Mating behavior of Psammotettix alienus (: Cicadellidae) Maja Derlink, Isabelle Abt, Romain Mabon, Charlotte Julian, Meta Virant-Doberlet, Emmanuel Jacquot

To cite this version:

Maja Derlink, Isabelle Abt, Romain Mabon, Charlotte Julian, Meta Virant-Doberlet, et al.. Mating behavior of Psammotettix alienus (Hemiptera: Cicadellidae). Science, Wiley, 2018, 25 (1), pp.148-160. ￿10.1111/1744-7917.12379￿. ￿hal-02628561￿

HAL Id: hal-02628561 https://hal.inrae.fr/hal-02628561 Submitted on 26 May 2020

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating articleThis protectedis by copyright. All rights reserved. 10.1111/1744 Science but has yet to undergo copy This anis Accepted Article that has been peer gendersBoth spontaneously emit species determine ofvibrationalsignals therole intra in investigated the of matingbehaviour includeshould methods on theuse The Abstract Mating behaviour of O * [email protected] A Correspond T Running title: Maja Derlink 34398 Montpellier : : twoThese authors contribute equally to work.the 1 foritle authors - riginal article National Institute Biology, of Department of 4, 49 Mnplir France Montpellier, 34398 54K, 1000 Ljubljana, Slovenia W heat virus dwarf

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating (Heady mating leafhopper detailed more and vectors leafhopper of importance economic Tishechkin 200 (Weintraub diseases plant of vectors important most the among are and species described 000 22 than more with al. from Substrate Introduction virus Keywords: mating viruses transmitted by vectors. insect essential forapproaches trying new indeveloping understanding of performance vibrational songs, Although with male receptivefemale with wingwing flapping were andrecorded vibrations from the substrate also immediately prior to pulse trains time. regularly inpulsetrain pulse anddifferprimarily repetition pulseduration repeated trains and ,

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Females conspecific preferred male the calling song. & ibrational order Hemiptera order - - results suggestthat results borne vibrations play an important role in mating behaviour of many insect pests insect many of behaviour mating in role important an play vibrations borne

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating ( wheat a in 1960s the in time first the for described disease, dwarf wheat The oat. and barley wheat, in symptoms yellowing family the which of pathogens different by infected Ti in as well as signals, emitted of complexity and structure repertoire, the by emitted stereotyped coordinated, a in thanthosemales emittedcomplex by females are more that (a) showed at describingstudies aimed t environment. the in unpredictablyscattered are that females receptivethe find to order in Groot 2011 Cocroft,& Luca De 1991; Nault,& (Hunt Auchenorrhyncha many in foundstrategy „fly/jump/walk the with associated is formation pair the in male the of role leading The 1991; established been has contact with begins formation signals Kuhelj (Mazzoni shechkin In eel, motn cos n h wrd o bt hmn n dmsi aias cn be can , domestic and human both for world the in crops important Cereals, Hunt et al. et et al Cica Geminiviridae n the and et al et , a , 2012 ,

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating an increased attention ( ofhealth,years theexploitation vibrational management pest signalshas in inrecent received Due to environmentallycontrolfriendly practices can developed be on mating behaviour speciesdelimit (Schlick and identification of challenginggroup behaviour lacking. genusis inthis the informationaspects onother of 1868) and 1758) vector (Zhang reproductive behavio chemicals theleafhoppers vectors. against area the on anti based is WDV of against used absence strategy protection the main the currently and sources resistance genetic of lack the to Due Abt in (reviewed literature the in documented poorly still are pathosystem (Lindblad grassland plant Psammotettix from transmitted is WDV production. (Vacke Republic Socialist The main objective present ofthe was study ,

P. confinis increased of awareness P. poecilus et al Dloehlne, oatc insect holartic (Deltocephalinae),

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating 2 a on literature ( (Sadeghi Henry virus While day/night 16/8, RH, 40% 50 populations leafhopper These progenies. selected [ methods France South grassland and vi and North in fields cereal various in 2013 and 2012 years Insects Materials Methods and performance. information whether included virus Polaj WDV × Based on Based Adult Leafhoppers collected in collected Leafhoppers

nar 80 cm) in a temperature controlled chamber (24°C during the day/20°C during the night,duringthe day/20°C duringthe temperature chamberacontrolled(24°C in cm) 80 , -

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w ef eel plantle cereal leaf 02, WDV 2002), et al ] et al. et [

reviewed in e Splmnay Material Supplementary see P. alienus P. n WV aly stain barley WDV and - free .

