/.

23~ PLATE XXXllT. Lobed combs o i IIcspula I'ensylwllica . J

Fig. 195. Comb from 0\ Qut in Itony gro1lDd; FiO. 196, nth from 01 nesl in homogonous .011

PLATE XXXIV. Queen nests of 11 "'I»<la pcnsyk'fJnUa . 236 Fig. 1'71. Nes, built in dark shed; Fig. 198, Same, ventral view; Fig. 190, Same. lOngitudinal sec~oo; Fig. J'OO, Nut. frO!D roden~ burrow: Fig. 2IJ1 . Same, ventral view; F,g. 202, Same. longitudinal . ecuoo

PLATE x..'{V. Dol ic/ltnltspula nests and n .. t details . 237 Fill. 2OJ, QU«:I1 O<li. 01 D. 1tI4&.lalo (1) ; Fil/. ](H, (rr<gUw \'lUI ?I c.u. from I1efl or D. aTIMN; Fig. lOS. Queen fld:l of D. ar,rtarr4: Fig. 206, Same. ventral view

( 1881

lor 01" lLLlt"STRATIO NS 189

,"". I' l.AU XXXVI . esls of Dol;c1lo'l"s/mla and V"spula 238

Fig. 207~ Queen nest ~f D . . ~"nari4. longitudinal sectiOll: Fig. lOS, Small n<>t 01 V. "'Myiw. ",a: F.g. 21)9, Sa ..... ven.",] view ' Fig. 210 Sam<: ve:nmt \~iew with I e.r oan o( 1NilH removed to Ihow t1~ COtJ1bs • •

P LATE xxxvn. Nests of 11 tsf'lIla spp. . 1.19

Fis· 21.1, Small nett of V. tm.tjitWlic(l. longitudnm! a.eclion· Fig 212 Worker-laid 0Ji$ of V .... llIonl in romb edit : Fig. ZU. Ou~ I~' oi V. f~/g4ri.s; Fig. 214, arne. lCflI'itudinaJ Jectloo

(' !.ATE XX.x VI! r. ,\!:\ture nest of Doiic/lO'Ucspula are.loria Fig. 21j~ I!xttm:aJ view; Fig. 216, Nest lnte.rior

. 240

I'UTE XXXIX. Mc'<lium-sized nest of Doliclt".lrspuJa orellana Fi . 217,. Side YiC\Io ; Fig. 213, Ventral view

. 2~1

IJLATE X L. Mature nests oi DoiicJlovu~ula 1nat'lIlata .

Fig. Z19, Side view; Fig. 220, r t:n .... ith lower p.1rt of wall CUt away to ,bow allOmaiQUS Jtructure:

PLATE XL I. ~ralure nest Df DoiicJ,,,,,,s~lIla arMaria .

Fig. U1,Side vicw of nest built touchina- the ,rouod; Fig. 222. Same, ventral Vlew

242

243

PUT£ XLII. Entrances to Vrs/'IIW nest burrow . 244

Fig. 22), ~tral1ec to nest of VUfulo 114Iph .. ,~o, sbowing e..'tea\"ated pebbles ; F,g. 2U, Same, close· up, showing mud p!:utering; Fill. 125, En­trance to nest of V UPNta pmtyl::o"i(a~ Ihowin& mud plulering

r UTE: XU II. Mature nest of , 'upula wlgaris . 245

Fig. m. Nat in situ ; Fill. ! 27, Combs of same ntSt. with mve.lope removed

I' • .ATE XLIV .. faturc Dest of f'csl»<la f>~"sY('lIi1mca 2~6 Fig. 228. N61 in iitu; Fig. 229, Silm~ with half of nest wull muO\'ed to 1b0\\ the ""mb.

I' , .\ TE XLV. Ne>ts of Vt$/,u/a spp. . 247

Fig. 2J!1, N~t C!f V. ltdph",~a in '!iitu; Fig. ZJJ , I rn!'guJar ncst of V. tlN­.I'JIlwnU'Q bwlt m nony a-round

Pun XLVI. Misccll~ny . 248

~ig. 2JZ. Dried peUe~ of mud e.:~.c3\'t1ted by V u/'lill)- Jl<Nsyil'OlI;ca . Fill. 211. Earthtu buJ~rk btuh by workers of 1/ uf'J4/a wlgans to enclose spac:c boleath rtL."ld, tlm~r under which th~r D.ef.1 was built ; Fig. 2J." P()l1XIn of same in detail : F,g. 215, £ntr.mcc to nest of lIupwla i't1ll)'/t 'UJli{'o

P l.AT" XLVn. Structure of wasp pa""r 249

fig. 236, Paper from the wall of VtsftuJa rul!lal'Lr ne.:. t ; Fig. 237. Pnpc:r fr~ the wall of V",!JJ,/a ",.1,ltu."(Q nest; Fig. ZJ8. Paper frr,)m the willl or Do/t(/fm. "{'~ttJID mac.lald n~l : Fig. l.J9. Portion of I1e'tt wall aJ-.d topmost comb of nest of V titulo ':NlgarU

[ !89]

190 NOkTR AMERIC." N V .SPINE WASPS

.... C&

PLAT& XLVnr. Comb structure . 250 Fig. 240, Upper suriOlCe oj comb of a mature neal o( Volerilllt't'sft!14 ,"(KN~ lalo; Fig. 2-41, Second md third c.omb. or a mQttlre nest of Dohchowspula armona in edge ,<iC"o'; Fig. UZ, Portion of comb faa: from mature r\t:.U o f

olit:h(n:~lftuJa GrntlJrW shO'A'inr p;l~red-ovc.r 111a.tllinal cell,; Fig. UJ. Portion of old comb of V tspur. t'WIUaris showi!!&" cut-aw.) abaodon<d cellJ

PUTE . LIX. Features of nest wall 251 Fig. Z#. Inner lurfae:e oi wall of VtJffda 1'~"ly/'ro,uCQ Obl ; Fig. 245. Inner lurbce of v.'2J l of ~iant V ut,,/o t.'J4lgorU nest

P UT L. Nest structures 252 Fig. U6, Comb from !al'\le ne.t of V"pula ""'garis , howil1j{ 'UJpen.oria ; Fil/ . 2-17, TOpolOlSt comb aud 5ec:lion through cap of mature DoUtho'll,t:.t/tU!o md,ultJla nest

Pun Lr. Combs of Vupula Sl,lphr<rea nest Fig. 248, Upper .urfaao of iJUt two combs from oest shown in Fig. z.;o; Fig. 249. Upper surface: of secood contb froni same Tldit

253

Pun U L Comb structure in Vrspula nests . 25~

Fig. 250, Fact: view of lobed comb front large nett of Jlt"JfHJ/J j!tnJyh.aNlca showing large and ,mall cclJ ; Fig. 251, Faa: vi." of combo from DCst of Vtlp",ltJ stdplmTttJ Ihowing lara"e and small cclls

P LATE LTII. D istribution of brood in comb 235 Fig. 2.52, Comb from middle of iarrc f/ tsp.IID ,.,.n.sylw";,a ~l Jho'\lo'ing «gular Wiling of brood; fill. lH, Ohler comb from same DC I ,rowing brcalcdown o f brood zoning

P LAn: LL \· . Mi=llany . 256 Fig. 25-#. Hibem:uilli QUC!eJ] of V l'sftt410 rmsyiL'OlIi'Bi Fig. 255, Gipnlic nest of overwintered ,;:0100), of V IS/lv.JO l-~II/aris, built beneath tim~r iUp­porting edge 01 roadway

[ 100 1

NORTH AMERICAN VESPINE WASPS

PL TES I -LlV

KEY TO PLATES I TO VTl : THE H EAD AND ITS APPENDAGES

CUTICULAR STRUCTURES

,aN, cra nio-cudioal (:Ond, le of mulJl.ry articu&.tJoll

Ac, acetabulum of mandible Ac. ac:r.tabWum o( po.looei­

put (poltcrio r ameulalioD 01 maodible)

/feB. aero~los .. 1 button Af antenDlfu ALiP, &Gteno r linruat plate AMS, acu.ory mandibalar

Klente oNJ, llpode.me of mandibu­

Ia.r- Idducror APII,. anlerior pbarynx dpd6, opodeme or ma.odibu­

Lar abductor AR anteonaJ ridle A;.c, .rticulu cup or the an · .. .".

AS. aDtenn.al K lerite III antt!!no:.l tulun: AT. an terior tentori.1 a.nn5 dt, anterior tl"lSlorial pit BS~1 bosal ad.rit. of ... -

.. nt. C. a-ank.-l eonrlrle of mAD-

diboJl.r articulation Cd. c.lrdo Clp, d~. CSt. pnncipill arioa o( the

atipe< CSt', feCondary carina ai

the .ape. DPJaB, do ..... l pha17nfell

bono DT. donal tenlori.1 3.fm1 E.BIl, cplpbafJ'D.&eaJ bal r Ei'''J. ~plph"rrn% E.R.. epi,~omaJ ridae n, cpl.tomaJ Illtun

ob ... d, abductor of mandible adn,,'. &dductl.r nf mandjblc: Qd,nl', anterior deftt' DOr "'{

lhe maxiUQry palpUJ ,,{Ti. &nte:rl(Jt' 8«nr of the u.ul. olim. anl~rior lnlrinsic. ph,,·

rynteal cnu!C:te cdntllt. eltpelll dilator of

mouth dim . dj,"r.t1ieulu UlUIClu dlbe. ditaHun butt.h, or

bucal t!ibtll'lr m~les dlb,. dtpr~T or labial p~'"

ldr;h. dilalQt'd pharyngis. f rDDtabs or (ront:11 diUaa ton ot Ibe an'c:ril)r phar-

'''''

Pt flag-eUum Pew, foramen m&pum Fr (rona ,~ coroual .mturc:~ ,. , frontal .uturCl? Go ..... Iea C.P. plcal pec.t~ G«. lena GI.gl_ C .. p, lnatbal pouc.b fie, hypodomlll eari.oa Ht"" b,poph.arywt It.t, hJ'Poltotnlll &UNrc H.lI hypo tom. H78, b)'l'lopblryngeal button H,P. bypoph.....,-n,iul pec.ten ;G/, uncrantc::nnary trlanalc lAS, lateral .ulc:uJ of the

po."' .... fAT,., labr'" tractor LbP, IalMaI pal""" Le laeioll LeB, lacini.1 bar LrP, lac.inial piVO[

LIJI P r IiruJar binlrc plate L~, hnaua L,", labrum tml. l.bio-maxllJary j tJPm U labral uturc LT,..labra l tM;IIIJgle Me, nandihulu moo,lc jldt mandible Mlh moutll M.f'P muil"'ry pal"", O. ocelli O~ .. occiput Oce, oeclpital cODd,le .. la, oC'COl1ilJ I 4Uturc O~~ ouoph"IlU!I OJ/, OCulo·UJAlar "!IIpacc

MV LES

~Jlr", Jllatnra poarpbaryn­gc:all.s or donal dilator. of the- po.tenor pba.ryux

3d/pi, . po.cerror dilator of Ibe anterior pharynx

-fdl,h, v(nln) dilator of the: lIte-rlor phuylU'

d,n.sW, dll.aton o f 1'hc All­,""anum

Iud, externlll an tennal de·

r.reuor-'(1. rxlcrnaJ an~ Ic\'a-

tAl, ''''''', f'XletaOr 01 tbe maxilill fAil, flnor Ilf tlll: 1Ia.l.ca fU, Ruor of the 1,ciol.l flIt. n('"or of tbe .tipn /p,I, P.uur or we: 1U~lEtoua

OR, orolar ridllt OrP. oral pe1:tcn (JI, ocular I.uturc ~ed, productDr of cardo P., p.dlcd P,«. po.tgc::na P,~Br. po",cGaI bridge P,/, parqlo.A P,~ parlillos .. ) seltnlC!! P(II. pua.loaaaJ tria.D41e PIID1, pharyngeaJ di-vcnlcula PItPI. pbaryngeal plate PLiP. flOlUeriot" lil.l,!URl plate PtJe. pO.loec:ipu~ POR, paslocCipital ridg~ pos, post.occlpital .uture P; pl"",. .... 11><1I PPIt}, po.lt-rior pbryax p,.."", premtntum /'S. plc-urolfomal .mture "',po!lterio r t~lOria.l pit SAN • • uba.otCQoal ridae 141, sutJaalenn..a.l .-urure Se. ICOpe S.Sd, .msUJa b.Jiconica S,R, .ubgen.aJ d dge SIS .rubgen-.J ftItor~ SID. ,.lhoat'Y d UCI Sltl. aaUvarium SPnn'. apaiuJate preQl~ntaI

orocua $. stive-TS. ren tori.1 bridge TL, tf"llncatal lobe. of an ·

la-iOt lingual plate r,.s,. tnlctoraJ su_pauDr V, \""cfte.x yj:l"B. "etllnal pharyngeal

ban

lad. lnltnwl anlt=nn&1 de-

l , ..... , jIJ. irl.le.rual antennal leva­

tor lpltm. lateral 11b.aryo1UI

UKka ftdHtft~ po.te:rior dcprra501'" of

lbc I'JlaJtdlatf [JaipLU It/m,It, pharyolcal dilator of

the mouth ,/Ii. 1"IO'teriol" Auor D tbe

I' ... ,lor. pMlcrior l.abial adduc­lors

,wult. .. potterio r m\llda of the aaJi ... anum

,ti .. , po.l~nor intrin.Jic: ph.­rynaeal mw:de

'hOtbl.ILAL SoL«£"e. Vnl \ lit. 1\"(1. I

"

•• Y .' 0 ' ........

'" '''. no t.I

"·("SIMla ~-lrJdlti(d (SaUJ~), qu~: f'lg I. J..at~ntI a.!lpttI or )lQ.lJ FiR_ 4 0aUib.11l!iP«1 or heattomouthpart.J rtlnO",rtJ Fig. 2. C<phAl;c .. pen 01 head Fig. 5 Do ... l I15pen 01 t...d Fi 3. Antf!lUlil Fig. 6. VOltr.1 a.s~l or heac1. moulh~rt. n:JI10ved

BltlLOG.I L ~IUcn. \'tU .. Vnl. So.

.. ... .. • AT

.. AT-f-t ', ," ..

... FUl 7

••

FI(;' 10

PuT.l: II

1'14.. 11

Ytl~ ft'nyiCIQrtKG (Saus..~), quten: (Key 011 po 1"1) F'1I. 7. Innd :L!JI«t o£ btc.nI hiM or h~d .'!Ikddon. "illl ntou1hparu rtmo\·ed Fig. 8. I nn~ UPK\ or pQSlr::rior Mil o£ head Jcdclon. with out ruantJiblr Tno(wcd fir 9, Ot"bJl of .ledion lhtollJ:::h ",uJt of anttnnal ~kct Fig. to. L..tt=raI upetl o£ land skeleton w1th ,jde CUI aw.a1 10 mow lillCtill .~ of

ttnloriulIl. ;100 Mar mandible. remo"etI Fig. 11. Inner alp«1 of front half of head I'oke.leton ;and nlOUlhpan" cxcqK. ri~t min­

dIble, rClllQvcd

VOf \ nl. S,. ....

FICo. 1M

I'un. III

Vutola /'<>U1Ivu';", (Sa ..... ). qu«n: tK<yon p. 192) Fig. 12. CaudaJ aJp!Xl of ~brulll, epiphuynx, J{l1.;a.tIla! pouch, anti ph~rynx Fig. jJ, I4saJ ~ 01 antenna. with a"IC11,"J lUu.Kloet. ollUlthet.l

Fi". 14 lnna ASptct of lateral hili o( head ,Jcelcton. and Cfftain of the mur.t.lc5 fig. IS. f>t:taJ 1 ar :M:ruoria ,,, .. pleal pectm Fia_ 16, Ce-,ph3Iic ~pcct of I;abrUlII. cpipharyns. Alld phuyn:.: Fi~. 17. Mts:.1 ."ftec1 uf m:uuUa and iB ntwcultlttU't

F'ig. 18. !lad ~dan. a-ph.;tlic tonlntbOlnt. R",uthpArI .. and Ih,;.oaIU~ mu k:s as ~n In AgIlW soctlOQ

PLAn. 1\'

flo. 19

Flf. 20

.... . ' -...

FIG. 2J

FIG. Z2

Vupwa rr.sy/uonico ( $;ouu.). queen : (Key on p. 192) Fig. 19. [n"cor ;J..Spc:cL of right h.d£ of he.-.d i kelt1on. with. Ilntenml mu.~et. m:uxhbular

adduc:tor mwde, and lateral phar)n.g(al 111W!('le. 10 Ilu Fig_ 20. Mu<\CUliuurt of antennae, mandibles. na..'C..illae. lind labium 1n $l tu Fig- 21. \fus(:Ullture of ~n,ennae :lOd C'ephalic \t~um, :tho ceruin nllw::.Ies or

rnaxi lillC and lahlum, in ilw ;u 'lIrtn rrom bdlillU Fig_ ZZ. u lcn.l ullect of right nwclll:l :uul i~ mU.iC,J1:alure: ri". 23. Deul:il1 of nndculatu re of the anLftU~('_ pharynx. :md Cl'tUin U1oulhp.1rU it' situ.

U Icc-n from the tide. with part of th~ head wull cut away

Fie. 2.

' ''-... ,

)17·

........"" .. " ., ~\~ ...

'- UJ{P

'" ... F,. 25

F,u, 26

V .. ,1Il4 "0S)~t ... i<4 (5 .... ,.). queen: (Key on p 192) Fig. ~, Cephl.li~ llomodaeum. U'IOUI.hlQJ1.l (except m:andibles). lind &.n«iaJN nlU~le,.

Ia l<"" asp«< ~irc. 25. CephAlic ~tornodaeum, mour.hr'-lrtft arid a.s~.ued muscles. in .... rill .. ' ,«lion ·iK· 26. ~ph.lic 5ilOnuxlatum. I"hrunl. cpipJ1JrytUC, gnouh.::a l pouch, labiuln :wd auo-

Cll1ted muscles, Illttral Uj)t:Cl •

IOu ... 21

.. ...

... Fu;. 31 Fut 12

Vul'wo ~rt.I)Ilw."i("o (S.:auss.). quten: (f\ey Oft p 1(2) Fi¥,. 27. D~a.ilj or mUMubture of m..txUb Inner .sped fi ... lB. LabIUm. ldt nl.:&xilb ant! mU.lo(')et. C4llfl:d a pttt

Fi,. 29. Detail, c ( mu..cul:lture- o( lU.u:iU .... illnt:r ulled. fluor of lacmill r'nuo\·C'{1 .Fig. 30. 1.>elail. of l.tbiulO amI ill 'llU!\(ul~IUr~ wnh half of h)'pt)pllarynx. nJ:ht ~ra­

~105sa. llnd "lJilUiule T'roct~ n nltht i~f('." pr~n\tllluUl re111O ... rd Fil. 31. Ortail, of I~blulfl .nd it. IlImcuIOltU«~' wilh hln of hHlOpharynx illOff of prcmrn­

hIm Gnd ;til uf ligula C' ~,tt 1~£t ~raR'I05~ rr"luovt'tl fig 12. J,..abio H&;LJtillary .contplc;: (e'(c~pt ticln l1t.u.iIl.lj ,...".\ LU-.ocilltw IUU.c!C' in

ccphalic a".pn;t

VQC VIII. NCI I

.... '-

l"tS'1I11J /l'lUy/wl'lira (Sam!!.) . '1uccn: (Keo. un p. 192) "'IR" 33. C~ph}.lic UfJtt"t of lefe mandihle and il 11Il1.'<uLllUn: ' .'g. J..4 . C.UtIaI altpttt of Idt 1I1.1.IIcJ,blc: IllH l lt. l\IulK:u~turt= Fir· 35. epmfic .. ~ of .. blum It1l1 dl"L1ih of iu mwculoature -",lth .. nlrnOr waH or

h;ypopltaryruc 1·t=IUOVN foig. J6. ~I~~l ol'"JJ«t of lell mandlhle IIncl itJ ntIDCuJaturt= Fig 37. Dcta:lIs of '",bium Ilnd i IT1ldCuI;ltur~ in t:rphaiic :tiP«l wilh hypapharynx.

I P~lluJ~le p~;a <Jr pnmcntum, pwf11;~I()-.; ..... c. .}nd lrft half of a.ut("rior ...... U of Igu .. renlDVai

SURTII A IEKIC,\ VE:PIS.E \\ A"SI ' ,s

KEY T O PLATES VIII, IX, AND X: THORACIC STRUCTURES

Cl/TlCULAR STRUCT\TRES

AF~. AF J. rural arm. of mHO- and metlUtcmal apoph)"act

AFS. apOdemal rold AIS. anterior InOeetioo or

scutc.Uum AIT,. anterior IoIrDnolll.l in·

flection A..V-P~ . ANPJ. anluior no­

tal winK prottUH IfrF~, muofurcat bridge ASPt"# apical Kalef or the

propodC\lm l ifot', '6111 axillary Klni,~ Bru, BaJ. bau'lrn eM, c.uvie;al membrane CS~, Iran.lC1ltdiar carina CT J. lranJimetano'al anna C,,_. conJunetiVll C-,_ COll. DIT/ , dt:)f'JAt pl'Onotal in-6~lion d.,P. attachmcnt pt1( (or d i-tator muscle of .11Ira.df!'

EfJlfu. m*pime:ron II", 1';1. //IJ • • 1c-maI pib FStll , po luiOt ani~l.,

tbnie or i«oad Ilf-mum HFI. bon1:onl.1 pl.te of

mCllOlttrnal lpoplly.l. INP, kCoodary proplll""Uril

r1d4< ISL, IOlenlemll hK.meat LTr. pottuior pranolal lobe MCI. anterior m~.ternaJ.

anna IIIIIS m~lan notal auturc M P~/u. antrrior procc.u or

metopostphrqrna

1G.dII, 11141.1. propodeal ole-­nlor and (onion mu.te:let of the abdqmc:n

Ildl,. il'ldu'ecl and principal depreuor oC lbe Co-re Wlnr

/lldl. relr .. CIOt of Ihe- meso­pojlpbn.ma

ufJt. ;u,II, dill tor rnu..c.Jet oClpinclca

114",. tndirw principal elc:­vator o( {ore winl

IIthu. furea t rctr.u:lor of the m~f,()poStl'hrailmll

IIp, III~', pro- and muo­IDoraeic furco .. ple'\JraJ muJdc:.

lUI. lu~. iantr :.nd ouu:r IlrooolaJ r("tnew", o( tbe "",tum

I lin. rc:lntctor of KutclJum JlIU/ , plLur.1 dC'prrslOf o(

the abdomtn lIiu, iDI(rfurc.a1 muaclc IIlin. m~I.Iurcill dcprdltO r

of lh~ .. Wom.:n II.1J. 1a'14. li'j, mt:totCemal

and ht IDd .!d mHO­(ureal 1"l"lraCl6'" or pro·

Ilru~~IC'rnal 10r'lon m~ de 01 Ihe .MomC"n

""~/ . "",.' , "".J, median atunal rrooVef

N]S, mc::hlnoLaI ~lImelit OP, oeciplul Procell of pro--

pI ...... OTJ. mCUlootaJ dtpreuion Por, parap.lidu Pd, Ibdornu\l1 pedicel p~'J. 'econd perit~c /d. parap.ut.1 furrow. PF;I, al'llc-:riOf ptOC611 of (he

mao{ural bridll"t" Pill, ml'!lIOpn-pbraama 1'11.1, mf!:lf'poIlpb"KmA p~J, pro(lO~ .. 1 p, .. pb"J!D'~ PIS, PITI. poIlenor inlIec:-

lion. Q{ IIw tcllum and nro"otum

PII, proplt1lron PIlB, trafUl\erk lH"oph~1Inl

brace Pll. O1C'1Cplwron tlu. me.aop'turtl 'uture PI]. meaplcufon P/J~ fint mnaplC\ln.1 pil PIJll I«ond metaptrunl pll "JI. metilpltural Kcon~ry

.ptUt e-F,vJ, mctapoalnolum Pro propocieum p,.Cz. prC'CoxAI tc:lt:rhe PROP, po.urior ranlw nf

or.cipitwJ proc:cu of .,ro-­pleuron

''''. pro,~ t~lh PT J: po.Ue~ula P'~ P. plroral win. proet:D Ro# rod from pronou;) pit RT. reduplle~nion of tera\l1D

MUSCLES

it, intenlt:mal ~xtc-ntot" Itsr, inner 'erco, ternal rc~

tractor ItJ,., lateral inlt:rtH,al re­

tractor lJ"u. /flu . IILf'n&I t'OU,IOt

ud plf!Ur.1 pTodm:tar o( rorc eo ...

