Journal of Orthoptera Research, Dec.
2001,lO (2): 263-269
Diel and seasonal mating peaks in grasshopper populations (Caelifera: Acrididae)
(KR) School of Biology, The University of Leeds, Leeds LS2 9JT, United Kingdom. E-mail: bgykr@leeds.ac.uk
(JS) Swiss Federal Institute of Technology (ETH), Applied Entomology, Clausiusstr. 25/NW, CH-8092 Zurich, Switzerland. E - mail: joerg.samietz@ipw.agrl.ethz.ch
(GW) CLONDIAG chip technologies GmbH, Löbstedter Str. 105, 07743 Jena - Germany.
(SO) Max Planck lnstitute of Chemical Ecology, Winzerlaer Strasse 10, D - 07745 Jena, Germany. E-mail:Opitz@ice.mpg.de (GK) lnstitute of Ecology, Friedrich-Schiller-University, Dornburger Str. 159, D-07743 Jena, Germany.
E-mail: b5kogu@uni-jena.de
Abstract
I i i most ceritr,il I:iirope,iii g~-~~ssliopper pop~ilntions, iii;itiiigs ,Ire
very rnrcly obsc~rveci. I:or five specit,s wc hert- quaiitify Iiow frt-queiitly nintings '11-C obst-I-vcd. In tot'il, only 7 8 iii,itiiigs wci-e rt-coi-decl ovcr n puioci of 1 1 00 observ,itioii hoiirs i i i fivt, ncridid gr,isshoppei- species. We forriinlly defiiie thc tt'rm m'lting 'ictivity (MrZ) as tlie proportioii o f iiidividunls nintiiig n t a giveii tiiiie. MA i i i 0c.riiporlrr scl.nlrrriicrr, biit iiot t:irtli)lsrir-rr Iwrrc.li)~pir~~-rr, took plnct, preferenti,illy '11-ound rioon. I r i SlrnaI~otlir-irs linc~rriirs, ( ; o ~ t l p l i o i r ~ - i p p l r s rlrfirs ,iiid (:hol-tllippirs prrl-(~llclirs, niatirigs possibly extcrided into tlie iiiglit. We also f o i ~ n d eviderice for a sensorinl decrense i i i MA iri S. l i n e n i i r ~ niid
C;. 1-i!flrs but iiot i r i (:. prrrrrllclirs a n d 0 . gc~-r~lrirlic-rr. M;itiiig feniales were yoiiriger than the populntion avernge in S . lincriiirs, C;. rirfirs nnd 0 . ge~-rtiiir~ic.rr. LI review of the literat~irc revealed n sensori,il decliiie i r i MA i r i othcr species as well. Diel niating peaks are not very ciistirict, tliough soiiie species seem to mnte excliisively ,it night, soirir liave pe'iks during the hottest p'ii-ts of the d ~ y , anti CI third groiip seeiiis to nvoid the hot riiicitiav periods.
Key words
fiiiding. I:or iiistaiice, wlieri males visually search foi- fe- males, niatiiig mny b e restricted t o daylight. Aiiother ex- ainple would b e t h e deci-ease i i i s o n g activity duririg t h e hottest parts of d a y in tlie katydid 1)cc-ticus iic~-~-ilcivor-us (l,iiiiiae~is). T h e lack of singiiig nctivity was related t o a low souiid traiismission rate (Keuper ci r l l . 1 9 8 6 ) niid m a y well lead t o a distinct b i m o d a l MA patterii ovei- tlie ccourse of t h e day. Biotic h c t o r s like predation pressure niay also play a role in shapii-ig die1 m a t i n g patterris. For instance, i r i katy- dids of t h e genus l'oec-ili~norl, predatioii is nssurned t o b e t h e inairi cause of occurreiice of eitlier rioct~iriial or diuriial siiigii-ig a n d lience m a t e attraction (I leller & vor1 1 ielversen 1 9 9 3 ) . I:inally, length o f c o p u l a t i o i i may iiifl~ieiice die1 MA.
Species that m a t e for a l o n g period o r late in tlie afteriiooii may iiot fiiiish niatiiig arid separate hefoi-e niglitfall, b u t c o n t i n u e t h r o u g h o u t t h e iiight (I:edorov 1 9 2 7 , Whitrnari 198.5).
Seasoiial MA peaks caii occur w h e n envii-oiimeiital coii- Cailifei.,l, copulatioil, feiiiale age, niatiiig hehavior, iiiatiilg ditioiis delay iiintiiig i)r resti-ict it t o a Iiiiiited pei-iod o f tiiiie (e.g., reproductive d i a p a u s e Orshari & I'erier 1373, Peiiei- &
frecluency
O r s h a n 19130). 'l'he synchi-onized eclosion of adults in c o n - lntroduction
I i i gi-nsslioppri- field popul,itioiis tlie occurreiice of i i i c i t -
ing m a y change drcimatically i i i space a ~ i d tinie. 'l'his cnii sonietirnes lend t o deiisc iiiatiiig nggregations (c.g., Wickler
& Seibt 1 3 8 5 ) aiid populatioii iiiatiiig peaks ( e . ~ . , Ellis &
Asliall 1 9 5 7 ) . I r i oi-dei- t o quaiitify this, we liere dcfiiie tlie psoportioii of iiidividu,ils in,iting at n giveii tiint' '1s mntiiig nctivity (MA). MA niny v'iry over botli t h e course o f t h e d,iy nnd s e , i s o i i a l l ~ ~ .
tliel cliniiges in b1A iii~iy b e :.issocicited with tlit, tlier- iiioregiilntioii of iiidividu;ils ((:linppt'll & LVIiitinnii 1 0 0 0 ) . 1't';iks diii-i iig t h c \v,ii-iric,st pcirts of ttic d,i), c'iii b c cxpcctcd
~iiicit'r cold coiiditions ~vlit'ii iiicii~iciii;11~ iit~txi t o lit',~t u p for ,icti\~it)~. Iliidt3i- Iiot cliiii,itcs, lio\vt~\~t~i-, iiidividii,ils nccd to ,ivoici tlic Iiiglit'st tt.riil>cr,itiircs ,iiid, tlicrt'fort', ~ i i , i ) ~ iii,itt, iiiniiily in tlit, riioriiiiig, tlit, I ~ t c ,iftt'riiooii, o r , i t iiiglit. Al:\
pc-nks ~ i i ~ i ) ~ ~ l s o I>c i-txl,itcd t o sciisor)r iiiod,ilititxs of iii,itt'
iiection with a n age-dependent decline of mntiiig freclueiicy inay also lead t o seasonal MA peaks, niost obvious in rno- nogarnous species that matt, oiily early in t h e seasori.