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neyards in West Slovenia inAugust Slovenia West 2014. neyards in 2000) according to according 2000) and WDV and sanitary status of status sanitary

and viruliferous individuals in order individualsand in toobtainsomepreliminary viruliferous ;

viruliferous statusof insectsviruliferous Polajnar - leafhoppers were collected using a sweep net in June and September in September and June in net sweep a using collected were leafhoppers DOI :10.1111/1744-7917.12379 b Abt Abt Comment citer cedocument:

Psammotettix t period).

& - oee b a micro a by covered w

Jacqu the field in in field the et al leafhoppers were maintained on wheat on maintained were leafhoppers ., Fresh potted plants were added to the cages every month. month. every cages the to added were plants potted Fresh o

the

the 2016 [ t , WDV

2015) ouain wr mitie o bre cv. barley on maintained were populations gravid female gravid ] host host ) 2013 were were 2013 ; Korinšek ; Korinšek This articleis virus , - b .

specificity of specificity ]

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perfora re n vrlfru (D wet strain wheat (WDV viruliferous and free has aneffect on were reared in large plexi large in reared were et al. maintained s protected by copyright. All rightsreserved.

collected in 2012 (determined by PCR by (determined 2012 in collected e clohn bg in bag cellophane ted , 2016

the WDV strains strains WDV

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In also thestudy we the

pre cv. - copulatory mating Sunstar - glass cag glass described in the in described

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating software 2.0 pro Edit Cool and Singapore) (Creative, card sound ZS 2 Blaster Sound a using (PDV vibrometer laser leafhoppers from escaping. vial plastic transparent a with covered was plantlet the of part lower and upper the separating and with covered was hole the and tripod woodenmade custom adiameter) of cm (10 circularaopening within positioned was plantlet The substrate. artificial moist with filled jar a into upright placed 40 at height (approximate Recording behaviour and signals vibrational determined method by PCR above described as Material). ofsanitary status adults. virgin obtain to order in above described as reared and plantlets cereal young grav 330 the of larvae. and eggs presenceof the for weredailymonitoredrearing individual systems chamber controlled Individuals collected in Slovenia in collected Individuals irtoa sgas ee registered were signals Vibrational % – Experimental set Experimental 50% RH 50%

overlapping incised paper circles surrounding the plant thus creating a platform a creating thus plant the surrounding circles paper incised overlapping id females collected in field, u field, in collected females id ) the 330 330 the . Individual plantlet Individual .

14 experiments. the in used until s ecie above. described as DOI :10.1111/1744-7917.12379 - Comment citer cedocument:

100, Polytec, GmbH, Waldronn, Germany) and stored in a computer a in stored and Germany) Waldronn, GmbH, Polytec, 100, cm - gravidfemale up

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were were h plantl the 10 larvae were transferred individually on individually transferred were larvae 10

maintained in individual rearing systems rearing individual in maintained a h sntr sau o te insect the of status sanitary The protected by copyright. All rightsreserved.

(de Groot Groot (de plastic vial filled with vermiculite and vermiculite with filled vial plastic t 1 et ro at )

m eo te platform the below cm om temperature (20 temperature om et al. et wheat ,

2012) cv. Sunstar cv. . The upper part upper The . – to 25°C

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating with stimulated were females single of activity periodstime USA). Ithaca, Ornithology, of Laboratory (Cornel aut were signals vibrational recorded and platform the above plantlet the to placed was leafhopper single A free, hours) 08:00 and 18:00 (between ( individuals Male emitted vibrational registered signals at ofrecording. the point program. Pro2 Edit Cool by card sound mentioned above the via computer the from driven was exciter vibration Denmark Nærum, Kjær, & Brüel 4810, (Minishakertype exciter vibration a of head the the via platform the below) (see recordingbelow theplatform. 1cm location the at stem the on placed was tape reflective of piece small a reflectance, increase to order USA) Phoenix, Software, (Syntrillium aln sgas we signals Calling set The S one and female beun experiments ubsequent