IIIJtfI, 111m.!. prnduCf.or" and. QUler rtductor or midd1.e eod

Illiml. 1111,"1, .Iernal pro· dLKlIOr .mel r~lucttJr of hind con

!JUlJ. pleural dt"(lfcuot or (rot:hiilnlet' .nd primilt)' IIdductor of (urc.lq

IJllltJrJ, IIIff1Jb{ meso(ww derre5JOr 0 lfocbuter and Cldduelor o( middle I",

1111,,13. depf~r or troch.an· ler Ind adductor of bind le8

II",.,. IIml. IUfal and di ... ~ onlll roUltQU of fou ron

111"'4. loner ntiuctor of midc.llC' coxa

1/",6, pronollil lu.ItJUf:tor (1) uf the 'arc coxa

O,¥t , I'Ileutal elf"'C'Ataro( h,. .. d

Set. "'Hum Sn, lCU.eD-urn ScuP, IoCUt~lIar pocht SnS, tWlell.r ercu. S...s., axil" S,u. S(lj, tbondc: 'Plrada $1'1. I1ropodeaJ -,>inelc $1'11. 2d alxS()ftHn.1 .piraclc: SU. SUo S'J. pn>-. moo· ,

and mcwt.ern, SilT, mUOftcnW tooth Stll . I«ond .ternum StlJl tubalar ph SIfR • • ubatar ridge T r, prOnOlulQ r,c" Ilt:llc-rio r prouotIl Q_

rill. Tlf' pronat.J pit TI • k"C:ond .bd.Olnin.l kt.

fI\Im T J. melllnotum T JI. melllDotaJ lilltcus tllU. trllnSmt:'~olal suture t/#lJI , tram-plalntl ... ItU~ T,." I rodwuer t,. tnlnMCUW 5Utu~ ld, '''lH4tU'llAI .uture TV F.1. ttansven;c plate 1)(

mctuttrnal apopby i. I'Fz, I'Fl. J0l1ailudia.a.1

platQ o( m~· and. met;)­sterna] apopbylCt

f'Pd, "-c:rticaJ 'Plate of ab­dominal ~c:el

VPFI, .. ·erueal plate of pro-atemal .paph,.-;..

I'PT. ventral 9ronoLiI loolb rR. entodor .. ' rid,e rl. ~1<HC'Utdlu utuJ"t

Dill, furcal cTc",..tor of head OisJ. OU4, fir" and lCCond

dcpreuorl of bead MI.Jt. JOlt, oc:dusor' oC tbo­

n.oe .."racles 'tNtl. anterior Dotal elcn·

lor or the pt"opkuroD and hud

IIpm/. IIJ,,,.,# bu:abr mu. d ..

'''ttI~, lateral no t.1 elevator and rOIAtOr of proplas.roo

"pm, . lI~mJ • • c:cond and tlllrd plcunl mWlclc. of third .axiltan 111/'''''(1 and 11",1.1", pltu­ral uaucln or tbird &-'Itd­lu,

IrmJ, aolmor and i.n,," no· ,.1 protrllctor of Vropleu.­ron

1'111.,. "Iff', I~Jlf6 , Inlenot:' · dia'c. 6.,." .nd .«ond po • . lenor notal prOU';Qelon of ptoplnlron

IIftlili. plnt,..1 tlC'fIrrcor pC lCulC'lIum

IJlpnq , deprellof or the mctl;aotum

1I1"'~, ,ub.alar m\lsdc 111',,1$, cox.o·trut.Jar mu.cle

~I ., .,. -.... ...

' .. F"", 10

,." ,

.., -.:21 F lU. Jg

... ..... ...

Vujttl.!O /,NUylWHKlJ (Saws.) . queen : Fir. J8 Cephalic IUIpttt or Ihoru

Fir. J9. CiI.udaJ a~J)ttt or tilOr.tJC

F'ilr' Ventral upttt of thorax

r n\ut

... "

II ~I

Fig. 4 I, Oo,..] ... p«t or ,horu Fig. 42. ulrr.tl .... pttC oi dmr.u

fllSPll/o Pl' Jt.lylwl4.;ca ($a;Uj5.) . '1UttD : Fig. 43. [n~r up«< 01 rigt, . holl 01

tuUKln ~n j,u

rrn. 4-3

(KeY' on p. 2(0)

til rOl'l( 3.nd base of .-bc.Ionttrl. following tho~oc

Fig. ~\4. inner ,,~pecl. right hall of l hgl'l,1~ i!ml b::IfiC. of .hodmen. v..-ith ent.in 1Tt~1~ in 51tu~ vertlol plaJeof pro3tem.,1 Olpophysi!li ilTnl both "~n.ic;al uti hUr1l:()nl;J1 J1l.1.~ ol n~tern.tl apoph ., .. sit rCl1Ioyed

Fu;. 45

F'u;, -i6

! '('It",O f("ltS',l·/tvllic.g. (SaLlS .) . quetn. inn.er Q~pec:t : (Key on p, ZOO) II". -IS, rugb t hiJ..lf or lhora.x allli li.;U,e uf abdomen .. ,·ft11 l;rrt.Qin mu; dQ in .jtu : plane of

!!".tClion 1A"'l~ IJigJuly 10 r igiH or cenl~r , t:lltin: pn)!,lemum a.nd iu :tpophYii , al!n but of mer,,()!.U:.rnaJ iJlK)physis. n:mo\'ed

'jg'. 46. Right hJf of pn.motunt, nJe<'Onchnr:uc (except t,.CUlunl). lttt1a1hou.x, and 1la.Se of ilbr:kmlc:n. YlQW'1og drt:ail.s or mUKulalure. pbne of ~ctigll lying- JUSII!) ri«b t of center

2().I XOKTlJ AMERICAN' \'ESprN'E WASPS

KEY T O PLATES XI, XII, AND XUI , THORACIC STRUCTURES

CUTICUl.AR STRUCTURES

JA flnt aoal von .AF~, (urcal arm or the

met()ltltl"Dal apopb;rd. AFJ. {urcal arm of the

metul~rnal .pophyli. rlFS, .DQdrmal {old of the

KUtum AIM, ahlcrlo r metaQotal in­

RC'ttion AIS • • nlmor infieaion M

tbe JlCutellum ANPj, a nterior notAl wing

p roecM 01 the tn~t:.a lborax 'lb . . th. J.".r, tint. tee-

ond. .n.d tbint ullb,ry :Klmt"

A.rC. lL'ZiJiut (() rd 041,8113. blisaluu-or mUG-

thorall: and metathonilJ: C. I:olull ven ex, coxa ,ml, lIIttu l CJruIr(in:.Jotiod o f

:llcufUm £,"'1, lI'\eupim~ ron FSIII. J)Of;terioT a.rncuJar

fbni!!t. or (be. ,rtOod ab­domina l sternum

TIP, bumc:ra' plate or .ub· cotta! IICIIllt

LPPll.I, latua l Ccrut ofrnc:lO· pmf'pb"'lfttla

M+CIfI. (UKd .\1ed.i1 And CubitUl I

MPPIII, ,Ullmor ~\Ulck­bearing proc:as of the mClOpOltpb.ragma

JlJdh , prDpode~1 cle'C"&tor of lbe alwlomCJ

IQdl.l~propodca l lonion nlus-. ele of the abdomen

IIdll. indirect and prind~ pal d~TUJOT or tbe ror~ winlll:

1//(11, I"f: lra~to-r or thl!: hU!.IO· poslpbraama

IIhl, fureal rc:tracto r of the muopoltphr.llmoll

Il/fjlr . mdOthortlcle (urco­pleurat mUJCle

111.11. rt:tractor or the Kutl!'l­lum

tllUl, pleuoU depr~r of the abdOll1cn

/lUI. inltriurea.l mw;clc lin$.:. mc=.tafurca:l dCJlfU.lOr

~f tbe abdoUten

NJS. lale.ral nu:tlqO~ kl~ meat

P"r', pa,rapridn Pd. abdominal pedicel PlrJ, ~riltc1Tle of 1«"000

thoracic Kpiraele ,f. par.paidal furroW" Pb me:sopo.tphragma Phi. detatht"d Lip of mQO·

pot.lpbra,amal nttlUI PbR. r.lmu,. of mf$Opolt­

I,bragma PltJ. proJKKltal prcpbniIDa p"JC1 amcrior carina of the

propOtieaf PTCfIhf'B,Im.a PIM. po.terior mctanolalln­

HeedOD PIS, po.tll:r1or Intlc.:tion n j

the leutdlutU Pl~, muoph:uron P/~."l . auterior m~leurB I

apodemCl PbAI. f«Qnd me.o(1lcuf'Q'

IlPOdrme Plu";, potterior me.oplcura l

.podeme ,',ul rnclOpleural suture PIJAr mual .... metapleunl

.podC"tDC P/Jrl.l • • ec:ondary mrt4lpleu-

ral apodeme PN;', muopo"LnOwm ~ PN 1. tnf:ll3POSlnotunl Pr propodcum p't J . mu.1de-bearina J:lrDl:t=::!II

or the roetanO!\IDJ

MUSCLES

ucr. Inter.kTJl.If elllm,JO r Illm]{l and Jb

l muorul't.:ll

dcpn:::Pur () Irot:banW and .dductor Or middle I ..

IllJmJ. dep~r of t tc ... ch::llftl.e.r anti at.Iduttor of the: bind lei

/llIm~, pleural produt;tor or bind con

mJA.r. )nlria.le: mUlM:lc or fil'lt pleural mu»C.lc or the third &;!tillary

m.l . dll:p.rdlor of the he~ula 06,1 t Ottlu.or or PTOpodcal

lplnde IIpml , baaalar mo:r.cle o. tbe

Ult'AOtt\onx Il/~,rtf, basalu mUlC!lc or

rbe mctathorax

[ l!).I I

PT" po.tt..,w. R, radIUS S~t , lC.uturn Sn . ICUlellum Sc,.L, lam~1IirQnn ptOCes.1 o(

the kut~llum Scw.$, K'U teU.r KIlt SntS~. IltU lellar KlIment or ~il1a

S,"cJl • • piraeolar atr1UtT1 Su, mC'SOnc.nlUm SIU, kCoGd abdominal atc::r·

DUDI Stu . StlJ. 1Ubah.1"d o( mes .. •

thorax and meuuhonx. tdp«liveiy

SNH. ,ub.ta.r ridg~ T J . mct::anolum T i, humeral tclerilc. or tbe:

ml!'l~IJOru.m

TJB . iute-Mal IUetan" ..... brace

TJP, md.notal prominence T ]I" mc-unolal .wcu, T • • tegul. TPIR. tr.I.n.fl ~ra' ridlic Tn, traehc.a huiru; Tro, ,rochanlO' It. tna olUCUl.IIIl \ltur~ rFI, nrlieal platt' of me.JQ ­

. Iernal a.pophYIII J'Pd. \C.rt.ica..l plale of alt·

domina! palicd VR. cntodorsal rlrlgt' 1.'.1. scuto1C:utcllar lutu.re

JJ/tIfU. I«ond p1eun l mu., de of thir<l u.illa.ry

l11pM/a and 1I1("12b, plc.u , ra.l mU.!Ic!CI 0 tb~ tnt"ta · lhonac:.ic tbird uillary

II""J. thim pleural maRlt". of third aJO.ll.ary

J//rrnJIJ and IIlplftJb. 1l1e-u· ... , musc:l~ o( subalue ot t.b(' mdat.horax

IIlNn4. dc-prtuor of the.. m~Ulno[uUl

lip,"!. ."w.,lar muade of meteJfbor::lx

1/1,I1IS. (o"o-su~"r mUlde or lhr. ltIetat1'l rax

8WlJ)(;lCAI Sn n Pun. Xl

47

:;, .. ,.u. "',oryIVflni<o (SIlU!I$.) , que.n, II'. 47. f:;,..nedr~~.Pl'Ct., I rialll h1aJl of mesOP«lus. rneblhoTu. !lDd but' of .:a.bdomen &bow.

'l' Q..otl S 0 mUloCU. ~.ture Filr· 48. D~agonal J I«t:lon 01 right tJl,?taCic w.:a.1I palling ju I anterior to pro_I •• ,

5plfllC to and t:r"iUbIRCt.anotat canl1~ ~

Fir, 49. ~:~!'~IQOI' of Idtdl~~[~c.ic w:tll pil$5rnc jU:1Il ,:lIIlerior 10 lhorac1co.propodc:al ~ •• _OWJ me, an ~'lIld pornqul~

FiC· SO. ;:Ute1J'!m. n~phral[lna. hOi$( of front wmg, and associated mwcles rt­(J"ed In Q oIrfoUp ftom tborax :md vie-wed (ram ft:H .ide:

BIOLOCICAL Scn.lfcp", VOl VI fl. N(II I Pun ,\111

• ==::::::::::::!!\

FIQ. 60

r'tz,wa f'ttl!U)!lw,.i~Q (Sal145.). 'jUtttt = (Key un p. 2(4) Pig, 55. Ex-lCrnaJ ,-jew (If articular r~()n of Idt ror~ wing Fir· 56. Ventral view of regula aut! depre5Sor nlu.scle In 'itu Pig. S7. Vi~w or Icrt tllird uillary or Lh~ rrool ""lng. viev.~d from withm and rot.lted

upwnd 10 exPOS<! OUter surface .Uld ,how intrin.!l.lc nlUSClc or lIxill:nl")· Fig. 58. rnl~mDl ~icw of .,trcul.u re,rinn (.1 1 (ore winjf, !!.]Iowing imtrttl:uiou-, of iCUttun,

lCulcJlunl, fin t :ucilLtry Klerit~. I.ncl indir«l wing hnuclea F~g 59. Lncr1l1 IJp«C, p.J11C'rior I)"rl ojf maolhorax, lohowing maopolJtphrngrull in silU Fir· 60. J. nntt ,-,r~", ~( ilrtlcular ~glQfl ()( left rQrt .. yj~; sclucllum rt:0I0\rd. showing

n:Jl.t1Dfl.JihiJ1 of anterior OOLaI ,ving pl"'Ol;eS;; :;arid rirst a.:cillary sclerite

SORTH AMt:RICAN VESPlf'I;P: WASi"S

KEY TO PLAT ES XIV, XV, AND XVI : THORACIC STRUCTURES

CUTICULAR S1·RUCTURF..5

liPS. apodr:m.l fold of the sculum

AITI, anterior pranotal in · flectioa

APP. antalor prace. of ptollernum

AS!.} • • ,c.b of ,plraeu1u Ye .. ubule

• co_laJ .. ein d card. Pt,l palletfor venial platt! 01 pr~mal ,popbya.. Lt , ventral ce1"'Yic:al ",a­menl

Cu, lalenl cervical lip. ... nt

CAl. crnicaJ membrane CS/lz. apc.tcvlum of 6 ... t

thor.de .pinde CSPJ, orcr~lum 01 itt'Cond

tboraCit 6Plrldc Cr, coxa CrS • • «.ttaory acleritc of

(on: coQ. DITr, donal prenotal in·

flrctloa P" lemur (/II pro"tt'mat pit FPl.. furco·pt~nI lla-ammt HF,. horizon, .. 1 plale of t\lc.

pf'O!iu.rnlll apopb,lu H P. hUlllcnll plate or tub­

cottal lCale ICFJ, iantr CTHI of fint fur-

eal """

.wel'"", dill tOIl!:;! po tpba[")'o .. .-... 1. or doru.1 dilator of thc posterior pharynx

Jdl,lt, posterior dilalOr of tb~ anlerior phal7lUt

oIflJ,II, 'Ven t .... dilator or the po.tenor philryn-K

IIdIJ, indirttt and principal dcpruaor of tht: Core wma

Ildvt, mdiRCl IInti principal clentor of t.h~ (ore win,

,od. ex'~nal .nt~nDAl ~

p''''''' "nl* internal &nu'.nnat de­prdJeOr

"p, pro'boracic (ur'CO-pleu­r.J rnWlde

l&slt inner pronuta:l retractor o the Kulum

lIi.1/~ rttnctor of tbe KUU!..I -10m

lIul

outc.r pronOlAl runctor o Ihe ,lCulum

h61. mesollcraal retractor or tbe propc.ctua

IPFt, inner prqce .. of fint (ureal arm

IRP. ACQDndary proplc-ural rid,.

ISL. Inlcn-tc.rn.l hpmcnt LPP. lat(!m proce&a o( pro­

Itemum MAP, mar](inal apodeme nr

~roplcuton Mel, Animor mClo h:rnaJ

carino ,"I~'. median tfe-rna! arOfJ'e Dc, O«ipLlI Occ QeC:lpJtal c:ondylr OCP" outrr aat or the fir.1

furcal .rm OP* oeeil1iml proCeA of the

propleuroo OPFt, ouler proCUI of hrllt

(ureal ann OIJtP, auachmt:nl pq (or QC­

dwwr uC finl ~o~ac nnrllCle

Pu parapsida PCFt, poatuior crest Dr 6f't1

furea! ann PrrJ. pailrcme at !I~nd

thoracic ..-piracie pI, panp.dal furrows PI.8~. po_U/;enal bndae Pili mcsoprepbn.ma PtTI. ,POltcrior pronow in

Ocellon PII. propteutOn PitA . pl'OfllmraJ arm

MUSCLES

lUI, tint mc:.otun:al rC!IRe .. tOt of Lhe prO~IUf.

/isS, aeeond au:to(l1ral r'e-­u-actor u( lhl:' propKIUJ

Ilmi. Itcra.al rotator ot Ibe fo~ co ...

11111.1, pt~1 produclor of tbe rQI'tI COU

I/"'J. pleur1l dtprrnor of lroth,nlno aad pnraarr adducUlf' of forc leg

/1*/, furc;at rotalor o( the (ore COn

/lIttS. lI«ondary adductor of fore C:OXI.

"IIIt6, proDotal adductor f?l of lbe (orc coxa

/1"11. dlqonal rotator of tbr forc co~a

011 •• [lleu,.1 elculor of the head

Oin, Cureal ek!"A'or of the bead

OIlJ. lint dt:pn:nt .. r of thr head

(208)

PlrR, tnlU\'cnc propleuul brace

P/~. m~pJa.trOT1 Pulf I, antcrior IndOplrur:l1

apodCUlI:' PlnlJ, poltenor mcaoplturaJ

apodcmc: Po~ P'O.floccieut pok. po locc"llt.a1 rid r ~I, pctStoccipital .. utun P['It!1. po.ttrior ph.,..,.nx Pr,,,'. prmlCtllum PROP, fIO!nrrlo r ramWi of

tbe: occipltlll ptoc:eu Qf Lhc propleuTon

PT, prorurcal troulb R. ""diU! NF'.lJ"&I1a,'rne ridt;e of tift'

(ureal Arm Set IA=Ulum SID • •• hvary duct Sl • • ti~ SI" proslemwn T,. pronG,um T,Ct. anterior pronotal CIl-

nal TZ. ",ula TN, lrac.bcll Tro, troebaoter J.!~. wenc VPFI. YCrtical plate or thc

prouemal apc,Dbysl.

Oil." IICcond lI~re550r or .hll:: head

lOS', oc:clusor o( fint tho· racic 'Pi IIch:

JDSp. oeclU.lOr of lCCood lhOrilt.1c tpinlC~lc

pJ(fd. pone.rior labial .adduc­t.!'>

tpWfl, aJllcr1tJr "OW cl~vator of tbe proplcuron and bead

l1t"'I, talc~ notAl clt:valDr and rotator of lhc pro­plnroll

J,""J, anlcrior .atld innet no­tal protracfDr of propltu· ron

I,,,...,,inlumedial.t notall)r~ ,,'aClor f'lf propleuron

rPm5. first ponctior nutal prorr:t.olor (I( pro..,lc-uron

11',"6. RtOl1d po.lertor notal rrotractor nf prop1cuton

Pun 1\

FII (\oJ

V<I,.I.I'<"'YI_«a (Sou, .. ), q""",,:

Fig. 61 ,Po5tef'o~hJrAl I ;upect of pro~t~mum, propleur., /ell bbes ancJ A.»O(j.ued OIUJ;CU­IIture

FiK. 6.2. P05fC'co-\'entnJ IflC'\\ of, protborax, ",ho\\ing inltrrela.tioru. b<1wetl1 pmplcurnl aml, Pru-.lcr~1 a~phYI I <;', nJlti I~ mu.~cles; riRln I~ rC'lIIllvtd

Fig. 6.l ~l:~c:d~' r~£ht I ... ~r u( 'pr()thnro1~ f~~('l:fJ.( pnKlntUI1I) IInti jlOi'lC'ritlr Willi of wure • OWlOC anlculalJQn of hurl With prothorax, and prothor.lC'ic IlllUCU-

ot- \'Ilf. SCI ru.n X\

Frr.. f;S

Vtsf'M/o f'nU11t".ir. (Sou .), 'IUffiI: (Key On p. 2(11) Fig. (,.I. lnnt:r \i~ of rirtll pmpleuron, filhl COXA, and .u~iOlted P1U3C.le Fie. 65. Debil; of dmin.g- n~h.a.nitm of fip;ht ('jot IhorlaCIC tpm.cle Fig 66. Left a.3opt'Ct of pr05ternuU1 .. nd Itt '1)()I.hy III, \\Hh mu leil origill."11in( the:renn Fie. 67 P~LCrfHlor....J Ol.'ipect of rruthulOlX (C-){CtJ)c 1)n)tIO{Um) " -Ith cl!r1.iIin ""add

fdM\-uI: c.ampart' with F,g_ 61 Fig. 68. OeuulJ ot dOll-I", meclumhrn of ri~hc IC(."UrK1 thoracic .. piracle

P'.An ,'VI

VU,.../4 /J",syIf1(IfIl~G (Sauu.), quC'en: (Key 011 p. 2(8) Fir. 69 Lt:h up«1 of ant.crior ~n of thorax with Del!" h"lf of rJronotum removed Fil" 70. Righi HP«t. anlc'rict!" pan of IUGOthor.u:. showing lndirect wing muscles Fig. 1 1. Donal upec:t of riA'ht fore co~ howin .. in-;eniol1J or tlt'pro or or troch.anter.

and the ~yel1 mlhClH alUchftl '0 rlnl ul CbX;l ; CDnlpoa.rC with Fir n Fir· n. J..)orlal Obpttl of right fore con And mU!lCle' or Itg h~.)Ci comp:m~ with F;It . 71 Fig. 73. Rigla lI.!ipcct,,£ prolhnrax (UCepl pronocum) anc.la.ssociated mu.dfS

212 SONTI! AMr.-R'CA~ \'I::Sr1S£ 1-'-ASPS

KEY T O PLATES XVII, XVIII , AND XIX: THORACIC STRUCTURES AND NEST STRUCTURE

I." uf , anal \"cinJ ,.F,. AFJ. (ureal ann. of meso-· and mclbtunal apophytel

JlFS. apod'tntal (old of the lcatum

Ai t, .Ionlenor mcllnolJll In­Rterion

A.\'P2, .4...\'PJ. antmor Dotal .,..11'1, procc

APP .Interi!')t prOCed of pro~ atcmtlm

.!IIpF,. mdf,harcallJrld.eo ,fIf,Pr, aplal ICllc of fJl'Opo­

deum IA.t~ nfr'j~-4.r. firat, M'C"ond.

and lhir axJ.llary tcleril~. B. 00"" oi propodeal .pirllc.le 0" B.t, baAlarH C. CO". \'cln eFt, po,tcrior \"c:rtic.t.l vlate

of prMlt'rn.1 apophpi. CV. clo.inR Yalu Cr coxa DB, d .... 1 ba r tI~,P . • 'bellmenc pep: lor di ·

lalor of propodcel splracll! e,nt.'. mncpimernn Ep31. Il1I!Kp'lu:murn F,. (('.M1ur IN. pr~ltrQ.tl pit II, hamuli or IraQI hooL;, HFJ. HFj. horiz.ooul pl.~.

of mclO· and me.utero_' apopb7"dl

HP. hwnC'f"I' llt.t~ or aub· c:olul.cale

lad)I, lodlJ. propodul cit'­vator and \or"oo mu-clep: of u,t= abduruen

"dil, indirect Il.nd principal der.repor of Lhc r ore wma

Il1d. rc1nctor of the m~ pottph r:lgma

upl, dilator o( propadal .plrade

fUrl. indirect aoLi pnnopAl ~1cnto r of 1he (orr wing

IIdt·~ {ureal rctraClOrof tbe metopo:stpbraa.mA

1I1,1t. ml!lLOlhor: .. :~e lurco.-1'1",",1 ususcl~

IIfpJ~. m~CllhQnclc lureo­pleunl ml1JC.lc aud IIp­men'

/ls/, rei rae-tot or the seutel­lum

IIIis1 pleurA' drltreuor !;If tbe abdClmtn

IfU/ Outer pronotaJ rrtraclor o tbe KUlum

niu, IDtuCurcal mu cle

CUTICUI.AR S1 RUCTURES

LPP lAteral proc:ra of pro­ncrnum

LPPA.· , LItChi etetl or tb~ me.oposlpburma

",. C"I . (Ukd media .nd ~ItUJ 1

JIC I m~stctn:ll canna mILl median "QUlI .. Ulur<l lIPP/u . anterior prou s oC

metGpottpbra(II1.l ",SII. ""I~. median J1ernal

iJToova N.IS: mC-UDolal Ulllllf!ol OCr I Opr-, . QUia crut

a.nd Ouler 11rocHI of fint fural arm

Por, pIU"iI_det Pd. abdumia .. 1 pedi:cel PN"J. pcritf"C'1t1~ III 'Pir"ac:l~ pl . .. ",p,idal CurTo,,". PFI. Inle-rior rrocus b( tb~

IDCIOCUJ'CIII britlac Pltl. me lprrpbr..",. Pit.!. mc:.a(lO:!tphn,m. P4J • .JI'ropodeal prq.hnl,UIOI. P.JC~ ""Iltriol' e.n.n.l of he

propadc.1 pr~bra~m. FI.!A,. Plu') • • n{ttlOr and

po.$tt'"nc:tr m~plf:"\'r1IIl .po. d ... .,.