Periods o f iiicreased MA ai-e importaiit in populatiori ecology for t w o reasoiis. First, MA peaks m,iy restrict t h e in,itiiig success of iiurnigraiit individii,ils because imiiii- graiits ni-e oftt,ii old (Kiihler 1 9 9 9 ) a n d may miss o p p o r t u - iiities t o niatc early i i i tlie season. A decrt'ase i i i geile flow mny, tlierefore, b e expected (Reiiihnrdt & Kiililer, forthcom- i n g ) . Seconci, h o t h sexes ndjust tlieii- iiintiiig str'itegies t o nintc nvnilnbility (Aiiderssoii 1 9 0 4 ) . 'l'lie tiiiie betweeii rn,itiiigs is tht, niost iiiiportniit comr>oiicnt of tht, -.diffc.i-ciice betwt~eii tlit, st'x I-ntio niid tht, opci-ation,il sex rntio of ,I p o p ~ i l ~ ~ t i o n . 'I'Iit' oper,itioii,il sex I-'itio is oiic of t h e niost iiiilxut'int predictoi-s for tlit, dirt,ctioii of st,xii;il st~lectioii (I<\~~iriicriio
N
:\liiicsjij I O O ( 7 ) .I i i t l i c fit,lci, vci-y Iiigli ,\t/\ l i c i ~ ~ t ' t>t,t,ii I-t,portt~ci (\$'icklt.i- N St>iht 108.5, \I1liitiii,iii 1 0 8 5 ) . 'l'liis ~ t i - o i i g l > ~ c-oiitr,ists \vitli
'I'able 1 . Siiiiiiii,i~-!J of t l i c ~ioliiil,itioiis stiiciit'd ,iiid tlit' i i i i i i i l i t ' i - of- iii,itiiig\ rt-cordtd i i i rt,l,itiori to tlic s,~iiipIiiig cf'ti)rt Slwci txs Stiidj. sitc 1't.riod \vlit,ii ,idiilts t i ~ r s I 1o~ii.s of' Nciiiibt~i-
(coorciiii,itt's) \vc~i-t> ol>st,i-\rtd o f iioctiirii,il of' obst.1-\,,itioiis siii-\lcys iii,itiiigs
obsci-\~c~ci
,5 1 / 1 1 ~ ) r l t l 1 \ I t ~ i t i ~ i t ~ i l 111 J c ~ i ~ i , 7 J ~ i l y - I 5 Nov I004 4 0 0 3 L O 3 4
( k i rnnny 2 0 July - 3 Sep 1335
(I'it 50" 52'N, loiig I I O 34'1 )
I It,ntli i i i - Velii\ve, 1 1 J ~ i l y - 4 Scp 1036 100 Netliei-lniids
(Int -5 1 " 08'N, loiig 5 " 04'E)
t t - tI J 22 Aug - 25 Nov 1934 200
(Iat 50" 52'N, long 19 Aug - 12 Dec 1935 I I O 34'E)
Kei-ribei-ge in Jena 18 July - 12 Aug 1996 5 5 (lat 5 0 " 92'N, long
I 1 O 62'E)
0. cq~i.r?znni~rl gravel pit Steudnitz 6 July - 27 Oct 1994 350 0 6
(lat 5 1 " 0 1 ' N , long 1 1 " 70'E)
o u r observatioiis i r i ce~iti-al Europe, where matiiigs are hai-dly seen i i i aiiy grasshopper species a n d hence MAs ai-e low. We use large observational data sets in ordei- t o quantify this rarity arid d r a w s o m e conclusions a b o u t die1 a n d seasonal changes in t h e MA of five central Europeaii grasshopper species. We review t h e literature for data a b o u t temporal m a t i n g Patterns i i i t h e Caelifera in 01-der t o provide a base for future research.
Methods
F3etween 1994 a n d 1996, w e conducted intense p o p u l a - tion ecological s t ~ i d i e s of five caeliferan species a t four different sites (Table 1 ) . All sites are riutrieiit-poor m e a d o w s o n l i m e s t o n e o r sarid. Detailed descriptioiis of t h e study sites are given by I leiiirich et 111. ( 1 9 9 8 ) for t h e nature reserve Leutratal (Gei-many), by Wagner ( 2 0 0 0 ) for t h e gravel pit Steudnitz ( L e r n i a n y ) , by O p i t z et al. ( 1 9 9 8 ) for tlie Veluwe area ('l'he Netherlands), a n d by Reiiihai-dt ( 1 9 9 8 ) for t h e site
i r i t h e Kernberge region ( C e r m a n y ) . Lift.-history data o f ceiiti-al Eui-opean grasshoppei- species a r e reviewed by Iiigi-iscli Q I<öliler ( 1 9 9 8 ) nnd Köhler ( 1 9 9 9 ) . Det'iils for tlie studiecl popiilations c ~ ~ i d m e t h o d s cari be fouiid in Samietz ( 1 9 9 8 ) for Stctiot~otl~rits lirlerlt~ls (Paiizer), i i i O p i t z et (11.
( 1 9 9 8 ) foi- (:llor.tllippirs prlrrll1elir.i (Zettei-stedt), in O p i t z ( 1 9 9 6 ) ;iiid O p i t z & Köhler ( 1 9 9 7 ) for (;orrlpliocc~ripprr~ ~-llfils (I,iiiriaeus), aiid i i i Wagner ( 2 0 0 0 ) for (leriiporirl ,qerr)lrlriil-ri (I.ntrt-ille). l r i tlit' eiivii-011s of Jen;i, tlie yt)ai-s 1004 ;irid 1005 were wnrmer, but \vetter tlinii ~1vt.i-;ige d u r i ~ i g J ~ i l y , ~v1iere;is
I')')(, \v~is coolt'i- ~vlicii coiiipdi-txi to t h e pei-iod o f 136 1 tcj 1000; ,111 ycnrs li,ici ,i 1iiglit.i- tlinii n\lt,i-,igt, I-ni~if~ill diii-iiig tht, siiriiriit,r (I lt~iiiricli r't r i l . 1008, M1,igncr 2 0 0 0 ) . No siicli d;it,i ,irt, ,i\r,iil,il>lt, for t l i c Vclii\vc '11-c,i. .l'lic pliciiolog), 01- tlit,
species is highly synchronized witli rnost iiidividuals eclosing over a 2-w period. Marking o f y o u n g grasshopper adults was started early in t h e season w h e n 3 0 t o 60% o f t h e individuals were still n y m p h s a n d marking was c o n t i n u e d for a further period, varying with respect t o species. Tlie d a y o f marking was regarded as d a y o n e of their adult life. This type of age estimation has ari advantage over others (see Köhler et iil.
2 0 0 0 ) , in that individuals d o n o t rieed t o b e killed, ari essential for populatiori ecological studies.
Individuals were I-esighted over t h e w h o l e season by meaiis of b o t h nocturnal surveys of individuals marked with reflecting t a p e (I leller 24von I Ielversen 1 9 9 0 ) , aiid diuriial surveys, except for Eutllysfirn b~-rl~-hyptern (Ocskay) (sec be- l o w ) . For instance, in S. lineatus, t h e average resight rates dui-iiig iioctui-rial surveys were 7 0 . 5 % (s 1 3 . 8 % ) in rnales, a n d 7 5 . 4 % ( s 1 4 . 1 % ) in females. Duriiig diurnal observa- tiori surveys i r i this species, resight rates ranged between 1 0 a n d 20% (Samietz 1 9 9 8 ) . In C l . g c r n i a r i i ~ a , t h e average i-esight rate of marked individuals d u r i n g t h e night was a b o u t 65% a n d t h u s a b o u t 1 0 % higher d u r i n g t h e iiocturnal surveys (Wagner 2 0 0 0 ) . For logistic rensons, night surveys (if niiy) were iiot cari-ied o u t i i i successioii t o dav survevs.