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating our were from taken stimulatorysequence inthe male songsincluded All intervals. 30 ssilent by separated order random a in times five repeated was song each sequence, stimulation the advertise male and songsfree (WDV and ofvirus viruliferus b of preference the test To Female preferences virus Vibrati lineages.separate eggs laid had they after and plantlets wheat young duration duration, viruliferous included we 12 vibrational signals, stimulation 0 sequence wa recordings ] . 01 wr placed were )

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating the to experiment the of beginning the from XM2)camcorder fittedwith DM (Canon with together sequence mating ma however until on placed were virus pairs Eighteen Mating behaviour pulse train) latency recordings call preferred the as call male the minutes, several starting the trial each for ( of call male the while plantlet, wheat a on registered of signals Vibrational male. calling the from away cm 3 than more not above mentioned vibrometer laser the with recorded been had and recordings of library Trifolium pratense Trifolium e n te female the and le In all experiments all In - the end of end the re efopr, s el as well as leafhoppers, free tm fo te beginnin the from (time ,

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating Terminology and stopped theplayback30 sintervals.for s ± 0.56 = SD ± mean (duration: recording calling female recorded male b males, Rivalry determine they whether deposition laboratory the in eggs laid female) the of location betweencourtship female durationand (time of thebeginning the thelocation ofcopulation). the to duet a of onset the from (time duration copulation of duration the the of beginning the from (time latency timulatory sequence wa timulatory sequence female), msin f h frt ae r eae us tan o the to train pulse female or male first the of emission In addition In , 5 pairs of pairs 5 , 5 To

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating in used sequence stimulatory to responding females Wilcoxon nonparametric Shapiro with normality for checked virus and field the from individuals par these sex each in Similarly, progeny). (WDV leafhoppers viruliferous all between compared were sex each in frequency) dominant and time repetition th individual each for us cumulative the night, the throughout leafhoppers single of activity calling ranges. with together medians as presented are data frequency Dominant from leafhoppers signals immediately emitted tocopulation were attempts prior later inter was bout the of end the regular, always not was time repetition form. or sequence al. et ese periods. ed to represent to ed

The parameters of calling songs (pulse repetition time, time, repetition (pulse songs calling of parameters The numbers the with together deviations, standard and means as presented are Data fin of sound of parcel homogenous unitary a as defined was Pulse - , pul close

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either of rank sum test sum rank the activity female preference test preference female conspecific aig sequence mating - b) and virus and b)

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were compared with compared were and accordingly analysed either with either analysed accordingly and pulse train duration, pulse train pulse duration, train pulse recorded termed termed t determined at the start of the of start the at determined

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating 3 Table 2, (Fig. p regular less and train the within time repetition Male calling song (Tabletime 3,Figs. 2 pulse the in was song rivalry male and song calling female of consisted males arecalling am, 6 and midnight between period the in low is activity calling female while that suggests below). (see s) 8522 and s spontaneously signals General Results statistical analysesconducted wereversion usingR their and using calculated females were songs calling the male for preferences of status sanitary between correlations while proportions, given All recorded vibrational signals vibrational recorded All recordings Overnight Male calling song is characterized by short pulse train duration train pulse short by characterized is song calling Male

observations a series of series a Although calling activity varied between individuals, the emerging pattern pattern emerging the individuals, between varied activity calling Although than ). Pulse trains span broad frequency range frequency broad span trains Pulse ). throughout

in males in ,

. DOI :10.1111/1744-7917.12379 5 T Comment citer cedocument: ). ). single pulses pulses single he

of single individuals individuals single of overall calling time time calling overall

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during the night was longer longer was night the during eeld ht oh ee ei vibrational emit sexes both that revealed was a result of longer pulse train durations train pulse longer of result a was le train ulse 3.2.1 the

protected by copyright. All rightsreserved. earlier stages of the mating sequence mating the of stages earlier .