PI,JA ( PIJA~ lIW"IiD.1 and ..eeood mc:'aplwral apo. de-met!

P\"J. meQPoslnfltutn tri. propodeaJ tDOth PT, troulI'b or pro(uTcal .rm PTe. J)OIttcaula

ItuSCLES

II/iS.! . mctafu.rcal depressor or the .hdomen

lis],. m~ltIfrn.1 retractor of the propecUIi.

JlluJ, lIem.1 tormon mUJ­ele Qf IIbdom~n

1"'-1. "Ij. fint and srcorW mOtJ(ureal relracton or tbe propectll.!

IIlml , /llttI" productor and outtr reductf)r 01 the mid­cUe COd

I/Ilm" .tr:mal I~roduelor I. f the hind coxa

flllml , r~uctor of bll,<1

"' .. Illrn]d Jnd Ill; Jne...alurcal dt=l)re.lIoOr Of Iruchantcr and .dduclor oi middle I.,.

I1I1'"J. depf'CVQr " f tra cbantn and adductor uf hind Iq

Ifln,." jnneT rttluctor DC muldlC" c.oJ ..

(~12 I

PWP, rlrural _ina proc~. R . radiUJ Nr radSut I RF I. tran.U~11I11! rid&~ of 6nl

(urcal ann R-S~. fwt'd radilU and IUb-

COlli RI, radial 1«\01' SrI. :Kulum Sn. aculcllulQ SnL. lamellirorm procH.! or

tbe aculcllum SCII.f • • cutc.Uar Ie.le Sri. propodC41i .pit,ele Sl_~. mao5tc.rnuna SuT, mC'lIOatrrnallooth SN'. SIIJ, .ulHlAra; S.R. _..,bal.r ... idle T J, meta oDIUm TJ. humeri' tclune: of met ·

anOlum TJP, mtl.ano,,1 promint=:Dcc T P. me:tannta' .. lieu. 76. tqulol 'pbs, trlln.tlJ1leur.J Jutwe Tr, trochantin TrfJ. trachea T"I). U'oc.hantc:r TI'PI. U1lnlVel'N: verticlli

plate or the Inetaltern.1 lpofIhlli ,

I ~B \~ml bIIr "F]. lonliludiaal urtic::aJ

plate or Ih .. 1n('liL51crnal apophytj,

.·PJ. vutkal plate of ab­dominal pc'dlc.d

r R, eatodonal ridac

1111"'--1, pleur.1 oroouctor of bind ron

JOs,. .:tec.Ju r of Rcand tboracic Iwiraclc:

IJlpl. occlusor of promol pirac.le-

11,"'1 , 111, .. " twalar mDti· des Qf mHO- and melli­lhonnt

11/,1111(1 and Illftm..!b. pku. ral mu c.lc.§ of the mCl :l­thontic third uilLu)"

IIlfJmJo and 11t1IlJ;ljhr.,lTCU. nl mUIICI~!'I of .UN rt: ill lbt metathorax

"/,1"4. pl~u",1 Llcptcuar or ~ul('IIUIR

IfI(I"q . .j~pr~"a 0' the Mrt llnOIUm

IIp u'J suhaJar fUulClc 01 m,....,lhun.:"

IIlflUJ. tUJl. O f,uhalar mll.ac:le of Ibc mtUlbunx

OIl

Flc. 76

F.d n Y:IINla p.:"sylT'dlt;ClI ($aWlS,), cJUttrl :

F~J:. 7 ... Ctphlllic UP«t oi meJOlJlorax. and 11 10 mu!o("u'Olture F~B'. 75. DeLlil or ri~llt JlropotlGl ) )l i r.ldr anti mU"f'Ie. IIptrating It F~Jt. 76. Ventl1l l yiew of J't\)<I,lltnlUffI and ib <&ptlllhv it

••

"'11 n. uudAl pea I I . ptace 0 II~ IUr.lx ant.! II .• IUUlC • .d:tlure. with ft'""po .. qJU'<l:C1H.' In

Fig 7&. Right propode .. ' tlpir'il.cI~ it "IUKI , anti lht'.i r nlt.adunetltJ

;;;

• "

0:

'" -'" ~ ~ a ~ ~

£ " ~ § 0 c;

;; ~ .. .~ i.i: ""

~ b~ ~ ;;--0 ~ • to· <: ~.= 0 > ~ . - 0 .~- ~ a."7 E e:K~

§ ~~.:: --~- ~ 0.= ,

a 3 ~ ~ B l'l !:! :: = E . =;;e, )("!:::i ... -. e ~y c )(

~:2~= ... e - .. :::: :::I ;)

I:""~-=':~': ':'~5~~~ "1_Y~cc: :::J ::: '" - ..... ..-

", ... ,-:!!o: ..;. B ~~ !; ~~i! ~~ - 1:- -;:: -I; ... 11"1: . ;; ... .. c= __ ;:;..~

""E. ';: E e " ~ .! c ... -;;: ::s It ;:"i"": i::: '" l.<d!!ra~

~R g ... " ;: .E;C :;: -"-

Itun . IX

(Key on p. 21Z) 8.\, 8-t, KS. 86. Oi~lIPOIlU ' 11" Ju"l1 .. '· of p;I{l'!"f" (rum wall of 1"I'.s,w/tJ 1't'(I~UJflj(iJ

nes.l. '111e dou~r lina ""fII.\ridC" thr irwj"dual sHIp" or p.:&plCr wlllch muke up tht: ",hen.", F.o.ch IrIp It l1J.1dC' hOln I hJjul of J.Wll'tr flUlp 111-=-,>< ·'.hdL ,. tOIbU.t t"!!Ipe..:'t1vdr uf SO. -45. 171 ~Ind .Je) trip See IIl~ Fig ~ ~tld .lZlI.. Plii.l.OI XUlJ .ud XLIV I itllf ''''' • . ',-Ir./Ilhol l~ ' 1. 'IUC"<'1I u1hm of hlnfl "'"Ill( hOl\IRU il~UlUIt or I... t I _

2 1C> ;-.'"Uk r-lf ,\ MLW ICAI"l K.'i I'I.hH \\'A "S

XPLANAT ION Of2' P LAT E XIX (Ctuu itJf.l~d)

( Kry tm I' 212) Fig. 88 t.:Iucbl p«t ~r mcMICho,,",~ .,r PiaJl.I~ And p.1rt ~( Its mU5CUb.tuno ; $Cutdlum

iuwrtd dcpreuor1o of \\ inc and «""am olher Ulwclt remo\'cd, And postcrio; m;u"'1I~1 ~pleum.llIpo&nl~ CUI 1\0011

Fi . 89 Ddlfclun,-'tl/,u/O Gr~"(1rlfJ (Fabr. , inallal cdl of Ai Ilew conlh :md rudimenb of stcOnciand third cda

Fie'. 90. DoI;cllDt'upuJo JUQclifUltI (Linn .) • • id~ vitw or comb from q~n oe:u. showing intenel.:uiw of cdl bbo: the «II arc: numbe.,td HI tbe. sequenC'e in '\\'hich they "ere (,OO!lilruct~

Fil". 91. Donai upect of comb in FIC. 86, showing dluKTilJ11matiCillly lhe inlc:rrclui I.!I or cel l baJea

Fig. 92 V,~ftJo /!~N.r)'lf'OlticQ (Sau I .) . qu~n . a.nU:l'o-dors.ll asptd or mesolhoTu and part of hJl. musculature; utum and Inchrm wins: muscl~ rono\'ecJ

KEY TO PLATES XX AND XXI: LEG AND HEAD STRUCTURES

C1JTICULAR STRUCTURES

ANPJ. anlwo r notal wieg

l 'Den, ot the- mclllboraJII: 1 r , uf..t'. JA.r. 6rat . 'eC'Otld.

and Ihird axi lla. ry KI~ntel 80). bualare o f mUlllborax CEo compound eye Cit. ely,...._ C,. O)xa Ft, femu r FI, fl •• cJluQI Fr. froM

I II,n , . produclor of middle ..,n IU".u, atemal produc-tor o f

hind eoxa 111"':1, oU'" r r:duclor of

middle co.u II JlIn~. teduc-tOt of hind

co .. 111"'30 :md Jb{ IIIcsofurul

dcpl'c:Mo r 0 trochanl.C'r

fl'. coronal .uture 1 Is" . (rontal auturc:t ? M J ruandtblcJ .liTo mC'!tat:ll.nu. N JJ'J 1. lrf.1 melanotal IC'~

ment O. o~1lI OM. oculo-m.ab r ".ce P" pedic~J Sc, .capc Scr. Kuper

MUSCl.ES

and adduelor or midd le leg

[{11m]. dqlrellllll"H or 11'0 ' chante r and .l.ddu(IOT f bind I .. "'JII/. mnu reduClor or mid­dle C(lxa

fI 11111./. pleural prodl.lCl0r of hind con

SIIJ, .ubawe: of m lalbor,n: 1 J , n\t1anotum TJ. humC1"al scJerile or lbe

metanalurn TIP, metan!), .. 1 promlnentc T 11, rndaDotaJ tul~ To Iln\lJ T,. tibia T . tibial .pur TrD. trochanter

JII/ttJlI. b.u.alar muac lc 01 mC:l&tbonx fI''''"'(I and /lfl",ub. pleu · nl mutC'ICI of mCUllbonu:. Ie th1rd uJlI.ry

JlltmJd and IlJ/lmjb. ple.u· ,...1 mQ54:!lft of mctathor-.c­ic .ubot.re

IfJ,mjt coxo-wbalar mUKlc of mcllthor.lllJC

KEY TO PLATE XXII : WING VENATION A D ABDOMINAL B ASE

rS.e p. 2 191

(Numben qtLllIH,inR' the- dr!ll~illioOAl of \ tin, and crlls Cu.: C., IR/) h.ne. bnn millrd here for ... Ile o f buvhy. It i. auumed th.t lhe aianifieance o{ 50eb nu."b~n i

uodcrstood.)

1,:4 an<l l "dn or cdl a • anal furrow

P • • vital «II .-fr, axillary c!c,;te BDl~. unal \'~Q B(,IA . h ..... icubu.IIl cell Bm . baJlmcdian cell Breit. brachi.l \c:!1n C, eNtAl vein or cdl CAS. cubho-an:ll .c:.le

.... cubital '10-('.10 or ttll rll · tlt c:uIJJlo....:t.n:&..l Cto .. " ... ·clfi Dj$~, dl.tCoidal nm or cell He. humc:!ul pl.te o r aub-

co .. ,,1 SCile I-("w. IOC('tculiuall.:IUU __ oei(1 , 1Jl. i,uenn~i.an crOll \'-dn

'f, hnc alon, _bieb Win' fold. In J"C'iiCU

'" medial c:ro5. \'~i n iI. III~U.J "elo or cell ttt-CW, medio-alh,t,1 (roM .... ,}I·Cw, In~io ·cubitlll cdl Mri;o. mt:tIGllfpU' ll~P. mrdian rl4.t'. mI. mC"di:llD rurroW' X rn-. rte1'VulU) Pd. ahdomlnal pedicel p"on , "U':m,1 Inh .. p"lr. pr~jjI,ry ,· .. d_ion PI. plt'rOlfijttllA or .1I"m;a It , radi.1 \C1n or «,11 r, rad'AI HOD Yan

"JI ndlaJ forrow RN'. rrcurrCll l vein r - tII. radlo-medial (1'OU \vn R-I, radial s~clor Sc, liuhQna,l n :in or cd! Sm. aubllle<han ('tH. Till ·

yanl'. nonlendalure S.tJJm:.liuIKli~(Ildill .... cin S.bm. ,"u'llrlf:LJI.an \'C'in or ctll Til , .IttOnd abdomil'lll ler ·

Rum TrCIf . • ran.l.u:nc-cul.!ltal

'rrin tl umbel ,'PJ. \('rl;c,1 pIA ((' or ,I,..

",unln.1 rr.diccl

Uf01.Qt'.U l.. \ nL II. "'~ 1 Pu.t' XX

Fit.. 4,16

V SItJ.1J #KJ.,Jva"iCG ( S.au~,.). qtJ«ll: .. ~g. 93. Cc.ph~ljc upec:c. of Jde hind eon 10\\il1'; 111luclo ul leg b3~ l·lg. 94. Inner UpeCI. or b.ue of right hint! wing. ri~11I f llUW or nlel.aootun~ .nd direct

mtbdet. 1)£ the: winl'

Fil" 95. ~I)halic:: .upec:t of h:h mkhllt:' ('OX;I. 1.howink It~ mil I~ ; fIlu,cle spre. ... d to th"Pby Lhenl Co Mlte:r .. d\'.nt:tt:~ Wh~ In norm.al pMilion UlllScle: IIlml . IfI.z. rr!lf4".exl~nd forwunl and 111e''''!! ( ttl th~ left in the figurl!}.lfI",J anti/lim" lYII)r: III the Iroo,flh 01 IIIm1

fig. 96. O.uter upel:t of 11ol..fI.~ 01 ri;ht hllKl Will~ ... llImine rnlcl"rd.aliol1.!i bt'"«n dU''eCt wmg mtc.Cles IUId il.X.illu)' tek-rites

FiR tl7. Otlrg, I :up« of 11oiL.~ ul Idt hind I~ with ",lfq"l~ pre>t.d

FlU. t01

Y,'two /u·HJj.h'O,.i(o (SaU$S. ), Clu«'n. 1;'". 98. ud; 1 a..'pcd "e lcfL fnrc Ic, Fig. 9'J C1~IIl1ic 1L!op«t of lelt rb~ 1~ File. 100 Cephalic a pttl of had 01 frulIr FiR WI. CelJhllllic CUop«1 uf ,Ii .. ul

lwu thiro... () riJtht hind lea:

F'K' IlIJ. fig It).! Fig. tu~ Fi~ H)I,

l"LA.u X. I

(I>:<y 011 p. 216) Cepl\Dh .... ~,pcct ur ris:hl antenna clr.ln~f ( nigil) cl-!ll .. hc 3~J-«t or ri$;itt .mlcnn3 oi Il\ol.It

C~U'laI a f'i~1 (,r tlglll Ullt lolle Jqc iq,h..l..lIe .a.:.r..n:1 Ilf l.a.\oC: uf It'h 11IH~1I1!' It"~ Caud11 ..-eel "I ri .. llt hipl lC'g

II!

. ......

~(r.J=::::~::l:::~;;~::~::~~~~r~. ~~ u< IA .

TO ' Fu .. 108

110

IG·!II.~C.I·C&I' u~u.u.

Fu III

.... Fu, I I J

(Key on p 216) File· 107. t'n,a uGbra (Linn.) , ritcht forr wiDe \Iollh 'tins aad cellJ;; rulmtd .1camhn.E

to the nomend:uu~ of Rulmer .. nrl t;ill!ao. l);l"\C't1 0" Ihe Jurinc:m ~p.ltll1

Iln/M/a 'rIIs,hlCl.~lJ (Sauss.) : Fjk~ lOS :md 100. RighI rOtc and hind "inlo;"' \\-illl ,·rin., ill") ceU. n"Utt'fJ ~C'CQrdil1g (0

Syt;trtn pf"OptJsed by R~ Fig 11 0. Ct'ph.lic: A PfCl. DC ba)C of abtlolnco or 'IUIeel l

Fig. f I I Dct.li l or Ihe {ure WlnJr; 1huwlng 'Iull., flO ('uhau. Jb Ju.,t hc£orr. It' Junction ~lit.h firit Ukl

Fig

F.". 112. rtsplf.la NI')rurnl (s.,'H1~). rl.Kh ( ron: \\linK Wllh HIPS oiIrnl cclh. ~1~lctl ;u: ­

oording w Till)~nl', ,> &"("111

I 13. JJoUch~",~ .. tJ"ltr4Jna (Fabr ), flf;hl (ore wing with ~· e-in and cdl:i I:lbtfed !J.ccnnJ.il1~ 10 (om rock-N«'IlhoJw ,y~t(11t ... illltr.,rct~1 IIY R~dl('}

II

2..'0 S KTlI AWER r CAN V £'s P I NE \0\0 1"5

K EY T O PLATE X X II[ : TH E ABDOMEN

CUTICULAR TRUCTURES

."c, .nt~lA A,St. apodeme of a1x.lomllu)

Ilemum 11/11. apodcme of abdominal

tct aurtJ ASS, anterior tlernal bncc ATB, anterior kl'1i111 brace CO". i.nte~gmcnta) eon·

lunetiva DSB. dtqonaJ .teroal brae~ DTB. di-Ional le:,.,41 bnacc FStl/. POSI~riOI" ..nirutar

di'r, doraaJ inln1~aJ;l re-­In(to r

dSt/IO ds"VIII. dilaton of abdominal _piracies

U~ and i.u~, Intcrltcm:a l r metDr muscl6 ,U. jn lcrtaval eItcli£Or

ftansc or I«ond abdomi· na.I II HTlU QI

FStfll, &n(C"rior ;u1icu1ar Banet' of third ,tcrown

FTIII. anterior ftance of tbird abdominal lcrgum

US, lateral ster n.1 bNlee Pd • • bdamilla1 ~djec l Pre. pr~.'a RS,. reduplieuioa or ahdom I~ sternum

RT, reduphc.atlon of .bdardi nal Icrf{Um

),(USCLES

"S" &nd " sr' . i nnt.r terKO­nemal rt:lracto,..

lur. btual Ullt'nlcmal rc· trl'Cf,o r

UI ,. and I.fr' , latec.1 In,et­tcrc~1 retn.Clon

mUr. median ib'«J:l~["nal re .. rraClo["

t 120 1

S,II. S~III. t:tc., ilbdol1lin:.t .pir.des

SliG • • UftJ aui~ StIl. 51111, ~tc., oabdomin.J

Itona r/l~ Till, t.tc.., abdol1linl'

I«p 1,1, tnnl'-cne Jroowe of

Ibird abdomin sternum 1'Pd. nrtica l plate of ab-­

dOtlu.n.a1 ~_h;c.l

tUP' 10 O$flVlIl. otc1usors of abdomIna l splnda

01..1", and oUr. OUler tergo· nernal retractors

I" and tU'. Ie:rl'>-!llunal e-x · t c:niOU

Da;'l.(1(.1 ~t. Se,t' . \ I!l 11 So. I

1-lt '-If'h/d t~'''X-,JfAHII-C-'' SOIIl1n.), 'IUt.Tr.·

F'!K". 114. hm~T a peel of .. btJ\1WlIUlJ It'rn.l fig. I J5 Ventnl ._j><'Ct or 4bt10l0(-11

r .... .. , 'Ill

F'ig. 116. Right i.al("r ... 1 UPfC'1. 01 ilhc.lUI11t=I1

Fif:". 117. tuner asP«' of leil h.;al( ,illbdull)C:ll

222

K EY T O PLATE XXIV: ANT E NNAL AND ABDO MINAL STRUCTURES

Ar. atlleco ..... An anut .,,~B. arm. of stin, bulb ASS. anns ar atinc 5btalb bl~ berbli of JUnl laoc:et "'. diverticulum 01 bodt

cavity in _tina lancet Cit". COif? ClC'. , col'lJunetiVllC' D7VII1. dorsal nrip or

d,blb tcraum DTI X . dorn.1 .trip 01 ninth

lcrrum LTJ'II1, lateral lobe- or

eiah1h tUK\lJ.tl

LTIX. latenl lobe. of the ninrn lel'cum

P D .. , jlOiJoJl dud. PSo. poiaan NC PSI. ova l plates o( ninth

sternum RI, rectuDI S8, I,in, bulb SL1.adnK lancct S/tt' III . eiahth ahdominal

" inac.ie SPo • • tinl pIIlpi SR,. atiDI r«Cljla . S. stin, ,beafh

SSt/X. su.tkll .,£ " lath .ter­num 0 r (cmaJe

StYI, S,VII. t!tc., ahdominal Item.

