On the biology of inhabitants of oak galls of Cynipidae (Hymenoptera) in Britain

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BRITISH TRUST FOR ENTOMOLOCY LTD.,
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TRANSACTIONS OF THE SOCIETY FOR BRITISH ENTOMOLOGY
GENERAL
A Now Creptpn rN Zoor'oetcl:'
Nouauourunn: Tsr Rprotus
rNstlrurnD BY tm Trrntnpxrr
Ixrraxntroxlr, Cor.rcnnss os Zoo-
r,ocY, Pelrs, Jur,x, 1948. By F.
Eomming, 1950. B pp., ls. 6tl.
Tsn PnosLnM oF srAlrr,rrY rN
Spnorrro NoamNcuronn, wrtE
SPECIAI/ B,EFEIENCE TO CASES WEERE
TTPE MATERIAIJ IS NO I,ONGER IN
ExrsrENcE. By F. Ilemming,
1951. 16 pp., 2s. 0il.
Sorur loarrarroNs oa rNsrcrs ro
ENVIBONMENTS AEAT ABE AIJTEa.N'
ATETJY DRY AND rr'OODED' WrlE
SOME NOTES ON TEE EABITS OF THE
Srnmrourrpln. By E. E. Ilinton,
1953. 20 pp., 3 flgs., 5s. 0<I.
Tus rsn,Mg ttLABvA" AND ttNYMPg"
Tsr Lrru I{rsronus er.rp Mren.l-
TroNs oF trp EpmunBoPrEBA rN
A sroNY srBEA![. By T. T. Macan,
1957. 28 pp., 12g. 6d.
TRANSACTIONS OF THE SOCIETY
FOR BRITISH ENTOMOLOGY
v()L. l4 NOVEMBER I96I PART XI
Oru rrrp lhor,ocy ot rrrE INu,lsn',rl'r's orf OAK (+At,r,s oF
CyNrrD,\r (IIvlrnNorl'rRA) rN Bnrl'AtN
I31'R. R. Asxuw
(I)epartrncnt of Zoology, IJnivers.itl, oli Manchester)
Gall u':rsps (Hymenoptera. Cynipidae) induce the gro.w,th o{
galls on a variety of plants, but oak (Quercus) is host to thc
gre:rtest number c,r-[ specics. The immature stages of t,lre gall
wasps are passed insirle the galls. Il:rny other i;lrecies of
Ilymenoptcrl feccl eithcr on tiie vegetable, tissue of the gnll, ol
on tlre otlrer inhabitar-rts, ol both. Tlrese species arrr confirred
aimost entir'ely to the super'-falnily Clialcidoidca or to the tribo
Syncrgini of the Cynipidae.
Rearing of tire inhabitants of Cynipir-l oak gnlls has been a
popular cntomological pur:;uit; the nurrtcrous lists of parasitcli
and inquilines of Cyrripiclae in the litcrature, particularll, zrbout
the beginning o[ tlie prcsetrt ccnturv. beat, rvitne-qs tcr this. Un-
forturratt'lr', largcly bet,ausc of thc dif6cultr. inr-olved in correctly
narning urany of the Chtricidoidea, thc vnliditl- of these olcl lists
is very suspect. llhe salrrre clilficulty has probably discourng4ed
rnany, cntomologists {rom investigating Iultlier the biology of
the c.rak gall inhabitants. Blair (1946), iron,ever, erstablishecl rna,ny
of thc inter-rclationslrips bet,u'een the inhabitants of g:rlls of
Andricus koll u r i (H:rrtig).
The present paper inclrrdes rlia,gr:rnrs oI tlrc inter-relationships
of the inhabitants of a largr-' nulrber of oak g:rlls. The factors
lr,hich detennine thc launa of galis of oali Cynipidae in gr:neral,
and crf Cllnips species in particrrlar, aro discrrssc.d. anrl a key to
the identity oI adults of the 0halcicloiclcz'r associzrterl lvith oak
galls in Britnin is provirlecl. Str.rrlies rrn the biology oI the
Chalcidoidc:r, inclucling dsscriptions of the irrunature stagcis, s'il1
lle publislred eiservhert-..
The inhabitants of oak galls may, for convenience, be dividccl
into tluee categories, as follows:
l. 'Ihe gall inaker. The host Cy,nipicl rvhich alone is capahle
of causing gall deveioprnent,.
2. 'Ihi.inquilines. Tlris terrn is applicd to spccies o[ Cynipidao
rvlrich do not inrluce lrritrrnr'1' gtrll {onnation tlrcnrselvcs (althorrgh
tlrel- rrurr- cirrrsr. thc gnll tr, lrr. lrypcrtroplritrl nntl rrrallonncrl; lnrt
live in galls formed hv tlrt. 1111c g:rli rrraker,s. Tlrc spercics assor:iated
rvitlr o:rk grriis belong to tlri: gonerit S,yrtergtts, Suphortecrus antl
Ceroptres. 'I'hey Ieed on the gali tissuc arrcl therir presencc rnay
or rnay not prevent the deleloprnent, of the gall rnaker.
rN DNToMorocY. A summary of
the views of W. E. China, E[.
Ilenson. B. IVI. Hobbv. II. E.
Hintonl T. T. Ilacari.' O. W.
Richardo. T. Southwriotl, and
V. B. Wiggleswortb, followed by
a review of Tm TusurNor,ooY or
Juvurrr,B Pslsrs or INsEcrs by
.8. G. Davies, 1958. 10s. 0d.
ENTOMOLOGICAL FAUNA OF
THE NEW FOREST SERIES
Introduction bv J, Cowley, and
Plnt 1. Odonata, by Lt.-Col. F.
C. Fraier, 1950. 12 pp., ls. 6d.
Penr 2, Neuroptera, by Lt.-Col.
F. C. Fraser, 1961. 72 pp.,
ls. 6d.
ORTHOPTERA, Etc.
A Sutrrareav or rEE Rrconoro Drs-
TBIBI'TION
TEROIDS.
L952. 16
oF Bnrrrss Onruop-
Bv D. K. McE. Kevan,
PP.,5s. Od.
H EM I PTE RA.H ETEBOPTERA
CorrnrnurroNs fottrAnDg lx Ecor,o-
crcer, Sunvnv or rsn Aeulrro
aND Srur-Aeuetrc Enurprpn.r-
EETEBoPTEBA oF TEE Bmrrsg
Isr,ps.
Scorrrsr lftcur,lNos lr.ro Eest
lNo Souru Dxcraxo. By E. S.
Brown, 1948. 45 pp,r 7s. 6d.
Tuo Rrssr,r Yer,r,uv (Lexce-
ssrno Sourr lNo lfro). By E, J.
Popham, 1949. 44 pp., 1 map,
8s. Oil.
Nonrx-Elsr'Wer,ns (Dnxrrcu-
sErn,E aND Mua.roNnrrsrrnr). By
E. J. Popham, 195I. 12 pp.,
2s. 6d.
Tur Hnurprane-IlrrnnoprrnA oF
EENI. -By 4. M. Massee, 1954.
36 pp., 7s. 6d.
Tup Broxouros AND Iuuerunp
Srecns oF TEE Tsrgrr,n Lecu
Bses (Tingi,s amph.ata I[.S. lxo
T. card,ui L.: Eru.. Tnrornln).
By T. R. E. Soulhwood and
G. G. E. Scudder. 8s. 0d.
COLEOPTERA
Aeuetro Cor,noprnne or Nonru
ATJEB. _ PV U. g.- Brown, 1948.
pp.,1 fig.,ls. 0d.
EPHEMEROIPTERA
DnsoaJprroxg or soME Nvupgs or
TEE Bntrrsu SProrns or rEE
GhNus BoZfis. Bv T. T. Macan,
1950. 24 pp., 6 -figs., 2 tables,
3s- 0il.
A DpsonrprroN or rEE NYUPS or
BaEtis bucerallus wrrs Norps AND
A KEY TO OTEE& SPEOIES IN TEE
cENUs. By t. T. Macaa, 1957.
8 pp., 3s. 6tl.
Tsn Aeunrro Cor,Boprru or Woop
'W.lr,ror.r Frr, wrrn souu obu-
pABrgoNg wrru Wtornr Fux axo
sorfg otEnn, Elsr ANerrlm X'nrvs.
F^y F. Balfour-Browne, 1951.
36 pp., 4s. 6d.
Tur
\r
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I
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Tlrsuz CERCPTRTS
ARATOR
1
ond
EUPELMUS
UROZONUS
EURYTOMA
BRUNNIVENTRIS
lO golls exomined
Goll Tissue
--) I
ANORICUS
SYNERGUS . +CURVATOP,
sp. t. COLLARIS
273 golls .xomiEd
lO golls cromin€d
TORYMUg
CINGULATUS
MESOPOLOBUS
AMAENUS
. '.?
MESOPOLOBUS
FASCIIVENTRIS
or JUCUNDIJS
. o. TIBIALIS
or FUSCIPES-
or AMAENUS
oTALBITARSUS
DIVISA
MESOPOLOBUS
AMAENUS*
s
EUPELMUS.
UROZONUS'
460 goll5 axomincd EI.JDECATOMA
*VARIEGATA
1
Goll Tissur
f
ANORICUS
CALLIDOMA
I. CIRRATUS
34 golls .xomin.d
coll Tissue
13 qolls .xonincd
MESOPOLOBUS +
22 9oll9 cenln.i
?. -.....
MESOPOLQBUS
JUCUNDUS
ANORICUS
INFLATOR
*T*
t9
Fig. 1.-Food rrebs in galls. of Andriats species.
I lO golls aromin.d
i EUPELMUS
. .UROZONUS
169 golls .rcmin?d
Goil fissu.
i
ANDRICUS
. OSTREUS
I,'. EISUNEULUS
a
MEsoPoLoBUs MEs@ooBUs
TIBIALIS FUSCIPES
L--iI
7...
MES@OLOBUS
XANTHOCERUS .,'?
MESOPOLOBUS
XANTHOCERUS
?..
MesoPbLoB,s
TIBIALIS *
?..-
CEROPTR€S
ARATOR *
?....
EURYTOMA *
BRUNNIVENTRIS
Gqll
Goll Tisa.
1
ANDf,ICUS
FECUNDATOR
OLYNX ..EUPELMUS
TRILINEATA UROZOAT,,S
7l gdlls cromincd
Goll Tissu.
I
I golls
Goll Is{. -.-? Goll fTissue
*-ffi6,., Ftf')iti}fu*:.m.
r_8IL!!!EALUS e _
2 oolls adin.d . SYNERGUS zue LMUS
- rNcRAstot* uo*o,l?L1?"-.0
1Jas.y)
Goli Tissu.
I
ANDRICUS
FECUNDATOR
f. PtLosus
l?
i€sofu-oBus
XANTHOCERT.,S}
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eoll Tls$.
I
ANDPICUS
OUERCUS{OPTICIS
30golk@ilEd
Goll Tisse
1
Tissue
,l
36 golls exomined
t
Goll Tissu.
1t€
l{r
*
ANDRICUS
SOLITARIUS
,. OCCULTUS
4 golb €rcnlild
Goll Tissu.
17 golb aoDhld
*
s
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qUEFCUS{AMT'LI
.,,? \
ISEXWA cftIliluu
4 golls axmh?d
or
HOBBIA
-/srENoNorA
TETRASTICHUS
, / DIAPHANT[JS*
4l golls cmincd
Goll Tissue
1
DU
EUPELMUS
UROZONUS
EURYTOMA
BRUNNIYENTRI t\,tESOPOLOBUS
TIBIALIS
ORMYRUS *
PtJNCTIGER :Eunvrova
BRUNNIVENTRIS ET,DECATOMA
VARIEGATAT
lTgolls aomhed
Goll Tissu.
1-_/ I
i alronrcus
I sEM|NATloNts
."'J**7.-'ro*,
GALLAE.POMIFORMIS AURATUS
od }IIERVOSUS
-*:ffi..rs*o#,F*,,.
EUEDOR.€SCHUS \ARSAMES
^raaorl.orra ,t trorrr*
XANTHOCERUS I AURATUS
I golb €xoilirEd
Goll Tissuc
1
CALLIRI.{YTIS
,.-,.? GLANOIUM "
MEGASTIGMUS EURYTOMA
DORSALIS* BPUNNIVENTRIS
Goll Tissuc
-"I
I TRTGoNASPTS
SYNERGLE r SYiITOMASPIS
THAUMoCERUS FASTUoSAt
Goll Tissuc
1
I
ANORICUS
GLANOULAE
3 golls eroined
(Durtrom e lrgyU) z
4
o
o-
e
4
HEMITELESSPI a,golls cxorincd
245 golls ?roDinsd t27 cctB .roDirEd $yrc rorct)
Fig. 2.-Food rvebs in galls of Anclricus, Riorhi::ct, Calli,rhytis and. Trigonaspris species.
? ---
MESOPOLOBUS
JUCUNUJS
MESOPOT.oBUS
TIBIALIS
77golsroiId
...TORYMUS
AURATUS
Go[ Tist
MESOPOLOBUS
TrBraLrs
EUPELMUS
UROZONUS
liESOPOLOA(JS
FUSCIP€S
,7"2.
." rorivr*,s .
, AI.RATI.,S .MESOPOI.OBUS
TIBIALIS
LIESOPOLOBUS TETRASTICHUS
FASCIIVENTRIS AETBiOPS
lO3 go[s .xqnh.d
Tls$r
1
?...
MESOPON-OBTJS
FASCII\€NTR'S
?..
I,ESOPOLOEIIS
OUBIUS *
MESOPOLOBUS
TIBIALIS
8l golli.rominrd
Gotr 'Iisiuc
I-3
"IHTSJ,HHS
T FUMIPENNIS
382 grdls rEnin.d
Tlis
1Gq[ Tis
S.rTGRGUS
ALBIPES
o. PEDIOBIUS
! | l"YSlS
TORYMI,S
AT,RATUS MESOPOLOBUS
TIBIALIS
67 gollr axodrio.d
C*0 Tlssuc
?...
r/ESOPOLOBUS
FASCIIVENTRIS
.7
TORYI'US
AURATUS
".EHftY'Ifr'*t,.
3l8 gons crqniRd
Tissuc
f
eu'Rrrotra
BFIJNNIVENTRIS
I
NEI'ROTERUS
APRILINUS
i. SCHI-EC|.fTeNDAL| NEUROTERUS
a+rNrrs
MESOPOI.OBUS
FUgCIPES
53 golls cxonin€d IS gols q@incd
Fig. 3.-Food webs in galis of Ne.uroterus s1>ecies.
t\9
lts
30 golls erohincd fB goXe rchin.d
\
EUPET-I,TUS,
UROZONUS
MESOPOLOBUS
FASCIIVENTRIS