Sui-vevs were riiade by walkiiig i i i t h e study sites 2nd iiiteii- sively seni-cliirig for riiarked iridividuals, t o maxiiiiize resight iiuinbers of marked iiidividu;ils.
il slightly different npproncli was used i r i 13. I~~-rlcl~)'pte~.(l.
At vzirious times of t h e dny oiie observer (KR) wnlked tlirough t h e iiie;ido\v <111d coiirited tlie riiimber of m , ~ l e s , femaIt>s arid copulntioiis eiicoiiiitei-ed pei- 10-iiiiii iiitervals (11 = 4 0 ) . All copulntiiig pnirs \vere collt'cted, so tliere \V,IS iio o p p o r t ~ i i i i ty foi- coiiiitiiig tlie s,iiiit3 11i;itiiigs t\\ricc. 'l'lic i2-li\ pcr 1ioi1r is tliiis tlie ,ivt~r,ige ii~inibt,i- of iiicitiiig kiii,iIcs fociiid ,it n givcii lioiii- of tlit, d,i)., di\ridt-d by tlic ii~iiiibcr o f ft,iii;iItxs ciicoiiii-
li. l < l ~ l N l liAlll)'l, J . S1\,4tll.'l'Z, C;. \ l r ~ \ C ~ N l ~ l ? , 5 . OI'I'I'L /\NI) C;. li<jl l l , l ' l <
265
'I'ime of d a v
Fig.l. Diel variation in MA ( t h e proportioii o f females eiigaged i i i copulatioii) i i i tlie grasshopper Eutliysrir-(1 br-iiiliyptei.ii i t i
relatioti t o t i m e o f d a y (black litie) a n d t h e average iiumber of females seen in a 1 0 - m i n iiiterval ( b a r s ) . 'l'he nunibers in t h e bars d e n o t e t h e i i u m b e r o f 1 0 - m i n intervals sainpled.
tered h o u r o f d a y , as from t h e O-miil c o u t l t s 1 0 4 5 niid 1 4 1 0 . T h e sixth feniale copulated for 5 0 miii i i i t h e ACross species, w e coiiductrd L-o 1 1 0 0 h o u r s o f iilves- l a t e afteriiooii wheii a m b i e i i t temPeratures ( 3 4 0
tigation, cri 60% o f which were at night (Table 1 ) . exceptionally high. I r i a further 4 y of less intense investiga- tions, o n l y oiie iiocturiial mating has ever been recorded.
Results Because t h e resight probability of iiidividuals is a b o u t 10%
higher d u r i n g tlie night t h a n t h e day (Wagner 2 0 0 0 ) , t h e o n e Overall, we observed 78 tnatiilgs ) , Thus, we o c c u r r e ~ i c e o f a tiocturtial iiiatiiig is i n d e e d a n exceptioii i i i
iieeded oii avei-age inore thaii 14 h o f obseivation before (I-
s~')'"'"~'~'.
s p o t t i n g CI m a t i n g grasshopper pair, despite m o d e r a t e grass-
liopper densities o f 0 . 2 t o 1 . 4 i n d . / m 2 . Seiisorfril clzrirfge
(!f
MA rirzrl rige of'tilriting inrii~1iliuals.-During nocturtinl surveys, S. lineritzrs females s h o w e d a decrease in illlnse b o t , , . u , s , 1 o i l y MA o v r r t h e seasoii, especially i i i 1 9 9 5 (1:ig. 2 ) . 1'1ie secoiid iiocturnal iii,itillgs were observed arid tllr majority o f s u r pak i i i results Irom siiigle m.itiiig feniale O u t o f ' 0 vrys took ,,IdCe d t l l i g l l t I ) , l i l (:, I>illilllL,l14s tWelvt, still alive arid itidicates t l i i i t tliis fetii,iJe Iias nidteci a seroiid systeillatic searcll days c a r r i e d o u t betweeil 1 5 'iild 700 tinie. Note tliat iiiales erlose before feiii,iles (S,iiiiieiz 1 9 9 8 ) . (tot,ll 70 li) i.evealed two i l i d t i l l g s o f L i i l i l l a i . ~ e ~ i i l d i v i d L i n l s All niatiligs b u t o n e w e r e observed hetweeii day -5 aiid d a y 2 0 a t,
400, Dui.iilg 6 iloctlii.ilal sui.veys, cari.ied olit betweeil o f 111e a d u l t life spaii. I i i 1094 tlie oiie exceptioti was a fetii,ile 2 2 0 0 ,itld 30 30 11), oilly ,I siiigle illatiilg was niatitig .it ' i r i .ige o f 5 1 ii (sccotid peak in 1994, I:ig 2 ) . I i iobservecl, this o n 1 G Augiist at 2 2 4 5 . b o t h years h1A in females pe;iked ,irouiici nii ;ige of 10 d aiid
i,i.il~.ifl,piui.ir iliatitlgs rvcillv d i s t r i b u t e d ovcr dccliiicil .iftcr~v.ii-d. '1s tlic i i i ~ ~ i i i (i s ) l ifcsp.iiis o f fciii.ilcs t h c d'iytinie (l'ig. 1 ) . were 2 2 . 8 i- 12.7 d i i i 1904 ;itid 1 9 . 3 $I 17.1 d i i i 1005
SLi o f Ci,i.iifii,fii.il wci.c e\lclllv c,isti.ihutcd bc- (S,iiiii~tz l 9 0 8 ) , tliis i ~ i i i i r ~ i t c s tliiit iii~itiiig f c ~ i i ~ i l c s ~vei-C tweell
ooOo
citld 1 8 0 0 ~ ~tiilli, ~ (tot,ll 350 11), ~ , ~ C ) t l l y l yoiiiigci- t l i c i i i tlie popiil.itioii .ivt.i-.igc I i i tiinles tlie ~ i i ~ i t i i i gi l l ~ i i , i l g s wert o b s e i - v c e ~ , ~ : i \ , ~ t ~ l e i l l t o o k ,,lclrc ~ , c t w c c l l 1 ~ " k occl~i-i-~cf .ib(ilit 5 I c i t t , i - , ~irolilid ~111 ~1gt. 1' d . I'lic b1A of C;. r i r l l r s dt~crt~nst~cl ovcr tlit, st~,isoti. I i i 1004,
I),ite i ~ i July tlieii August
Fig. 2 . Seasonal v'iriation in MA ( t h e proportion of indi- v i d u a l ~ eiigaged in copiilation) of marked m a l e a n d female grasshoppers S. Iirreritirs in t h e field. Recordiiigs were stopped wheri tlie nuriiber of marked iridividuals was less t h a n 15.
iicss, \jZt, siiggtxst t l i , i t tlic' iioc-tiirii,il iii,itiiigs in tlicstx slic,c-itxs '11-c, tlit, rc'siil t o t' pi-oloiigt>d ,i f'tt.1-iiooii oi. c\,t'iiiiig i i i , i t i iigs.