SIX, I tt':rna.1 loUe. or leath 'ermeot

TI'I. TVI/ , ete .. abdominal IUp

ToY. tt'rpl lobe of tenth Iql menl

TSYIII. triangultl r tckrilc o{ a,blb alunum

EXPLANATION OF PLATE XXIV

II tltNi" "vyJ..,.i<a (50 ). QUC"" :

FiK. 118. Lrft lAli:~rIl18sp«t of sting o.nd as-.ot:ioattd Imdure" ( xc.~ Jl1ulIClcs). in nor­m:a! pcbltion. ret r.tctttl within Min' f«C$J,; 1ell halytS of "ilC"lh w ~"'(nl1t ,tewa and tera-a rCfl.lOvtci

Fig 119. Oorql oup«.t of .unjit and :U>oci.lted ,Iructurn in nnnna.! rcJ.-,ionship WI re­mu\-ed (ram the stin!( rK~~

Fig. 120. DOrN] il.'pe(:L Q( unt: and ~btfd ~trU(l ur~!> aftC"f rmtO\·aj of rectum. eichth tergwn. ri cht hllH of ninth t~flt\Il11, :lind n,ht half of temlt rgT1tCJll

Fig. 1_1. Ventral -upect o[ !.Iing IItHl a.s5OCi~ltd "Iructurcs 111 noflll;tl rd.itianJhip bUI reruo\crl frcwn !.ling ~

Fig. IU. Victw uf ting me<:h ni4Tl1 with p3n~ 011 Idl side mno\ed, 10 1110'" rebtiou of poi1OTl duct to wng bulb .

Fig 1Z3. Left bttnllt :L pKt of p:a.rl fi of "'-in~ llwhilnisU1. winch He devdope\J (rom ninth ~nl

Fig. 124 I. • .c-ft bIenl npect of nin~ IlUd Of tlCialeti UructUIts 1I her rtmo\ .. J uf eighth trr~m

Fig. 125 Tip o( 5tiOif lancc.1 fir. 126. Extremity of irq; 1:anat .lre..atiy cnlOlrgf'd

\.ut \ III u I Pun XXJ\

1:.1\1 c.i. ... , Ful. II~

1., 'Y1ll '1'C

, I

F";,, JM

~II ,I LTIX

Flf, III

'" .... .. F" l~':

Ftr.. IZJ

Fn .• I15

KEY TO PLATES XXV TO XXIX : MALE GENITALiA AND MISCELLANY

.-Ie- anl(co.tA Aces apica.l earina or 1.0Ro-

atipu AIt, AlJU-I ArP. apodcme (lr pcnu A"sa'lII. A"sax. opo-

dcrneJ or abdominal tter­.,. A,T, apodeme of .bdlJmIQ~

let""'" AsP,., 'Pleat .aId! or tbe

pt'Ol-ooc:um AT8

1 ilhtctior Icrpl hrace

ilL. lUll lohe o( f"no.I'peI BLP. bucJ 10M I) ptlliJ C, cc:.ll ceo COO'1poUlId Of!' Clt,ci'llC'W Co,., cOIlJI,lnli,". C ... , toU Jltr, donal inten.cI'I:al fC' -

tr.ctor ULI', donal Iobt- or "oleella DTB, d)'aonal u~ry.1 brut' EpJft.l. mCfcJ,11mcNn lipu. mCklU lrtnUIQ J-,. r tOni ct, lonourdo " .~ as .onosbpu

He humrral plale or '00' co.'al acall:

11" , Ifttcranlcnnary Irian,le I'~, intt'rtC'rpJ C'"" lc~r I.,,. l.ucnl inlt~1 re'­

LfiletOr LTH t.1C'rallcrpl bract: L 71.{', btcnl lobe:ll of the

ni nth 1C'f'lum

oW+-c." fused media and cubitu. l

Md. mandible (J j oulli GU. o«ir,it&1 wlure OJrl. exu Q-m,alar ,lace P"r, ptlrllp"idCt Pd . abdominal peei«1 p~. f'CtJJccl Df anlmn. Pn. pmi. rl!rJ. prnlrcmc of ond

thoracIc tpir.cle-PI, para!) ida! rW"t'Ows P,l. proplcumn pb." mctoplcur.I.u1urc riJ. mcul.Ilcuron P/j" 6 .... t lU~.pleur.1 pit plJ6, mel.plmral ,C'condary

.utUfc PSj~ 1J1d.po.etnotum Pr, propode-um Pt'~. precot'" PT"J'IOtUCIula R. ra iw R" rectum RT, r~uJ.lieahon of abtldm·

inal tcrgum Slt, UliUA SaS, .... iu.aJ pine S<, ",.pc S~"II{. tule. of Ihe tlPU1 te~m

Set. fCut\lm Snt t tculellum SOlS .. , 1.ltraJ tctHcllar RK -

t11cnl o r l:!tnt. S,.X, I~lb l!(01C',nl S" IIbdomlna' '{lIncle: Sf'}, propodcal .pinelc

Str. Ipiculum Sq, Mluama Sl~ meto,ltTnum sat. 'ceon .. 1 abdomlnalltHo

num SIJ, mdUl.trnuru SI.' , f:le., abdominal 'Iunl s,' 'lJh-/X, fUJol!'d eighth

and ninlb ItCra..:t T l . pTOnvlwn TICI , .nterior proDoIli1 Cit

<ina T,,'.I. wcoodary (1 ... onolal anna

T tej. ttrtiary pronot.i&1 (::11 0

nna 'r,p, prouol.lll pit til, «Dod abdomi nailer.

rum Tf Qlctanolum T I, 71' 111. tiC., ;ilidomlnal

lerp T I la T'f:,.1!!tAI It'rmlnal polnl

of gonOllJpc. TP.:, \~J1Int1 terminll I'OInl

Q( lono.upes 1'0, troc.hantrr 16, Lntn .. eutal .lutuee rl. ()'plaid. "CGS. \c.-tral emn. of go.

DOtupeli

"", \ 'C'ntx 1 '~1. "',,r , \·olkll.a J'Pd, ... ~tal place of ab-­

domina' .,wittl 1'1. tlCUto-.&t:ulcUa .... ulun~ ron', "crur.1 tul~rde or

'rolaclla

KEY TO PLATE XXX , STAGES IN LIFE HISTORY

I co J. lhorac:ic '\CIPt1a1tJ flt Ilin-'

I 10 X, abdomlrt11 kl'met1l5 of larva

• .f antrnna ~"lo. ;lnlennal foramen CE, tompound eye CI" cl,pan CP, Clud.1 Ilapin. er, coxa F,. rcmur FI, ft.I(~UUIl1 F,.. iron. /,*', fronbl IUlurc Ce. i!cna Glt aloua LbB, ).ahiohllM:': LbP. labial pil ip UP,. . labtAI pn)mlDc:nc~

LPL., Jllc:unl lobel LIP', hnR~ LIII, 12bruns l.TI, po.te-rio r lo l~ ur pf"~

llUt\lm AU, mandihle IU6J, "l6rJ. medJ'" .tC1"nJll

ITOO'CS of me.".fernum and II lelVlernum

.'/.r, rub I tla [ttlt' , pu'.p ules Pd, abdominal lK'dlccl "', p.al'aj».uJaJ (urrow 1iIJ, muor,tcu(on P,_ flNpot cum St', tcapco Sri, IleUtUD) $l'.,s" .C\,IIell., 5C"KrDent or

uilla

12241

SO, IIli"-UT orifice SIt . • pu""cJes S,U to S,t'U , abdomirlal

,1t=:rtI.I T I, ~ronotum r J, met.anoturQ Til 10 TI ' i/. abdomirual

Ie"" Til, t.nu. tIt lc,"PQ,..1 fOPa rl, Iibla TIS, tihtallpurt T,o . lroc;banlcr U , ttanucuut aulurc I 't', \ertu rJ', ..ctHo...eU1~,n.r .utuf'~ U PI, WPJ. W'lnl padJi

Fi" 127 FIg 1.2N F'R· 1;!9 Fig_ 130 l;oig, IJI

\ . '"

E XPLAN ATION OP PLATE XXV1

V"pNl. ~.S)i,,,.i,. ($oIUS-), mat.: (Key 00 p. 22~) Fig Ill. ~tnlJ aspect of pc».tuior ttkI n( abdomcn Fir U4. Oorgl aspect o[ rmimlia still ~osetJ by ninth anti (~tllh (crp and the (used

eighth oa.nd ninul sttma Fig US. Dorsal RSpect: or pc.qerior end of abdomen Fig, 136. Ldt lilt.e.r.tI upttt of t.re.niUllilil end())('(j hy ttgmcn~ u\'tn to tm . left hal'f'e,;

of uXlh and seventh tcrp bnd Jtr:Tf1Ol removed Fig 137. ent lAl iUpect of fused eighth and ninth .. temoa with Klcrotixetl vdliCes of

ninth tcrcum mltilcbed: Ipic:uJum poinLJ cephalad Fig. 138. en.ral ... pcct 01 Renltali •• ,,<10..-1 by "'P""''' cigh! 'm Fig 139. 1)o,sol .. pcct or c<m~ ,emoved r,om .btIoma. fig. 140. Ventral IiUpect of ~illliia. removed (rom abdol1l("n Fir. 141. Left tatent aspect of g~itab" cnclo:Kd by lug-ite 9. fw.ed Slttnllo 8 UK! 9,

and oq-roc:nl 10 Fig. 142. Left l:ltenl a!pect of genitalia remo",ed from lhe abdomen

lImu I.U St IUf"flI. \ I ", lit fl I

ITaX

= ,"', n.

\ \., r~ So

Frr. 1.18

Fm. 14 1

\, \

'\ 1w;\lr. IS. . ' . \ \ \,.,

Fm. 13~

s. :;.

0'

I'un \:X ,

I ..; t ,C. u s

Flu lJ7

••

So

F ir;;. 141

EXPLANATION 0 1' PLAT& xxvn Va,,,,. ~.JyI, ... ic. (Sou.u.j . =1<: (Koy 00 p. 224) FiC". 143. Jnnd' b:.JJ of gon05aTOO tlnd of right &,oaotlipa wllh volJcH. and sqilU At·

,1Kh«l Fig. l.a.4. Inner aspect of leh haH or gonocardo lind of IltCt ~on~tipe:!, with pcni~. If,aJ;" itttl,.

and \"Olsdb iillUu::hed Fi&,". 145. Inllc ... aJptct or righl '-rlua and volJdla remo"ed from I'onoslipes Fig. 146. Vcnlrnl tl:!prtt of ~jJ .nd portions or gonotllipes 10 which It " :attached Fig. 147. Dorsal Ibptrt ot pmiJ uod portions of eonOrotJpeI to which it is attached F,g. 148. Inncr "'pea 01 riBht hoI! of JUnoa.nlo and or right Ion ip"": .<>lJclb and

&&gilt. removed,. but .aquama in plsce 1'1,. 149. Proximo! upe<. 01 genitali.a

Dolidctn;!~ltMl4 4"rw,;" (Fabr. ). male : Fi~. ISO. [nnc.r upect of right tWf of &'OOocardo and of right &OOOStipa. with pcnl!;,

voloella •• nd .ag;.ta in ~ Fig. 151. Don.al 8$ped of g~iLl.li.a removed frotn abdomen Fie. 152. Left latent ~pect of gmilaJiA rano,'cd from abdomen Fig. 153. iDll<r .. pea. Icft b.oIl 01 (O,,_<do and Icft """"up"". with ,OlJelJa and

53Kitta in piau Fir. 154. tnner aspect of lett h~lf of gonocardo aOO oi teft gonoslipes with \"Obe1t~ and

acin~ mrtOved. but "'oith MJt.WnOI in ,.laclt! FiB" 155. Ventral aspect of gtnitalia

Flc;' 14J

FIG. I.f()

, .. ,res

Fit. lSJ

'1'1':

PLA1..., X.·\ n

" ... i Tn '" h l __ 1.r; 1

.,--.--

~n XX\UI

Ftc, 157

F,c, 161

F,o. 158

(Key on p. Z24) Dolicltovrsp;Ua arrn0:rf4 ( Fabr.), queen: Fir. 156. Left lilern..l upect of thotiLX 'Ig'. 1507. Cephalic u:~ of had V"'J'G crobro (Uno.), maJe : Fig. t 58. ~bJc.j ccrhalic ASp«1 of he:ld Fir. 159. Dorsal bp«l of "eniblill . Fig. 160. Ven tral:u.pect of 6:wiuIU., Fig. 161. Ldt Luen! :,l.tlpttt or gtnlUlha Fill" 162. L,.{t h';llf or g'OnocamQ and 1'0000tipH . ...... itJI yol!Jefla ;;J.[ld sagilta in II~ . • F lC. 163. Rtaht hill r Qf ~nocanlo and I"OfIOIItipo, with pen,,- vob.ella. and lakJttQ m 31(\1

(I' \ lit. ~"

r." ,'". , .• I -

FIll. 167

I' 06$/IU t rabrD (Linn.) : Fi._ 164. Left I:uend 8.fjpea of Ihorax . queen Fig- 165. l\oIale. Qudl:1 UpeC'1 o( ri~hl

.wt..:t,nal llA.atc:num

Flta. 168

I ,

"~TJt , 'XIX

(Ke,· on p. 224) Fig. 166. Queen. cr-pbalic iUptef 0' hu.d Fig, 167. QUttTJ, daClaI a'/ipecl of h~d Fig-. 168.. MalC'. ur.~liC' ilIiprc1 0' head

fl!OLOGIUl ~11'''fC. • \ Ul \"In.,·tl I Pu,1' XX.'

Ftr. lift (Key on p. 22")

F[.r 1(.9. ,., n,Jll(J I"'tLtylucH,iro (S3~.) qurrn. \"'I!"ntnl 1110.11«"1 of pupa Fi~. 170. Vc:ntral upect oi b.n.-a Fig. 171. Cevh.:.Jic: l.ot.pte;1 of the 1ant.lll hCHI

F ijf 17l. l)(lual hpttt of tiipterow (1) ~ Iltt;1C.hetl 10 ru:'~t papa or l',sp.illa ~pp. I·'r. 173 Uler.ll 01 peel of tt:'K aho\ i"~ m .. nn~r ui its an. ehnkt1t to "" H of ~II Fig 174 Queen, '~ft latend ""Ipcd ot lJf'Vl& ut 6amc: F'iJt. 17~. I )utlinr 01 ITQ.(ItlVe,rw 5«:lIMl of IhJMCfOW tmr hDwn m Fil" 172 FI~. 176 S~ 3""'" /'tfUyl. <Jllit. ( u~ I 'Iuet'n. Id t htt'raJ J'C'l..l CJl VU~ ~'-----~

l'ut XXXl

Fip. 177 to 179. Va/ltd" Jk'".f)"W"I·~41 (SAus'!;.) . • ,bdol'l1inal c:ulor p3t1trI1Ji Fig. 177 rl!pr~lIlAI the I,.tl~ !11.a1 I C'OnulIont:!l ill .... ·orkt!Ni, F ig. 179 the po.ltlem CUtll­rrtmteS1. In Q1.!ctna

Figi ISO to 182: J.' u/tw". ,wla ~ Unn.). '''r. 4l/rO/,I/CtItJ ( .. Iadl::n). Mmot wtlrkef10 h;4"e a.bdottun .. l1 IliI!ten 'Imtlar ttl lh:lt of fiR' 180. mo"t (Jut!CJU ha\e. pattt!nl. simi· .... r to Im..t 01 FIR" ut2.

I·ir 183 '-" 185 S"o,,,I,, "'II (Unn.). '~r tlfdJlD (Sa.~.) Q.bdontllw colur p;lu~rns

PU1 1 '\:\.\.11

.... , .. 190

Figs. 186 10 188. D(J/lcllof:u/"f'/4 4I't'",lnd (Fahr) t abdoom\1J color partuns : that In Fir. 186 tYf,ial for Pacific ~O<ut ~·ork.er!>, FiJ;. 188 fOT l:).3.ci6~ ~l queen.t

Fir, 189 and 190. Vts/,.IG Iq!lll'HDIOJ (Dr. ), ( -:- " ~a,."Ii,.a (Li,~ »). abdominal color pallerns of a tYPlc::t.l work(:r aoo a tylncaJ Iluc~rtt rCitKctndl

Fip. 191 and 192. Vts;"lo SMlr#n~"nl (Sad .). aWominal color 1,;Au("n\! or .. Iyptcal worker and a typical queen. ropa:1i\C'l)'

rip. 193 ancl 194 Vup./IJ ~/IJ Linn). :lbclnminal color p.merns

UIOLl~1 At. ~1"Jn \ m._ \ ilL ~u.. I PlAH • xur

Vt.r,"la ~llS)'Ir'O,.iC(J (51 Fir. 195. DillgTllm or lobed comb (rom. ~l built In tony rround (Fig-. 231 on PI.

XLV). ~n:uwa tllClw axel of Krowth: lohing in th is nest ¥o"U dUIi!: to rcstritti<lm on IrQW\h I tuJKl!Rd by .. tones

Fie· 196. I..o~ C'Oft11, IMm. not builT III bOl1'lOJC1lOUl soi l : lobing d~ to difficulty of ClddJnsr. 'A-Ifhout lou ,of p&m:m uoi fOTn,ity. lure cell for queenarearinc to m::~ o~ ~fUb comn(lntr: of 'nall cell, i" which ""orx(':u aNJ males are de­~~ (0 .......... f' kC"1 bne i l-.ticalu!, bOUJ1doal"~ bet"'etn rt,ionA of ~31111..Dd luge «U,

\OL \1I1. :'<.:n •

Flo 200

Ftt. 201

rig, 197. Side view of qura\ n~1 of I'tr,"'tt /,NI.ly/wninJ (Sauu.) builL in dark .hed near I{Obeq(fI, La~ Co\lnlY, ~lirornu.

Fig. 198. Ventral "ew Fig. 199. LonjCitutli".II1 a.ccthm fig. 200. 'ide "itw of queen no' of Vt'J/,HIIJ /,~'U)-I~lI~I'(a (Sauu.) , ftOm rodtnt hurro""

four roches below the crouttd ~urr"te Otl trilbltte nrM \1Iddletown, Lake CouDt)', C&JlCcmia

FiC'. 201. Ventnt YlfiY

Pun X. '\"

FIe;. 205 Fu:: .. 2f)}

Fir· 200. Side view of queen nc.. .. t Df DlJlidorJ,s,llla 'lulnJalQ (Linn.) (~) built en a nett of J·oli.nts ,-". of the JJr«et.li0it yrat

FiC'o 2tU, I tn:!J;tiliar m"ss or cd,ll huilt all'lng-aide the comb-l in G Y~ry large not of OO"(Ir~#"IdQ arnot'W (Fabr.), Suanfortl Uni,'ersity. CaIHomi.

Fig. 205. Side rirw of quren n~l of Dolld€1'CX"z,tJ(J I1rt'lI"m'" (Fabr.) Fig. 206. Vmtral view of Mn)C

Ftc 207

F,o.2\)')

FI, 207 Nf:st o f DoJirlt""rs;k!G ",.,,""M (Flbr.), ,-hDwn in Fig. lOS ~ longltuwlII:l,l ~tion

Fig. 208, Soall nc:~ of I'r,jtlf/tJ t~"~yl;.,a,.tra (S.u's.), about thrtt inches in dinmdtr; the fbrk 'hiny •. reu on the 'col\ .ff' r~lC'hclI nr ~hcH~c: u.ct! til repair thr: \\, .. 11. d.un D.r.:~l in ttl.Kinl (lU1. tht' n~t

D'ltloo..I\'At ~.UCL" , \ ' rll", 111, ' ". 1

F,e. 211

rUl . 11.1

Fig. 211. Nesu of Vut~IQ tr"~!TV"U{1 (SauS\.). thown in f'i~ 208-210, ICH11f111J(fl11al 'loCt,ion: the 'Kontl cumb had been lou ·" Ilt'n ti'is ph~rtlpb ", .. 1' takm. nOle intrnr:bljou~ ut :talea. af 11o"lJOirt in· Db-I wall "hovt tht

Fil(. 2fJ9. Fir. 210.

..... ,uI.XXXYI

'Fla. 208

foUl 210

Ventral vic',,' of ... me:; n(lt~ Irn"f:\tlir entl'lln't hot~ Ventral view with lowe'r PIn of wall n .mo\'«J tl' I~OW Lhr COfllbt; inl! ,,' comh tfltar.:tti And !'«and comb ~1art~; I. ring Dr cocoons in fh~ lint zone j)( \\orkef'­built cdl~ ~urr(Juntf' Iht' C'~I' ttuill h.r the- f4urrn

I', "n XXXVll

FI(O. 211

1-1 ... 11-4

Fig, ll.l. Superm,mcrary ClI' laid by ,,'nrll:tn In l"Omb or Ift..­".IIJ l'tfllldM', (Linn .)

Fi",. tlJ Quern oot of Rnlt" buill in mlwtJl)tl IllImp Fillf_ 114 l-llomriluli inal Aft1'rftn n( _n ..

c~ _0

~~ ~'5 ii'

" - .. ~-= 0 ei: ~1!

'" N .. ~

fit., 217

Fir· 217. Mrdium· ... ilftl netl marlr hy n younli: ~iRI')rl1U" ('(lInn,)" Irf OlJlrrlimV's/".Ja 17rnl-err"", (FlI.br.). in Q lJnme tree.U ·upc-nmo. Sant..l Clam. Coumy. Ii forni,

Fill'. ll&. V('ntral view or $8nllt, II! redueed

c z ..: =

"I~n XU

Pig. 221. Mature: nest of DoJid.IYJr1ftUa lJrl1ffori4 (Fabr.). huill in a p.uch 1)£ Uec.h:tnu\, a~ l~tJ'~r In" herLJgt; hrl>t tuuching 1M fTTJUltd

Fir· Z22 Vt'ntraJ \it\\ or 5a1l~. "",11m liltro ('tlhl the: J,:rdu:rWI. f'nm&n(r allop of photo­I,fraph I "fIfe dl.U If"" not "" .... " il'lt"~",~C"l1! f~UE:.llb. Ihe .... '1.p' Qf rhl gfonu. Iaclr-1"1' the: l Lince. II) CXC.l,,'lltC And n ke room (or ntlntml t:':K( .111 inn ot tJl~ 1. t

~

" ~I

z d ..: 0:

il " ~ ~

:; l. J < ~

~ 5 0:

.. on '" N "'''' ~bC

c.i::i:

';-5 &0 ., ~O

oj] ~

~:! cn _ -.:

~1 .-~! l~ cO .. :I ... .. --20

• --:= OQ.

ii~ .2 ~ .. • ~:;; ~ g] ~ .5t:t: - . - Col

~ ,. t::

F" 13..

FIt, 21't

Fi~. 226. Mature nClltt of Vr;,.lo vw/guris (Linn.]. in .itu, in KIll an hillside Fijf 'l27. Climbs of lAme nest WltJ, em'dope: rtfllovC'd

Flo. 2JI

Fi". 230. N'ut o r I'I'S/'Ulo Mr1JtU'('Q (S;aun .}, budl in marl-like !\(Iii in hillside at Cuper-tino, SOlnta iIJ":jJ C.(tunty. ..liforma.. "Ole initiAl w,perooriUlTI .. ' top of nest, plA tered t fllr.ltu:e Kalltl)' al upptr ri}:t1 1. and fragile chal"01ctt1' of 1taper

Fif;'. 231. Corr,hI; of \~~ irrqular ~t or I'nptda ,,:uyltfrHtKo (Saujf.). built in stony rroond on hill .. ide AI .. 1.a..IIf(lru l'"nio,cnity, u.lifoml.3.