t9
*r
lc
CYNIPS
otvrsA Gol Tisru.
CYNIPS
ouERcus+ot.fl
t TASCHENg€RGI
Goll Tisu
1
CYNIPS
LONG&ENTRIS
T SUBSTITUTA
?,
i/€SoPoLOBUS
FUSCIPES
MESOPOI-OBUS
FUSCIPES
22 golb.rortmd
Gol 'Ii6sr
3gS golls .xomin.d
;? i.
12 golls aodn.d
252 golls .xomin.d
t\lCSOPO{-OBUS
UJBIUS
SYNERGUS
PALLiCORNIS 2772 gols erahin.d
Goll Tis$€
T
E
CYNIPS
DIV'SA
t VERRUCOSA
TORYVUS
AURATUS
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- Goll TB5u.
/// I *-..-.-
"i['EtEti. nfiiiE'nFfiiHEfffut
MESOPOLOAUS TORYMUS EURYTOMA
JUCUNDUS AURATUS BRUNNIVENTRIS z
o
v
o
E
29 golli .xomin.d
(C!nb.rlqd)
Irig. ,1. I,'ooii rrol.r:r in galls ol ('tlnips slrci:ios
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=.,.i-:,'i - p ] i. ';i. = t.- ia:_7I i5<:ir:i=?, l== .:.==-! 3l=.: i3 =;
a: lZIrr:: =a'=i-.!'=" 7;=_ 46.^r ='-:5=- -i: =:.=:. + =-:1 r= r == q 5== gf, 5 -.
+:-='r= = i =;:-'n-iGoI.=il12Zl^_ i !^qii::li;==.=;*n,.2
^ !, n j=.-I' =-- i 1:'? - -' =-=:i"-'-= - ..D ='-t _?,a = i:.= == i. lt;*.. 4'_. ='6
1: ;;,!-:: E.i?is : ; !-r !11=1'n'io 1=;E =; ;?=1: :i::i ;n': z4 ii;'
€€ :i::; =:1i; it: ; l.i' -: r ": 6 6 --i =) -=.-!, + =.=1':.:+ii i t:-i3
:; !a+;a { i5r::}iiF;: +zAtl=:E:a1ii=,slii;:1a; a ti3=ia
l;! = I =S =:': ]: + ;",': i + =q! ?i?; +1+ 3eq -=:i,'=?ii1i'i =: *7; i3
=r -;:l: i";E 3.: if 1a= er.i=i':1il:_21i,: =iia^'=;':; i=: 1a:l
;;?: tl; ;l9 =;1ii I* = aTlistl=u1l_=:=ti;a=_21==l .; =?- 1ii
i*:i?,i_3 ; i i;ii;i=;1 ;;=; i?:21=;Iz;T i liii; i;='Li= i;
g:s.;-d. 9=?^_i=_o i..:: o !:r=, ;._l?6i=!=. Z==i3r=11::;2li-=
;; r;?r)fr - it; ;i1.3 +, i-= = : :-=' 1--:-p o -;o: ?.;ri
==: = i t -* z^?E3 =li. ; :- = ;1: =L:=4'. - -=' ^ -1jj3 :+a;;?7:E=
ig*;1rf lgei;li = 1= i:;! i;z:;;|!; :=/i;-ii:ii=t*-
:':-^ =' Y Z'-a-i ^ a ==
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=---"-A 'i*-'"- Snre '; n^a =:i., E=:En- -353- !'=?ii=+.
a * P 3 +;
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F: q =.i 1;1; i-i;1 B- 7 +si ;; l; J= i:-, =:lzig ;=ii:iiiA E
.r; . i.:Jl

244 [November
5. Possible exceptions to 4. are Cecidostibct lencopeza (Ratz.) and
C. senti,fuscia (Walk.) rvhich are probably polyphagous in the:
gnll of lliorhi;a Ttullidu (Oliv.) (i.c. thcy are gall specific).
6. Spccics ol LlesoTtalo{.rzs anrl 7'onlwtus risr.ralll, attack any of t}re
inhabitants available in a r.aliety of gails.
Frorn an exarnina,tion of the food wobs a nurnber of othcr
generalis:rtions can be rnilcle.
A sirnilar p:rttcrn is fotrnd in the foc<l u,erb o[ the rose gall of
Di7:lolepis rosne (Blair. 19,14; Aske,x'. 1960) although in this gall
the po,'ilion oI n ,\riltctgus is oct'upierl lrr- /'r.r icl isl tts btut,dtii
(Ratz.) (Cvnipirlae, Svnergini), ilut'tltottttr bt.unniuentris by il.
rosoe Necs anc.l a Meso4lolobus species by another Pterornalid
II abr ou 1 trts tt ede11t un' is (Ttronrs.).
196r I 21it
(iv) Forrn ancl positir-rn on the tree are also oI irnportzrnce in
rlc-rternrining the parasito curnpiement of a gall.
Galls oI the contenrporaneous sexual genelations olNeurote'rus
-'N. rlrtt:tcus-ltaccururn. (n-.), l/. numismalis f. uesicator (Schlccht.)
anrl Ar. albipes (Sclrenck)-have foul species of Chalcid pnrasite.
'l'anlrttus ctttt'ctt'us (Fc-rrrrc.), Mesopolobus tibialis, DurqLomu
brunniuenLtis and Olynr arsarnes, in common. Galls of .4,/.
n,unti,strutlis f. uesir:ator and N. ulbipes, however, ha,r,e anotllr:r'
tlrree species in comrnon. Tetrastichus aethiops (Zett.),
hlesapolcbus luscipes, and "[/. Jasciiuentris \Vestw., which do not
attack galls of N. qt;etcus-ltuccarunt,. Galls of N. numistnal'is l.
tesi.cotor and N. ulbiprs are srrrall, thin-r'alled and situated on
le,rrves. G:rlls of N. cluercus-ltuccurtnrt, on the other h:rnd, arc
large, ttrrick-wzrllcri and o{ten situated on the male catkins.
'i'[re contc.nts of (a
quercus-bQccaTu,rn
Taslo 1.
) catkirr galls ancl (b) leaf galls oI lieluotetus
(L.), gathered in Wytham Wood during I\[ay
an<l June, 1958 and 1959.
(i) Onlv a ferv specics of parasites are gzrll specific
nraioritv attack a lirniterd range of grlls. The
195ti 1 959
(u)
ltrtli n,
oal,l s
128
(b)
.l,eu..f
g a.l.l.s
117
(ii) Tlre semcr parasite spccies :rre not usualiy found in galls of
altcrnate gencrations oI the same Cvnipid species.
For exarnple, the single Chnlcit.i, llesoXtolobus fttsci,pes (Wtrlk.),
found to att:rck galls of Cqnips lonqi,t;entris f. substituto I{jnsev
and C. quercus-folii f. tcLschinbergi (.lchleclit.) is not includeil
arnong 1.lre clevc.n Chalcid sperir:s rccorclecl fronr galls of C.loncli-
uentris Htg. and C. quercu,s-lolii L. Similarly I'orqrntts nobili,s
Bolr.. the only Chalcid bred fr:orn galls of Bio,rhiza pallida f. apterct
-(!.), has not been found associaterl u,'ith g:r,lls of B. pall,ida (Ofv.1.
None of tl're three parasitcs in galls ol tlnclyicus ltollari f . cira.rlo,ns
M;.ryr llesopolobus fuscipes, M. ronthocerzs (Thoms.) ancl M.
tibiuli,s (\\restw.)-are includecl in thc list of tr.velve Chalcicl species
knovrn to at.tack galls of A. hollori.
Tho tirne f:rctor, discussecl bck.rlv, is :rlone insufficicnt to
account for all thers-" rliffercnces.
(iii) The food lrebs in gal1s of corresponrling generations of
close'ly allied species are often s;inrilar'.
For example, those in galls of Aqnips dit,isa and C. longiuentris
are alrnost identical, and in the spangle galls of Neu,roterus
querctts-baccurTl,trl f. le.ntictilurls (OIir'.), \7. nunisynalis (Fourc.)
and N. albiTtes f. loe,--ittscuh^rs Schenck they are very similar.
Since only very feu, species ol parasites arer in flight continuoush.'
frorn early srpring to latc antnmn, it is obvious that the tirne at
r'r,hich the gall is on the tree deterrnines, to a large extcnt, whiclr
par:rsites may attack it. The sirnilarities in parasite complernents
in galls _of corresponcling generations' of altied species are partly
rhre t.o the galls concernerl hcing contornPoraneoue,
(a)
(.lutlti t(b)
LcoI
galls
tlull s
Nrrrrrbcr of ls ox:rrninotl 59
4l
63
l)c(:rcs I)t cs0ll
(inruratrrlo si;agcs)
,\i. qrrergtrs-L e.( cu.t 1t.rtt.
S 11 n e r g rt s 11 ol.h r.e- 1tr nr.i ! r t r ttr i ;
or /t(]r't0sl.ls
()l.t1n.r: tLrstrntes
'1'ot yttt,tts*'
'1,) rt t' t 1t o nt o. lt r r r n.n ir a n tr i s
ll eso polob us ti lti.al.i s
trt j trr tu, d ,t s
9{)
"''l'ht: 'l'rtrytrtrrs s1x,cir,s u'ct'o tnostlv rritttrttts, brrt 2 lrtlrrlts of ttigrit:ttrrt.i.s
clt'r'olr.rpotl f tont Iitt vll' forttrd itr lt'ali glr,Jls.
'I':rblc I gives the results of anah,scs of tlre contents of g:r,lls
ol N. quercus-baccuru.rn. lrorn leavcs and frorrr catkins. Therc art:
hoth cllantitative and qualitat,ive diflercnces betu'een the contents
of catkin und le:rf galls, rrrost obvious of v'hich are:
l. TIre greater survival of the gall maker in catkin g:rlls.
2. Tlrc largr,r number oI SrlnerElzs in leaf gaIls.
3. Tlro absencc of llunltorrtct brunvr,iuenflis trntl Mesopolobus
species {rorn catkin galls.
Wliern on lcaves, galls of Neut'oterus querclts-baccet"ttltt. slro'vr.' i't
sinrilarity in folrn to thos:e ttl Cqnips cluercus-folii,-which apperrt
on the leavc.s lat.er in the year. The {ood rvebs in ieaf g:rlls of
these t'lvo species slrorv parallels in the importance of llut'qtomo
brunrtiuentris and in the presence of T'orEm,us nigricornis Bolt.
ti 24
I0!J
34
8
15
12
lt
1)
;J
rJl
t2
:]
7
2\l
t7
6