two and have and train train repetit - tailed Fisher‟s exact test. exact Fisher‟s tailed duration and pulse and duration o tm wti te song the within time ion clear harmonic structure structure harmonic clear the the , highly regular pulse regular highly , male calling song, song, calling male in female in

repetition repetition (13020 (13020 All 12

Version postprint 305 304 303 302 301 300 299 298 297 296 295 294 293 292 291 290 289 288 287 286 285 284 306

behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating t Test, between significantly differ not did song ( variable highly however, range frequency broad span trains pulse pul regular more within time repetition pulse regular less and longer duration, train pulse longer Female calling so characterized (Table time train repetition S2 bypulse long of eight However, longer significantly was time repetition train pulse a from males free longer time train pulse repetition males the songs was with - Test, ae aln sn dd o dfe sgiiaty between significantly differ not did song calling male dominant frequency dominant P ( P

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se train repetition time (Fig time repetition train se t ng the progeny produced in the lab the in produced progeny - Test the 10.5

dominant frequencies dominant harmonic structure harmonic DOI :10.1111/1744-7917.12379 nine F1 males tested belonged to the same lineage that appears that lineage same the to belonged tested males F1 nine Comment citer cedocument: , P ±

between 13 between

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Version postprint 328 327 326 325 324 323 322 321 320 319 318 317 316 315 314 313 312 311 310 309 308 307 329

behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating aa male female and regularly exchanged in pulse trains male. by train pulse emitted 1 = latency was signal vibrational registered first the copulation, with pre and courtship approach, recognition, movie Supplementary the in shown maximum duration sequence mating the between Mating behaviour = viruliferous d differsignificantly( not femalesdid viruliferous song. male ecific females these of e n N h Female preferences observe anyobserve us one of the females responded to responded females the of one the 1 )

, however the number of, however the thetested number individuals females started In general, observations did not reveal reveal not did observations general, In conspecific male calling song conspecificcalling male In the

11 min 101 = 9

9 ± 25 ± 9

correlation virus tested P. alienus P. Preferences

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in preferences for the male calling songs emitted byemitted songscalling male the preferencesfor in males

15 = (excluding copulation) copulation) (excluding The first vibrational interaction was a coordinated duet in which in duet coordinated a was interaction vibrational first The for

during the females the emission of calling clearly associated songwas wit of theemission females

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givenproportions Broadly protected by copyright. All rightsreserved. P. confinis P. . an P. confinis P. n 1 In

M a behaviour mating in differences - , male callin male 7

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Version postprint 350 349 348 347 346 345 344 343 342 341 340 339 338 337 336 335 334 333 332 331 330 351

behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating 1 a of concluded ( with copulation sequences mating all in registered were flapping wing by produced signals Courtship they flapping substrate the in recorded signals vibrational (Fig. leg D however usually associated with however calling, stopped courtship stage females most lasted for (Fig direction same the in facing female, they calling stopped coordinated precisely less 5 female. 1 uring the uring

min, min, When When A C only sporadically emitted single sporadicallyemitted only pulse trains have have opulation attempt opulation tr irtoa co vibrational fter - s og us tan (buzz) train pulse long s 3 On average, the approach stage lasted 2 lasted stage approach the average, On , , N

pn ra feuny ag and range frequency broad span S1 we were not able to determine whether they were produced by produced were they whether determine to able not were we ±17minutes 34 male

a the 7). = close C la hroi srcue wi structure harmonic clear )

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. tc w ntact w continuous emission ofcontinuous emission pulsesemitted withirregulartime; repetition Males were signalling only when they were not walking and often and walking not were they when only signalling were Males ( and ere minimum minimum

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= 10min, = 2; (Fig. This articleis h oiat rqec bten 1 between frequency dominant th S1 . maximum

ihn the within continued to continued ( Table 3, Table most malesmost rubbing

A , S1 maximum 4 the , . D

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= 25 The latter latter The 8

protected by copyright. All rightsreserved. On average, the the average, On Fig. and

main min ( min emit calling songs songs calling emit continued ,

= 64min tapping the fe the tapping N 4 ). Pulse trains trains Pulse ).

rqec band frequency minimum = 16 resulted in distinct courtship distinct in resulted vibrational signalvibrationalcomposed ).

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating and masking signals emitted male which determine to possible not was it leafhoppers, 2, = callingsong male the dominant frequencies between 400 regular. not is time repetition train by characterized parameters spectral and temporal by emitted train pulse the of part least at overlapped song, calling female the in trains pulse between interval the in signal this emit WDV a included that pairs two in of type another emitted males song, p a with presented were that pairs male tested 25 of out seven In Rivalry behaviour stimulated that showed laboratory the in of88 s copulation was 522± flapping Results of recordings of recordings of Results which in trials all In maximum pairs of virus of pairs and vibrating and with the

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emitted bya emitted DOI :10.1111/1744-7917.12379 one Comment citer cedocument: akn signals masking the

wing - b

virus ( putative masking signal masking putative N male

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4 w (Fig. 5) (Fig.