... ~ ~~ .... :Z~

Pu.n XL\I

F,I..:,. lJ7

Fig. 2M). OQ5l"--UP of fhlJ.tC:'r from the \\'~:ll of the: 4!normQu!I ne- t or Vl'StIIla 'l'lll!}r1rU (i.Jnl1.) •• hown ill Flb_ 2.55. PI.3,tc 1 ... 1 V, Sho" , r"'pial ~uudurr ur,· udUjJrl,.r pg.~f'". Lbuu.gh "itlt n IIUJ~ mort:" ("Qntra",llng p.1l1trTt th." h. ww.l

Fig. 237 L1Oa~-up "r ~pu from the \\'311 of the !lOt 1)( fJutwff Iw1rh,.,.m (&uu .), .hlO"'" tn Fi~. 230. Note. mUllCmu nlinule SpiCd. 111 lk1.PU.

Fig 218.. Ob:.e--UP of flJptr ire.n lhl!' \,~Il Q(.:J. ne tor f),,J,trlHl1 ~J~tJlJ Irm(' .. IQf~ (Lmn.) NClle C''V1I'Il tr.xwrl!' :tint! hnrbnntal ('llUt~ Ilf the: o"npOflj~Dt lolriptl

fUr· l.lc) P·'Jlum Qr l1e~t ~'all.nd lopmo t C'1'1Il1h ()I tt'lr PC' 101 "'ut'''ll odga,u (Linn) ~Clf" {lOlJ!"lhl,- IIi r),l;l~ ~n rl1f \\,.11. ami n nhU in \\hi(.'h edl~ JJ1 WAIlI!iJUI BJDI!

"0

G • l! ~ . =

~ .; .. z oj

;: -Ii ~

~ , = v. .. ~

§ $ "' '" 0

,;; i.i:

t ~ ]

;

t •

:;: ., .. iZ

Fig 244.

Fig 245.

I'L..\U XU."

Fl(o. ,H4

lnn~r .uri;acc uf lower part 01 \\'ollll)£ neq of ,'oplIlu "lUyl:,uw' r:1 p~.)~ J. ~UIC opo~l f;t\'. f'tIgn of p.lJ~:r" lItH .... du~ to u:;mng ,Iown of .... -.111 on l1I"il!~ [nncor iUrrOlec of c('lIullr moth ur r:-1lC'r ft,rminx I,,\\('r l:.an nf ~U Df kiianl nut uf Vot./a toW"I,rrs (Linn ~, IloIl0\\U in l'-iX'. ZSS upo!-Cll CfiR" of I,",per arc ,we 10 learln, down of .... 1111 uu In~idc

Fig:. 246.

l'ig. 247.

•

P'f. 147

Upper ,urface or comh from I"r~~ nC1l tlf l '('$~tlln t'tdg(lriS (Linn.) . {rom SJim.1 Cruz., Califomi;a. Note pll1:tr-like fonn IIT"I IUUTWlgaH1tJ" texture tiC Ihe. nUnlC:fOUJ ,,"'pen~"iJ. UP{)(:f uria.;;t or .opmo..:: (mllo uml K'Cuon Ihruu;;:h ClIp (lr lIIUUI't: nt'~l of Du"thut~lrullJ IJklruln.I" [LlI1I1.). rou" n '" FiR' .. '!zO. Satt" cellubr dliUllCICr or cap. oil"' Iht Ihin lamin:&.tl! "u pcruorla. em h>fJmu t coni',

"cpt. \ Ill.:" .. 1

Fig. 248.. Upper ~uriace of 1:L'fit 141111 IItxl In l.,~t ellull fruUI not o' , ','.s/w/ll t/'dplr.~N CSauy.). ihU\\n 111 Fig.!lU ~ote tllt'ei'liUH ar paJK'r -tnrh in rtll bJI,("). dt[· (C1't"tlce. from their \t-ry irK'l"'ptlOo. bttwn:n .. lIl.i11l! cdl in which wfJr~t'r • .1r~ ftUtd aud a.rl:C cell, in .... 1111.'1. "U","" arc- rl'ar~l. atM), hoi-ling ut LJ.r.:t"f cDmh ,Iue tu imlJO JI.ihly of .. dctillK' lan;f: cdl lo u&OLl"J:ill Ilf cl.mb ('QlIIl'm«=,1 of _Ill 111 uti, kut! 1II4i1u.J:inil1I: pailI'm

Fi, 249. ~Jlf~f -urian:. ~Qlld cOmh inm1 IIbt <'I( Va/.,/tt Sllll'ul",Q \.s'lU~ .). flDwn In Flit. lJO. Note rini; uf lI«omL:.ry ,U.,pt1liUr1;.l

1.1

J .... '" LJI

FI 251

Fig. 250. I~ace .. :irw (If ccmb in w''o-C'r 111f1 of lan:c 11~1 nf r·C'.It.ID P"u.\'f·I(J"~f" (Saw. ). BDdy o( C'ocnh cofllpollef\ of ~11l;lll «II. for runDI: \\urkert. lube 01 b~ (CUI (Clr qLlU11-rt'nillg

FiIr. 25 1. Face ,-iC'w o( cOfT1b~ (rueu lt,mer ~n. uf Ith-t of "'3Pfll!l lw/,hrta (S.:t.u ). 6hC' ....... In FiK. 2JO. f'otc .tn"IflK di!rc:I~IIt'C In i.r:II. 1iC;' ..... «1l sm .. n ",('ih u..coJ (or r(2tmf .... orlc:c:1'" .1lI1 I r);'c t"dl tllr 'Iueen rr.:t.,rin . .'J'Jlc at ) edit of .lmonu",1 lu r~ .ilof'll; IUIji!' 01 OJUt..art bn.,,«n t.,.o I no DI I;C'U

DtpulClrAI <":1:..-'. \.'fII. .. \ III. No I Pl.-\fII; LI

fICa. 252

I'f';' 15.1

Comb [roln midtllc of l'K'~t or 1.;afJ::1= aCli\(' colon,- nf l'fI/' lIla ~.SylVGII;('(I (Sau ) Note: regulu loninlt' of hruod 01 \"ilriou afle Ohler crotob rft"tI ",",me nt,,",:1 th.u frnm wh1ch comb in Fig 2.51 \L'n t~\tirlj Nrne 11IC'tnlc in ,lrIJpnnlof1 of brooo.l Mil o r their ~ J:Tou(I

U \

f ,Q 254

"'11.1 155

INDEX Abandonmt'Dt: of combs for brood .. rear­

ing. IZS. 141. 142. 144. 145. 148. 156, 172. 173: 01 n ..... 128. 129, 133

Ahdomon : color pattern 01. I 7.l-1 76 : dio­colontion of. in queens, 161: manory musc le (= propodeal ele .... lo<) 01. 85: of fmr.tJ~ 13. JiI, JS, 72--79; mUICU­!;,rure. 65. 66. 74-78 ; of male. 81h'14; 01 1Mn. 162. 163; pedicel 01. 62, . 65. 66. n . 73. 74 ; 01 pupa. 165, 166: spJ­",el o~ 76. 71. 78 ; .1« 0[. 65. n -78, 81: ter.ltoIOJric. 176 : te .... oi, 66, n-78; 01 Vut., 89 ; o i V'srffoa" 5; 01 11"/'1110 . 92

udar muscle, mandibular, 21. Z2 Abiu nmCq/fl r . tOol Abnorm.allUea: in ~st. lei Irrccular ities ;

in morJ>ioolOlP' . Itt T eratolov AecC".SKlry p~oxa l p te, 40 Acccuol')' IClerite : of the fore coxa, olO ;

oi the mandible, 22 Acet:J.hulum ; of the er.uuum. 21 ; or the

nlA:nd.ibIC', 21 A trO£"IOi 1 buuon, 27 Adductor : AlIterior. of the hbiulll. 24, 28 ;

of me cardo, JO: pnlOOotal. of the. fore coxa, 46 ; aecoudary. of the {ore wnp

46; pleural d("pruscr of t.rochan ttr" and primary, o f 1he. fOJ"rI¢. 45; of the mandit)Ie'. 2 1. 22 ; mt"Sofureal deprasor of U'oc.holntcr limd, of the middle Ie ... , 57 ; dCllrcs50r of U'Odartter awl or e hind Ie ••

Alder, 101 Aliment.ry canal. 79 Amr'/o~sis qwinque/oliu t 104 Anal lob<. 67. as A n;U veins. 66. 67. 69

iK, 104 Ant, .;6. 38. 107. 114 Anteromol ridge, 73. 7-1. 75. 76, 71 Antenna : a rticular cup 01. 19. 20; b:!$<: 01.

16; bulb or. 17: cleaning 01, 71 ; f!l.gc1 -lum n f. U, 88. 91 . 93. 95 : J!1'1"".1 lca­tura of, 19. 20; in DoIidmvlpultJ, 9S ; in mal ... 11 . 13, 88, 91. 93. 95 ; 1.,.",,1 . 164: r "scltlOI, 19,20: pupal, 165. 166 ; pedi~1 01. 80; position of. during hiber­nat ion, 101: O«I t.>e of. 19. 80. 101; in Vuta.~; in V~.s/Ji"a#. 85; fn V • .", .. Ja. 91. 9~. 9J

Anlenna-cleanu. 71 Anttonal f ramen. IIH Anlenoal rid,e, 16

"tennal Icleritel. 14. 16. 17. 19 n":",,al soctee •• 16. 17, 19

Antermal lulcus. 17 AntetmJ:t Autute" 16, Ii Anttonif.,.. 19 Anterior proce!." : of mesorurcal ann. 47 ;­

I meooluoul rid\:e, 54. 57; 01 the ""'!IOJIO'tphragma, SO. 55: 01 the pm­.tcrnum, 38, 40

Anm my;.a. 107 A US: 01 fenuJc. 78 : I malt, 81 A phids. IQ.I Apica l """I ... r the propodeum. 61, 62. 8S rlJtis. 60; IU 01<. Honeybee Apod""," : .nterior moopleuro.!. 35. 37, SJ.

55; antt'r1nr IMtOlple1u'3.1. lS, 6J ; margi· nal met. pleural, 61. 62. 63; margin;U. or tbe mcbuternum, 35 ; marginal. of the prOllleuron, 39. 42. 44. 45: of the pm;" 84. 89, 93. 96: .econd .uuopitunl, 52. >I. - : s<COtld .. otapIe"ral. 61. 62, 63. 64. 65: .. crl ~l. 73. 7-1. 76. 77. 7S ; ttrpt. 73. 74. 75

".4,orhy" trlgltollTJl,." I AfIOpIoY"is: mCSOl'trnal. 35, 37, 47, SJ. S~.

505, 56, 57. 58 : mctaslernal , JS, 36. ss. 61,62. 64, 6S ; pr ttt ' I, J8, 40. 41.45. 47: <Iornill, 81

Andmids, l06 Arlie, J9 Arthr opods. mUc.ltalll!OUI, 106 AnicuL,\ f sc lcritc:t, Sri xillary tcleril6 Milid, 107 Asi/Uf 'r)'lhro~iMS. 107 A I. nta butlcrRr, mI, 106 rlItO('" ty"lhia, 108 .E111acu.s JCllllo~lai, 108 Austr1lian bottle tre:e, 1 Autotropha1laxis. 117 Axilla. 48. 49. 50. 5S ; I" oh. Scootell", _I> Axillary cords, 49. 59 Alti1lary en.i.ion, (il A~m.ry Kleri, ... 48, 4? ;0. 51, S2, 56.57.

59. 63. 64. 66. 70: art Icular sc:Jerite. 3

&ceJraris dONgtGl.tii, 1().t BaJ' trap>. IQ.I. 108

1257 J t 257

T 8 ~ ' O RT B .UU:' ltt CA N VESP . \ AS)'S

Bald or bllcl·faeed hmnd. ,I< D.r",no-t:tlill/a 'If<<JcuiatlJ

8;aldu l. W . • 140. 169 Ilol ...... 41. 4Z Hark ;J.J building m:Lt tti.1 1. J21 Biln : of propodw .piracle, G6; J'lharyu­

,eal. J2 Ba .. la r<, >1. 55, S6, 60, 62 Ba:Wll:ar mmc1e . of the rn . thor . 55.

56. 62. tAl ; Ot the m btbo""" 62, 63 B_ 1 luI>< 01 Ihe goood ipel: an Doli..-ho­

'('ut"la, 96 ; in V t'spo, 90 j in V u/IUJa, sz. 93

0 ... 1 I.lb. of I ),. pen; , 84 Ba,al H : In, (/) Bmro 11( ''!I, 132. 154 11«. Zl, 38, 52.78,79, 11 4 ; 'f' <Ih. Atis;

HQlttyhoe ",· l1y - bombilid) , \oi, 112

ntt'b iv~. 103 B ... plant. • li£omia, 104 U .. tk . 15. 10 1. 106, 107 s.,'IU.<rI, JW<Jlh. ,83, 84, 85. 87, 88,

Il l, 170, 175 Bt-rle ~ A", 40 Uirch, 104, 121 Bird, 1 tmr) . lOi Bi >Cl,ofl, II. 87. 1216 UI"" IIi." 1117. 11(1, Il l. 11.1 S"mbil;d. 1lI7, 112 Bvul.nil<. I I.. 81, sz. 8J Brace : Iljf'tanl,l tal . 59. aternul, 7 • 74, 76,

ttl'RaJ. 73, i4. 7S: t nnl\'c~e IJfopleun.J, J" . +I ~5

Brll.-l')'flljltln (= St~rttllia) . 104 S <y, J. ",07 Br~'IlIHn (rt: tur.ttiou). 13, 7.l j n. 101 llri~g< hypo"om. s. 18 ; m_{ural, 47,

- • ;, /; (11 ht: nl(:S(),11un.al :l.papb}"!lJ, "1, ~ • l)01otgf'naI, 14, 15, 18; tc.ntcrria1. 16. 1'), !, JJ, J.I

Hrlll,' . W. L.. 12l liT ,1111 )', . W., 10:, 106, 107, 108, 11 3.

11..1, IZI Bf"OIJd, JU £ J: .. ; l..ar\' jI ; Pupa Brood p' od\l<li u. 0)8. I-IS, 149. 158, 160,

161, 170. 171 Arowil hamel (= r ,,Ita (mbro) I\u 'u l Ciu;ty, 31 I:,,<khur I, A. .• IJI Uuc::k .... beat. ",,, 1((, IC>4 Iluih!m.g OlCti"itit.5, '!1. 70, 98, 120-160 n ' lild ilJll tml1lrl.il. 121, 12:8; .u 0.00 H.ark;

Cc:xpon: Po1l)(;r: Fiber Bulb: t)t the f\lCJUlII pedicel, 11; ninK',79 Bull t h isll~. J().4

Burns. Ed .... rd } .. 107 BUllaRy. red aW2ob. 108 Button: ac r 105 I, 'Z1 : hYPoPharynceal,

25,28 Buy,,,,". R. du, 86. 88, 117. 157, 161. 165.

Hit>

Cabbage wornl. 108 Cacl .... prickly p<ar, 104 California bee pl. ut, 104 CaJlith.,.. t'7,Moc<phaJ., 1Il7 ConnibaUtm, 96, l~, 117-120 Cepito"IUI, J5 Cardina. muilhuy, Ilf Cardo Card[J14)oltipiu l hinge. 23. 29, JO Cru-do, 23, 29 ; adductor'S of, JO ; aanio­

c:artHnal articulation, 29 : productOr of, 29

ear. of brood, 70, 98, 103. 107, 1I2. 115-H7. 119

Cari a:a : ~ntt:rior maostenUI, 37. SJ: an ~ tenor (lroaD' I. 3«); mcwtemal, 61: secondoI')" pronotal, 88. 90, 9-, 01 Iho ti~. 30 j lenia ry, of the pronotum. ;

tno mf"t:ano~ l , · 9 ; tr.ulJscutcllar, 49; tnn<l\'er)l!. of the pronotum~ 11. 91

r..arpenl.r. . fl. 68 Ca ..... IJ ; brood cell. for dllftrCTlI, 139-

141, 167; dilf<,..." i' llnn 01, 116, 139. 165. 11Ii-l i O, liS, 176 : Irnrth 01 immalure . Ul/:C_ 111, 165, 166' rmrphQI "'Y of, IJ­&4. 87-89; origin 01 workerl, 139; rea­ti," I' roporlions ,,[, 1.?9, 159, 160, 167. 168: tn,ion of ~mtlltfl!C'e rtf, 1 170

CalcrpitJan. WI!. II!, 116, 118 Co',,/o.ecvs, 60 Caudal papillae. 16.3 c:.uoidin, 4 I

II. ; additlon of, c:omht. IJ I, 1J7. 138. 1:Jl. 140. 141, 142. 15;, 156 : cuulq,; do ... , 0 olt!, 125, 1+1, 148, 149; initial. 01 «Ia,b 130, IJ I, 137, 142, 171 ; method ot tnl.rairuf, 131, 133, 137, 138. 171 ; pooilion I brood in. 142. IfJO, 161, 162, 164, 166 : renovatiou 01. 21. 1I8, 119, 166, 167. 170, 171 . 172; .... pc ai, 130, 131, 1l7, 138, 140. 142, I~J ; ,IJ"lIC'ure I. 1'0. IJI. 1" 7.138, 156 ; type. 01, 139,

140. 141 . 159, 167: wing. 6741, 89, 91. 9J Cdhd.3r lnlctUr"I! in t \\ II .. 147, 148,

152 Cellul c, IQJ CctnCllt. ali\-ary, 1 IS C""'GMrl'O "30"1"0;]" 105 ..en tj~d'. 106

Ct'IIh31ic sWttuklacum.. JI-J4

(258 1

' . ' E:l

Cuvial Jig.am~nt : 1a.te:nll~ 38: vm1r.aJ. 38-40

Ct.rvical (= n«k mmtbrane, 13, .lS, 42 , ial sclc.rites. n 'ico-proplenron," 40

CluJeid, 106, 107. I CI,.,nbulin. J. . 13. 109 Chapm:1n, 176 ChirOnQmids, 109 OU'Y anthcmulll, 108

il ,.. 100lh oJlf I1'''-T, 104 Cit~rium. J2 C",ado, 107 Ci<kr, 105

imbidd, 50 Cirsillm laflCtolalli,ift, 100l a~<1.nlng: of antmn&. 7J ; of body. 25, i O,

71; o{ """. 116. 11 8, 119. 1 4~, 1;0, 170, 171, 172

I .. cm,r 1 OYJ'CIII dilalor of the mouth, , " J4 Clypcus : 01 Dolich.,!ts~.IQ, 95; o{ the

lo:mal<, 14, 15, 16. 17. 19, 22, 2.1, 29, 30, JJ. 80 ; of the larvo, 163, 1601; 01 lhe lIlalr.-.. : o( r ""/10 . 88 j of V Is,-imu. 85 ; of f'ufl'k1. 90, 92

Co<roth.'llU, 60 Uxrxltl. 118 : fragm flO uJCd in ~pe:r

makin~, 120, 167 ; I ,iliDn of l'Upo. and knc:r:al arl1.11t in. ]66: 5rrinnin~ of I 1M; t.J;m iug of. in comb., lil- ti l

Coltonan, G. A, 105 COlffiJ)lCnI , 68: 1(( also Hcrtle

loui.· : .!I;ondoM'<I1t of n"" 128, 129. 133 : 12l1nibolism ' '' . 98, 108, 117-120; <l.tlin.ol, , 97. 98. 116. 117, 118, 119, 128, IJ7. HI, 142, 168. 172, 17J; d\l 'O­,ion of, 94, 97, 98, 99, 118, 1\9, 140, 141, 1"1, 1100. 167, 168. 169, 17U : ioundina 01, 98, 10.?, 130-1 . 100, 170 : ovU\\ inrer­i"" ~I, 99. 118, 140, 14 1. 159, 160, 170, IiI : q=o1 .... 100, 161 : . i", 01, 15S-160

Color; cute. :tnd Sp. diffu(.nces in. 79, 80, 173-176; clUa>luralinn . old quw • 161 : of the egR. 160 ; of the lamo. 164; o[ papa, 12.1, 114, 1 : of l (IU l ~ . 1G6; of renaal adult . 166 cW·· ; ;tb;mdanmrnt of. for bmod~rC'U~ ing. 125, 1~1. 142. 1#, 14S, I~ , l4" . 156. In. 17J; inlildi", of, 122, 124. IZ". 130. Ill. 133. 14l , 14J. 148, 151 : c<11 Iypes in. 139-141 , 159, 167 : In Doti<h", "tf'td, 98; <:1r1y >1lI11t"S uf, I . 131. IJ7. 142, 171 : <l1ll1rgement of, 137- 143, ISO, 151, 171, 17!: r.te 01. when old. 143-1 SO i

irrqular ,-truC1 urc in, 140, 1-47, l....s, IS". 156. 157. 158; ""per in. 12;1, 114, 125 : shape 01, 138, 142, 143, 155, l56. 157, 158 ; Jlructu", 0 1, l ZJ, 124, 1:16, 140, 148, 157 : in Vat""', 94, 136. eight of. 141. 142, 14J. 144, 145. 146. 1;0. 17J ; .oniog o f brood in, 171-17.1

Ccmpolilet. 104 ComoouDd qt. Sit y • Com.tock, John H enry, JI, 67.68 Conutodc~Nec::dhaJn "cin and ttl) nomt't1-

cI.tu .... 67, 68 CO"iVHI mocllitUum, 104 ConJunctiva, i Z, 7..1. 78 Corl<, IZI

mflowc:r. 103 "Corn .nake,'· 106

nronal .utnrc: of adull I i : of tno rn, 164 rr , Erne" N., 111, 116. 134 tal v<in or ceO, 66. 67-$, 85, 89

Co,o"",",'", 104 Couper, William, 11 7. 119, 127, 130, IJI ,

133 Coxa ; acceuory tc.lt:rite or fon. 40: ar1ic­

ulation of, 40. 045, 47, 52. 61, (H; fort'. 37. 40. 45. 46. 47, 70. ; bind. 60. 61,

, 65, 66. 70; middle. 53. 55. 61, 70; moo .. nf. 45, 46. ' 7. 64. 65; 01 Pill ... 165

CoXI ... :uhalu muscle o · the mt::t1thnriU.. 64

Crampton. G. ,40 Cnme ni .. , 107, 1I6, II Cruniol ar"",'. 16-18 Cr.'"w.....tyl .. , 15 Cranial flexors vi tlit=: pal'2glouu, ~3 ,r.mial uture-~ . 14-19

Crnnio-arditQI ardcul.:ation. 29 C rv'u~DtU, I ().4 Crnu ' 01 th fin t lurtal (= pro-Iurca l) .rot.. 9. 41, 01.1. 46, 47 : "' •• r;d, 01 tbe

m P<>I'1>b....",a. 51, 6l ; ><utell ... 49 rop. 115 ~y.iM, 68. 6'1. 89. 91. 93, ?S

ubital run or c<11. 67. 108. 6'1, 85 Can; , John. II I

utic:lll4a (::: cuticlr), li S; pupal, IG6 ; 'f" <ltlD In1e«umcnt tworm, 108. 114

C)'ftnmym ",lIal/urm. 107 Cypr .... 101

Dahlia , 104, l ZI · 'Damlmu~keJ." 46 Dm.,-", curQ/in". t()8 lltllJrul Nfl'lll . 1M

2591

X tlRTIl .l\MEIUCAS VESPINE WASPS

0 .. " . William T" 1()4, lOS, 106, 107,111 . 121, 128. 154

Dd.me and oll.n ... 13, III . 112, 158. 167; su allo Sting

Ooo'cn. S. B.. 107, III Ikpr....,.. (mu",I.) · .",un;,!, o f tM an­

' ''''' ... I ~, 20 : 5 .... ' . I "'" hea.d, ~J : inditt=el nd prindp:d. or the fotc: winrt

...a i , :;: I -- , SS, 62: hllxr pronot:l. l, u i the scutum, 44; IntcmaJ. of the •• Blenoa, 19, 20; m~furca.l, of the trochanter and adductur of the middl 1q. .57 ; Dl~· iurcal, I the , belomen. 65 ; of .h. labial palpus, 5. 26 ; 01 the """,ilI.ry paJpus, .n ; of the mtl1notum, 6.3, 64; of the t gula., : oi t. trochanter 3J1d ad­d.,""or of .he hiad!<g. ; oi the tro­chanter am pri ry addUc.t r of the fl1rei'll . 4S : OU'U proootal, of .he 5CUJ\l:Il, 4·J; Il le.u r ill, o i the abdomen. 6S: plru ral. of the lCutel lum, 56i &rtOnd, o( the head. 4l

D ... romyi4 ,w,bri"" , 101 Dllalr r : o f an abdominal .piracle., 66, 77 ;

dyp<>1. oI.h. meuth. lJ, 34 ; dor>a1, o f tM- pcd:t nhr ~rynz, 33, fron 1. of the ;mter-ior pha ro , JJ; pbaryR e.a i, o f th~ mout h. J4; pcscc rio r, of the an ... trrior ()barrnx, J4 : IJi the al iva r ium, 28; ntrOll , of the posttriar ,mar,. IU, 34

Di/otorr.r bur oUs, JJ n italern ~ibo1ri~ JJ Di/iJICi~U ,Jr:Jr)'"g is fr(lff/a/ltl, JJ f) i/a/ort's l'~I,hrY'flJta IUt 3J V im 'lIQ(l;iil,60 Dit,lop,.ra dlploptorw.) , as Di ptCGJ (dipuruus1 etc.), ~ 107 i tU {llso

i lia Di~J Td.JjOIl £ the body in old '1tl

161 D istrihuhOfT: tit .pedes st ud ied. J.j; cc:b­

logic:d, 11. 12: l~raphiC3l. 86 ; lea­:'t'1oa l, of eutes and coJaniu, , 97. !)S.... I . 116, 117, li S, 119, 128. 137. 141, 14!. 16S-1 70, 172. 173

DoIi,hu;..,.s/lu/a: biology of, 96. fJ7, 98-176 (nwncrou.s refercnces); combs of. liB. 130-133, lJ7- 148; diajmo .ie ehu­••• eriitia 01, 11,95, 96 ; diltnbutiOl1 of ""II t",.. in ..... of. IJ9-14 1: durallon

I coloni", of, Q7, 119, 140, 167, 168, 1"9, 170; etntI'XC'Dce d;.teJ of ff\:.ll iI.Dd i,uttns of. 169. 170 : erratic budding by, 14,. 157. 158. III q ollolOfO' 01, II, 12. 85-96: neJ.t e:ntran<::e o f. 152-1 : 1lUf$ of, 96, 91, 120-160, f'o.,uiWt , ncatil1l ;Jhc.