2.1(i [Nt-rveurbt,r,
'fhe 1;:r.r'as;ite ccluplelrcnts of sinril:tr g.r,lls oI spccios belonging
to diflerent genera are rnore alike than tltose of galls of congeneric
species when these g:rlls rlifler in position or forrn.
Tire food webs in g:rlls oI Cqnips diuisa antl A. ktngit-,entrfs aro
sirnilar to Lhe food rveb in the gall of Atriricus ostreus (Htg.).
Theso three galls are :rll contcrnpor:ineolls, hard and globular, anrl
sitrratecl on the lower surf:rces of leaves. The gall of A. ostretts
slrares all its six r:halcid paras,ites rvith C. rimrisa and C.lonqiL,entri:;
galls, :rnd is rmrch closer, in this respect. to these CtTrzlps galls
tlrnn it is to other Andticus gzrlls, for examplc those of A. soliturius
(Fonsrc.), A.lecunclutor (Htg.) anrl ,4. callidornu (Htg.), r,r,ith which
it is conternporaneolls but tvhicir ar'e of differcnt {ornr zrntl posit on
on tire tree.
Tlre gall of And,'ricus ltolkr.ri is another harrl, globular gall, anrl
its parasite complernent has trruch in corlrnon r.vith those oI gali:r
of A. ostt'etts, Cyri.ps dtrrisa ancl C.lonqirentris, but it develops
frorn a bud ancl this positional differencc) accounts for the fact that
Mesopolobus fasciiuerfit'is, rvhich has beern found to attack only
leaf galls (.Askew, l96ll;), is npparentlv abscnt from tlie
gall of A. ltollu'ri nlthorrgh :rlrunrlant in galls of the other tlrrce
$pecles.
Poi;ition is also an irnportant factor in host. gzrll selection by
'1.'<ntlrrtus ytobilis Boh., 7. erucarum, (Schrank), 'f'. annoertus Bolt.
:rnd 7'. pletuul,is Thorns. r,r,]rich attack only galls growing on ttrrc
oak boles or roots.
Galls of Anilrit:us ostreus f. lurunoultts (Bev.) an(l /. l;oll.uri l.
circuktns slrarc tlteir thlcc p:rrasites, Mesopolobus fusc.ipes, nI.
tibirilis ancl 1111. runthocct'us; the galts are contornpolaneous, very
sirnil:rr in form, :lnd both are situ:rtecl deep :unong bud sc:rles.
I-lre fact that..1. kolktri f. circulans galls only Quercus cenis L.
w'hile tlre gzrlls of A. astreus I. lurunculus wcre taken lrorn Quercus
robzr woulcl seern of minor irrrportance to the par:tsites.
ft rvould therefore appoar that, the f:rctors lvhich determino
the parasitt' corrrplerncnt of a gall nrc:
l. lfho tirne o[ year during which tlrc gall is on tho tr.ee antl
contains avail:rblo larvzte or pupao.
2. Tho position on the tree lvhiclr the g:rll occrrpies.
3. The forrn of the gall, particularly its size.
Using this inforrnation it shorrkl be possible to pretlict which
lrarasites zrttack the galls of spccies of oak Cynipidae not yet
investigated.
TIre gall of. Andricus nucl,us Adler is a srnall, ovoid, spring gall
grovring on the male catkins. and :rs such is probably-attackecl
by ltlesopolobus runthacerzs, u,hich also attacks the similar catkin
galls of Andricus fecutrdalar f. Ttilonts Adler.
_ Tlrc gall of Andticus nu,dus f . molpillhii (Adler) is about 4 mm.
long, spindlc-shape('l and thin-l.valled, and grows in axillary buds
in Septernber zrncl October (Connold, 1908). It may be eipectecl
b t tlrt t'i-
247
sirnilar
colluris
in th<:
.btr.s
ntris,
If the
species
specles
Icv'er
thzrt no
at corrr-
srnall
in their
causecl
frorrr
attract
of
tcs
to
COIII-
\['as1)
Cqniy;s
lls
detailed
lvork
winter,
o:r,k (Q.
the g:rll
the
singie
1\la-v.
contents
nlatn
also
figurer.
trndant
lIl
to
galls,

NUMoER OF GALLS PER SAMPLE
50 62 78 95 81 40 68 AA 19 20 363 ro 2
60
.r !)61 1
to be:rttar:liecl b1, titt, stttttt, prtt'rrsilr
l-rrtrl g:rlls of Andricus xslitrttitrs (l'r't
(Htg.) arrrl A. ccillidcnnu (Hlg.). slrir
yoiu. A. solitu'ritts galls rtrc krro\r'n lo
.irtcttttd'us (\Vrilk.), flL tnnuert.tts (\\' tl
rtrt.Lris, A. t:ullitionut. galls lry Xl . jtt,,t
nnrl ,4. c'urrutot'f. collaris gnlls lr.1' ,l/.
The partrsites of thc gall \y:lsl)s r.rr
corrtents of each typc of gall u't.r'r. rrl
of parasites in tho sitnrc pr'ol)()r.liorrs,
rvorild be it-i cr.rrnpetiti<rn lor 1lrr, :rt
lrr,t'atrst'anf inrproventeltt in orrl I
parasites to attack its contcrrt,s. rvor
r-rther gall rvasps in tlrc sarrrr irlr,rr.
would fit the thirrl casc of \\rilliirrrrlo
trvo species of gall rvasrps have i<k.rrt
petit,ion of this sort l)et$'ecn gtll u,rr:
diflerences betr'veen g;rlls ale r'csl)olrl.
parasitc corlrpk)lrients (uide bekru').
b1, gall \{,'asps r.vhich all derivo tlrcir.
thc o:lk onables a lurrge nlrmt)er' ol' (
persist, togetlrer in the s:rnrc localit r,.
The Iorrnation oI rliflclent 11 1rr's
generations of rnost specics of girll rr':
rliflerent paraFrite compiements to r.r
petition ancl probably err:rbles a gr'
species to co-exist. 'Ihe significarrr.r' r'
species is rrrentionerl below.
Sequence of Parasitism in Cy
Tlre galls of Cqnips dzoisc rv<,rr. r
stucly of a rjitll cornrn(rnity bec:ius(, (rl'
u'its dorle in the riulirrlrer of l95fl, rrr
in a mixetl plant:r,tiorr (OornJnr'lrrrr
robur) planted in 1949 in lVythrrrrr
population rvero cxamined 'rveeki.i, r.r
first, two w(reks of Arrgust., fronr llJl
galls lrere lirst observerl) to tlrt, t'rr
sirrrrples rl'ere collected in Octolru', i
After collc.ction 1116r galls wer() ( )
identified. The ciranging pattt'rrr r
lanal charnbr,r is slrou,n in fig. 5.
cells of Syrtergu,s ytallicotniLs I{1g.
t:xarninecl. brrt thesc, rcsults arl r r
Parasitism in these cells u,:x,l forrnrl I
\ that in the main ?o11. Tlrc fliglrt lrc
parasiites of dittisa gzrlir. conipilorl l'r'
W1-tham Wood tlrroughout 1958 :rn'
\Vithin a l'erv clavs; of tlie lilsl
TORYMUS CINGULATUS 20
or NIGRICORNIS
MESOPOLOBUS 20
FASC IIYENTRIS-JUCUNDUS o
SYNERGUS
NERVOSUS ol
TORYMUS AURATUS ,3
SYNTOMASPIS CYANEA IO
50l.-l ro
ul
o
J
z
o
UJ
J
o
a
o
v)
J
J
u
Z
z
a
F
F
z
E'
tu
d,
CL
t!
d.
lrl
u
F
z
U
U
d.
r!
o-
EURYTOMA
BRUNNIVENTRIS
70
EMPTY, ETC.
EMERGED
(uirDrilrrflto)
JUN JUL AUC SEP OCI DEC MAR MAY
#1958
-1959,-..."--.._
Fig. 5.-Tho chartging ('ont{'lrls of thc nrain cclls of C11ni1ts rlltisrr gulls Ir'orrr soorr aftt'r'tlrL,ir apgrr:ar.ance i1
Jurrr. l1)i3 to tlrr' follorr'irrs Jlal'.
CYNIPS DIVISA

r): re6l | 247
to be :ltt:rckerl by tlrc s:rnre prrrasites which :rtt.ack the-. sirnilar
lrrrrl galls of ,Tndyicus solitorius (Fonsc.), A. cut'L,etor f. t:ollari,s
(Htg.) and A. cti.liclonttr (Iltg.), .w.hich irlso devclop late in thc
yeal'. 1. sctlitut'ius g,alls are kr,orvn to be attacked by il.leso7-,ol.obtt.s
juctttrclus (Walk.). A,I. utnaenus (\Valk.) antl Euttltc,ntu lniitu:i-
t:tntris, A. culLidomu, g:rlls by XI. juc:undtrc ancl Il . brurtniren!,r'is,
irncl !. curtut.ot'f. collaris galls by M. antuenus.
The parzrsites of thc g:rll wasps callse lrmclr rrort:rlity. If tire
corrtents o[ each t1'pc of gall n'erc attacked by the sarne species
of parasites irr tlrc sanlc plolrortions. tlren the gnll wasp species
worrkl be in r:oinpetition -[or the ayoidance ol these parasites
trc-cause any inrprovernent in one typc of gall, causing fewcr
parasites to attack its contents. r,r.ould be to tlre detriment oI
other g:rl1 w:rslls in tlrt, silltre 2lrefl. Ttris t5,pe of comltetition
vr-ould lit tlrr, third cisr(:) of \\rilliaurson (1957). The fact that no
two species of gall rvzrsps have identical galls errsures tJrat corn-
petition of this sort lrt,trveen gall rvnsps ii reduced. Jlven strall
difl'erences betryeen galls are responsible for differences iu their
parasite cotnplcrr'ents (.-ide belorv). The varictv of galls caused
by gall wasps'ivJrich all derive thc.ir nourishrnent :ls lar,vac from
the oak enables a large nrrrnbc.r of otherwise sitnilar spccies to
persist togetlrer in ther szrtrc locality,.
Tlte Iortration of r.liflil.ent types o[ galls by thr: a]ter.nate
ger:erations of rnost specics of gall wasps, r.vhich therefore attract
tlifleront parasite complements to cach, furthor decreases com-
pr'tition ancl probably enzrbles a greatel nurnht'r' of gall waslr
species to co-exist. 'Ihe significance of tlris r.vith respect to C,yrtiTts
species is rnentioned below.
Sequence of Parasitism in Cynips diviso Htg. Galls
The galls of Cyrtilts drr.,isn rvc.re sclccted fol a rnore rletailerl
strrriy oI a gall conrrnrrnity bcc:rrrse of their abundirnce. The rvork
s,:rsi done in tlrc sunllner of 1958, and in the succeeding rvinter.,
in a rrrixe<l plantaticrn (Cornp:rr'tnrent 25) ccrntlining oak (Q.
robur) plantcd in 1949 in \\/ytlraur \\roorl. Snrnp)t's of tlre gall
poprrlation were cxarninecl u,cel'i1., excopt for a break during the
first two rveeks of August, frorrr l8th Junc (thc date on u,hich
galls rvere {irst obsen-c.tl) to tlrt: end <rf So.ptcrlber, arrtl singk:
suruplcs u,t,r'e trollecterl in October, Decernhcl, March trud Nlzr-v.
Alter collection thc, g:rlls \I:ere opcned and their contenls
identified. The ciranging pattcrn of inha,bitants of the rnain
larval charnber is shorvn in fig. 5. Inhabitants crf the inquilinc
cc.lls of S11rr,e'rgus Ttalljt orrris Htg. in the gzrll u,all rvr:r.e also
erxa.rnined, brrt tJrese ros-.ults arL: not inclrrdecl in the figure.
Parasitism in those cells w:rs {ountl to follo.u'a sirnil.ar pattcin to
that in the m:rin cell. Tirc fliglrt peric-rrls of the nrdst abundant
parasites c-rf rlir.'fsa. galir:, r:orrrpilorl frorn rveeklt ficld captures ir-r
Wytharn Wood tlrroughout 1958 are shown in fig. 6.
Witlin a fer.v days of tho first appearance of dit.,isa galls,
sl
T
si
r:1,
tt
(r
it,
OI
itr
ol
ft'
Ll
Ie
gr
sr
tr
..r1
,lr,
AI
oi
c1
t:i,
i;i I
1'l
w
ra
th
I.
2.
t
1,,
In
gr
br
ga
lo
in