( , ' o i ~ l / ~ r l i . i s o i l ( I / i c ' j i t / ~ c ~ i - i l l i i l t l i i i l s /~r'il/:.\ tlc.t.oss s/~c>c.it>s. -'l'lit' pi.t%stxiit ficld ,iiici litcr,itiirc c1,itn d o not s u p p o r t tlic li)rpotlit.st.s tli'it tlitxi-iiioi-cgiil~tioii oi- 131-ed,itiori ,irc c s c l i i s i \ ~ c l ~ ~ I-t~spoiisililt~
foi- tlic, oc-ciiri-t,iicc of tt>iiipoi-nl inntiiig pc,iks. A t t w stiidit's siiggcst t l i ; i t h11\ iiicre,ises dui-iiig Iiiglitxi- tt.ii1pt.i-ntiii-txs i i i
moctt,i-,~tc clini,ittxs ,ind t l i ' i t t1it.1-e ni-c distiiict lo\z!t,i- i i i ~ t i i i g ncti\~itit,s ,iro~iiid iiooii i i i Iiot cliiiiotes ('l',il>lt' 2 ) . '1'liei.e ni-t., lio~vt'vcr, esceptioiis t o this: ii,tmely N o ~ ) i r l r i r l ~ ~ . i s s c ~ p i c ~ i i i f i i s i - i r l t r i
;iiid t h e t w o 1 , i ~ i r r o t i ~ i i i . v spt~cies ('l'c~blc 2 ) , OCCLII- in very hot clirrintes ;ind it d o e s not setxiii likely tlint t h e reason for '1
innting peak nroiiiid iioon is I-el,itt,d t o n pliysiologicnl iieed for wariiith. 1 Io~vever, giveii tlint somt'timcs mntings rnay iiicrense t h e v u l i i e r ~ b i l i t y to visiinl pi-edntors (blagiiliagt~ii 199 1 ), i t niay bc thnt grasshoppci-s pi-cfci-t.iiti,illy matt. ,it ri t i m e w h e n their riiaiii pred,itors are least nctive. Yet, there a r e iio e f f e c t i v e t e s t s t o d i s t i i i g u i s h b e t w e e n t h e theronioregulatioii o r p r e d a t i o ~ i liypotheses.
If seasonal MA peaks wei-e inerely d e t e r m i n e d by teni- perature w e s h o u l d find peaks d u r i n g m i d s u n i m e r in colder areas, o r peaks in t h e beginnirig ,iiid e n d of tlie suiiirner in warmer habitats. In additioii, iioon peaks s h o u l d occur early a n d late in t h e seasori arid nooii depressioiis in t h e midseasori.
No such data are available in t h e C'ielifera. 17isclier c7i (11.
( 1 9 9 6 ) could n o t find distinct singingactivitypeaks ovt,i-tlie Course o f t h e d a y foi- s o m e o f t h e g o m p h o c e r i n e species m e n t i o n e d here. rl'l~erefore, inatiiig peaks niay n o t occur in these species. In t h e iioiisingiiig species 0. s e i - n i r i r ~ i c a , h u t tliree o f t h e f o u r matiIigs were obsei-ved duriiig t h e first t h i r d n o t in tlie siiigiiig E. b i - ( i c i ~ y p t e i - ( i , there was evideiice for a n o f t h e sui-veys, t h e fourth in t h e secoiid half. In 199.5, 7 o f 1 I incre""'d aroiiiid nooii t h e
nlatings ( 6 4 % ) were observed irl tlle first t h i r d of tlie In field oi- iab poliulations MA decliiied over t h e scasoii moiiitoriIigs, (820/o) iri t h e first 500/0 of t h e survey Iiigllts, in S. lirreririrs ( t h i s s t u d y ) , B i y o i i e r i i ( r t r i b e i - c i ~ l i l r ( i (Fahricius) r17hese d a t a a r e sul,ported by t h r age d i s t r i b u t i o I i of ( ß o r ~ l l l a l ~ n 1 9 9 1 ) . ( : i l c i i - i i l i p p u s b i ' q i r t t i r l i r s ( L i n ~ i a e ~ l s ) inatiIig (;, ,-iifirs females aild males t h a t were hetweeIi 2-68 ( Kriegbauin 1988. Kimüller 1 9 9 1 ). M i i - ( i r i l e i l i l f o i . r r i o i ( i n t ( ~
aiid 2-55
of
age, respectivelv, Whei.eas t h e age of iliatiiig (1:ruhstorfe) (Köhlerei iil. 1 9 9 9 ) aiid I f i c i ~ i ~ q l y p i i i i s r i i g t r ~ i - e p l e r u sfeInales (ineaii 25 d ) was illdeed below t h e p o p u l a t i o r l Bolivar (Siddiqi 1 9 8 9 ) . A s s u ~ i i i ~ i g that thermoregulatioii is averagr o f 3'2 d (illdicatillg early inatiIlg), males arid fCmales l l i # l i l ~ ' p t i m i z e d ( C h a ~ ~ e ' l
'
Whitinaii 9 9 0 f Sainietz'
fouild m a t i n g iIi 1995 2 u aIid 25 d , respectively) l<öliIer 1 9 9 8 ) iridividuals may b e a b l e t o be active (aiid wcre olily slightly below tlle popu]ntioil menIl of 2 8 aild 2 7 "'"'C) du'i"g a w i d e raiige o f tempei-atllrcs t h r o u g h o u t t h e d , i-espectively. Marked fouild matiiig iil , 9 9 4 (meail d v Y While tliis p r o d u c e die] patteriis o f MA! o ~ l l y a age 2 3 d ) were 'is o l d as tlie meaii of the pOliLilatioii (2 4 d ) , iiegligihle seasoiial variatioii is p r e d i c t e d Season.il diffei--
N o cleai. seasoilcil pattei.ii was fouild iIi prll.(lllelirs. .I.he e11cew11ay lie diie rather t o different predatioii pressures oi- iliarkrd C,'. prli-il//e,i4s f e m a l e obseii.ed inatiilg was 2 3 d d u e to age-relatcd c l i ~ ~ i i g e s iii i h r lllatiirg frequeiicy.
o l d . While in oiie laborntoiy s t ~ i d y , female age d i d n o t iiiflu-
I 'qcl.liI(l riie.ri six IliatiIigs were Ilot iced i i l t h e secoild a i c e MA i i i C. piri-irllelils ovei- tlie first 8 p o d s Iaid (1-il 3 0 d )
third oftlii. inoilitoi-iiig pei-iod. H~~~~~~~ feliiales have liigll (R'illhai-dt
er
( 1 1 1 9 9 9 ) . i t dccreasfd i i i f e ~ l i a l e s o f M . cige l l l d t i l l g tl l i S coLlld llle<lll <1 SCasOI1ill deeliiie iil / i ~ i i i i o s ( i r i i i l ovei- tlie se.is«ii desliitc coiistniit tt.nipcr.itures blA, lIldeed, cq<.,. „riirIii.ll feillales iiiediail age ()f (I<öliler er i i l . 1 9 9 9 ) . Such .in age-I-rl,ittd st~asoiinl ilecliiit. i i id; tIiose ~ o i i I i d matiIig Lvere o i l I y as o l ( n l e d i a i l 2 2 MA iiiriy b e cxplaiiied by .in iiici-ensiiig reluctiiiice o i iiinhil-
d ) . ity of females t o reinnte with iiicrensiiig age. I:or exnmplc, C:.