I I, I l. 125, 126, 127, I ,129. 130, I ; ....... 11 o~ 96. IJO-IJJ. 150-152; not I rue rnw, IZ; paper or, 123, 124, 127. 128 ; r lathe propt1T1io." of the ctJtCJ .. 167, 168; quem _IS oi. 98, 126, IJO­IlJ, ISO, 1.; 1 ; fU pmooria of. 96. 143-H8; <a.'onom of, I I. 85--9-1

D D/{clitn'Utu/11 ad"'ttnrao, yae, grctko, 87, 97

Dolic~_"whJ om","" , -oI",y of. 96, trI. 100-176, 'OSStltl : color paUun of. IN: distribution of ..,U .ypes U1 nos. of. I ~ I : eombs 01. 138, 139, 141 , 1>16, 148; du ro tion of c:olonl .. 01, 1\9. 167 ; etnttic. buikJlng by, 148, 158 i foods or, lOS, 106; lungi altacking, 171 ; hibcf'llll­cion of, 100, 10.1; bun,,", behavior of. 106. 117 ; irregular nell of, IS: ; mor­phology 01. 69. 95, ')6, 07 : noslS of, 121, IJO. 13-1, 138, 139. 141. IoU. 146, 158. 159; n ... ~ill: 01. 126, 128, 129, I . nat wa ll of, 1.52 ; paper 01, J2J; qu«n De_II oi , 130; repair of dam. .... "es" by, ISJ. 157; IU'penlOri> of. 143

Doli,hrn.:rspulfJ rnondnta: abandonment oi nes •. 128 ; beaked .... 1S f. I .'. 154 : bi logy of. 96, 100-176. /'<U"'"; ccllu-1M .... ' 0 ne!l of. 147; nmb of, 142; d uration 01 c:oloni of, 169: etncrgencc. datt o f ma.leJ and quemt of, 169 ; malic bu~dina: by. 1-18; rood. and leed­ing o f. 107, 108; hihernation 01, 101, 10.1; huntinc hehavior 01, ]07, itO, III. 11 2. 113 ; neru of. 116. 13tl. 1 . Htl. 146, 1.10. I SS. 159; n ling oi.e 01, 127, 128. 129; p3[)e:r o i. IlJ i queen n~tJ: of, 130, 131, Ill; OUlIpOlIIO!'ia of, 14.1; type , pe­eics 0 I ~<lIIl , 87

Oo/ich(1f'f'sp"to ,Mdio: 97: cdlul.u tap of n<JI o~ 147 ; .rra.ic btUldi"ll by, 148; nb' .ad nu, bUilding of. 130. IJ I. 143; Ilest e.ntrance: of, 1·3 ; nest .. .,. 11, of. 1S2; ",building- f d trO)'ed nest b,. I "'l*>r of datnaj<d De t by. ISJ

DoUchtlt'Vsfnt/a nDfUVgiraf 91 Dol"hot'e¥~t(J .ty/wstris. 154 Donnanc:y. 98-103; I ngth of em.nl pe-

riod. 102j ob1iaate type: o~ , 100 ; Slt aJSQ fI ibr:malloD

Dron. ny, 107 Duncan, C. D .. 121 Ow.ean, F. larlin. 100, I II, liZ. 117, 169 Ourori ,n of ooIueln. 9-1. 97, 98. 99. 118,

119, 140. 14 1, 159. 160. 167. 168, 16? 170, 17 1

Duration of larn1 lifo with ut rood, 111

1260 J

INDEX 261

E4c1ES i,nrrrialis, 1 E , . 13, 160, 170, 171 ; laid by ,,"een, 98.

131, IJ8, 160, 161. 168. 170, 171; laid by .... -ornrs,. l SS. 16 1; mala from unfuu· liw l. 100. 158. 161, 168, 169 ; ili ... o f, in ttll .. 160, 161 ; rUng of. 171-li3

Elttaror : antuior notal , of till! pro· pleuron .nd h"'<~ 44. 47; diree. and ttCOfIduy, of the , re wiD¥, 56 ; furcal, of .h. head. 42, 43, 44; indlr-ert and vr incipa ], of the fore wing. 35 i talc-rat DOtal, ;lI)d rotator of the proptetlton, 44; pl.ur ... . o{ the h<:ul. 42, 43, l)ro!Irxleal, ul ,Ire .bck>ll,,:n ( = • . '<lemMy mU5cl. 01 ,1\ abrlo .... n) , 85

E lcn, lOS Ema.r.-in.iIrion 01 the lCutum, law aI. 4 ,

I, 59 £mug.""" of adult .. 120. 166--l iO 1l",,1I0 y ""'. 108 Enetnitt oi wasp , 128, 129 Ef"I;'lI ivy. 10-EnIa'1lanu >t: ui ""lis. 131. I.JJ. 137. 1.38.

IiI; f (·"",bo. 137-143, ISO. 1,1, 171, 172; 01 1. 21,9-1. 9rS, 97. 98, 128. 132, 134, 143, 147, 148. I~, 1.10, 1,-1

· nt ow innie ane., 71 Enladortal (V -.haped) ridg. of tho 101-

lum, 49. 58 Entra.ou: to tJx n~~t burrvw, 136, 137.

152 ; I" the ~I proper, ~. '1l, 152.-1 Entranc screen in V t.tfulo nats, 1-48 · ,oIopes 01 I " 122. 130, 131. 1.12. IlJ,

1.10. 151, 1, 2. ISJ. 154: Sf, aim N ... wall

E"iaanial suture, 14 "EpUl><:ral ann," 39. 40 lipi""""". 39. 40, SO, Sol, sa. 60 : st, olso

MeIC'pimcRlI! Epiph ... ynl .... I",r, 22, 31 Eplpharyn . 19, 22. 24, 32 Episternum, 39, 40; IU also Meupil1u.

Dum Epi to lIlal rid ~, I " Epiuomat 'tlm~. 14, IS. 19 IZri(l!JVIUHH lo.rricilialum~ 104 Eriot/olll"" ,,~b,Urtlllm. 1().J Ensto/is ta,.rDnun., J07 Erinalis I,.IIZ. 107, 113 • ophagu., JZ, >16. Ill, 114

Eui ... E. 0 .. lOS Eueolyp'U>. 9\1. 101, 105 F ut()Jo(':!.n honx. l (= Vu/'6 P'UHo. q,v.) 1':x<ovation, 21. 120, 128, lJ4-.U7. 15S E.xcrrtiou, J

Exten-.ot: intcnll"mrat. 75. 76, 77 : i ntc:r~ l~rvl. 73. 74, 7S, 76, 7i ; of the n.nxiUa, 30. 'ClTD-"mlOl. 75, 76, n . 71!

E.xt~n y muscle of the abdotnm (= pr podell ,levalor oi abdWtlcn). 85

ExUtlatcs, gt..nrluhr, 105. 10? F-x",,.., 162. 164 Ey< : eampound, 13-15, 16. 17. 21 , lUI;

lan~ 164; pupal. J6 , 166; HI Pu/'C, 86.87,88: in. V~s(tWI(J. 90, 9Z

Fabre. J. Henri, 107, 114 Fa. bodi .. , 79 F""I pelle " 1-48, 173;.It also Mc<OIrium Fcedinr, 27, JJ, 103-109. 167; ulth. IM-

n • . 70. 98, 112, 114-117, 160; of .he mal.,., liB; of the qu«m, 10_

F.mal •.• " Que ... ; Worlcer Femur, I~, 70, 71 , 102, 165 Ferris.. G. F .. 13 Fiber. vq:etabl., liB, 12D-122, 125, 13-4 Fi<ld moo ... 106 Pig,·ta.35 Fil!Wor, . , 102. IIi "F"mi hinc" of l'I~l parts, ISO, 151 i by

Dolichm"'l'ulo. 96, 11]. 154; by Vu,",I., 04, 156

Flaccllum, 13 , 80, 88. 91, 93, 9S, 101, 165 . (xor : of the 111'1 • • , 30, 3 1 ; of the ,10<­

De, 'Zl : of the i:u::jnia, JO, 31: of the ligula, 24, D, 28 : of ,h. paragloua<. Z5, 27, 28; of tM Olipcs, 2l!. 29, 30 ; "f tM 'jog., 56, ;7, 63. 64

Fli.., 106. 107, 110, Il l. 11 2. ll3, 1.?7 , igh., IJ, 36, 49, 55. 56, 58. 62. 63, 64, 66,70

F1OWUs. 98. 104. 169 FOC'"ictlillln t'ul!1d1't~ 104 1"01 m. Justu. \\ . ....,.,. 122 Fooci chann<.1. 20, 22, D . 31 Fo"d_ kindt 0( '

Appld, I Apri.".., 1()4 Bacon, 106 Berf, 105. lOG llIood, 106, 114 Can.doupe, 104 Carr",. 104 Qiclc<n. 106 Collontail rabbi" 106 Fish, 106

1'lIpe, 1()4 Grapclrui~ 1()4

reeny;r,ge plum •• 1001 Ham, lOS Lamb. lOS

[261 1

262 NORTu A)d£Rl \ N V P I NE W 5

F • klrub nl (C •• tinued) U...,r. 106 I.inrd, 106 ),l.,.tJ, 105. 106. 115

<ach." I~ P .... , 105 P ickl ••• I()'I PI • I().I Pru_, I().I R2b1Jj~ 100 RJi ., 1011 _ 1"",0, 100

n.akea (raruclllWce. watcrJnilice, com ' nak.,. 106

Sparrow, 106 Sw«", I().I, 1l4. liS, 167 Tomo,o, 1M \ 'I.tumclon, 104 \ lener-w n;ts. 1

Foods. I ' . 21, 25, ?1, 10J-109 ; adol ... , IlJ- Jl5 : cannibali 01. • 108. lIi-120; Olnd stc oricin&, 139: of lu\:' e, 9 11 5-117 ; ~r "'ri"" of. US. 167 ; o r tClle:rai aduJ", 167 i undilC.tted rem· WU1I of, 116, 164 ; , erlebrate, 105. 106, 114

Fono",ng behavior, 10.!, 10;, Il'). 153. 154 Foramen magnum, H . 15. 17. . • . . 4J F rlx., \\liUia1l1 T .. ti8 For CClG, 3i . 47. 70 ; ace ry Klcrik

ot ·10 : articula tion of. 45 ; m V tiro. 88 ; muscles of, 39, 42, 45, 46

Fo ... Icg., 36. J8. 7.). 71 Fo \\;Ill . 6o--7u; axilla ry cordJ of. 49j

axllJ.,y . lerit .. 01. SO. 5!, 5.!. 56, 70; Illr~cL arId Jt'COOd\l.fy ckvlt ref the. 56: Ir~1 fold Qi. 66; indir«t Olnd prine;· potl dtl1f"C5 r oi the, 47. 54, 5~ . 58, 62: inrfircc.t ,u ld pr incipal eleViitor or, 5S

Ft'l : uf the th ird I , ilIart, -1,52: oral, 15, 16. 18, 29 • .

F o",:. il ill~t', us F Jt1 ndi ll l{ c:tf col ny, 98. 102, 130-134, 100,

1711 Frenal fold. (l6 Fr"'lw! hook (= h.:tmuti), 63, 66. 89, 92 FriJOfI. T , H" 1(1) FrOM. 16, 17 . .13 Front.1I sulure : of tl.e a.dult~ 14, 16, 17.

80 ; 01 Ill, 1Mva. 1 Fru' .... I~ . 116 Furw, i. 101 , 171 FUI'QI arnu : of the nldO.strru.1I1 apoph­

y.i (== lC<ond fural, JS. 53, 54, SIi. 57. 58 ; 'J! th nlCD,U:rn.l1 .paph,.si" ~5, -J6. ;8, 61. . 65; oi 111. p ..... er I ,-"y-

ili (= fin t fu rca.l arms. = profural a rms), 41, 4J. 45; UelU of lht-. 39, 41. 4.3. 46, 47 : procesJ<' 0 r. 41 . 4.3 ; U'3 .....

\ Cr$C ridge- of • .oil, oIJ; ltouab oi Ike,

41, ~ 43 Fun::c-plwral muscle : mcsolho rQ.cic., aoo

I"''''''nt, 53. ' : of the nldOthorax, 35. 54 ; prothoratic:, ~3. 44

Fyi.., Thcmu W. 107, 128

f'Co"'lst'ilnerrluslltl," oll, « Galwl, A. B . 67

alea. 22. 30. 31. 163 lcal 1'fCI<J1. 3 I

G. 'cr, 35. n , 74, 76, 77 Gosl'rNptiem. 3S Gmt, t4, IS. 17. l l , !2; pupal, 165; o f

Vu,.. 8)' , 88; 01 V,'Ptd., 90 Gmitll chamber, 81 Genitalia, I'n~le, J. 1-1, 81-84 ; In Doij,/to­

Vlsp.,fa l 92. I)S, 96 i in YU/'D* 89, 90 ; ill 1I11/"dQ, 92. 93, 94

G .. "ll1er, l... G., 101. 1a2. IOJ I..; irard, 108 ~nd labial. 115. 1 landul~r exudiltet, I 5. 109

Glandular sccrttions, 161, 162. 167; SI, oIs. all va ; SilJ,:

GI ..... , 26, '0 Gna t, 109 Gnatlml poodt. Z" 71. tl3. 114 Goldenrod, l(M Gouapoph,. .5, 78. i9 t ionocordo. 81. II? Gono,tipH, 81-84 : "outer edgc." 8:2;

.erm inal ,' '_ 01 lb., 82. 89, 90. 93. 96 i " upper ir..nu edge," 82

<berry. 153 ~ A\\fly. 107 Gopher, 132

rO> hoppen, 14, 15. 108, 111 reon bottl. 8,. lQ7

(''';IIJ~lia. l().l roo,'C', rt'k.'Ciian stel"fa.l. 40, 53. , 61

Growth ; ... ul <umb>, 14?, 143, 1;6, 162 ; form of €;oml),. 1551 1511: in DtJlichm stll1a neds, 151. 152: in YeJ ­pula lI~tJt 04, lSI. 152 ; of 1a.r \'~~ 34, J5. 137, 156, 161. 162

Gular ~U1Ur('5 . 1.5 Gum plant, I~ Gy", .. ,..g'lD. 107

Hacmato:<}' lin- io. 41 nair., ~ , '5cttt.c; abo l'l~nt h"in namuli (== Irono>t h""ko). 63, 66, 89, 92

ISOEX .!6.1

H •• th ro. 104 Had : anterior notal clc\'alor or pro­

pleuron and, 44, 47 : of Dof"hu<Ylp"fa, 11 , 95; 5,," llJId _ond dep.....o.-. of. 4J; iuro" emtCr Qf. "'1, 4.3. #; in htbernation. 101: o£ tat'\... . 162. ltil, I ; m"'l'hoio,ry of I.n."", 11. 13. 14-l 4 : mo."hol0l1 of mal •• 80; pI urolelc­," lOr 01. 4:1. 43; of pupa. 165. 166; 01 Va,... 90. 91 ; 01 Vat_la.90. 91, 92. 93

Hear~ SO He1iotro wild. 104 1l,liotroffum Cttf'dSsm'&1H". I H iber",,,,,I., 99. 100, 101 Hibernation, 98-103. 1,1); ,u .f,. Dor·

m:uu:y, OveN'intf'rina H ind coxa. 66, 70: articulation or. 00, 61;

mu!ldes oi. 64. 65 ffi od 1"11, 70, 71; IJIwcl .. Qi. Hind willg, ~. 63, 66-70; anal lobo: 01,

67. SS ; axliluy _d 01, 59: axillar')' (: xcisioQ of, Q7; axillary cduite.t of, -1,63,66. 70; frena l hooks 01, 63, 66 ;

tcrior lobe DE, 85 Hing. p13l .. Iigu~r, 26 "lilll',,,, r""t1t1litfr. ," 32 Homopt~iI, 107 Honty, 10J. 114, 116 Hone) bo •• '0 . .16, SO. 52. 72. 107. Ill, sa .ls. Atis Honeydew, 1114. 1 H H izontal ,.late ~ of the e50Ikrnal

IIoph) ,;', 37, 47. , .1, ~. '7 : of th. lJrU5 lermd apopb.)o .i , .;1

Hornots, 3, 11 , 12, 45, 7. "7. 103, 1I J lIo,".A,., 107 Hou ofiy, 107 Hu<kltberry. I().I Humeral a lii . 89.91 Hwncml scluite of !~ O1et"aru)(um, 51,

59.60 Iiunlin, behavior. ;0. 106, 11G-1I . 117 Hymeoopun., miscdlan('Qu, ~. 2,~ . 31.

34, 35, Ji. • 40. 60, 66, 67. 68, 69, 7 . II? IIJ. 86. 107. I~. 127. 161, 163. 164

Hypoph:1r)'nGul bUllon, 25, 28 HW'IJhar)nlleal pet.o:n. 25, 31 If ....,pharyn<. PJ. 2;, 27, Z8, J 1 Hypo1lO''''', 1 '. 29 Hyp<»lan ... 1 hridg", 18 lIypo", m;u carina, IS, 18 ityJlO'.umol ,"Wro. 15. 16, 18

J ft«tioo or the: m~tanntum. S9; or the J'rvl1(.1tl1m, 35, 36, 31, ~. 4( : of Ihe. iC;U" I<Hum, 49

Infrabuc:.c:aJ chztmbcr,,Ju l1athal pom:.h lnquilinea, 116, 117 ltutinctJ". 156, J60; '" DUO ·annibit.li m;

3r~ of brQOd. Nett building'i Sanita­tion; Dden~ and offe:tUe

Tnu~guman. IJ, 166 : sr:, Glso Cuticu'" i Vestiture

inll:lIi,ence. 100, 101. 116, 127-130 Intennteonal a rea, 11, SO, 9Z. '15 In.enon.ennary tM3o~h; 16, 17. ,\Ill Intu!tIJ"CI.l m~et 58 inte:rltema l ulmk'l r anti runtctor. 7S,

76.77 latutcrrat ute t'I and rCI.r.lctor" 73.

74. 75, 76, 77 Irregutaritid: in nest rtruc:ture, 123, 124.

128, 129, 1010. 147, 1'\8, ISS, 156. 157, 158 : In brood ,,,,,ing. 171, 173

rvy, Engl"h, 105

Jon". rl .. , 25, 63. 107, I~. 114, 115. 11 7. 121. 129, 130, 131, 134. 138, 148, Is:!. 153, 154. l57. 167, 17l, 176

Ja .id,. 107 J rin<, L., 67, 69

Kirby. W illi.",. 115, 116, 117, 11 , lZI Kirmayer. Roben, 17, 23. U, 17, 32 Klu ., ~ . II. E .. 81 Krugh •• ,.. A .• 121 Kuscl,., J. A., 170

bini ~dduct()r •• posterior, 24, 28 Lab;'1 J13lpu,. 2J: deprusor I. 25, 26;

f .he I.rva, 164; of V,s/'U, 88, in VtJ/'iJtu." 85: oi 0,14/0,91

L..abia l promineru:;. 164 IAlii ...... , IQ Labionwrillorr cmnplex, 18. 22-J1 r ..abioffi3_:..:i llary jugum, 24, 29, JO L.abium. H. 2!J, U-29, 31, 109 ; 01 the

larva. 11 5 La raJ. suture, Ii Lahral tractor, 24 Labral ' riafl!lle, 22, 24 Labrum, Ii, 19, 22, 163 Lacinl •. 22. H .10. JI. 16.1 La",n iaI bo.r. 22. J I Lamclli fonn prcuu oi tbe scutellum. 49,

SO Lan«I, ,'ing, 78, 79 Larva. 97 ; Illl lo,rophallaxil of. J I;; bfo­

hoivlor nf, when hUIlII")'. 117; Citonru­l i ~m 01, 98, 108. Jl7-12fl, col~ oi.