:2.18 | Noverrrb,:r
Eut'ytotrttt. brutt,rrit:<'ttttts harl 1r:,trasritist:rl :rborrt 20',. ot'the galls.
Totqntus oltiul.rts rv:ts also 1)x(.!rer)t in tlre gnlls at a vety earlv
s:tagc of tlrcir citr-r'loi)rrlcnt. fitlrrtcitnuspis tllotlce (Boh.), r,l.hich
is ir specific parariti cr{ thc qnli nraking larvae, apl)eare(l in tirt:
grllls a fetr. riat'sr latci', tlrtts a\'oi({illg tlre iliti:rl heirvy zrttack oI
L' u t yl t,ttttt lu,, tr ti i t'c tt I ris.
Tlre effocts of 1-.lu'asitir.trtby ll'c;rtlttitrs auytLttis ancl SLltLtornus';is
cuatrce rnake trlr. intt-re:ting cL)lril)irrisolt. At tlre tinrc of ovi-
lrcrsit.ion, tlrt-: fr,rrrirle'l'. ut;tutrrs pillalys()s thc first instar dir,isa
lillva, rvliich i"s tlre,n crrten l--y tirc plu'asitc larva. The death of
tbe C1tniy:s lllrv:r ies;nlts in a ce!:s,:1tion oI gall gl.o\,yt,]r, the gall
rerr)aining airorrl 2-3 mm. in rlianurt.er. In coltiast, the {emale
S. ctlnrteu, clocsr not peralyse the 1'oung Ctlnips la.rva, and both
lzrrr-:re feed togctiic.r'for r,orrrc tirrre on the 6lall tissrre; tire gall
contirruersl to grou' rl<r, r't:aches tirr. sl'tnre clinrerrsions ts a gall
containirlg onl\,:,. Irt'iiitlrr- C11n.i7;s lan':r (tablc.2). The Iull5'-grc;r,r'n
gilll confers tlrr,plr;tet'tion oi a hllrl, rnodct'atol1- thick wall on its
inhal;itarrt, sho[u by'ttrre fact tLat no 1'. rnltttt?rs \1'elt: Iound in
small di?.'iso galls altcr the lirst \r'eek of July, rrll apparently
lrnving been rlt-:siroved by litter parasites, rvhile S. cqaneu in the
larger gtrlis r'r,;u,,i onlr' retlucercl to altout ono thir<l oI its original
porcentagc incriclcncc by the {ollou'irrg spririg.
T.lBr,u 2.
19611 249
b runniu entt is, S 4 n er glus ne?\: os u s a:nd Suntonl a spi s c,t1 u,rtect, u,hich
exlrlirins ils r.:tpid rerlrrctiorr irr nrrrrrLrers rlurit'rg thc flr'st feu'rveeks
oI gail gro.rvth.
Eurytotrttt ttrunn,iuentris adults u,ere in flight, cluring rnost of
the sumrrrer, but their later attacks on .liuiso galls $'ere con-
centr:r,ted on the srnaller, rriore casily pierced, thin-wailed galls
conta,ining Sqtt,erqus rleruosus or tireir ov'n species, or on tltrr
f,,
L:Pcc'|mc -(
+.> -{>-
n3
@F- <::'-e
-a+
MESO POLOBUS
FASCIIVEN TRi 5
MESOPOLOBUS
JUCUNDUS
TORYM U S
AU RA TUS
TORYMUS
NrGRtCOSNls
TORYMUS
C I NGULATUS
EURY TOMA
BRUNN IVE NTRiS
EUDECATOMA
BlGU TTATA
SYNTOMASPI S
CYAN EA
SYN E RGU S
PALLICORNIS
SYNERGUS
NERVOSUS
<> -o
€-+
->-<D>
MAY JUH JUL AUG
+C-
-..-O.---.1>
-:ft
---D=+
Flize oI Cyniy.;s dllisrz gnlls in r\ugus;t antl Septerrrber.
containing larvae of S tlni,orrut,*pis cyu.meu zrttain the
tiiatttrrter as unpalasitised gall:;. rvlrercas galls contlrining
par:trsites :r.r'rr r:tunterl.
Galls
SAIIIE
ot]rer'
(.iall rnalier'
,tr7 ii torn.tr.spis
Other palasitcs
l'otttL'ttts ol
gulls ^\' tt iri L r'r
Il:tlsil tt(l
1,1
20
lI r'un.
tlitt nttft r
(iruri.)
,\i{t ni.ti ttr rt r:,:
(l') ll-trll th.icl;ncss
\.rrr.tn.) 0.t (teIl
rri.th m co n
tLiu rttetar
0.9
I.0
0.2
4.7 + 0.15
4.7 + 0.l;i
2.5 + 0.26 (().01
A few aclrrlts c-rf tlre fir'st, gcner:rtion of lllesopolctltus juctutdus,
Tcn'tlrtttrs cingu(.atus Nt:cs rnrl 7'. ttigricrtrui.s nre in the fiekl
rluring the c:rll;' days of rli'ur.sa gtril glorvth, but the,qe species onlv
lrarasitise the contents of di'L:isct grrils Llter in the srunmer when
the galls have attained a, larger size.
Synergus neruosus I{tg. rvhicir, unlikc S. prLllicornis, clestroys
llyJ ('urtills larva, attacked the grrlls {a-irly e:rrly, but, as .lvith
Stlrttctnruspis c!/urtcu, it lnissccl the peak ol Ilurytottto, brunni-
uenl.ris attack. Galls or-iposited in b\- ^\'. nei'r,osrzs rertrzrin small
nnri thin-wallerl ou'ing to earlv tlcstrur:tiorr r,rf the gall making
IalvAe, cleath in this t.a,se bcing rluc to the rapicl growth of thl
S11trc.rott.s lilrvae u,lrich :rpparentlv crush the slorver growing
Cgyrr,ips I:trv:re.
It is tlre eall;r attack on the Cynips larvae by Eurytonta
S t,P 0cr
Irig. 6.-Thc fl ight Pcliods of tirc rnain parrsitcs ol 0yt,i7Ls d,tti.srt
ga,lls.
I
)

250 f November
similarly accessible inquiline cells of Sqnergus pallicornis in the
walls of fullv grown €ialls.
Torvalds the end of July adults of the second generations of
hle_sopol.cittrs fusciir:en,t ris and M. juutndtt er,,"rgeri from spring
galls of crther Cynipidae, and fuIesopolobus cggs and larvae
appearetl in dit:iso galls at this tinre. 1.,[. fasciirentrris, rvhich is
tlrc mosL abundtrnt of the tv'o species tn cliu.isrt galls, attacked
nrost frerqrrently Srlnerqlts ltert,oslls ot li)u.n1tott't,cr, brttnniuentris
I:rrvae, cspecially those lllunltomo larvae in tlrin-rl,alled galls that
lrad plevicxrsly att:rckecl S. rtertosas larvae. As a result of the
deprcrlations by Lleso'polo(tu,s anrl trXnrt1tornu bra,yntit:entri,s, tlte
nurnbers cf S. n,er.--osas dccreasecl during July. Il. bru,nnioentris
r,vas itself redrrced in nrunbers by Xlesopolobtts.
The first erriergences from thc-. galls occurrecl during August.
Tlrese were of Il u,nlt otnu, b ntnniu enttis, !v[ e soTtolobus fo,sciiu entris,
M. jucu,ndus and SqrLerot.Ls rleruos'us, rno:rtly frorn sm:lll galls which
lr:rd been attackrrrl in ,Iune. The rS. lt,e.tr1ost.ts elnergence .v\.':rs onlv
partial, sourc of these .S-37ler'grrs overwintorirrg as larvae to emerge
the following spring, :rncl the forn:rles of the ptrrtial autumnal
generation probablv failed to oviposi-'..
Tlrc secon(I gener:rtions of Torqntus cinElulatus and T.
n,iglticorrtis u,er"e in flight from tlre end of August, and their eggs
anrl larvae were first forrnd in dit,iso gall's zrt this tirne' I'hev
fi.rrther rcdrrced tlte nrirnbers <tf Sqrtery1us n(rt1os7.ts and Eurtltomu,
bntnni,terftris :rntl also the gall rn:lker r.nrl SyrttomusTtis cqurtet.
Tlte Tcr'11nuts species in their turn 'wele attatrked rlting Septernber
:rnd C)ctobcx bv hlesoXtttlobu,s fusciiterttt'is and ]11. iucuntlus of
the thirrl gencration, some of u,hich h:rd emerged from df'.*i.srl galls
carlier.
f,i1ilf, Qrlnips dirti.so emerged during November from 2-3/o of
galls collected at this tirr.re.
The f:rctor 'Emptv, etc.' incltrdcs all those galls which did not
contain living larvae, except those frorn r"-lrich adult insects had
ernerged. Galls rvhich fililecl to grovl' after tlieir first appearancei
presrrin:rbly owing to the very earlv de:rth of the gall rnaking
larvae frorn a causc unknown, accotlnt {or rnost of the galls
recorded as 'ernpty' in the first sample. Later in the sumrner, the
I:-lrval cell in rnanv gnlls'was found to cont:rin a fungoid growth,
and this accounts for the increase in numbers of 'emptv' galls
tlrrorrghout the period of analysis, ultirnatelv reaching 63"/" of
ga,lls examined in IIav. The uails rnost aflecteC by mould wcre
ihose of sm:lller sizc lr4rich lrad contained a, seqlrence of parasitcs
during tJre surnrner. It is possil-rle tlrat incornpletely consunred
lzrrval rernains creatccl conclitions suitable for fungoid growth.
Larvae in cells containinq nroultly lentains were therrrselves rnori-
bund, and it seems likelv that rnoulcl can calllse rnortality.
The contents of galls gathered frorn thc gronnd during l\{arc}r
:rnd Nlay cliffered in cornposition front tlre December salnple (from
clead leaves still att,aclierl to trees) onlv in having smaller nuln-
lrr,rs of MesoTxll;ltrts larvnc, :rnrl less particrrlarll' 'l'rtrtlrnus lan'ac.
19611 25t
The reduction in numbers of these specios during the winter was
probably or'ving to att,ack by mould. Chalcid parasites ignored
the fallen galls and only a few galls were destroyed on the ground
by predators. IIo'wever, srnall manunals may have eaten the
contents of the few di,risu galls found on the ground during the
winter with holes torn in their w:rlls. A captive water shrew
(Neomqs locliens Schreber) rcadily tore open galls of ditrisa (and
also k;ttyliueizlli.s and clu,ercus-folii) with which it was presented,
anrl rlevoru'ed the contents. Titrnico (Pu,mls spp.) were plobablv
responsible for the opening of a srnall nurnber of galls fountl with
Iarge holes in their walls while still on the trees.
Parasitism in galls of Cynips diviso, C. Iongiventris and
C. quercus-folii
The g:rlls ol C'yn.ips dit:isa, C.longiuentfl:s and C. clueruts-folii,,
abundant in Wytharn Woorl, were selected for an investigation
into the effects on the parasite and inquiline complernents of
small differerrces in form and in growing times of the galls. The
study was undertaken during the summer of 1958. Galls of these
tlrree specics are sirnilar in many respects, although the gall of
lonqit:entuis resembles more closely the ga,ll of di,uisa than that
ol quercus-folii.
All tlrree of these galls are globular, those of quercus-folti
being splrcrical lr,hile tlrose of tlit:tsa ztnd longittentrts axe slightly
flattenerl. Ilach g:lll, rrnle'ss attacked by Syrtergus pullicornis,
contains a singlc, central larval chamber. The surface of the gall
of diuisrt is srnooth, that of longliuentti,s bears low ridges whilst
tlr:rt of que.rcrts-folii sornetintes bears low papillae.
The galls grow on the unde: surfaces of oak leaves, attachecl
to veins, and tencl to be aggregatecl. Connold (1908) records 35
dh:iso ga7ls {rorn a single leaf, rvhile the maxirnurn mrmber for
quercu,s-folii is givon as 12 and for longixentris as 8. Correspond-
ing {igures frorn personal experience are:. diuisia 18, quercus-folii
3, and lon(tit;enttis 4.
Dates of lirst appearance of the galls were 18th June for
rliuistt,20th Jrrne for longitentris, and 2nd JuIy for quercus-folii.
tr'ig. 7 illustrates the grolr'th clrrves of the galls based on the
mean rneasurernents of samples collected in Wvtham Wood.
Tlrese include parasitised galls and are similar for dfuisa, and
lonylir;entris, growth ceasing towards the end of August at mean
diarnetcrs of about 3'2 mm. and 3'7 mm. respectively. The gall
of quercus-|ofti has a rnuch steeper growth curve since, although
the grou,ing period is of about the same duration as in t'he other
two galls, thc final sizo attaincd (mean diameter about, l0'4 mm.)
is rnuch grenter. Appearing some trvo weeks after the other two
galls, it reaches {ull size some two weeks later, towards the rrriddle
iif September.
I'he sizes of the larval charnbers (fig. 7) also show a similarity
betu'een galls of tlir:isa and longiuentris, the larval chamber of
tlrrr gn,ll of clttcrcus-foZr) increasing in volrrme at a, grea,ter rate hut,