.

mated in the field and have and field the in mated pn bod rqec rne with range frequency broad spans and Fig. S1 Fig. and ere

the the male Although males usually started to started usually males Although

recorded, female tre signalling started . and a WDV a and la These signals were recorded were signals These yback of the female calling calling female the yback of the E

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Version postprint 395 394 393 392 391 390 389 388 387 386 385 384 383 382 381 380 379 378 377 376 375 396

behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating and conditions field under strategies reproductive and systems mating their about little very & also Dober behaviour such leafhoppers, in rare females maybe high t male, the by initiated formation also Mazzoni (Claridge signals calling male of loosely coordinatedduet simple in species leafhopper signals. vibrational alienus other in as that show study present the of Results Discussion

P. alienus P. Žežlina For leafhoppers, it is it Forleafhoppers, by ecological aspects of the environment, including population density ( density population including environment, the of aspects ecological by hw o lvl o sotnos calling spontaneous of levels low show spontaneous spontaneous let

structure of male and female calling song calling female and male of structure

is & et al. et ,

(Saxena

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2009; de Groot de 2009; equally likely to trigger the exchange of vibrational signals. vibrationalof triggerexchange the to equallylikely ). ept cnieal eooi iprac o lahpes w sil know still we leafhoppers, of importance economic considerable Despite i eaeclig ciiyi oengt recordings overnight in activity calling female ) so far so ,

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aken together, aken Comment citer cedocument: Kumar . the pair formation is initiated by either byinitiated is formation pair the studied in detai in studied

generally accepted that mating sequence is initiated by the emission emission the by initiated is sequence mating that accepted generally Mazzoni Mazzoni soitd with associated ,

et al et 1984 ,

1985; Hunt 1985;

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2010). lo revealed also he unique features of vibrational co vibrational of features unique he &

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and msin of emission protected by copyright. All rightsreserved.

, Cicadellidae ( common in common

one older study older one iii 1991; Čokl Čokl 1991; and can can and ) progression from a patterned to a to patterned a from progression )

motn dfeecs with differences important mating systems are shaped in part in part are shaped systems mating gender; species rarely , suggest that in that suggest planth

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating the in signals vibrational trains pulse c contrast In 2014). Doberl (Čokl females by emitted those than complex more are males by emitted signals identification in that location its determine identity the shou and mate to readiness and presence their advertise individuals when behaviour mating refractory period that and multiply(H mate field femalescan leafhopperthe in observedthat whether clear not is it conditions receptivity. sexual lifetime. that accepted them locate cannot of strategies different communication inAuchenorrhyncha. provide should studies future omplexity aln sgas r long are signals Calling I n contrast to males to contrast n et

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2003; Percy 2003;

. In particular, female calling song in this respect more respect this in song calling female particular, In . n i bt gnes aln sns r sml ad omd y eual emitted regularly by formed and simple are songs calling genders both in and (species and gender) of the signaller and also provide information necessary to necessary information provide also and signaller the of gender) and (species of the partner. the of in leafhoppers in ,

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2008; Mazzoni Mazzoni 2008; & & , However, s However, more insight into mechanisms that are involved in the evoluti the in involved are that mechanisms into insight more

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2005). 2005). 2003 ,

2009; . alienus P. a ms lkl mt ol once only mate likely most , protected by copyright. All rightsreserved. ). natural calling

This is in agreement with o with agreement in is This oersosv again responsive come de Groo de

P. alie P.

situation. situation. t al. et mating song t

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Version postprint 440 439 438 437 436 435 434 433 432 431 430 429 428 427 426 425 424 423 422 421 420 442 441

behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating mechanoreceptors detect that also vibrational signals. leg same the stimulate likely movements leg since walking, while signals vibrational female detect to inability from or walking, and production signal in involved muscles of activity the t inability from result may stage approach the during observed duet coordinated alienus ( 3 the test of ability localization to enough complex not was study current the in (Virant used environment experimental difficult is partner the locating communication, successful for D partner the by emitted oscillator endogenous the resetting needed are studies (Polajnar leafhopper the in sequence mating the of stages early the in described structure al ( Cicadellidae other in described signals vibrational female Consequent bugs pentatomid the (e.g. partners between Polajnar – uring the approach stage approach the uring ., 4