164; d.pooillO<l of fr:<u~c by, 156; di'JIO"'I 01 dud. 170. 171 : u"rati of

[ 263 J

• OIlTn AM£RtCAN V . "PINE. \ 'ASrS

Larv;a (CDnlil1lucf): life of, wlth .. 't lood, 117: leeding 0;, ro,98, 112, 11 .... 117, 160, foodl ol 98, II S-1I7 : growth and mctamOTJlho.io 01, , 35, 137, 15(" 101, 162 : length of Ih. d".lopmentai \><rio<! 01, 165, 166 ; mO'1'hnlng), ni 16Z-1M ~ poIitioo oi, in .... 11< , Iii.!. 11>4. 1(06; .anlni of, 171-1 7J

Lo tr611 • . ritr .. Andri, Jo! Le.ch. I,ll Led·hopper>, 107

. : I11hcrJlilnrou!, IJ. 36. 38, SO, 52. 56, 95, 101, lOS, IIJ, 120. 122 : mo<­phDh,1J)' 01, 70, 71: mu.<l.- of. 35, 45, 46, 57, 04, 6 ': pupi l. 165, 166

Lepidopltra, 108 L.ttogrul .. IIm;~<I, 107 Lnla j(>rf'lfOrfU, 108 Le\"'l.tttrS f the. anlC11~ 20 Lewi • . IV. H " 97. 1l 7, 122 Life cycle, IJq, 1W-168 Lipment· furt'tI·plrural. :w; 11:1 rill (:r;TT­

I I, .18; n hc:,racic: (u pleural mlliclc Ind, 53, :: : \ tra l cuvia~ 38, 4 1)

Li 'ul •• 24, 26, 'll. 2l!, 109 LiguJ.ar hinee Jllate. 2b Lil"", 121 Lingua , 16. 'll LinJ.tual plate, 26-2"l LiuJl;;l..(u~ arc!u ... (Linn '. Cllrt), 86 Liff"' " .J.II .. , . IU·I Liu, ChunK Lo. 2t Zi u ,k ' nn:al, of tilt· hllld ing, 67, as;

dor ... 1. Clf tI · \ 111 ih. ; lateral , of the cighth I rglJDl. is; median. ul the ei hth tc=rgum. 81 ; plcural. of the larva. 162- 163 ; POd t'rior, of the pranowl f

,16, 411. 51, 1. 165. 166 : sternal, of Ihe t(mh alnlornH:1:I.1 ~egmt llt. 79; tergaL of the tentli abdominaJ scament. 78, 79

Loc. li l)' Jtudy, 110, I L Lowland. tir, I Lubbock S ir John, IIJ, 168 Lu(ili •• 107

Me ruken. lube-I. 121 MacGilli 'ray, Ai"" 0, 67 MOCN1I' '/10. 87 Malaxalion, Ill. 11 2. 114, 167 M.Ie, 12, 150: beha, or 01. 98, 16') ; can­

nibalism of, I I ; edb in .""hic.b rared. 139, 14t, 161, 168; in Do/ich(N~#.la, 92,95 ; fr<xn unrtrli liz.fll eggs., 100, 1.58. 168, 169; leedin!{ of. IOJ; I,,""rnalion

I, 9!l; lil. C) I. 01. 139, 16>169;

mO'l'hclOttY of, II , 12. so...&!. 85, 91, 93, 95, 165. 166: pu~ 01, 1~S-I68 : . •• 1OtI&I diltributi n 01, 09. 100. U9, IS9, 167 : lerotolosic, 176; in l 'rlp,87, ~ : in Vntw/o, 91, 921 93

Mammal, 128, 129, 153. 1-Mandible : 14, I~, 19, ~22, 24. 101 , 101,

100, Il l ; in Doll</.Ilt'tJ/'lII •• 95; of the Ian· .. , 11 S, 163; of the pupa, 165; we. 01, 105. 120. 122; of V""..de, 85: In V ,.~wI •• 92, 93

Mandibt.l1ai abduc:tor and add.uctor muscl •• , 21, 22

Monuni"" 128 . 1I1p l~, lOS

~r8il.J It)Od~ ; of the mctapleur n. 61, 62, 6J; oi the mt:talternUI1l . 3 ; of the propleuroo. 39, 42, .... , 45

Mouiq, 98 . I .xill., 20, 22. " 2], 28. 29-.31, 109,

122, 163 ~faxmary cardin .. , Ire Corda M :udlJary labu ' of the adult. 27 ; o j the

I...-va. 163 y.,..il lary palp ... 23, 30, 31. 85, 163 .. laxillalobial compla," l

ealybu ... ,I04 foconium, I . 16:, 171, 173 : .u ouo Fe<al pellets

Mffi>1'1 ran, lOS M«01Jt~roid inJeCb, 69 M«Iial Yei" or «U, 67, 68, 69. &9, 91 , 93,

QS lietiio-cubital c.~·vcinJ, 69 : in Doii(ho­

wlpuio. 95: ill VeJ/>C , 89, tn r"tllula, 9 1. 93

Mt:g3.secoptcra, 68 Mentum, 2J M~!ltCpimuan (mclCpimerum), 53, 8S; Sl~

uizo Epim~o ~CIC'pisternum. as; IU olso Episternum Mc.socoxu, su ~{,ddJe coxae Mesofurcal "it, Sot U...,.,owm, 13, 55, 58, 59, 85 M~tw, 51- 54, 55 I""r hragma (= m<OOpoitphroOlOl')

_ t50Jllcural apodcme : 1.11tt:rior, 3';. 7, 53, SS: potl.rior, Sl, 53, 54, ", 59; sccand, 2, 54, 58

Yeiorleun.1 a.rm, 53 "Mesoplcul1ll pit, .ubaJar, 52 M .. ""leunl ridjj., 35, 53 ; ubola r, 51 ,

52 I pleural 'ulo .... . 52, 53

M .... pl.uron, 3$. 37, - I. 52, 53. 55, 56, 60; I"'pal. 165, 166

I NO£X 265

)JC'tOJl'PItnahl1n~ 51 M. _"pitta,,",", 41)..,51. 55, Sr" 62 :

(UJ"(":lJ rdf"Xtof f the, SS (<JOpr.~hrall=. 47. 49, .).1, SS

. f CJoQscuLum, It' Scutnm M ..... I<r I al"'Pbyus, 35. 37, 47, S3, 54,

S5. 56, S7, 58 rcsostcroal lOCi l b , ~ , 5~

MCJO ... mum, 35, 37,51,53. 54. 55,57 Meso,honuc, 14. 34. J5, o18-S9; mulCl .. of,

5+-.19; I.ril .. 01, 4&-5~ . !mmorphocis, .34. 1W-167: .. also

Ontoge y Mculnotnl pro incnce. 60

lctaootal ramUJ. 63, 64 ~i ... """'1 leen-t . 59. 63 M .... IOtum • .IS, 49,51. 5S-6O, 61,62, 1>4;

anterio.r iLlRcctiou of, 59; dq..r r of, 63, : hwnc:nll Idt.rit~ of, SI. 59,60; ulmlatJllnal ,ok:ta 01 til<, sq, 60; ...

oIso Pmucult:llum M ... p<CIw. 1iU, 61 Metapltural ilpodemc. anterior, 5, 63 ~

_rg'inal , 61 , 62, 63; seoond, 61 , 62, 63, , 65 ; ....,.,ru,ry (= '..,OIld), 35. 36

?Ietapl.ural pits, 60. 61, ' ~!cupkural I«Drlduy .utun:, 60, 61 i\ lelapleuron. 60, 61, 62. 64, ~; oj pupa,

165 M.t.pol urn, 60 "Mctute:rnal pit. 61

ldasternulrt , 35, 53, 60. 61, 62, !J4 ; marginal opoW::m. 01, 5

M tatarJ1.LS, 7-1 Metath rax, 14, . 58; musdet of. 62-

66 i 'uites of, 59-6.2 Mete.H, Z. P ., 101, J02

... !toW of lIudy, 5, 4" 42 Middlr (~ae (= me.soco.xac) , 53. SS, 61 ,

70 ; .=",16 of, 57 f.iddl. 1<If, 70, n : adductor o f the, 57

MischocSIlI1.IU, 102 Molting, 162. 166 ~ont.~· pine, l ().l J.forpholOW}·: of the <gi', 160; of 111< I ...

_Ie, II , 1~79 ; oi the 121" • . 162, 163. I &l: of th =1., so...&! : 0 f the pupa, 165, 160 : Ie:rnlologie, 176 ; 01 Ihe " ric ... , 79, 80

MorttJi(y, 101, 170. 171, 172 ; ". aU. .uonal history ; Weaut('r

MOlh. lOS, 109 MOlae, C.old. 106, 13.1 MQUth. 20, 25, 'll, 31, .13, IIJ, 114: ely­

peal dilalor of. 33, 34; piary. ",I dila­l r l "I, J.4

·'Mouth·field-.... "" (== Mun '1c:Jduc:k). 1« Oral fitIJI

Mouthpartl: 01 Doliehcn'ts/'lII •• 95; 01 the lor".. 163, 104 ; mot1>ho~y of, 13, 14. I", 20-31; <>nIOKmY of. 26 ; of Ih. pupa. 165; role of, b. ckonina, 71; rol. in feeding, 21, I().!, 108. 109, Ill, 114; role in Dt":It·m:altin,. 21, 122; of Ytl/'G, 88. SIO ; 01 V"twla, 93

MlIGUJ, 161, 162 MDlbcrriCl, I().! Mulo rat , 104 "l!undfeldsac:k." 61. Oral field 'usea dDtfll·Sti~fJ., 107 ~hlKl ,13, 14; abdominal, 6S-66, 74-78 ;

IJrtmnal. 19, 20: D16othoncic, 54-59, 66: n><tathoroci., 62-{,6 : 01 mouth· p rls, 21 , 21. 25, 2IS, 27-.'11 ; 01 pr tllaru:, 41 • ; 01 nomodaann, 33, 34

fu.llIrd. wild, 106

Neck I1Jt'mbnnc:, Ut CuvicaJ membrane Nettar. 103, I()'! , lOS. 109, 114, 116 • edary. I , 109

Needham. Jam .. G., 67, 68 N ...... t .. (= Pt ...... ). 107

.. t: >hand""", •• t 01. 133, 134; ""ak<d, 132. 154; Im;tding oi. 2 1, 70, 98, 12t)... 160: caun" lSI: combl. <)4, 98. 120-160, pa.uifft, t61. lil-l i J; concealmcllt. 127; of DDI;clf(n,#lpul(J~ , 91 i en­IlI11It:mf.nl. 21, 94. 96, <Y1. 98. 12l!, 132, 134, 1043. 147, 148, 149, ISO, 154; en­trance, H 97, 136. 137, IS2-I54: ir· r<cuIM truttur" in, 123. 124, 128, 129 ; m"<rialJ, 1)8, 120-122 : ItO..." 98, Il6. 1.lG-134, ISO, 151 . 1>1 ; rcbuildil1ll of d"lro)'«I. 130; rcp.air of, 128, 12'1, ISJ. 1>1. 1.7, 158; sanil.don, 116, 118. 119. 149, ISO, 166, 167. 170, 171 , 17Z: ohape, lJO... I33, 154-157: ,it<, I I, 12, 21. <)4. !)6, 08, 116, 121, 125, 12&-30, 131. 133, 157: il" modification, 21, 120, 1.10. 134-137 ; .ire:, 134, IS2, 172. 173: .... pcn$KIP, 94, 96, 130, H I. 141, 1013-148, lSI. 155, lSi : taxonomic n1 .. of. 92, <)4. % ; ty.,.,.. <)4, 96. 125, 126, 133; 01 V<sfo. 1211, 157; oC Vuft.I., <)4. 122, 134. 154, 156 , WIlli . 94, 96, 97. 123, 124. IJO...IJ3, 134, 136. W , 146, 1 ~7, 148, 149, 150-154, ISS, 156; w<i"b~ 141, 142, 143, I~, ISO, 173

Nixon, E. J .. 132, 133 Nom~nc:lat.ure of wiq veUu illld cell~. 67 Notal toture. rne(]ian. 118 Noru .. Ioe Pt1Xl:>', 48, 49. 51. g 56, 59

l 265 1

2 ~OIt.TJ( AUE1U

1\ot;LuJI, ~, s... ~5 .' tum. 55

0 .... 101 . 10; O·Byr"". narold. 108 Occipital arch. 18

ipital condyle, 17. .19.42 Occirital IUture, 14, 21 . J01, in Doljrlto­

,vs,ula, 95 ; in V,sto, ts8 ; iu V('spwo , 90. 92. 93

OcciPUI. 18. 21 . 88 ; lubl of the. 18 OcdlUOr : of the abchtminnl pi cle. n.

7 ; oi the.: 6uI lhorack ,-piracle . .17, ; of the proJlOdw . pirael •• 66: of the 5<oond thor. tic ",ifacie, 59

Oedl,r lriangl • . 17 OcclLtr lub.,cl., SO Ocelii. I I, 14. 16. 17 : 0; V"/>O, 86. 7.

88 : of V .. ".I. ( • ./,). IlO cular ridge, 1 Ct, 21

o ular Id efhes, 17 Ocula r sutures, 16, Ii' f)culo---miJ lar space! , 11 , ]~. 17, 18. 21,

86. 87; in Dalichoo<#w/a, 95. in 17<1/>0, 88 ; in Jlupuiu, 90, 92

Od"""ta, 108 Odon/aulactu, 60 "Orcalr,rh.bWsis:· 115 Offensive behsL\'ior, IU Ddmse and or­

fense O nt rwtny oi mOl-l thparb, 20 ; U t aJ.ro

MctamorphOli.s Op<.: rculum: of the fi rst t ho~ci,c: tpin.c.1e,

-18: of the ~ t,.; ~nnd thoracic 'Ipinclt. -9 Op". tia. 104 Oral r.<ld (= U ",14!.ldsuek). 20, ZJ. 24.

.10 Oral I", ... 1 S. 16. 18. 29. 30 Oral margin, 20 Ora] pecten, 25. ..., 11 Orb it~ AnteriOr aDd pos teriOr, 17 Ormerod. E. l. .. 170 Orthoptcrn, 108 O ... ·cr willlcri o f coloniet, 99, 11~ 140.

141 . I-Q. 160. 170, 171 Ovi{lO!'iti n. IJ , 160, 161

Poipu.: labial. 23. 2- 2!i. 85, . 91 , I maxillary. 23.30. 31.85, 163 : . tinll'. 78

P flIIorftd ~01I'U.H.,.i.r , 108 Pdllorfa (OlUVrJ"dill iJ .. 41

per: c.hancte r ui. 12l-1Z8. 133, 143, 144 : collecting fabcr for . 70. 98. 122. 123 : ..,lor IlDd p""'m in. IZJ 124. ISO; male rial for. 120-122. 148. 149. lSI. 107. 171 ; maki .... f. 21 . 70. li S, l2U.

1 .... 2. 12 : pulp. 94. 122. Iz.l. 1;'3. 1.10. 144. 14 • 151 ; .. rentrlh of, 123, 124. IJ8. ISO: tD.(morn ic diff~mces in. 94. 96. 12l-12S

Paporin .\·or of comb I:u:es, 149. ISO 1'. ",,1_ 216. Z7. 23; cra"i,1 flexors of.

25; lI"""rs 01. Z7. 28 ParallflolSat eclt'nte. 26 P.r3.lossa1 tri.n.le. 216. 28 Parap idal furrows. 48. 35. 85 Parapsid ,48 Parasites: IOCial, amona th~ VtlpiltDt, 12,

87. 98. lOll ; 01 \' .. ~inc """'PI. I , 164 Parietal ocl.ntH, 17

arke r. ]. B .. 163 Parth.n~ .. io. 168, 169 Pecte,, : hypopharyngw. 2;, 31; or • 23.

33. 113 Pedicel : of the Ixlomen, 62, 64. (IS. (,6. n.

73. 74 : oJ tbe 3ntenna, : u~· ais(l u.spc.nsoria

Penis. 81. lW; apod.",. 01. 84. 8<), 91. 96: haJaI lobo of. 84; of Do/i<hot'u".I., 3. 96: 1 VU/>O. 89: of 17",",. I , sir.) . 93

P.tzl.r. SO. -Pcr istomc, 15 Pennian inieCt.s, Ptrom,sttu, 106 P hA r)"ngeal bar •• 32, P ha ryngeal <li l.tor. 01 the mouth. 3,1 Pharyngeal diverticula, 2, 3J; intriosk

muscles 0 r. 34 P h:l. rrl1~tll musc.lc-.;: IIn(tnOr and poste­

riw intri nli:. ,J4 ; ln tcral. 13 I'ha~.1 plale. 32, J.I P harynx. JI-J.I . 1119. 11 3. 11 4: i ntal

dilators of the ilDterior. JJ fJ ltornc;" n OtM, 107 P hfl/{nlll , phl1lllUlata. 60. 61; nl ,,/so

~rc""l\re~hrajllO : ,~ •• opo.tphr' am", I'ropO<leat pr<phrajlma

I'h),I",<n),. 68. 69 PiRl11 ntnlion, dcn.lopment of. in pupa.

166 Pine. \01. 104 Pi",." rudiatd1 104 P its : anterior tetltOrial, 16 ; fint md RC­

ond mttapleural . 60. 61 . 62; meoolurc"J . ; me5O!.teTl I, S4; Intwlunal, ~ I ;

pOllcrior tentarial. 1 ~ 16; prnnoUiI ,17, 44. 88, 91, 93. 95; pros","",I. 40: ull­ajar III pleuraJ, 5 ~

P lanl hain. 121 Ploutc-rlng: o( cntromcc:.J and burrow

with mud, 136: ( I F n t Yt.--:t11 and comh

I Z66 J

I>lDU

I.CH with pape r. 94. 96. 97. 125. 149. ..rior lobo). 36. 52. 88, • ~I , 165. GO; 01 .u.pcruoria. 143 166

PI=. 12; IU .ls. Propleuron ; M .... · Pro""tal p;~ .17 ...... 88. 91. 93. Q5 ptt'Uron ; ~{e IJJarron P ron()lU.IJl. 11, 35, 36, 37, 38, -4-1 .046, 47,

Pieuni arm. 3'). 40 : canna. oilhe. II . 36,88.89, YJ. 95: Pleural depr< ... >r : 0; the ahdomon. 65 : inHectiOllS ~I. 35. 36. Ji. 48. 4 ; of

of the srutcllwn. 56; 1 the ocbanttt D.I.charvl"l.J. 95; of the pupa. 165; aod primary .dduclor of the lo .. ICC. paotcnor lobe of. 36. 48. SZ. 91. \(is. 45 166; of Vu,., . 88; 01 V"""l . 91. 9J

P leural produd.Qr : QI the lore "'''''. 45 ; Prop«tal ... "' .... 85 Qf the bind con, 65 Propo:tUI. 37 j m9QwraJ rttrKlcorl oi.

Plcun. l suture, 39, 40 47, 54, 86 PI.., ... I 'in,<r I rocas. 51.51. 56 P rop\cllral arm. 39. 40. l ...... 45. 016. 47 Pk uro toma. 17, I Propleural <ldae, 9,42 Plnlf'OlItmrnl lbturc, 15 ropl~uron. 36, 38-«), 42. 45, >16; am..r.rll r dPOCM tpfiJtJuJl~:+ Uf.' Gnatha l pouch and iMcr notAl prol r ctor of, .)9, 4-1 : PolsoD dUCi, 79 ilrttulo r notal tltntor of, and ht"ad. 4-4, PoiJan hemlock. 104 47 ; fint pottrnor n bl protraclfJr of Poi.JOQ .sac, i9 4.1, 44; intc:r'IDrd~tc: nobJ prolractfJr 0': Polis" •• 71 . 1Ir.? 116. 121, 158 4-1: 12tenl nota l d .. at" r and rotat r PoII<n. 116 01, .... : I1W'ginal al'Ud<mc oi. J9. 42. 44. Polyhii",,,,. 116 45 : oa:i(lit1l proc:< oi. 38. 3? 40. 42. l>orlu, 105 43. 44. 46: «000 ~terior nnUl pro. Poot • 74 tncIor W t~.c, 37, .J.I. 45. POltcrior orbit. 88 P ropM .. 1 l'I'<JIilr.u<1n: •• 62. 6-1. 65. 66 P ... t~ .... 15, I • 11. 22. 29. 88. 90 P,opode>l piracl .... 62, 65. 66: cI",in ' POi~l hrid e, P I ] -, 18 .. 0lJ.. 0(, 6Ij; daual ar of, 66; ventra l POSloccipital ri<ll:<. 15. 19 bar. 66 Polltoccipital ri m. J6 Propod I lCcth. 61, i2 P O§toccipitnl dna, 17. P ropodeum ~ 34, 35, SO, 59, 60, 61, 6Z, Poolooxipit1l "'to .... l5. 16. 19 65. 66. 72, 77, 35. 102. 160 : apical P lOttiput. 17, 18 ICOI .. Cl i. 61. 62, IG P OIIorbil,,1 Ii .,.. &7. 88. 90 I',oolcmal opophy iJ, .38, 40, 41 .45.47 PO'bcutc:llum (:: mel>llOtum. q.,'.) . flj Pro,to,nol pi~-«I P uqul .. ,48 P ror.ernum. 38. 40. H . 47 Prea~llil.r)· exci ioo, 85 Protharaclc sduilC:S. J Pr ......... 73. 74 Protlwrax. 13. 3+-18: muscl .. 01. 41-Pr~ 1~late. accessory, .w 48 Pr~o);al !:iClc.rit~! 53 Protchymeooptf"r&, 68 Pred:a.l . in5«cs, 99 Prolr-att-on: : plrunJ. o f the (ore con, JtJ, Prem~lItum , 2J....26. :!7, . 29 oJZ; o f the prottJ uron, 37, 39, . oW. 4Sf

Pr""ral c~vily . ZJ 46 P repup:.. 17 P'O<"'lpa. 86. 8'l P r(y r "",pint was '. 21. 70. Hl6- IOS; Pammocharidac:, SO

capture 01. 109: treatment of. 106. 108, Plcwdau"/lf$. 126, 128 \I I, 112. tt3 P"odat'lllra. 87

Prickly p&r cactI! • 1 Pltwdm.'t.lfl alttt,iaLJJ (= flu,.~ltJ allS-P rodnctor: of the carda, 29; o f the middle tn'(Jcu~ q..t'.)

emera. 57 ; pleuTa~ oi th~ rore «XIn. 4Si PUUdfltt.tpKfal87

pte-uraJ, of tht hind coxa., 65 i sternal. Pl(!UJur'U/PIl adwl"riHd (=Dblicnot"'U-of ~ hind co:Q. 64 pula aoullm,uJ, q.v,)

Prolureal .rm ( = fin ' furcal amI) (= Pt"V""1 (N",,,.,ou) rib.';i. 101 rureal arm or the prosttf11ai ipOl,hy.il, PleroJtiJ{m:a, or Stiatrwt 67. 89, Qt, 92 q.fJ.) Pupa. 17, 34. 108, 117-1 20. 164-166, 17 1-

Pri.m:uor oi lh-.:= wi"lf. 56, 62, 6J li'J ro""tal "'''''" (= I.ttral lobes) (= POOl ' Pynmmtha , 104

[ Z6? J

268 NORT J1 A)dERrCA-N VUPI ~g \\·.ASP·

"Qtadratr plol .. ," 78 QII«'Il, 12, 80, 117. ,98, 140. I~, 159 ;

activit", 01, 12~134, 137, 160. 161. 170, 171 ; « It., IJ9, I ~O; color patiOI'D of, 115. 176 j diJColr)ration of old, J61 ; <ItB'-layiog by, IJ7, 160, 161. 170, 171 ; emcrgtnce of, 168. 169; fate t l f. i f! ahan· d oaU, 128, 129; rO<X! ai, 10J; hibi=.rnation 01. 98-t03, 169 i inkttillence o f, 100, 101, 126-130; lar .. of, 161-165, I of, by colOIl. , 100, 161 ; mor-11hol<>gy 01, I J-79: ne 1, 98, lJ~I 33. ISO, lSI : rhy.i .. 1 wc>rintI out of. lGO, ttd : I,roductlvity f . 1!iO: proponlon or Quttru 10 malt'"J, 100. 129, 16i' ; pupa. al, 1b.1; euooaI distribution 01, 100; ... lectlotl of n .. tina ite by, 126-130

(J wj"reus ao ifolw. , lOS (JUIf"(lU JtJtlUI ii, 101

Hab!.it brUsh 1O~ R,ulial cd l, kelOr, aruI ",Iru. 68, 69, 85,

.\\1, 91, ~3, 9S R:u.1io- iii ero Cil. 69 RlIu, Phil, and Rau, J-:cllic. 101, 100. 107,

I , Ul'J, III 11 2, In 12>, 1Z7. 128, 12'), 131, IJ2, I~ , 14.1. l' , 157, 169

R<>di , P hi lip A .. 100 Ro..':IIwnt.l r, Rc.n~ . F. de, 121 Reciprocal r~ing by adult", 114, 1(,7 j

.ftl' tll.$o Tr()phallaxis Rc:: t:ognfticn, b ! prey, other \\·~S. etc.,

109, 11 0, III Rtctum, 79 Rtdutturs of mi<1dl. :mel hind co""o, 57 /{ooupl lcnlion : r an obd nin:>I J"' m um,

73, 76, 77 ; of I.n abdom inal te:rwum.. 73, 74, 7~ , 77

Rttlw .. ;uJ l'~r.101S4' IIS, 100 [u 'pnxlucti\lt: ')' ttTll , 79 Re ~imt ion, 1.1, n 77, 101

(tractor: fir t al.ld econd mesofurcal . of te PfOpectus, 47, -4. 86; iurcal, of tl~c

m~lJpo.!> tph ra-llnu., 5S i intcntemal. 7 .... i fJ; intcrttr~, 75. 76, 7i : mtJOste.ml..1. ~7: of the mCf.Optphraama. ~ I. 6~ ; 11fOnotaJ. of the ICUtum. 46 . ..J7: of the ,ouldlum. 58 : ""II ternal, 76, 77

RI;vJiff'I, 35, (iO Rhodtoclcndron, 121 ROiIoch , J ' Ul/hhu nj~J, 1{j7 Rod.., .. 94 R",l '.f' , Tom, 100 Hbhwer. ~. A .. S.? , 53, 6'1, 86. 87, C).i,

116

ROOII, 136 H.t.J». HcrhC!" H.. • (I), 89, 91 RtJtator : diolooal, of the: 'ore coxa. 45,

46; of tbe pruplcur ,lateral ooIaI ele­,"at r aad, -44

Rothke, lAo<, 121 Rouboud, It, 115

$alP"'" 82-84, 90, W, 94, 96 SRlJitl I spine, SJ, 90 Saljva ( .. livary ,ocnlionl. 2'1, 10'1. 11~,

1\5, 117, 11 9, 120, 122, 123, 125, 14.1, I 7 ; stt also TrophaJlox iJ

Sal ivarium, 25, 26. 28. 29: mu~dn of, 2.5. 28,29

Sali""l"l' duct, 24, 28 Salivary ri6~. II ": of larva (= I ... bial

II'land , 1 Sanitation or Ih" ne<t, 116, 118, 119, 149,

I SO, 166, 16i , 170, 171 , 172 Sap. plan t. lOS, 114 . au, ur Henri dc. 86, t 16. 126. 12i , J 28,

IJ , 131, I , 43, 1<18, I . • ",8y, lIooocbary. 1117 Scale!. sa S~lae Scale ,DRCt. 104 Sca1e. . of the propodrum. a()ical. 61, 62,

85 ; oi ei~hlh la'gum ~f the mAle. 81 Sca~ 19, !IO, 101, 165 Sere\\: ·'IAo'onn ny, 101

"nt""lnMa ""Til ..... i, •. I!}+ Seuttll tt f'" un , 49 Scutcllor qpncn (= uill •• !. .IS, 49.