252 fNovember.
never becorning as large as in the other trvo galls. The first, instar
lirrvae of cliuisa and /onoiucnfris oc.cupv onlr- about one-third of
tlre volurne of their larval charnbers, but ir't cluercus-lolif galls thev
occupy ne:rrly zrll of the chanrber. Tlre snrall lar,val chanrlter
enrplrasises the tlrickness of the wzrll of the gall of quet'cus-folii,
rvlrereas the larger lar.r..al chanrber o{ drri.sn and longir:entris gztlls
nra,kes thr: walls of tltese galls thinner.
19611 253
'Ihc tliick rvall oi thc ga,}l ol querats-loiii ]rorvever is soft and
rr-rong], r.r'lrilst tlre tlriturer n.aiis of dii,isrr ruri loltgir:ent,ri.s gitlls
:-;'e [ri.rr'<l ;r,nrl rvc,,or]v.
11re lrrtes rrt r.,'liicli tirc gilli rrrirliirrrl- llu'r,r',r, -.r'cle rltstlovr'tl in
tirr-'sc. tlrree grrlis arc slllrln in Iig. 8. 'I'hc cirrve l'or tLc gall of
ILruti,t:cyttiis, <'olrsirrrr.ted {rotn rcltiir-oiy li*r, s*lrples, slrorrlrl
1.:.,Lr.rl,lt b,'r'lr'.;,'t lo tlt:tl rrl r/irisir.
In all tlrree r;pecier"r, the grt.ii{,.est rlt:s{,r.u<rlirin of t.[rc Cqnips
occuls aftcr the gali riiturreter excot.'lls rr,boui :) rnnr. and before
[u]l size is:rttnilrctl" A rlialtteter of 2 nrrn. is reacirerd bv dir.iso
t"nd lctntlit.:anIi'rs gtrllii in a leu. rlir.ys, lrrrt orril'after abbrrt tlro
rveeks in quit'cus-folii galirr; ltt-forc t]ris siz.r, is leuched parasitislr
is liglrt.
'lhrcc firctoris:l,r'o resprrnsil rl<, Iin tlrc giirrlrrirl talrcring olf of
rlcstruttion oll tlLe (lrlttipr; larvae:
I. (.lall rrratlu'ation.
il'lrt: lr:rrulness ol' tlre ur;rlls ol' rrr;rtrrro r'irri.so anrl kiltqit:t:ttl.ri.s
g,alls incle,rses thc tirne a pnlasitc takes to pit'r"r:e thern with its
ovipositor'. llhc l:esistarrcer of tlrc r';ali of yiilrng d,i,t,isu, galls in
late,trrine, As nloir:rilu..(l lri- thi::rppltlatns tlr.sclilred elservhere
(Askr:u,, in lrres;s). l;r,s forurrl io lrc oirlv 5-10 s.nt. ancl at this
tirrre Iemalc llur'ry!ttttt,, ltrrttttiit'citl,'is u-,,r'i.,,|>51,1'1-161 pierr:ing tltcrn
in lcss than ir rnirrritr'. 1\iirtrrli,galls in l:r'! r- srrrrtur,.r. lrorr6'1rp1 1r,-,,1
a wall resistauc'tr oj l50-:100 sm.:trtri itt 1lrir; tirrro /r,'. ltrt rtttirerrtt.is
neederi scvclirl lr;in,.ites to rllir.e its irvipl:,ihor tlrrouglr the gall
wall, on one ()ccitsi()xr 3(i irririutcs bc,irrg t.tlrc,rr.
Alt.lrorigh thc'ilall of l irrirtnr'o q:rll oi' 11tr.r:.rcus-fol,ii, is soft, its
l,lrickness rcnrLels it irti;nsrrrble for lrrosi, (lhnlt.icls to reach the
l:rrv:rl r:;ranlbr.'r' u jtlr .l,lrt:ir ovilrositors. 'I'X:c avelage wzrll thick-
ness oI a nratrrro gail is jlisi orr:r 4 ir'l;l]., rvhiic a large female
llurq{o:ntr, brunttii'ttt,r'is lias trn ovill{)sii{}1. only aboul 2.1 ntrlr. in
leng'rlr ; fcirlr!e ,\lc,:,tl;oloi.tits lasciit.c.ti!. t.i..r arrrl lI . jucundus lta..;e
ovipcrsitirrsi rrrrrle,i'2 rnttr. 1o1lg. atrrl rinly ll'ottt:rtus nillricornis Ttas
an ovipositor osr,r:crlirrg 4 rnn. in length.
2. Atrsr-.rrce of host specificity tou.alils t1v: (-lr1mt\ts in all but tlre
o:.tllicst pillasitr:s of tlrc g:r1ls. Thil; iru're:r:;es thc degree rvith
'rvlriclr ttrre prr,rrites attacl< t-.ntlr otlrel as tilcir nrllnbers in the
galis increasc.
3. (lonccritrntiorr of attack b1. later parasites on those galls whicfi
:th'e:rd-tr r:r;ritri;t 1;:rrlsites, and as a resrrll lrar.e ltc.corne stuntcd
,.nd thin-wrLllctl lendering tlient triottr trlsill, pi:nr:traLrlr'. Tlris
alrplics lraltictrla,rly t,at dit:i:;rt alrrl /,1i1-rit.r:lfr,als galls, nnd Iltls
l-reen rliscilssr rl ir.lrove il coni:et.tirin \rii,h tlle for.rrrer.
lflre sirnilalit,l, betrveen drr.,jsrr iur<l l.on(tiuentris g:rlls with
t'c{irt{l to [t,rtri. lrrtrtlne.,',. r:rouinX iitrrls :rnll ,]lr,\\iilq,ffllcs Inakes
for::-r, simii:rliti'in loorl y,-cl-rs. Calls of both specics ha,r'e the
r,:rrrie spe':ics of llllasrl.es :llrri illluiliirt..r u:,t:trr,iillr:rl rvith thern in
\Yvtlratrr \Voorl (r-xccpt for a {t'i''. {)rtent;ci:} {:li..i.se (Walk.) rccorrlerl
I't'ont (lqni.It's r/ili.svr, gails) nlirJ qrrantilrrtivelrr also thc p:rrnsite
Io folii
o
c
u
F.
ul
5
GALLS
o = folii
x: longiv€ntris
o ' diviso
longiventris
xeX.@'
diviso o
c
xa
xx
x
a
lohgivc ntris
I
I
LARVAL CHAT."{CERS
JUNE JULY AUGUS] SEPTEMBER
Jt'ig. 7.---'l'he rates of grol,th oro gails of ('t1ni1ts tlit:isu,. C.lort,git:qntris
antl ('. rlttt't't'.trs-Jttlii, haserl on thc rrrolnrs ol s;rrrrpli,s of l0 galls oaclr.

254 INovemtrer
Oo
re6ll 255
complements are very similar and the inter-relationships almost,
identical. Galls of longir:entris are slightly less lreavily attackerl
by parasites and inquilines than are galls of diuisa and a higherr
percentage of adult Cqnips emerge. This is probabl;r due to the
rather larger diametcr ancl thicker (but not more resistant) u,all
ol tlrc l.ongiuentris gall.
The parasite and inquiline complernent of the gall of q'uerclts-
lolii cliffers frorn that of the tv'o other specics in the follor.ving
i rnportant particulars :
l. Eudeccttonut bigluttatu (Swed.) replaces Sqntonutsytis
cyuneu as an irnportant parasite attacking only the gall rnaker.
2. Sgrtergus neruoslts is less common than in di,uisa or
longit;entris galls, but Synergus pallicornis is more nrlmerolls.
3. Tory'ntus cirt,clulatus and 7'. aurctttt.s are absent frort-r galls
of quercus-folii.
4. illesoTtolobus fusciit:en fris is lerss corrtrrron in galls rfi qtrerctt,s-
loli,i tltan in galls nf thc otlrer two specit-.s.
Tlre flight periocl of T'ot11ntus ct.u'ra.tus lvas over by 2nd .Tul1,
v,lren the galls of querclts-lolii first appeared, anrl by the time
the galls had rcached a diarneter of 2 nrm. at the end of .Iuly the
flight periocls of ^Surzfom,aspis clJenea ancl Synergus nernosus
(Generation ll) were almost over.
Eudecutotna lti.guttota .was frequentlv found in galls of
quercus-folii, but it, rv:rs only onct: recort'led as a larva frorn both
diuisa and fron lortglh,enlris galls, :rlthough it rvas obscrvcd drillin,:,
into both of thesc galls in the ficld, anrl a few adult insects ll'/ere
bred from dioisrz, galls. It appoars to bc specific in tliat it attacks
only larvae of the gall makerrs. It is an endoparasite zrncl this
rnode of life u'as not di..covr:rccl until aftcr the fuliy-grou'n larvae
had emerged florn rr-.rttaills of querctts-folii larv:re in September
1958. Cynips iarvae parasitised hy this Chalcid n'ere not taken
int,o account rrrrtil tiris t-irnc in constmction of the curves of
Cunips destmction ({ig. 8). It is possible that Eudeattttnra
bil1uttata attacks galls of riioisrl and lonqitentris as {rcquentlv as
it nttacks tho-.e of c|.uercus-folii, l;ut in the latter galls its survival
is glreater owing to tlre lesserr destluc.tion of Cynips lilrvae by later
parasites.
fludecutontu biquttat,ct and Srlntornaspis cyune,o are similar in
that tlier,- attack only the ga,ll rn:rking trarvae, r'vhich they do at
an earl;. stagc of g:tll gl'orvth, as do all other host specific parasites
strrdiecl (e.g. spccies of Ol,qnr, Pediobitrs, ttntl Orntocerrzs), antl
also in not destroyin54 thtir hosts until gall gror'vtlr is cornplete.
This ensures rnarirnum protcction frorn latcr parasitt's.
As many as thirtccn fully-grown l:rrvae of S'qner91us |rul,li,cotnis
were forrnd in a quercrr,s-lol.ii ga"\\, r,r'hilst the larlrest nurnl'rer of
such larvae forrnd in n gall of dir.,isa was four. I'his is clue to
tlrer larger volunre ol tlu qttercus-folii gall wal1. Tern .9. Ttulli,contis
eggs lvere found in a gall of dir:isrt.: rnanv lalvae presumahly
perish through shortagc of space.
TIre, rcason for tlre ahscncr: ol Tonlrnrr.s cin'rvttlnltts frctnt
o
o
xa
x
a
C
o
N
t
o
a
a
a
:o
(Jo
*!
=4
.i b0
6O
'r:
tF
.:5
:,C
Ed
.o
'Ja
.O
n';
.: A
JO
o
o.5
.iP
d-
U
o
A
Er
.L
oi
e
o
o
= F.-
6 E.=
;:!
oxo e
x
x
o
od
6
E
Iai
r
&
!-
J
E
5
-
o
6
a
5
{
t
8,
e
@
oxoe
>(
s9d:i*,r'3 AHr"lY3H HflA S-l'1Vg 3DVJ'N33Uld
I
;