I~i,qiriiirlirs femnles seein not t o reni;ite ~t all (Ki-iegl>iuiii
Discussion 1 0 8 8 ) o r rarely (Ki~iiüllei- 1 9 9 I ) . O u r field rt.siilts showiiig tIi,it iii,itirig iiidi\,idu,ils ~ ~ ~ c i - c ~ ~ o i i i i g u - t 1 i ; i i i tlic ~ x ~ p i i l . i t i o i i
f o l i i l ~ iloilt, of oui. species Illated d L i r i i l g tlie illoi.ll- v c - c in 5 . l ~ s. ,i.ri/lis aiid 0 g i ~ t - t i i r i i i i i - ( 1 , sceiii t o iilgs, i,s i l l t , i l t i o i l ~ , d ciho\lc, iIlt,i.e \j,pie i l l C i d ~ l l C e S I1oCtlll.- s~ip13oi-t tlir idc'i of<iii .igc-rcl.itctl dccliiic (;i\lcii tIi.it in tlic ii,iI c o l ~ i l , i t ioiis i 11 (;. r-lr/lr>, S. lirlc~rrtirs ,iiid (:. / ~ r l t - r ~ l l c ~ l l r z . field iiiost fetii,iles ;ii.t' nlilc to iiintc bt%foi-c tl1t.y Iny tlicir first I(c,cciiise i l l ~ l t i l l g s il1 t ~ l c s c slit.cics cii-c i l l i t i ~ l t e c i 1,). Ill'llc soilg g 1 ) ( i C g 0 1 1 t I ' ) ( ) L ) ~ i i d t I i . i t i l l c i l l t c i i r i i l g l l , L l r s O ~ ~ l l l . l , o\riliositioii is tciiipcr.itiiit' d t ~ p t ~ i i d c ~ n t (st,t, P.:<., S . i i i i i i t ~
I,ihlt, 2 . O l > \ t ~ i - \ . , i i i o i i $ o i i i i i c i-t%,i\cxi i i i , i t i i i g , I ( i i \ , i i ! ( 1 \ ) i i i I - c * l , i t i o i i to i l i t . i i i i i i . 01 ci,i\
d,i)r niid iiiglit L 5 ( ~ l ~ i s i o ~ ~ i ~ r ~ c ~ ( ~ ;{r.cyil-i([ ( I o r s k ~ I ) l.llis & ;\sliall 1057
;it iiiglit I<o\vt~ll iiiipub i i i I<itdt, I 0 8 7
( ~ ~ i - l > o i i t ~ l l I056
13111-tt 1046 iiiiddn), ,4t~l(1tlop11is (li/fc~r~c~~iii(llis ('l'lio~ii~is) Kyl 1038
N I Is ~p ~( S e i l e ) s 13uriiett 105 1
1,isirr-oic7iiis spp. GI-eeiifield I007
; / - pi i s -( II ) N icdzlek-1:enver 1 00.5 Oc~(lipo(f(i ~ ~ > l . ~ ~ l ( l t ~ i c . ( l ( 1,;iti-eille) prest,iit stiidy
~ ' / ~ ) ~ I I I ( I ~ P L ~ S 1cpr-e)slrs (l:c~l>rici~is) I<ölilci- t,t nl., iiiipub. dntn inorniiig and/or Aorloi-~~-~ts rlqiins ('l'hiinberg) Wickler & Seibt 1085
Inte afternoon (ialirfrie-/-is spec. Riede 1987
1 li~~)-clglypli~r~ 11 iCqrur-eplci 1r.i 1301 ivai- Si-ivastC~vci 19.56 Atrr~c-torrlor-pllii c-I-en~rlrliri (Fabricius) Srivastava 1957 Aii(rc-r-itliirr~i irpcq)~pii~r?~ (I~iiinaeus) Fedoi-ov 1 927
M~liriic~plzrr srirl~~tinipcc (1:abricius) Pickford & Cillott 1972 C:rillipiiir~lus plebc~jits (Walker) Köhler, iinpub. Data
1 9 9 8 for oiie ~f t h e stiidy specics) it seenis likely that ,itller seasoil
",-
tiint, of d a y w h e r e a s iio die] chaiige in relativelv old feinales fouiid niatiiig ai-e I-einatiiig rather thaii iiiatiilg duratioii hcls beeil obsei-vcd iii ( ~ ~ l o i ~ ~ l i p p i r s ~ i i - i l r l i i e L l smatiiig f o i t h e first tinie. Age-I-elated seasoiial decliiie i i i MA (r17hunberg) ( ~ ~ ~ l i ~ ~ 1 9 8 5 ) . o ~ s e i - v a t i o i i iil L~~~~~~~~
maY b e d u e t o cllaiiges feniale recelitivity througl1 t h e I,iigr-(lioi-i(r (1,iiiiiaeiis) (ßoldyrev 1929) niid 0. 'q e i / ~ l ( i r ~ i ~ ( i accumulation of male semiiial substniices which preverit ( ~ ~ ~ ~ ~ i l l ~ ) ( ~ ~ i& ~~ hö ~lfortlicomiiig) sLlggest tllai ~~ ld~~~ further copulatioii ( f l a r t m a i i i i & I.olier 1 9 9 6 , 1 9 9 9 ) . It is duriiig coolei- temperatures, copLllatioiis last loiiger, rlliis iiiterestiirg t o iiotr tiiat in C:. iiuus, where these substances iiicludes cases whei-e copulatioiis arc leilgtheiied low lrave beeil ideiitified (llartniaiiii & Loher 1106, 1 9 9 9 ) . wc llight temlieratures arrive before matillg is coiiiplete~ arid f o u n d a seasoiial decline in MA. rnating pairs d o n o t separate until tlie next morniiig, i i i
Anrrc.)-irliur)l ricsllpiiurn (Liiinaeus) (1:cdorov 1 3 2 7 ) , in N.