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localization. localization. 2008 2009; Derlink , ienus planthopper

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- protected by copyright. All rightsreserved. dwelling insects relying on vibrational on relying insects dwelling rhythm

uh oriae de rsls from results duet coordinated such et al. et (Mazzoni (Mazzoni to some extent extent some to e. g. e.

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating however leafhopper satellite use also males rival behaviour female toapproach the male. duettingwithanother whether and disruption to vulnerable is duet coordinated to or duet female determin 2015) ( species leafhopper other in described been have signals males vibrational emit signals visual and/or chemical signals. however, movement wing that show results Shaw 1974; close the during been (Polajnar completed location and recognition the after signals demanding more energetically that hypothesized been has it costly(Kuhelj is signalsvibrational of production leafhoppers, In behaviour. al. ( sequence mating the of stages earlier the ,

2009), in 2009), u rsls ugs ta i te rsne f rcpie female, receptive a of presence the in that suggest results Our As Preliminaryr .

The data obtained in the current study by playback experiments are not sufficient to to sufficient not are experiments playback by study current the in obtained data The in some other Auchenorrhyncha that emit relatively simple vibrational signals during signals vibrational simple relatively emit that Auchenorrhyncha other some in , our results also also results our , wehr hs pttv msig signals masking putative these whether e the role of these signals is not clear since wing since clear not is signals these of role the does not have have not does ,

1976; P. alienus P. - range stage range esults mask female calling song. song. calling female mask Saxena DOI :10.1111/1744-7917.12379 Comment citer cedocument: obtained in obtained

more complexity has been introduced at the later sta later the at introduced been has complexity more a significant a et al. show & s

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Virant itnt and distinct

repeti r used are et al et several - protected by copyright. All rightsreserved. Doberlet flapping ti Mazzoni Mazzoni . , on

1986; Hunt 1986;

stereotyped vibrational signals; signals; vibrational stereotyped leafhopper species ( species leafhopper y mtig oe ope and complex more emitting by

o irp a oriae male coordinated a disrupt to time in male calling song song calling male in time - & female r mance the viruliferous status of theof status viruliferous the

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2009 , of

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Version postprint 486 485 484 483 482 481 480 479 478 477 476 475 474 473 472 471 470 469 468 467 466 487

behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating 2013) (Eriksson management vibrational of re Until agriculture. sustainable in challenges main the of one is pesticides potential effects account. into co ability the affect turn in can that properties transmission physiological) and (phenotypic plant host feeding al et (Kuhelj signals vibrational emitting while tissue plant into inserted stylets feeding their keep may plants (Kuhelj demanding energetically mat with associated Behaviour 2015). (Ingwell physiology and behaviour on infection virus. the by testing worthwhile the individuals tested of numbers virus the in than viruliferous in longer significantly n specifics. . eeoig e apoce i ps mngmn ta cud ep o eue h ue of use the reduce to help could that management pest in approaches new Developing ,

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating alienus. Fig. 4Representative courtship pre (a) and stage. Fig. 3Coordinated male (M)

DOI :10.1111/1744-7917.12379 Comment citer cedocument: –

female (F) duet in Psammotettix alienus registered the during recognition

- copulatory (b) signals emitted by This articleis

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating Inset shows expanded section ofthe signal. Fig. 5Putative masking signals emitted (MS) by male Psammotettix alienus. (F) female pulse trains. in (a)Inset showsexpanded section ofthe signal. DOI :10.1111/1744-7917.12379 Comment citer cedocument:

This articleis

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behavior ofPsammotettix alienus(Hemiptera: Cicadellidae). Insect Science,25(1), 148-160., Derlink, M.,Abt,I.,Mabon, R.,Julian, C.,Virant-Doberlet, M.,Jacquot, E.(2018). Mating

DOI :10.1111/1744-7917.12379 Comment citer cedocument:

This articleis

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