256 fNovcrnber:
quercus-folii galls is not knou,n. The species would not he able
to oviposit on the contents o{ the rnain larval cell oI a mtrture
gall, becarrsc thc thickness of the gall lvarll (nbout 4 rlrrr.) excee(ls
the length of tlrc ovipositol (nhout 3'7 Inrn.), but tir.o ovipositor
wcruld be long ernottgh to rcaclt colls oI Sqnu'y1tts pall,icarnis in the
gall wall.
Mesopolottus luscii'L;ettt'i'ris ]ras a prcferernce for r:rna1l .g:rlls
(Askerv, 1961b). The pcriod o{ tirrie, ruring wlriclr thc galls
ol quercus-folii rvere srnall (l'lrrt ovcr 2 rrlrn. rlizrrn<:tcr) 'r,vas of vt,ry
short duration and this prob:rbly accounts for the scarcity oI
Nt. fosciit.ten,fi'is in these galis.
Fig. 8 shorvs that tlre destmct,ion of quet clts-lolii. is
proportionatelr,- less than the destmction of tli,t:isct anrl longi-
uenttis. The late rlatrr at lvhit'h the gall reaclrr,,s 2 rrrrrr. in rliarnetcr
results in the gall escaping the heravy, e:rrly prrasitism suffered bv
cliuiso, andlctngliuentris. Thc period of gall grorvth lret,r,r,een 2 mrn.
rliarrreter arrrl bc.fore full sizc is reacherl, during rvhictrr the gall is
rnost liablc to bc attacl<crl, is of sliort duration.
Thc gails of the sexual gencrations of thesc t}uee Cllnips
species :tre all early spring galls. lfhose of lon.(liuen.t:ris f .
substitttta nncl quetcus-lolii f. tasch,enbergi jn arlventitious buds
on the bole, are alrnost indistinguishnble pubesccnt. ovoirl galls
aborrt,3 rrrm. long. but those of di,t:isct. f . o-enttt'.osu, arc clr:rractelisecl
b5, ircing lnrger (nbout 4 rnrn. long), :rlmost rlevoicl of hairs, anrl
sitrratcrl cn )roung lcavcs or shoots. ps11, galls ol dit:iso f.
Llerru.caso rvere forrncl, blrl b1, an:rlogl' u,itlr the factors governing
para:ritisn in otLer g:rils, it r.voukl he experterl that the food r'vebs
in lonqirenttis t. substitrrfa anrl (frcr'c1ts-loli,i f. tcLschenbergi galls
worrlrl be verv sinrilar, and diffcr lronl thosc in galls of diuisct
f . r:errucosa. No p:rrasites or inqtrilines wc.re founrl associaterl
rvitlr rii'irlsa f . rerrucosu ealls, but l[esopolobus fuscipes anrl
Synergus gallae-pontifomris r'r'-ere both found in galls of the other
tr'vo species.
Galls of lonqi,t:entris nrost resernhlc those of ditisa, but in
contrast,, galls of longliuentt'is f . strbstitut u rnost rcsr:mble thoso ol
qu,ercus-lolii f . tuscttenbery1i,. If. as seenrs likclv. the populations
of tlrese thxee Cytti1ts spccies arc being controllec-l by their
parasites and inquilincs, thr:ir different palasite ancl inquiline
cornplerncnts (both gcncrat ons considered) probably enable
thcrn to persist togetlrer in the sarne arca, siincc thcy arc not, in
complete cornpetition rvith each othcr for avoidance of parasites.
!['lrere is another Britisli CutriTts specics, C. disl,icka Htg.,
r.vlrose g:rlls in both Elenerations, disfrcha and d,itticha l. indistincta
Niblett. are siruilar to thosc of di'--istt ancl dilisrz f. reyrutostt.
Frorrr tlrc Urnited data. avzrilable. thc agamic ga1I o{ disticha scerr;
to h:rve :r similar p:rr:lsite antl inquiline complentent t,o that of
d,itti.so, (fig. 3). Tlrerc appears to he nQ infolrrtation on tht'
1$61 I 2si
piLrasite antl irrc4rilinc crurtplcttrent o{ the gall ol distichu f.
i.nd'i:,tittr:.ta $'hich sirares, i1 v('ry stlong rcscmblance witir the g:tll
of rl,it,isu l. r,etrurt;stt lNiblcl1, 1948). It rnigirt be expcctnd thal
thost' trvo Ct1ni.1ts 1-,i;t',cies rvortltl conrpt-'te Ior avoiclance of tlte samcr
parasite spccies. Holvcvcr, tlrey lritve rathcr tliflerent gcographica.l
l'anges Ler,luse aistir:hu i:, tttost :ibttndant ttn Querttus ytetruea.
tr,iebi. anrl dirrisl orr t). rottur; Q. pel'raeu, is the dorninant oak irr
tlre rr<rrtlr nncl uesit of the tsiitish Isles, Q. rctbur in thc sciuth and
cast.
Key to the Chalcidoidea associated with Oak Galls in Britain
No singlc key to the ideltifica t,ion of the 44 species of
Lhalcithidea {onncl to be assocriatcd r,vith oak galls in Britain has
hitlicrto Lrccn publishecl, and it is to fili this gap in our
entornological lit,clature that thc following is presented. More
sp::cies prol-r:rblv at,a-i', lletertion, an<l :r,lthough the author feels
confident that these r'vill bo {evr in nutnber, this fzrctor should be
borne in rnirrrl lvhel using tlie key. Sotie of the characters used
in tho ke.y liave been t,aken, after confirrr:Ition, frorn other
airthcrs, zrnrl rederences in the key indicat'e rvorks rvhich deal
rnore fullv r,l,ith thc, genus llndcr considelation.
'Ihc liost gall list of sorne of the species 'r,r,,ill doubl,less be
extcnded. Key to species
A(I3) Tarsi 5 sogmented ............ (-l
I3(A) I'a,rsi 4 segrnonted ...... EULorrrTDAL
I Scutellurir rvit]r a pair of clistinct sutrrriorlian, longituclinal
grooved lincs; rncsolscutum rvith a Iaint, rnedian longi-
tridinal grooved line (Tetrustichus ({ig. 16)) 2
Tlrorax rvithout or occasionally (Olynr ursanles) wiLh
very faint glooYed lines . 3
Bnrlv black; only:r single line of hairs on each sidc of
rrricl-lohe of rncsoscutlrltr ... 1'el,rostir:hus uethiops (Zett.)
(ex g:rlls ,1n,tlt'ictts ostreus, Cynips clir:iscr, C. lonqi-
t:crr.tris, C. querctts-folii, Neuroterus oltsipes, l/.
t ntn'tisnto,lis l. u esicator)
Body cla,r'k Lironz.e-green; soveral icingittrclinal lines oI
hairs ovr each sitk: of uid-lobe of rnesoscutllln ...
1'. diay;h,cttttir,s \Yzrlk. (- terrninahs Thorns.)
(ex galls lliorltizu pallida\
Scrrtellulr with one pair of long bristles; octriput stronglv
rnarginecl (['ediclt[us (ilg. t5)) (''-id: also Grahau 1959) ...
1
Scutclhrrn rvitlr trvo pairs of long blistlcs; occiput not
marginLrd (Ol'ynr (fig. t7)) (uide ak.'o Crairarn, 1959;
Askr:r'v, 1959a) ...... 5
Nlala.r space alrnost tibsent; tarsi mainly rvhite .........
l'ediobius clifo (Walk.)
(ex galls Neuroterus a,lbipes f, laeuiusc'ulu$
2
1

:58 lNovcrrtl.rt,r'
ll
t2
l3 l4
l,'igs. 9-14.--\1, ()rntrtrc,us ret.nrt!is \\',lli., f.rrr.lt,: 1t), .llrxrtpttloi,tr.s
jtrt tr ttdus (\\'tlk.;. fc:nalt; 1l. .l/ jtrt.urr.dus, ltca{ irr f r.rtlt yieg-; 12.
(ttrrttq,ci.s rliuiso (\\rtlk.), hcad in florrL ,';,111.1 113, Aecitlostibu Ltu,rrtlrt,::tr
(llntz.). foleu',rrg oi ferrralo (hails r>rrittcd);14, IlttltItyrt stcnonotu
(llatz ), forervirrg of fernalt: (hails ornitted).
i
oo
oo
196r I 2bt)
- [1:rllr sparcc irt ie:rst, ecguri to rrrelxirnrul ]tre:xlth of
lla,geilurrr: tarsi trsuzrlly {rrscous F. hlsis (\\tzrtk.)
(ex galls Neuroteru,s 'nunTis?rtalis, N. ulbipcs f. laeuius-
ct.tl.ttr)
.i> Spet:ttlrrrn cios:t'rl l;elorv L,ry rr line of hairs . ....... ti
Specr.rlrrrn open below ...... 7
6 fi'irst {rinicie s,ogrricnt abcut equ:tl in lcngth to pedicel;
scutr-.lhun contrast,ing in colour with rnesosclt..irn; fore-
u,ing oI fenrale usu:r,lly ll,ith one fusc.ous spot ..
Ol ut uv et"seln es (\\ralk.)
(ex grrlls Neu,rotetus quei r,rLs-ttitccut'tu'rl, N. ;t:t..ttis.lrtal'is
f. te.sir:iti;or, N. albi,pes, Andrictts Lluudrilinc.ut,us, A.
cltrt) otor, A. i nfkrt,or)
- l-irst frrnicle scqrlent longel than pedicel; scutellurn not
conttasling in colour rvitir tnesoscutrun; folerving of
fe.ira,le lls'.;ailv lrvitline ...... O. skiatru:<t:; (lt,.rtz.)
(cx galls Ricrhizu puLt,idu)
7 }iirlclle tibiae with a brown line orr flexor aspect.........
O. gallorunt f. prrlclu'rr l\{:ryr'
(cx g.rils N e u i'tst t: r't.ts que. rc. u s -b itccu )' utlt f . le t tt. i t: ul uris)
l\'lliclrile tibiao rvitlr no snch lrlorvn line ... ......... 8
8 Srnall :rpercies (body length l'1-2'2 mm.) ; tibiae pale
r,-cllow ...... O. gu,llrtrwt (L.)
(ex galls,lndrictts qttct'cus-t'(n)tztli, -4. cLo'L'ut(n')
- Larger specries ......... ... I
I F3od.y length 2'3-3'4 rnnr.; tibiae golden yellolv; lnarginal
vein rrsrrillly lcss tllan tr,vic'e as krng as st,i.grn:rl vcin ...
O. cur:dorescfurrs (\\rzrik.)
(ex grrlls Atrclrictrs quucl t'ilirx:ut us)
I3ody lengttrr trsrrall1, greater thzln 3'5 mm.; tibiae p:r,le
vrrllou,; rrlalginal yein rnorc than twice_as long as stig-mal
vern ,.. ... o. triline(rlo Mayr
(cx ga.)ls ,lndticrrs lecundutor)
lfunic]e 7-segrrrcnted ...... E
Ftrnicle 4-, 5- or O-seginented ...... G
llind coxtr cnl:rrged. at leasb about trvice as long as rniddle
r.oxa; i{ rxft (fuIc.gu.stigrn.ris), tlren thor'ax and held rnarkerl
rvith 5,r-.llou' anrl forerving u'ith a distinct fuscous clourl
abdut eularged stigma; stigrnnl vein very sholt. rnrrcii
r;holtcr than post-rnarginal vein; ovipositol strongly
t xscrted anrl unicolor'ous . ...... Totir,lrrDAll
1 Hirlrl croxrl less tlrtrn tn-ice iis ]ong as rlicldle coxa
(Mt:ottstiorn.us (Iig. 24)) Q:ide also }Iilliron, 19{9) ...... 2
Ilincl coxa :rt loasb trvicc :r,s long as rniddle coxa ......... 3
2 S1;iqrna rnoro or less circulnr; ovipositor shoaths about
eqrrni irr lrr-rgth to head antl tltorax togetirer; srnaller
specit,s, length of lrcrdv (less crvipositor) seldorn exceerl-
ing fi rrrm. ...... Lllegosticlntus do.rscLlis (F.)
(ex galls And,rictls kollari, A. in'ftu,tor, Biorhiztt gtallitlu.,
C ul Li,rh ry t i s glontl i u.nt,, C y tti 1ts quer cus -f olii)
[](D)
i) (0)
U(F')
ti
t\,
9
I
l,
l0

260 lNovolrLtrt'
Stigrn:r elongute, rnole t-hitn l,r'r'ice :rs long as lrroad;
oviprtsitor sltr:atltlr ne:rrl1, tu,ice as long as he:rd and
tlror:rx i,op,;et,hcr'; Iirlgcr species, lcngth bod5, (les-.r
ovipositor') rlrruilil)'irJroul 6-8 rnrn. ... ill. sti,gmuti.zrnrs (I,'.)
(cx galls Alr.dri,cu:: ful!cri)
il Scutcllirm u,ith transverse fullorv rnalking off postcrior,
ligtrrtly sculpLirrcrl, :rrrc:r, (irenur:ln) (Sytftomuspis ({ig. 25))
('uidr: nlso Arl<rxv, I9{itra) ....... l0
- Scuterllurvr rvit,iiorr'1, t.t'arnsvt:t'se fttrrorv and more or less
rrnilorrrrly sculptrrrod (1'trytttrts (tig. ZS)) (uidc zrlso
IIoffmcycr, 193 1) . ......4
4 Lengtlr o-t innr:r' spttl oi Lilrrl tilria rrtuclt gleatcr th:rn
rvirltlr oI apcx o-[ lrinrl i,ibirt; borl1. predomirrantly bright
grecn -in cclour' ........ ll'oryrrnts utt|uttts (Ir'ou}c.)
(ox grrlls rVcu;'oJ.u'rrs tibiy;cs f. luetiusculrz..s, Ar. ulbiy;es,
ItI. nl.oriis*lrrli.';, N. n.u^titisrnulis f. ucsical,or, N. querczr.s-
ttuccrt r unt , N. 11 ut;t'r:us-ltoccuru?n l. lettticularis, A'.
tricrir; r, Llior l t.i:u. 1 t;t tl.idc,,ln d ricus curl) ato r, A. irt l l,u,tor,
A. quert:us-r'il;r'rull, .l" sit;nittrltio'nis, t . t1ltttdriLineutus,
Cqni'pa cl,it,isu, C. lori1;yirerttti.s, C. disticlrct)
Length of inncr spul o{ lrinrl tibia usually distinctly less
llrarr rlirll-[r. (,litl rJi,l' Iritt,t lil-rirr..jf irborrl tlre surn(' (s(]rrr('
il'. n r,,",lfii\fr.ffi,r Ir,,iI pr*St'€1!*H€;a+11' rer]dislr pr+r?}r.
i.*.rrr{r+s, ...(...ts.i"./s.s€t:ss. ......... ....... ;
5 IJorsal edgc o{ irinrl coxa rvith rnanY u'liite haits
proxinillil'; botly prertlornirtantly ltright, green ......... 6
. - I)orsrl erlge ol lrintl coxa, 'r,ith at rnost I or 2 hairri
proxirnallv; btidy dallior', ttsunlly'w'ith rcdclish plrrple or'
('1'. 7l.curo.!it;) lrronze piroon l'efloctions .. ..... ..... 7
(i Forrlalc s,itlr ovilrositor r;hcath nezrrly alll'itr,s shortc,'
tlrrrn irody (lterrld, tlrcl'ax, :r,Lrtkrrncn t,ogetltcr), a.xd gastcr
rvith. testaceous brnrl 1*rs*11r, '1'. cirtgJu.lattts Nees
(ex galls Or1tri1.;s il.irixt, ii. l,oit.lliuertt'ris, Ili,tsrhiztr, yNillidtt,
Arrrlrir:tu: ktiluri, .1. curL:tr{,or, A. ostreus)
- }-truralc rvitlr ov-ilrositor ulre:rth lrcarly :r,lu.a-r's ltinger, anrl
gastcr ll,itlrout testat'eous bnntl basrllly .....
'l'. rti.,r1t'i c ot' rris Roh.
(ex galls OtptiTts rlit:isu, C. lortl1irenl,ris, A. tlttc.r'cus-folii,
Ilio rLri.:o 1 Nil Li tlu,,1 rt.d, rit: trs l,:olh L ri, N etu'otct' us qTtercus-
(ttu:t:ttrunt\
lI:rrrr rrntrblc to separ'ate rvil,lr cr,r:t:rin1,)'the rrtalr:s ol r:inyltirttrts
antl rigrit:orrtis )
? Scape r.x-tcntling rrt le:rst to top of vertex ........4'........ 8
- $r.apc blirely r'r,aching nrcrliatr ocellirs ........4. .:"-..... g
3 Ovi1,,,silor' ;,lrc,rtlrs rlir,l irtr'l ir l,,tr,;*r llrrrn lrorlr':&:rslo
rvith tr:sta<'eous J.r,,nti l-':rsrrllvl. .. l!'. cruxotu rzl (Sclrrank)
(ex rl,rlls ..lt,tl,ictr" ,1',rr,",'-,'i,1:'1."1 \-
Ovi1r,,',iJtry.sllt,rtllrs rlisl irrclli slrr,t'ter llran borl..':'fs.rstcr
rtlrt',,,,,f1fiitf,'.,,',, lr;rrrrl l,,i=,r11rJ......... 7'. nnl,i/,-l B,,lr.
1,,x g,rlll llio,!ri:,t lnllirltr l'. ,il,ic,a, .4t,cl ,it'ir.' tl:!t,('tti;
radicis)
re6l l26t
I Anterior triangular, lmsculptured part of epimeron large
and broader rnediallv than tlre posterior rounded,
alutaceous part; ovipositor sbeaths :rbout ertrual in
lcngth to thorax:rnd abdorncn together
(ex ga]rs Attcrr.ictts quercus-rttdicis) 'l'' Ttleuruli's Thoms'
l6
f igs. 15-17.--15, Petliobi.irs /ysrs (Walk.), fernrle; tr6, '1'etrasticltrt.s
rl.i.tr.plurn,ttts Waik., femalel 17, ()l,ynt: utsa,mes (Walk.), ft,nralt..
t5
il
t,,l
,i' ,, i
\l 1,,
t7