Mlrtin<qir[-iiiliiyiin<i iiiiliiris fre(lii~rrcl~.-Aii importaiit applica- „,pi„T,!;„l-i„l(l ( K ~1<)5 ,1 ) aild ~ ~perl1aps ~ ~iil soliie ~ ~()f oiii-
tioii o f MA is iii t h e estimatioii o f feiiiale matiiig freq~ieiicies studied species, ~f t h e i n a t i i l g fi-rqliellcy of these s,,ecies in t h e field. O u r first niethod o f inarking iiidividuals a n d
„„
to be cnlculated froiil tlit.ii- MA, wc would n couiitiiig t l i ~ ~ > r ~ ~ o r t i o i i of iiiarked feiiiales i i i copuIatioi1 at overestiiilatioli o f t h e foriner brcalist. ~ r i l g t ~ l r i l e J sevei-al times of tlie d a y o r ~ e ~ ~ s o i i is very Iabor-intensive aiid c o p u l a t i o n would be inistakeii iliaiiy s h o r tt i m e coiisuiiiiiig. With inatiiigs averaged ncross species, we I : ~ , . feiliales, secoiid matiiigs were slightly I~~~~~~ thai,
obsei-vtld oiily oiie niatiiig iier 14 obsei-v.itioii h o u r s . O L I I first oiles iii P t l , ~ l j j P I L , s ( ~ ~ 1 1 ~ ( r i , 1 9 9 2 ) . ~ ~jatei- iil t i ~ ~ ~ ~ secoiid nietliod, exeniplified i i i 1:. iiiiiiliyptri~i, d i d n o t re- tlie secls()il sllollld t ~ i e i i he expectrd to last loiiger, an^ e;il-ly
quii-e r~iarkiiig (sei' .ils» Riede 1 9 8 7 for aii exaniple). Couiit- would
L , ~
iilore likely to reiilaiil liiidetectedwe
are ing t h e proportioii of rriatiiig iridividci;ils seeins a n easy niid tlierefore c o i i v i i l c e ~ that t h e seasonal reported I~~~~less oiierous alteriiative, niid it will he possihle witli renson- for S , l;ririitiis aild C;, ilr/irs reflect pnttcrils, despite tlle able effort to cornpare iiiatiiig activities, i) hetweeii species fact t h a t we kIioM, how iliating dui-atioil vnries
i i i tlic s,iriic Ii,ibit,it, ii) bctwecii different pop~il,itioiis, o r iii) o\,er tlie seasoii,
withiii r>opulatioiis nt differeiit times of tlie se;ison. 'l'lie MA
t l i ~ i s iiivestigatrd niiows .i c.1iciii,1tioii ~ > f ficld iii.1tiiig frc- Acknowledgements queiicy, n key pnrnnieter i i i sexu,il selectioii, 13iit i-arcly
s t ~ i d i e d (Aiiderssoii 1 3 0 4 , Ariiclvist & Nilssoii 2 0 0 0 ) . 'l'liis reserirch was i i i part supportcci by tlit, Ilt~iitsclit' 11s st.ited in tlic iiiti-odiictioii, kiil .ilso depeiids oii copiil.1- ~ : o i - s c ~ i L i i l g s g e i l i e ~ i i s c ~ l a f t ([)I:(; gI-'lilts sei 7701 1 - 1 nlld tioii c i ~ i i - ~ ~ t i o i i , ~ \ l ~ i t l i t ~ i i i ~ i t i c - ~ i l l y , iv11\ i i i '1 pi)piiIcitioii is t l i t , I < ( ) 140411-1
1, I , ~
[lle ~;t,rl,lcii, ~ ~ i i l i s t , - y ( ) f ~ ~ c ~ l i c , l t ~ ~ , i l ~ l I l c ~prodiiit o f ~ 0 p l l ~ ~ 1 t i 0 1 1 i ~ i i i ~ ~ i t i O l l .iild ili.1tiiig f r i ' ~ l l i " 1 ~ ) . 'l'l1t.i-C'- ~ ~ c s t . ~ l l - c ] l ( ~ ~ ~ ~ [ ~ ~ : g l - ~ l l l ~ ( ) 3 3 ~ ) 5 2 1 ciiici tIlc stclte o f ~ ~ ~ ~ l L l l - i i i g i , l ~ fortx, \ ~ l i t . i i boili \I[\ ;iiid t h c col,~il,itioii diiratioii ;ii-t' kiio\\'ii, \ V , t j l c l l l l i J , ~ l i ~ ~ ~ ~ - ' l l ~ i j f ~ , ~ f o r l)l-O\ric{illg lllll,ll~,lisht~d dclt<i.
t l i ~ i l l ~ l t ~ i l g f l ~ t ' ~ ] l l t ' i l ~ ~ ( - ~ l I l ~ c l ~ i l y ht' c ~ l ~ < ~ l l ~ ~ l t t ' i ~ . ' I ' I l i S 13l-OCt'- I ) [ - . [ ) O L l g \ \ ~ ] l ~ ~ l l l ~ l l l ~ i i l i { tIj10 ~ i i i O l i ~ ~ l i l o L 1 s i - t ~ \ , i t ~ \ ~ r t ~ r s g'l\nt, diirc. I-t>cluircs tli,it copiil,itioii diii-,itioii clot~s iiot tli,iiigc O \ ~ H \,~llLi,ll,lc. ~ - o i , i i i i c ~ i i ~ s tllt' i l i ~ l i l L i s c r i l ~ t ,
268
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Iiiiiiiillci- 1 . 1 0 0 1 . Iliitci-siicliiiiigt~ii ~ i i i i i I'ortl>fl,iiizii~ig\\~t'~-li,iltt.ii dei- I t~ldliciisclii-ccl\t- ('lior-r/iip/~ir' I~ixirrrirllr.\ ( O r t l i o p i c i - , I : ilci-idid,ic,) iri ciiieiii I.rcil,iridk;ifig- I:xperiiiieiit. ~ i i i p i i l ~ l i s l i c ~ d , I)iploni,ii-bcit I I ~ i i \ ~ ~ r s i t y of Ilrl,iiigt~ii.
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I3iitlin R . I<., Ilcwitt C. M . , W e b b S. 1'. 1385. Sexii,il selectioii for iiitermedi,iteoptiiiiiim size i i i (~lior.r/iippirs Ilr-irtrtirjlrs (Ortlioptci-'1:
Acridid'ie). Aiiim,il Rehnvioiir 3.3: 128 1-1 2 0 2 .
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I:edoi-ov S. M. 1 0 2 7 . Stiidies in t h e copiil,itioii ,irid ovipositioii of Anrrcl-iriiirrri ric>~ypii~1rtr1, I,. ( O r t h o p t e r d , Acrididne). 'l'r,iiisnctioiis of the Iiiitoinologicnl Society of 1,ondoii 7 5 : 5-3-60,
1:ischei- [:.P., Scliiibert II., I'enn S., Schulz 11. 1 9 3 6 . Dicirii'il soiig nctivity of gr,issland O r t h o p t e r a . i1ct;i Oecologica 17: 14.5--364.
(:reciifield hl. I). 1337. Scxcinl selectioii i i i I-esource defeiise polygyriy:
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Xi Crespi, I . C:., (Ilds), 'l'lie evoliitioii o f ni,itiiig systcriis in iiist~cts nrici ni-,iclinids, <:nriihridgt~ Ilniversity 1'1-ess, <:niiil>ridge.
I 1'11-tniarin R., 1,oht.r W. 1 9 0 6 . (hriti-ol niecli,inisiiis of tlie behavior 'secoiid,ii-V defeiise' in tlie gr,isshopper (;ortrplrol-ri-lls rirfirs I , . ( ( ; o i i i p h o c e r i i i n c O r t h o p t e i - ' 1 ) . J o u r i i , i l o f c o m p a i - , i t i v e I'hysiology A 178: 3 2 0 - 3 3 6 .