SO, 55 Sc:u",lIum : cntodorul ridge of . 49, .is ;

inftl!:Cllons o r, 49; brnellfform t'OC!~ ai, 4'), SO; latual ,=,It. I. 48. 4Y. ~.::; i latera l ut Ihu- pocket. 4 ; plcunl dcpre .. or f. 56 ; relractor of, 58

Sc:utoocutellor uturt . >II!, 49 Scutum, 35, 36, 47, 48. .~, 54, 5: , OlIO­

cIem.1 laid of, 49, 58; latera l ~r­(ilUltion or, 48. 51, 59; prnnotlll d..­pressor! 01. 044, prouo I r'e11"U1.on oj, 46,47

s.....w hi tory, 98-103, Ill. 16G-170 St-nK oryan., J 1

'uRl UI~l in katiu roods, 109. UO "Scn,ill. b iamica. " 31 Sef1!loria. Irl Scn!lilla buiconica: Tyloid 5f:rial v~in , oi. 68, 6Q ScUll!'. t.l, 14, :2"', 2'1, 31_ 32. JJ-, 66, ii ,

8.1, sa, 8 . \I(J, 93, ? , I I J, 160, 166 Sht:lllh, "lng, 78, 79 Shrui>o, 120, IZI. 128

illln, 109, 110, I II

[2M!

INDO 269

Silk. 11", 120, 164, 17J iu of coluniet. 1.58-160 ; I~e ailD _ est bi Z/:

kunk, 129 51.den, F. W. 94, I i ; Smell. 109, II

mhh, H. D .. 163, 164 S mith, John B, 116. III SnodlrfU', Robert E v:utS, 16. 18, 23, 17,

2l', 30, .11 , 33, :r7. 39, 40, SO, :2. 60, 66, 78, 79

Snowbcrry, 104 Solid.go, 104 Spcn .... W,lliam, 11 5, 116, 117, 118, 121 Sperm, 1b8, 169

pcrmathe<2, 168 Sphecoideo, 86 Spiculum, 81 Spider., 99, 106, 112 Spin niq:. 11 5, 164 Spiracle : abdonun:al. 7 78 ; I&rval, 163;

rppodal, 112, 65, 66; thon Ie, .]6, 37, 46. 47, ~8, 52. 58, 59

SpiraC1lI3S 'Oltibulo, J7. <18, 112. 66 ; arcb of, 37, 48

Sq'''=, 82, SJ, 89, 90, 93, 116 _ table 81, 107 Stan.ib nd . L. X .. I.it Slr"~"/ia Ji."n{ Ita. 11).:1 SternitCi t = ikrn~ ler iteJ . 1 i 14:

Sit 0110 Me OSlemwn; M.dutunum: Pro"n'mum; ~( en\um

Sternum : ahdo(uin; 1. 72- 79, 81 ; In •

35, J7. 51, 53, 54 , 55, 57 : lIIota·, 35, ~J, 60, 01, 02, <>4 ; OVIIL i plal< " I tbe, 78, 79 ; prc-, J • ~O, 41, 47 ; ilrO\)Odra1, J5, 6S ; pupal, 165, 166; st:tlla 01 nin th, is, i9 ; Iitl g/so S te:n li tes

Stigma (= plcrooti>rml'l, 67, 89, 91,92 Stinr, 13, 77. 78, 79, 1lI . 112. 158 51i"&, bulb, Sling rnidr, 74 Sting lancet, 78, 79: bubs 01 the, 79 ;

Yahc>, 79 Sting palpi, 78 Sting tecess, is, 81

ling ,hcoth. 78, 79 SbPtS, 23, 24, 28. 29, 30, 31 Stipito-colnlinaJ runic, .tn Cudino-.stipitaJ

hirwe S tomodaeum. sa Cephalic Ito-modatum St r1mo:r)', calolrans, 107 51on., S .. 107 Stone tli.CJ. 14 Striail, 71 Sub lar meooplcura! I';" 52

Su.b:thr m.c:tQpl~ r u' c.. 51, 5~ Suhllar mulde of the- mdf.ll.honax, 56. 57 Sohal .. e. SO, 51, 57, 60, 63, 6-1 Sub.mtc:Dml l j,utufU, )6, 17 Subenttal vein or C!.(:11, til, 89. 91

ubB=1 a""", 18 Subtr_1 ridge, 15, 16, 17 Subte.nat suture, IS

ubmonrum, 2J, 24 uoki", pump. 3 1-34

Sulcus: antcmw 17; of the occipital .reb, 18; lullmar.ll'in.ol, 01 tho. m<:tano­nom. 59, !iO

Swptl1'O<io: of the coo,l>s . 94, %, 12J, 12S, 130, 131, 138, 139, H , 14J-148, 173 ; .1 the ncol as • whcl 94,?6 lJO, 142, 14l , 147. 14 , ISS, 157 '

S"'_iUlU or tho hood and f.orclq5, 38, ~5

") m/J"Dr1€ar;Ju t"a(nnozus, 104 S1"/OltUUpir, 60

yrph i!lac. 107 Sj'IImI:lUC trc::umcnt. 8-1-97

T.w.u. .. losiophthal,.u.r , 107 TdbanlfS Drim" 107 TarJ;J' c:laws, 8S .. us, 1J, 71,85, 101, IOZ, 165

TaXonOmy , 84-9; Trcltniquc, 41, 42 Ttkula, 31, 49, ~ Td.·7fPM'u.00 T <mporal rou .. 164 Trnual adultS. lOS, 11>6, 167, 171 Tcnthrcdinid, .l9 'TeJUoriaJ DrmJ, H, 10. 18, 19 Tentorial br idge, 13, 19, .l2, JJ, J4 Ten ri.1 piu, IS, 16 Tentorium, 1.5, 18, 19, 20, 28, 33 Teratolosy, 176 TCf1tO""lt nul extcraar and rt:t~ton. 75 t

76, 77, 78 TOl'!fum, 73-78, 81 , 85, 92 Tlurtt..'O ,lAtota., 107 Thore, idac, to7 Third phr"""", sa PrOpOdcal pre-

(Ih!'3lr= Thompson, W. R , 163 Th""""n, C. G., 86 T lmraclc spiracles. 36, 37, 46, 47, 48, ~

5~ Thoracic trunk, J4-J6 Thcru, 114 ; in Doficlutt'up./Q, 95 ; or

th" femal .. 13, 34-72; of the !;arv., Iii?, 163 : of lhe ,nolo, 80 : of the po"", 165, 166; in r'j'JJtvJa, Il2

r 269 J

270

Tibi •• 13. iO. 71. 5 : nf Ddi ... htr.~s,."la. I)S; f the pu 165, 100; of J/uf'O, 89 : of V upuw, qJ

T;hial .pur. i l Ti{,ir~n ChiOrtJfttErtr, lOi T ibic(" tyr kNf. 107 Til'v~1 to}i, 101 TiUyard. R . .I .• 68, 69 7 .r.la, 107 Tood. 106 ' "Ofl.10H Iltwcles of the ahdomc.n. 65. 66 T ...... lla ', .... 010. 104 Trachea, 550. 66 T ractota l 'u ~pcI1.ll0rt 24, 2S T nrurnc:tanolal .suture, .59, 60 1'ra~pl(:u ral sutur 52 T rau scuta I fUture, 48, 54, 55 Tr~usst('rnal I Utu.re. 53 T .. p • . boi l. IG4. 108 T rj%'. 101 . 1G4. 10.;. 120, 13J, 134 Tri:n'1"ilar sch:ritc oi Lilt: c.il,{hth abdomi·

0\1 st("m um. 78. 79 Trflf~nkr. 43, 57. 64, 70 ; mc~{urcaJ

dt!prusor of wo, 57 TrOfb."&ntin, 40 TraphaUaxi •• lOS. 117. 11 9, 167 T)luiJ,. II. SO. , 91.93.95

li Jricb. \\""d'DU, 20. 23

J"'aail' nun onfl,,,,,, UN "/ '(Jft"O~ j"'lrrfal'," 8J

ah ub ot Lhe propo<lcum, 85 \·."cc. A. I.. 16J. I Vcn~t ioo of the \\ .tl,!::I, 6lr70, 85, 89 \'ClIQm , i'V Ve""",lf. . \ 1' . 81 I·ertex. IJ. 17. 21. J3 87. 88. • 164 Vcrtkal " lo t : ,'f tbe aW,.muw .,.,tiul.

62, i2,. ;3 ; tJ( the mda.tn-ml avophy· !'l i ~. ~4 . 5.7; oi d ie meta l~mal.rophYJ~,

S. 61, 62, 64. 65: 01 the p""'lerlUl "I"'ph) Ii .. <0. ~ I. ~S

f'ura : food and reeding behavior of, 10.1. 104, 1115. 107. lOt!. 114. 117 ; mor · phology o f. 11. ·11-1 . ~59. 6Z-6u. 67. 7-1)1 ; n13U anrl 1)OIin habilJ: 01, Il ,

121, 126, 1.13. 1J4. 15.? 157 : tnonUl"it charac",,. of. II. 85-90. ?:!

l ~tSta blilicl1Sf1, 81, 1; \.r snr'Nri, 90 1'est b.rtaii.r. 97 VeJf(! (iH,~/a~ 90 r·n,. ""hr.: nnihGli m of. 111 ; foods

and l«di,," I!cluovior dl . 10.1. 104. 105. 107. lOS, 114. 117: morphoi<Vl" "f. I I. ~ 1-18. 54-59. 6Ui6, b7. 87..00, Y2:

De1bi and ne.ti-ng ~IQ.\'"ior of. 1 I, 121, 1!6. 13 • 134, 152. 157 : "'xo""""" <har­ac:t~n of, I I. 8.S-90, 9l

('uta dUJ:tn. i)() Vupo diot10/ i' iJ (= D(llicho.·,.s/1wto art.

"ana, q.'V.) f' ",. !.,.,;aa. 00 Vupa gt""dHI'(C (American ~ns

= VrJp.1a _cNlil ' D .... q.t' • . lOtI, 126. 132

V u~a JlMnUallJ = DtJtic/un,atwl4 " lQ.CW­

lola. q.v.) V,zta rMlId", i,tid. 90 V fll'fJ medi(! ( = DoUd!M't.JltJA lll nu,JiD. .

q ".) Vt'sfa mo"goti(tJ, 90 1-' tlptJ tff'inua/is, 90 I't.pa /'Oroll,la, 90 V t.t/'fJ sozolt;ro, 129 Vcspo s.vt!J~nris (= Do/idtOtysl"la syl­

'VJl r i.s, q.;'.) Vt.,., udgDl'u (= V<lt~/a wlflDns.

q,{f.) Ve: pidac, 3, II, 'ZJ, .. -, 116. lZ7. 1-16,

1601 Vf pinae: MoIo n' oi. 11 . U2. . 96, 9S-

176. definititJ11 hi, J, I I· dlagnnftie e1aracton of. 85 ; morphology ui . I.l ..

: taxa.lOm)' I. 8>97 Vetpoid 1 J ~noptc.ra. Va!,uia, binlOCY 01 . II , 92, 98-17~.

fiUI''"; canuihDhm or, 108. 117-110 : c.hanl~ in nest ~hap<" f. 1 ~ - , camhi oi, IJ9, 1 .. 1. 142, 156 ; cUltinv. down 01 old edt. by, H8, 1 .. 9 ; t1 ~iD" tiC ehar;ac­ten of. 11. 116-94 : diAtrWut. " 01 ""II typet in n~l or. 1..l9-141 ; dU~lion of tOlum., ul. IJ • 119. 1~7. 168 : CJDCr1I­Cllt""t tb.les ftlt rnalc_ and qu«ns of. 168-170 ; e:nlr:mCts t " hurro~s of, 136. 137, 152 i mtnnee ttJ 1'1 t prolltU 01, Q~. 152. 154: Ofr.uic ne..( bll1ldin~ bl. 1048, l Si ; cxca\-.tico 0 ,oil by. l l , J.!t.I, 128. 13+-137, 155 : 1Tl<,<phoIOfY oi. II 96, 1~166 : nt_'" 01, 11, 12. H 116-124 12;, 1!6. 127. 121:1. 129. 1.10. 1.11, 132, 134.151.154. 156. " 9, I"": "" I 11 of 94, 136. 150, 131 : o\trwlnttrinK o[ coin· Ole ' 1If. \19, 118, 140, I ~I. 159. 160. 170. 171: produaion C) ttt,.,. que:enl Rnu ma or, 139, 1 : rruportJCJn$ of cash:, in, 16i, 168: qUl:rn nr I of. J31, 132, 151 ; taJeQl1Qmy 0(. 1 1,85-9-1

l 'ulultJ allSlna(o, 86. 93. !N. 175 Vtst~ /o tCU'tJlmcs (=: r'aJtuld .tq .. H' 111 ,

q.,'.), 106, 107. lOS. II I, 175

12701

1'.·.1/" .. /0 dUJh~/ird (= Dolidtot'ts/ltlla '"~. I na. q.,:'.)

V U /,u/a gt'""O'fj.'o ~AJl1etian i'il)«imms = Vt'Stulf). HlacuJif"ot$.l. q.t-'.) , 94

V~ltllla kctrulfsu, 175 1If'¥/,ulQ lmU;i~ 115 JI u/lrJla "'"Mia/a. (= Dol~htx..'Up",lD ma.rv'~t." q.t'. )

1'''1',,/0 HlOCN[i!r01ts: «Ilor paltnn and phyl""cny of, 175: IIXld •• 00 feeding ... . "lOr oi. lG5. 106. 107. 106. 1()9. I ll . 11 4 : hibornation of. 101, 102 : 00.1-inK si te f, 126 ; quew ne~ l of. 132 ; .prJnl\" males o i. 170

Vtsft1,t14 p.ensyhvmicn : cannjbalism of, lOS, 118-120: tolar l)IItlffll of, 17-!-170 j <k:r~h'e and offensi\ c behavior 01, 167; disposal 01 dt>d brlXld by, 171 ; .,.<11' Itllg of. 160-166: ...-g-Iayiug by

2 7 1

ent.c o f young I.{UttrU ilDd malt'I of, 169; foods and fcedioa bchavio.. of. I ll . 112: hi"'rn.hoo 01. 101 : II. o f. 121. 115, 1~ . IJJ, 1.34, 136, 152. I : Qvt'rw.intc: r­ing of colooi •• of. 10\0. 159, 160: paper uf. 12J i I perinI" over of old comb and nest .... 11 o f. IJ6, 1~9. ISO, J)OVul.t; orr of oat!: 01 , 159, 160 ; qUWt ru:\t or, J3J ; Juspe:ru.ori 0(. 144. 145: 1u:o­oomit Ii.ling of. 901

\ ·atibulr, £piracular. J7, 48. 62, 66; .. rtb o~ 37, -48 e m ure. 13, 14

Vir5dnia cr~t 104 Vol .. lI3, 82-34. 90. 93, 94. 06 ; ' <lnn I

tubercle of Ih .. 83, 90 .. V.NY" Ift",.lj~p,." 32 Vo ... F .. 41

r ke.n of, 161 : c.xca\ .. tiO!l u{ toil by, \ \ falke.r, F . \ r .• 110 21, 1.15, IJ6, 158 ; food, and lecding \\'alu t 135, 169, 170 Iuobi .. or. lOS. \(18. 1 H : hibernatlon of, Wa""". "- B., 107. 131 99. 101- 103, hunting bohavior nd Weather •• ff«1.l of : on hibonwion. 9'1. trr.atmcnt r captured prey (II , I I I- I IJ : 100, 101; on tnortalit)·, 99 : on nC"!t , "A. inquilincs of, 1101 11 1 ; i rr~lUlarilid in 127, 128 ; SU \\'intef "e" c nsl ruet i,,,, 01, 155, 156 : mO<- Weber, 11. R, .w. 41-18. SO. 51, .i4-SY, r liolo\<y of, I ' • 86. '14,95. 1bll-166; 60. 62-66 nc-!> 01.. I . 124, 126. 131, 135. 141 . \ e<ds,I20 147, 152. 155. 156. 159, I ii ; nelt ... all I 'ti h. of nC>t and c"mh •• I~ I , 1<12, 143. of. 152: 0" rnint<r ing o' 99. lIB, 141. 1-14. 145. 146. ISO. 173 159. 1; 0. 171: pap<r of. 123. 124 : W. t .. oOO, ). 0., lO.!. 115. 121, 133, 1 • poP' lhuie," or nel t of, 1-9 : proportions 135 fIr . and small .e11s in "eU of. 141 ; 0,""«1 ... W. M . • 5. 114. 115. 116. 117.

quo:c" n<>U of. I 1. 132. IJJ: spring IJ • 168 IIli1lu of. 170 : ' Hato loait pcotmens of. Wh;,,· fa"<'d homt: t (= DDlkhaN.,.I. 176 muculota. t,pl.)

I'tstuio rN/a. 901. 117. 1'2, 175. 176 W, ll "'. 101 l'tSfU(" ,"Wfa yat. (1. (H/,('a, 170. liS \ \ "ing,, : D.J1aI lobe of, 61, 85: axillAr)' ,'t1ttl/tJ rofll ' ·ar. -I",,,riftJJUJ (= Yll f . C(lrd.s 01. ",,0, 59; axil1a.ry exciJim 0(.

CU'ilJica, q.t·.) 67: u:illary scln'ilb aI, -48, 049, 50. 5.1, r'u w., ... /1l yar. DlrotiloslJ . 9-4. 114. li5 56, 57, 59, 6J. c;." 66, 70: cOlttll n12rgin r 13,.,10 ,.{o var. ''Ill,,! "'Ul. 94, IT, ul, 66. 67, 85, 89, of DDlif"hO't'('s/'M/g~ 95 ; f'rsl'l4/tl nil.] var. ",'ITPHtdiu . 170. 175 I~nal f~ld of, 66; fr~n:u hoolQ; Of

1'u/,ulll ,..-fa ..... 1' Jln/rn' . 115 h:UDUli o~ 63, 66 ; Rwenl morpholusY rl 1/,lf{IJ rw(<f Ya.r. ~,·dJ,uJ. 94, li5 (,f, fl6..70 : pO. h-rio r IrJ)c of, as: pupal. r"lIt-ul'ltl'UUHO.P 9-1, lU I, tU2, 106. 107, 165, J66 : \cnation of th~. 66-70. 85,

lOS. I ll. 17;. 1;6 89.91.9_.93; 01 Vo,.. 89: 01 Vlrtula. l 'tJ~"I. ,1 ... <lDr. 175 91. 9Z, 93 I '"fa.da AlplcwuiJ, 94; color l"auerll and \VillK-eJune:r, 71 rh~1 <ny 01.175: lood, ilDd ferlli. hc- Wmg ""d, 1M ~\·i(lf I. lOS, 109: ~ tinK" site 01, 129 ; \\'ina: IJf"CJ« J . antcriOf'" naQl. -48, 49, ~l, or: 1 truct.utt 01, 1 ~2' paper of. 125; 59, pleural. .51. 52. 56 : ~l(rior notal, populat1(JQ o f Dell o i . I29 ; u.se or WIIkr 4(1

b). 135 Wini«. rcl.tlOfl 10 donn:oncrand """0,,,,1 Vt:ll'~/jJ .'II/gam: tt'llor tMfI Uan ill1d phy. llutctry, 97, 99-103

IOKrny of. 175; comLs of, 142 ; cD'lCrC- \Voad fiber. 98. 123

[271 1

27'2 :-. OR1 H AMERICAN V£SP I SC WASP

WoorI , }. c;. 1011 \\~ onccr ; acuvi1ld and duli i 70, 98,

1113-16(1. 167, 170. 171; b<h.v;or 'M! en· terinQ and luvin,g Ddt, I SJ; cannibal­"',,, of, 98. lOS, 11 7-120 ; color It<ra 0 1, 17", 176, COCOOn 01, 167; <>.rty .~ .. oi, 16G-166; cgJr.l.ylnR by, 158, 11>1 ; food. and Iffilm. activities of. 1OJ-120; hll><rnotioo 01, 98. 99: mor" plIO OIl)' ai, 12. iV. 80. 85, 87.88; nclt· bUllillnlr b)" 120-160; nurs ina irutinctJ

of. 119, origin nf auleJ, 139; propor­tion of, to other castel. 129. 167 ; krato-10Iie, 17(); of V.s~. 87,88, of VIS""'., !l2

Worm, 106 \ nabt, William ., 107

Zabriskk. J. L, 146 Zander, E., 81 Zonin& of brood, 1611, 170-J7J