262 [November
- Anterior triangular, unsculptured part of epimeron
small and nArrolver rrretlially thzrn the posterior rounded,
trlutaceotrs part rvhich is inflated; ovipositor sheaths
rnuch sholter thln thorax t1r)d rlbdolnen together ......
'f'. ant,oenus Bolr.
(exgalis@
l0 Eyes largc, in lront:rl vier.v rvidth of cye greater than
half dist:rncc tretlveen inner horders oI eyes; frons vcry
concave; (ovipositor sherattrs aborrt as long as lread and
tlrorzrx together') ... .. . S.tptt<tttrus1tis crltuteo (Boh.)
(ex galls Ctlnips diuisa., A. Iongire.ntri,s, C. L1uercus-folii)
Eyes norrnal, in frontal vierv width of cyt, not greater
than half dista.nce betrveen inner borclers of eyes; frons
lessconcave .......1I
I I Head in dorsal view rnore transverse. post-orbital region
narrow with width less than lrrrlf transver.sc dizuneter of
an eye; rotxtst species; body bronzerl; (ovipositor
sheaths shorter tl'ran hcad anrl thorax together)
S. lastuoscr (Boh.)
(ex gails Trigonuspis meglaptera)
- Fleatl in dolsal view less transverse, post-orhital region
bro:rcler with width greatcr than half transverse
cliarneter of an eye; less robrrst species; bocly pre-
tlonrinantlv green .. ....... Lz
J2 C)r,ipositor slreaths at least, a,lrorrt, tirree tiures length of
head and thorax together; basal cell of male forewing
hairy; stigrna not edged by infuscation ....
S. upical,is (Walk.)
(ex galls Biorhi:o Tto.llida)
- - Ovipositor sheatlrs less than l'5 tirncs length of lread and
tliorax together; basal cell of nrale forerving very
sparsely hairr'; stigrtra eclgetl by a slight infirscittion ...
S. notata (lValk.)
(ex galls Andricus cu'ruator)
F(fi) Hind coxae snlall, only slightly longer than mirldle coxae'
thorax ancl heacl ,-,,,f ,rr*ik"ii rl.itir yellorv ancl forewin!
witlrout a fuscous trloucl about stignrtr; stigmal vein zrbout
equal in iengtlr to post-marginal vein; ovipositor less
strongly exserted and rvitir 21 llre(lian vellou' band .........
lluPEI,MIDAIi
One species (figs. 18, 19) ... EuTtelmtts ut'ozonzs Dalman
(ex galls Neuroteru.s tluerc'us-boccat\nrl, CyrtiT2s diuisa,
(. lonslosvltris, .4ndt'icus kollari, A. alboTturtt;tatus, A.
t:ttrrator, A. ost)'€?{s, A. l:estuceiTtes l. sieboldi, A.
fecu:tdator, Riorhiza pallirla)
C (H) Hinrt coxae rnuch enlarged, nt,arly 4 tirncs length of middle
coxae; gaster strongly sculptrrred; stigmal vein very
short . .... ori.MYRrDAE
One species (fig. 26) Orm,lyus punctioer Westw.
(ex galls Biorhizu pallicla, oak trr,igs)
le6ll 263
H(G) TIind coxae not enlarged, at most about twice length of
ruiclclle coxaei gaster never .strongly sculptured; stigmal
vcin rvell-rleveloperl .......... I
f (,I) Pronotrun sholt autl tlzrnsversel colour rnetallic green or
bronze PTEROMALTDAT;
1 Ncrtarrlices cornplete (Orm.ocerus (ng.S)) .................. 2
- Notauliccs incornplete posteriorly ...... .......... 3
2 Forewing hl,nlinel basal cell hairy; maxillary palpi
yellou, ...... Onnocerus la,tus Walk.
(ex galls NerLroterus olbipes)
Forewing lrith tr.vo filscous clorrds; basal cell alrnost
bare; maxillary palpi fuscous 0. t.'enlclis Walk.
(ex galls Neu'oteru,s albipes)
3 Antennzre inserted about or just a,bove the ventral edges
of eyes (Mesopolobus (figs. 10, Il)) (oide also ll,osen,
1958 nnd 1961; Askerv, l96lb) ......... 4
Antrtnnae insertetl lvell trbove ventlal erlges of eyes
(fig. 12) ...... 11
4 Antennal formula 11263 5
- Antennal formula, 11353 7
5 Female with body elongate, g:rster longer than head,
tltorax and propodenrn together': colorrr predominantly
gtecn; male N,ith zrntennae unifonuly yellorv ............ 6
Fernale with bod-v less elongate, grtster slrorter than
head, thorax anrl propodeunt {ogether; colour trsually
bronze; rnale rvith antennal c,luva fuscous and rniddle
tibiae u,ith stripes and rlark clistnl spot, the latter not
borne on a projection ... Mesopdlobus tibialis (Westw.)
(c,r galls Neuroterus qu,ercu,s-(tnccorTtrn, N. tricolor, l{.
nu'ttisntalis f. uesicator, N. ulbtpes, N. albipes f. kte.t:ius-
c?rlt{s, N. numismalts, N. quercus-boccerunr f . letfiicularis,
-lndriL:us cllrL,e.tor, A. ostt'eus f. furuncultrs, A. albo-
pttrtcto,ttts, A. kollari f. circularts, Riorhizct, pal.lido)
6 Clvpeus with fairly strong median incision (fig. lI); male
midclle tibiae simple ...... ... M. jtrcundzs (Walk.)
(ex galls N eurot e nt s quercus-b ur',ctn'7t'trl, An dricu s curt u-
tor, ,1. ostre16, A. kollari, A. soltt,atius, A. callicloma,
Biorltt:o pallid<r,, Cgttips dirisLr, C. lonrlirerttris, C.
quercus-folii, C. disticfut)
- Clypeus with very srnall medinn incision; male middle
tibiae r.vith a small, triangular, dark-spotted proiection
distally ...... lI. 'fusciiuentris Westw.
(ex galls Neuroterus numisntu,lis f. uesicator, N. ulbipes,
N. albip'es f. lueuittscttlas, N. rnttnismalis, N. qu.e.rcus-
bctccurutn f. lenti.culuris, Andrictts curuot.or, A. ostreus,
(1qttip's diuisu, a. quercus-|olii, C. Iongi'--entris)
7 Pronotal collar broad, about equal in r'l,idth mediallv to
distance between lateral ocelli and orbit; (male rniddle
tibiae sirnple) ...... ... li[. amaerzus (Walk.)
(ex galls Andrictts curt:ator, A. kollari,, A. solitarius,
A. curuator f. collaris, Bi,orhil.za palltcla)

264 INovember
l9
22
2t
l'igs. 18-22. 18, lltr1tel.tn,tr.s it1o,:on,1ts D:rlm., fernalr:1 1!1, ll-'utozottns.
rrralo irr outiine; 20, ]Ntt.r'ytr,ntu ltrunniL;entris llatz., female; 21 , b.
brunnircrt,ttis, male in outlinc; 22, lludecatotna bi.gtt.ttufa (Sri'ed.),
forc'wing of fonralc (hairs onribted).
1e611 268
- Pronotal collar narrow, median width much less than
dist,ance between lateral ocelli and orbit ............... 8
8 Fernale with v:rlvulae ventrales long, about two-thirds
length of gaster; male middle t,ibiae with both coloured
stripes and a distal, triangular, dark-spotted pro-
jection ... M. ranthocerus (Thorns.)
(ex galls Andricus ostrelts f. furunculus, A. kolluri f.
circulans, A. quatlrilineotus, A. fecund,ator f. Tilosus,
Iliofir.iza pullida)
-_ Female lvith. r,alvulae ventrales shorter, at rnost only
sliglrtly longer tlian half length of gaster; rnale rniddle
tibiae not adorned as above ........... '.. I
9 Fernale tibiae uniforrnly infuscate; stigrnal vein rather
strongly curved; male middle tibiae with coloured
stripei ... M. Juscipes (Walk.)
(ex galls Neu'roterus nunTismo,lis I. uesicator, N. a'[bi1tes,
N. u,Ttrilinus, Andricus cllluatar, A. ostreu,s f. f'urunculus,
A. k,ollo,ri, f. circulans, Cvnips quercus-lolii t. taschett-
bergi,, C. lortgitentris f. substituta)
- Fenr:lle tibiae testaceous or yellow; stigrnal vein only
slightly ctrved; rnale rniddle tibiae without adorn-
rrrent . .......... lo
I0 ]fotly colour bhrish; all funicle segments longer than
bro:ld. ...... M. ctlbitarsus (\YaIk.)
(ex g:rlls Andricus curuator)
- Body colour brrinze-green; distal funicle segrnents
brozider than long .. lll. dubius (Walk.)
(ex galls Iliorhizu pallid,a, Cynips diuisa, !{euroterus
q ttercus-baccuruwl f . lenticulttris)
11 Nl:rrginal r.'ein about l'5 times length of post-rnarginal
vetn (Hobbya (fig. l4)) (uide als:o Delucchi, 1957;
Askew, 1959b) ........-..- 12
- M:rrg-inal vein at most equal in Iength t,o post-marginal
vein (fig. 13) ... .... ' ' '..... 13
12 Head in frontal view 1'5 tirnes as broad :rs higlt; antennal
scape of fcmzrle attains level of rnedian ocellus; body
colour bronze-greerr ....... . Ilobbyu kollari Askew
(ex gails Andricus kollari)
- Head in frontal view I'2 times as broad as high; antennal
scape of female not attaining level of rrledian ocelllrs;
body coloul cl:rk blue-green .. '. ..... H. sl;enonottt (R:rtz')
(ex galls Biorhiza Palliclu)
13 Olyferis rvith anterior edgc straight or only- 'slig}-tly
incilsed medially (Cecidostibu) (ttide also Delucchi. 1957;
Askr:w, 1961c) ."" i'4
Clype'Lrs v'ith anterior edge strongly inciscrl rnedially
(Caenacis) (fig. 12)
One species'............ ...... Cuenacis-diuistt' (lv1t-t<')
("* falk Cynil,ps diuistt', Andricus kollari, A' albo-
Ttunciutus, A. testaceiTtes f. sieboldi)
l8
20