I I,irtiii,iriii R . , 1 ohei-L\'. 1 0 0 0 . I'ost-rii;itiiigefi-cts i t i thcgi-,issliopper, (,ortipiroic7rirs rir/ir.i 1.. iiiedi,ittd by tlit, spt~riii,itlicc,i. Joiii-ii,il of conil~ii-,iti\rc 1'h)rsiology A 1 8 4 : 325-3 32.
I Ieiriricli \>lJ., ~ ~ l ~ i r s t ~ i l l e r I<., l ~ ~ ~ l i ~ - i i i ~ i i i i i I<., l't>riiei- l., Scli;iller (11.
101)8. l1,is N , i t i ~ ~ - s c I i i i t ~ g e l ~ i c t " 1 , c ~ i t r ~ i t ~ i I " l>t,i J e n ~ - StriiIiti~i--
i i i i c i S i i k ~ t ~ s s i o i i s f o r s ~ - I i i ~ i i g i i i ( ~ r ~ i s l ~ i ~ i c i - O l \ o s y s t t ~ i i i c ~ ~ i N,itiirscIiiitzi-e1>01-t 1 4 : 1-4.24
I lellt-r K,-C;., \ r o ~ i I l c l v t ~ r ~ t ~ i i 0 . 1 0 0 0 . S i i r \ ~ i \ ~ ~ ~ I 01. '1 ~>Ii~iiiei-ol>tt~i-ici hiisli-ci-ic1ic.t stiidit,d 1)). '1 i i c ~ v iii,ii-l\iiig tc~cliiiicliic (Ortlioliiei-,I:
I ' l i ~ ~ i i t ~ r o ~ > t c i - i c i , i c ) . I:iitoiiiologi~i ( ~ c ~ i i e ~ i - ' i I i ~ 15: 2 0 C 2 0 8 . I l c ~ l l t ~ r K . - ( ; . , \.oii l I t ~ l \ ~ c r ~ c ~ i i 11. 1 0 0 3 . ( : ~ ~ I l i i i g I>c~li,i\~ioiir i i i I>iisti-
c-I-i( 1 i t . t ~ of tlic ge,iiii\ rlir/ic~rr \ \ ' i t l i dil~ti~i-iiig c - o i i i i i i i i i i i c - , i i i o i i -
\) \ t < , i i i \ ( 0 1 - t l i o p t t ~ i ~ ~ i , I e~ttigoiiioicit~~i, l ) l i ~ i ~ ~ t ~ ~ ~ o [ > l t ~ ~ - i c l , ~ ~ ~ ) J . Iiist,c.t lit>li<l\, 6 : 5 0 I - 5 ; : .
Iiigi-i\c.li S , IioliIe~i~ C , . IOOS. l ) i c x 1 I t ~ ~ i \ ~ - / i i - c ~ c - ~ ~ c ~ i i , ! l i ~ t c ~ I c ~ i i i - e ) ~ > ~ i ~ . .Il,igtit~l>~ii~g: \ \ ' c ~ t , i r p \ \ ' i s ~ c ~ ~ i s c l i ~ i l ~ t c ~ ~ i .
o . 0 \ I - l t ~ o o g i cS t i i i t i '111
I c~lcilicii\clirc(-l\t.ii. Lt-itsrlirift I-iii- I it~r~>s!~cIiologic., Siip1>1c~iiit~iit 1 : 1 - 1 8 8 .
I<t~iipc~i- . \ . , K'iliiii-iiig I<., S(-Ii<itr~l , \ . , I ' i t i i i i t > i - \ L r . , li,list\r \\'. I 0 8 6 . l3cti,ivioiii-~il ~ i ~ i ~ ~ p t ~ i t i o r i s 01 81-oiiiici Ii\,iiig l>iisIicri(-l\c~ts to tlic l ~ - o p u - t i c ~ s 01' \oiiiid l ~ - o p . ~ g ~ i t i o i i i i i 10\\ gr.1ss1,iiitt. Ot'c-ologi,i 70. 4 1-1-42'.
I<olilci- C;. 1 o o o . i j k o l o g i ~ c l i t ~ (;i-~~iidl,igcii \ r o ~ i . . \ L I s s ~ ~ I - ~ ~ ~ ~ ~ ~ - o ~ . - c s s ~ .
I ,illstiidit~ii ,111 I Ic~isclirc~clit~ii ((:,iclilCi-,i ct 1:iisitt~i-'1). I ,iiii-ciiti, I3ocliii ni.
tiii1ilt.i- (;., j e i i t ~ s c l i il., R e i i i h ~ i - d t ti. 2 0 0 0 . ilge rclattd iii,iiidihlc ,ibi-,isioii i i i thrt,e s p c c i c s ot' s l i o i - t - h o l - i i d gi-nsslioppci-s ((;;iclifcr;i: iIcriciid,ie). Jocirii,il of'01-t1ioptt.i-,i Kt,sc,ircli 0 : 8 1 - 8 7 . liohlci- C , . , I<t~irih,ii-dt li., h~Sliot't'1i. I 1 ) O O . Lui- I(iologie dei- Iessiiicr
(:ebi I-gssch I-eckc, hl it-ririrc~lltr /orrrrosritriri ( I:riihstorfei- 1 0 2 1 ) (ilcrididae: (;nt;intopiii,ie). hlitteiluiigeii d e r Scliureizerisctieiii I ~ n t o i i i o l o g i s c l i ~ ~ i i (;t~sr~ll\ch,ift 7?. I1 5 - 3 2 8
Krieghnciiii 1 1 . 1 0 8 8 . Iliitci-scicliciiigt~ii ziir "l.ebcrisgeschictiter' v o n I ~ e l d I i e c i s c I i i - e c k e n ( A c r i d i d , i e , ( ; o r i i p h o c t ~ r i n a c ) : I:ortpfl;inziiiigssti-ntcgie u n d ,iliiistisches Verlinlteii ini ii,itürliclieii I 1;ibitnt. unpiibl. 1'ti.l). thesis Iliiiversityof Ilrl,iiigeii.
Ki-iegba~im I I , , v o n I lelverst,n 0. 1 0 9 2 . Iiifluciice of ninle soiig o n feniale iiintirig beliavioui- in tlie gi-,issliopper (;/ro~-iliippirs Ilisiriiirlirs ( 0 r t h o p t t . i - a : Acrididnt7). Ethology 3 1 : 2 4 8 - 2 5 2 . Kvarneirio C., Aliiiesjii I . 1 9 9 6 . 'l'he dyiiniriics of opei-atioiial sex
I-'itios a n d c o i ~ i p e t i t i o i i for mntes. 'l'rends in llcology arid 1:volution 1 1 : 4 0 4 - 4 0 8 .
Kyl C;. 1 9 3 8 . A s t u d y o f c o p ~ i l n t i o n ,inci t h e forriiatiori o f spei-matophores i i i Mrlririopllrs r1iffc.1-c~ntirilis. I'roceedings o f t h e lowa Acadeiiiy o f Science 4 5 : 2 9 9 - 3 0 8 .
Mngiihagen C. 1 3 9 1. Predatioii risk as a cost of I-eprodiictioii.
'I'rends in 1:cology a n d Evoliitioii 6: 18.3- 1 8 6 .