266 [{overnber'
25
lrt;'
2t i
ff igs. 2;l-26. 2il,'l'orynius lul(rf lrs (lt'orrlc.). fenralo; 24, -llegustil|tttt:
d,orsulis (1,'.), lrt'ad Nrld thola\ of fornalo; 2it. Syntontuslis nt)tut(i
(\\rallr.), fcmalo scutelltrrn; 26, Ot'ttt.yt us pttn(tiUe,r \\resi,rv., ful:rle.
1e61] 261
14 First funicle segrnent shorter than pedicel; basal cell
o,[ forerving hairy in clistal tliird
C ecidostib u geganius (Walk.)
(ex galls Andricus quet'cus-rudicis)
- l'irst {uniclc segr}ent at least as long as pedicel; basal
cell oI foreu,ing bzrre or with only I or 2 hairs ......... 15
15 Olypeus with slight but distinct median incision; stigma
very large; ftons rugose C. leucopeza (Ratz.)
(ex g:rlls lJiorhizu, Txtllida)
- Olypous without nedian incision; stigma slightly
enlargod; frons lightly reticulate . .. C. semi'fuscia (\Valk.)
(ex galls lJiorh,izu Tnllido")
i(i) Prorroturn quadrate; colonr black or black and pale yellow,
nc.r,er rnetallic .... ...... IluRyroMrDAE
I Forewing with a distinct submarginal fascia (Eudectt-
tontu (fi.g. 22)) (uide also Claridge, 1959) ......... 2
- l'orewing irnrnaculate (llurytom,a)
One species (figs. 20, 21) ...
Eurytoma brunniuentris (Ratz.)
(ex galls Biorhiza pallida, Callirhytis glandium, Andricus
l.;olluri, A. ostreus, A. i,nflo,tor f. globuli, A. curuator, A.
<:allidotrta,, A. alboTru,nctatus, A. solitarius, A. testaceipes
f . sieboldi, Ollnips diuisct, C. quercus-folti, C.longiuentris,
C. distichu, Neuroterus quercus-baccaru,l't7, l{. qtr,ercus-
baccurum, .f.. lenticulu'ris, N. tricolor, N. albipes, N. numis-
tnu,lis f. uesicn"tor'\
2 Marginal vein rnore than 4 tirnes as long as wide;
stigrnal vein rrnch shorter than marginal vein
Eudecatorna biguttata (Swed.)
(ex galls Cqnips rluercus-folii, C. ditisu, C. longiuentris,
Andricus kollari)
- Margin:rl vein less than 3 times as long as wide; stigmal
vein usually longer than marginal vein ......... 3
3 Subrnarginal fascia large and,sprcading on to disc of io-r!-
wing; spines on hintl tihiae distinctly longer than width
of tibiae ...... Il.llauicollis (Walk.)
(ex oak twigs)
'- Subrnarginal fascia small; spines on hind tibiae only
rarely longer than width of tibiae ... t. uuriegctta (Curtis)
(ex galls f)iorhizu, pullida, Andricus kollari)
Acknowledgments
It is r.r,ith great plcasure that I thank Dr. M. W. R. de V.
Gr:rharrr for his lvilling assistzrnce in the identification of thc adult
Chalcidoirle:r. I rvish to thank also Prof. G. C. Yarley, Dr. M. F.
Claricige, L. 1\{. Oook, NL J. (lotton, I{.D. Eady, G. R,. Gradwell,
A. I,l. Le Gros. N{. Niblett, I(. G. Y. Smith ancl my wife for
invahrable assistance in thr: collection of galls, identification of
specin)ens or disrcussion o{ prohlerris. l\fuch of this work was
cornpleted whilst in tenure of a studentship of the Department of
\i
,l
26

268 [Noverrrber
Scientiflc and fndustrial Research at, the Hope Department of
Entornology, Oxford.
Beferences
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OhTrr:r lr'tir.stor (IIynr., lt)ulopiridae). lf,ttt. ttt.o,n. X[ag., 95, iS_Si .
--. 1459b. A neu. spccies ol llobbya (Hym., pteromalitlae) fronr
1}'itain. ljttt. rtott,. )llog., 95: 6g_70.
--. 1960. Some obsor.val,iols on Dilttotepis rosae (L.) (Hylr.,
Cynipidac) arrd its palasitcs. Iilttt. mort,. llag., 9B (19b9i : 1g1_
'I OO
1961a. On t}c palacirrctic spocics of Syn.tornaspis Fijt.ster ([Iyrn.,
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---. I961b. A strrrly ol tlrc biology of spcciers of tlre gcrrus
llcnpolobus \vestrvootr (H1'.re,optc.n : ptcror.aritrae) asso.i-
a,ted rvith (lr-.ipirl galis,n oaii. ,j,,rutts..Il. erfi. Soc. Lctrtil., 115
(B): 155-173.
-.
1961c. Sorne hiological obscrlations on tho J)tcr.ornalid (Hyrn.,
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a"fil llubl:,ya l)elrlsxl;i, rvith de,sc'iptiors of t*.o rrcl sPecirs.
lin.tct ntopltogn, 6 : ST-07.
llt,rrtr, I{. C. 1944. ,\ note ot tlre ecorroruy of thc rose bedcguar.gall,
llhoditcs rrtstLc L. Proc. S. Lonfl. c,nt. nat. H1st. Sor., iSAS_
1944: 55-59.
--. 1946. On tlre cconon-ry of the oah urarlrle gall (.C.unips L.ollari ,
1:lartig). 7'rot:. ,lJ. Loncl,. c.rtt. n.at. IIist.,Soc., tSa.S_f S16: 7g_g3.
or,.*rrco, N[. l'. Igig. A corrtrib.tio, to ilre biologv a,rl taxo*orny of
t'lre Ilrilish sPecics of trrc gcrrs fl*trecatotn.a *{shr.carr
(:I)ccutottto, nrrcl,t. .cc Spinola) (IJy,r., Jl,].ryto.riclac). l,rr,ttts.
,\oc. llrit. .Ant., 1t (9) : 149-168.
Coxxrrr,r, lI. lli. :1908. Tlritislt, Ool, CalLs. Irorrdon.
J)nruccrrr, \'. 1g.j7. f)cr: cacnacis-0ecitrostirta-Kor.nprex (oharcitr.,
lltetorrraliclac). 1,)ntortt.opltcrgo., 2 (2) : :137_160.
(i*.rrr,ln, Ni. \V. Il. ,r Y. 1g59. licys to the llritish gencra arrd specics
of i')l:rche.tirra., .rrruror-rhi.ac, E.tccro,tinae. a,d ]rr.dJrina.
tll;rrr., ( lral,.i,loirlr.a). 'l-r,trrr. Snr. llt.il. Et,t., tl (J0): ltig_,-.10L
'ITol'nlror:nn, Iir. B. 1gii. Ilcit.iigo zur Kenntnis der driniscr*rrr
(lailirrro,rirlc,,,rit liesti,rr,rr.gstabcllen cler e.r.opiiischerr
:\rtou. (Oallirnourirlcnstudicn. 5.) lint. Medd,., 1t : 28;_285.
rYtr,r,ttr,oN, .[I. Il. ]g,lg. T:rxorroruic *,cl biological investigatior,, i,,
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Nrur.nrrr', nl. :1948. rlo.o arto..i:iting generations in cynipiilae (Hynr.).
l'tttr:. fl . ent. Soc. f,onrl . (.R),17: 144.
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I)t.r'or*a1id.n-G...s,\rcsolrtrobtts westrvoocr, rg:l3 (Fryr,.,
(llralc.). Opusr:. l,)nt., 23: 20\-210.
. l!)6t. Zrr. Ji,r.t,is tl.s I)t.r.o.rarider-Gc,us il.sctltLtrornts
l\rt'st*.rd, l8:l:j (llynr., Chaic.), VIl. Ertt. Tidslir.,82: l_4g.
\\rrr,r,r.r'sox, ,1{. }I. 1957. A. elcrnc.tar.v ilrco.y oI i,terspccific cour_
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LEPIDOPTERA
Lrsr or rm LnprooptsBA or Donsrr.
Plrr 2. By W. Parkinson Curtis,
1947. 138 pp., 4 pls., 11s. 0d.
Posruur Ilelrrs AND Cor,oun
Perrpnr Evor,uuor rN Lppr-
DoPTEBA. By M. W. B. de Y.
Graham, 1950. 16 pp., 4 pls., 4
figs., 4s. 0d.
Tup Lrrn Iftgronr lNo Bpsevtoun
oF Col,eorfi,ora alti,colella Zsr,r,,
Ry A. l[. Jordan, 1958. 16 pp.,
10s. 0d.
Knvs ro rEE BBrrrsE Grrree exo
Sprcrrs or Er,eosnarrulr, Eulo-
PHTNAE, ENtpooNrwes ltro Eunr-
RrNAE (CEAr,oorDEA). By M. W.
R,. de V. Graham, 1959, 36 pp.,
12s. 6d.
A CoNrnruurroN ro rrn BrorrrcY
eNo TlxoxoMY oF TFn BBrrrsE
Spncrns oF rEE GnNus Eul,c-
catom.a Asuupno (Eunrrourom).
By M. F. Claridge, 1959. 20 pp.,
7s. 0d.
Trs Bnrrrss Srnorss or rHE GENUS
Alaptus Illr,roly rN Ws,rnn
(Crlr,croororAMrMABrDAE). Bv
W. D. Eincks, 1959. 12 pp-,
7s. 6d.
Norps oN Prpsouu,rnlr (Csrr,oro-
OIDEA) WITE DESOR,IPTIONS OT
Ntw Guwrnl AND SPEorEs. By
M. W. R. de V. Graham, 1959.
16 pp., 7s. 6tl.
HYMENOPTERA
Tsp llvunwoprrn.l Acur,ueu oF
Broronnsrrnp. By Y. H. Cham-
bers, 1949. 56 pp., 3 maps,
10s. 0d.
AN lNrnooucrroN To rgp Nlru*lr,
Hrsroav or Bnrrrss Sewrr,rps.
By R,. B. Benson, 1950. 98 pp.,
9 pls., 10s. 0d.
Norss oN Soun Bnrrrss MvsMA.n-
rDAE. By W. D. Ilincks, 1950.
42 pp.,5 figs., 1 pl., 5s. 0d.
Trrn Bnrrrsr Spncrns or rsn GnNus
Ooctonus Illr,ruv. wrru l Norr
oN sor[D RncrNr 'Wonr ox rm
Flrny Fr,rrs (Ilvu., Dfvnuln-
rorr). By W. D. Ilincks, 1952.
. 12 pp., 8 figs., 4s. 0d.
Tnr Nerunlr, Iftsronv or solrn
Pamphil,ius Spporrs (Hru., Plu-
rurr,norn). By Y. I[. Chambers,
1952. 16 pp., 4 pls., 5s. 0d.
A Sruov oF soME Bmrrsu Sproms
or SEnergus. By J. Ross, 1951.
16 pp.,4s. 0d.
A RuvrsroN or SscrroN I (Iflvn,
7872) oF rEE Gnxus Synergus
(Ilvar., CrNrrrnm) rN B*rtlrx,
wms l Sprorrs Npw ro ScrrNcr.
By_R. D. Eaily, 1952. L2 pp.,
4 pls., 4s. 0d.
lbu Bnrrrsr ANrs Ar,r,rmo ro
Form,ica lusca L. (Elru., Fonur-
orrxu). By I. I[. I[. Yarrow,
1954. 16 pp., 8 figs., 3 maps,
5s. 0al.
Tsr Bnrrrss ANrs Aranrn ro
Forrnica rwta L. (flru., Fonur-
oroen). By I. E. I[. Yarrow,
l3|l.uu.*t pp., 58 ngs., I map,
DIPTERA
A Ssont S:nropsrs
Solroprlcrur.
1958. 20 pp., 7s.
Ar.r Ourr,rxr or e Bovrspo Cr,lssr-
FrcATroN or rEE Svnpuroer
(Dretnu) on Pryr,ocprvBrro lrrNns.
By E. R,. Gofie, 1952. 28 pp., 3
figs., 6s. 0d.
A Ruvrsror or rEE Bmrrsr (exo
Norng oN orunn) Sprcrxs or
Loxcsrrrolr (Drrtunl). By J. E.
Collin, 1953. 28 pp., 3 pls., 3
figs., 6s. 0d.
Tap DrsrnrsurroN AND Ilerrrg or
rm Bntrrsu CoNoptom (Drpr.).
By Kenneth G. Y. Smit'h, 1959.
20 pp., 10s. 6d.
Noros oN TEE IDENTrrroAfroN or
Li,mnophila Irlnvnu (Trpur,roen).
By Allan Brintlle, 1968. 12 pp.,
5s. 0cl. oa rEE Bmrrgn
n.r...f. O. Collin,
A Rpvrrw or rEE Bsrtrss Sun-
BAIIII,TES AND GNNEBA OT TEE
Feurr,v Musorpep. By E. C. M.
D'A. Fonseca, 1956. 16 pp.,
6s. 0il.
ORDERS, accompanied by the appropriate remittance, should be adilregeaal to
Dn. W. D. IIINOKS, Manchester Museum, Manchest€r 13.

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voL. 14 PART 11
TRANSACTIONS
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SOCIETY FOR BR!TISH
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.World List abbreviation : Trans. Soc. Brit, Ent.
CONTENTS.
R. R. Asxuw
On the Biology of the Inhak'itants of Oak Galls o{ Cynipidae
(Hymenoptera) in Britain
Dero or PusLrcarrox, 30rE NovnMsos, 196l
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