Niedzlek-I-enver M . 1 9 0 5 . Crepitation, pair f o r m a t i o n , ririd female c h o i c e i r i (:lioi-loplrrrgrr iliridijrrsr-irrlii (De(;eer) ( O r t h o p t e r a : Acrididne). Joiirnal of 01-tliopter,i liesenrch 4 : 1 3 1 - 1 4 2 . O p i t z S., 1 3 3 6 . bloveiiieiit s t ~ ~ d i e s in t h e rufous gr;isshopper,
(;onrplior-~rirs f-lrfils ( I , . ) . Snltnbel, 1,eideii 16: 1 3 - 2 1.
O p i t z S., Kiihler G . 1 3 9 7 . Wie iiutzen 1:eldheiischreckcri (Caelifera:
(:oniphocerin,if) ihr Il,ibit,it?. Mitteiluiigen d e r I>eutscheri (;esellschaft für nllgeiiieiiie ciiid ,ingewandte Ilritoiiiologie 1 1 : 6 1 3 - 6 2 2 .
O p i t z S., Köhler C;., Mabelis 11. 1 9 3 8 . 1,ocal niovement o f t h e gr;isshopper Clloi-tlrippirs ptrr-rillr~lirs ( Z e t t . ) . I'roceeclings o f experimeiital ;ind ;ipplitxi t : n t o m o l o g v o f t h e N.I:.V. Anisterdniii 3 : 5 3 - 5 8 .
Orshaii L, I'ener ki.1'. 1 9 7 0 . I<epe,ited revcrsril ot' t h e reprodiictive ciiap,iiise by pliotoperioti aiid tciiipcrature i i i rn,iles of t h e gr,isshopper Oc~riioporlrr rriirririirr. I:iitoiiioIogia tlxperiiiieiitdlis e t i\pplic,it,i 2.5: 2 19.226.
1)t~iiei hl.l'., 01-sli,iii 1.. 1 0 8 0 . Revc~i-sible i-t.pi-od~ic.tive cii,ip,iiisc arid iiiteriiiedi,itc st,itt,s b e t ~ v e e n di,ip,icisc ,ictivity in iii;ilc Ot~riioporlri rrritritlf~r gi-~isshoppcrs. I'Iiysi0l~)gic~1 1:1itoiiio1ogy 5: 41 7 - 4 2 6 . Picliford I<., (;illott C;. 1 0 7 2 . (:oiiplirig betinviour of t h c iiiigi-atory
gr~isslioppci- ,211~lrirroplirz sritigirirripcls (Ortlioptt'r,i: ,-\cridid,ic).
(:,iii,idi,iii 1:iitoiiiologist 104: 8 7 3-879.
I<t~irili,irclt I<. 1 0 9 8 . Liir l.i,ibl,igt~1>1-~ifc1-t~1izder kle~iiic~n (;olcisclii-ccAc l.'~rilr,v.c~ir(r l~i-~i(.lr~ti~re,i.~i ( O e - s k ~ y ) - 1'1-t,iI'i~ici vcrsiis ~ ; c \ v i c l i s l i ~ i ~ i ~ . , \ r t i c t ~ l , i t ~ i 13: 2 8 - 37.
I<e~iiili,ircit K , I<tililci-, ( ; . , Sc.Iiiiiii,itlici- C:. I O O O . I't.iii,ilt.s of ilic.
g r , i \ s l l o ~ ~ ~ > c , l - ( ~ ~ 1 0 r - 1 ~ r 1 ~ ~ ~ l 1 1 . c p(ll(l//l~/ll.\ ( L c t t . ) cio I l O t l-t~111<1tt~ t-01-
t i . t > \ I i spt,i-~li. l ' i - o e - t ~ ~ ~ ~ l i ~ i g \ 01 t l i t % I<o\,<ll Soc i t % t \ , 0 1 I o i i c i o ~ l , St>i-itt\
li J ( > ( > : 2 0 0 < - 2 O O O .
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I , i ~ i ~ i s c - I i ~ i ~ t s ~ ~ l ~ ~ i i i ~ ~ i g (7c~its~-lii-ift I U I - ,iiigt~v~iiicitt~ OLologit,).
I<it,dt, K. 1087. .\ coiiip,ii-,iti\,t' \ t i i c l > ' ol- i i ~ ~ i t i i i g I>c.li,i\.ioiir i i i \oiiit3 iicotropic-,il gi-,issIioppt~i-s (,\e-i-icioiclt,~i). l'tliolog>, 70: 2 6 5 - 2 0 0 , 5 , i i i i i c t ~ I . IOOS. I'opiiI,itioiisgcl';ilii-diiiigs,iiiCiI~~sc ,111 t.iiici- I I t ~ ~ i s c - h r c c l ~ t ~ i i , i r t - h i c t l i o d t ~ i i , ciiipii-i\cIic (;I-iiiidl;igt>ii iiiid ,k1ociclll~ilcii11ig l>ci L5ti1~~ol)oi/11-~~.~ l ~ t l ( ~ o t ~ l . \ ( l ' c ~ ~ i / t , ~ - ) , ((~cjttiiigcii.
(;ii\~iIlit~i-), I 4 6 pi>.
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Srivastav;i I'. 11. 1 0 5 6 . Stlidies oii tlic col->ul;itoi-V ;ind cgg-lnyiiig 11,ibits of l f i r ~ ~ - o ~ l ~ ~ p l i ~ r s rligl-ot-cplrltirs 1301. ( O r t h o p c r , ~ , i\crididne).
Annnls o f t h c I:iitoiiiologic,il Society of Aiiicricn 4 0 : 167- 1 7 0 . Srivnst,iva I). I > . 1 0 5 7 . Ohscrvntioiis ori t h c brecciiiig Ii,ibits of
Atrlic-ic~~~or-pllci ~ t - ~ ' t l i r / i l i i i ( I : . ) tlie t o b , i c c o g i - , i s s h o p p ~ ( 0 1 - t h o p t e i - a : Acridid,ic). Aiiii,ils 01' tlic ciitoiriologic,il Society of Anicrica 5 0 : 1 5 - 2 0 .
Wngiier C;. 2 0 0 0 . Eiiit. Popiil,~tioiisgef~lii-dciiigsnii,ilysc dci- Rotflügel igeii iitiiaricischrecke 0c.rliporitr ~cjrttirr tzic-rr ( I . c ~ t r . 1 8 0 4 ) (Caelifern: Acrididne). Articul,it,i 9 : 1 - 1 2 6 .
Wliitnian D.\.Y. 1 9 8 5 . 1,nhorntot-y biology of 7'rrr)nropotlrr cr/~tr..s ( O r t h o p t e r n : Acrididac). Jourri,il of tintoiiiologic,il Scieiice 2 1 : 8 7 - 9 3 .
Wickler W., Seibt 11. 1 9 8 5 . lieprodcictivc bclinvioiii- i i i %onoi.clt.iis cjl~grins ( O r t h o p t e r a : P y r g o ~ i i o r p h i d ~ i e ) with spcci,il rcferenct, to iiuptial gift gii,irding. Zeitschi-ift füi- 'I'ierpsychologie 6 0 : 2 0 3 - 2 3 3 .
