Size and dispersion of urticating setae in three species of processionary Edoardo Petrucco Toffolo, Daniel Zovi, Chiara Périn, Paolo Paolucci, Alain Roques, Andrea Battisti, Helmuth Horvath

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Edoardo Petrucco Toffolo, Daniel Zovi, Chiara Périn, Paolo Paolucci, Alain Roques, et al.. Sizeand dispersion of urticating setae in three species of processionary moths. Integrative Zoology, Blackwell Publishing, 2014, 9 (3), pp.320-327. ￿10.1111/1749-4877.12031￿. ￿hal-02633992￿

HAL Id: hal-02633992 https://hal.inrae.fr/hal-02633992 Submitted on 27 May 2020

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article cite this article as doi: 10.1111/1749-4877.12031 which may lead tonot been through the copyediting, typesetting,paginationdifferences and proofreading process, Thisbetween has article publication been acceptedfor undergone and full reviewpeer has but this version and the Version of Record. Please [email protected] Fax +39 049 8272810 Tel. +39 049 8272804 University Padova,of DAFNAE-Entomology, Agripolis, 35020Legnaro Italy, PD, Andrea Battisti Corresponding author Roques Edoardo Petrucco Toffolo Petrucco Edoardo Size and urticatingdispersion of in setae speciesthree processionary of moths Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 . Institut für Experimentalphysik, Universität Wien, 1090 Vienna, Austria, 3. INRA, UR0633, Zoologie Forestière, 2. 2163 Av. Pomme de Pin, Ardon 45075 University of Padova, 1. DAFNAE-Entomology, Agripolis, 35020 Legnaro PD, [email protected] Orléans, France, [email protected] Italy,[email protected] 2 , Andrea Battisti Comment citer cedocument: 1 , Helmuth Horvath 1 , Daniel, Zovi

1 , Chiara Perin 3 1 , Paolo Paolucci Paolo , 1 , Alain 1 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Key words: bio-aerosol, , pollution,seta dispersion,seta size, to the infestedtrees and atlonger distance. to understanding the importance of the processionary setae as air pollutants, both close importance of larval setae against natural enemies the of processionary moths, as well as counterparts. This information is relevant afull to the understand defensiveof consequence in dispersion,as smaller setae can spread 5 times further than their bigger integument fields where they The occur. difference in the seta size has important T. pinivora state.Setae are highly variable insize, with bimodal indistribution in theairborne thesetae of properties dynamical of analysis (ii) setae; of length and potentially helpful in understanding setaeof dynamics inthe (i)environment: diameter Thaumetopoea pityocampa,T. pinivora here identify the traitsmajor of the setae threeof species ofprocessionary namelymoth, knowledge about morphological traits of the setae andtheir release by the larvae.We processionary some studies provide detailed information about the setae, particularly those of the Although distress. respiratory and conjunctivitis, dermatitis, causing outbreaks, during pests forestof and urbantrees andconsequently may threaten human animaland health released against vertebrate especiallypredators, insectivorous Afewbirds. species are Larvae of the processionary moths of the Palaearctic region bear urticating setae that are Abstract C91J10000320001. - 2010 grant Padova of University by the and 'URTICLIM’ project 013 n°07BDIV of ANR framework the in Agency Research National French by funded was work This Simonato, M.Stastny C. andVillari forreviewing an earlier version theof manuscript. especially F. Goussardand J. Garcia. Weare also indebtedto S. Larsson, D. Lees, M. We warmly acknowledge all the that people have provided help for this work and Acknowledgements Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 ; in these two species short and long setae are interspersedwithin the Comment citer cedocument: Thaumetopoea pityocampa

(Treitschke) (Denis & Schiffermüller), there is little , and T. processionea T. pityocampa T. (Linnaeus), and 2 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article that the larvae may actively open the integumentmirrors when disturbed. Werno and larval colonies treeson as well as at pupation sites. andmany are left with the oldskin, so that they occur in a high number in tents used by and pointed barbs directed distally (Lamy 1990). These setae are renewed ateach molt setae/mm60,000 urticating setae areproduced from the 3 The al. (1987). et Novak and (1982), al. Lamy et (1963), by Démolin continued (1900), Setal production andmorphology was firststudied in the pioneering work Fabreof Aparicio etal. 2006; Santos-Magadàn et al. 2009; Rodriguez-Mahillo etal. in press). recently shown in numbera ofcases (Vega etal. 1999; Moneoet al. 2003; Fuentes Gottschling & Meyer 2006, Vega et al. 2011). In addition, an allergic response has been conjunctivitis, andrespiratory distress in humans (Vega etal.1999; Maier etal.2003; (Matthioli Indeed, 1568). there arenumerous reported cases of resulting dermatitis, ),have been well known since ancient times for their ‘burning’ effect this stage there areabout 1,000,000 setae/larva in molts until the last larval instar, in which they on eightoccur abdominal segments. At owing to their property to reflect light. These mirrors increase in number with larval the species, on integument areas ofthe abdominal tergites that are called‘mirrors’ processionary moth al. 2003). the environment, often far away from theinfested trees (Werno& Lamy Maier1990, et (Specht et al. 2008), setae are readily dehiscent from the integument andcan disperse in to cause the reaction (e.g. the larvae ofSaturniidae, Megalopygidae, andLimacodidae) to spines that are firmly attachedtothe integument andrequire a contact with the larva health (Specht etal. 2008; Hossler 2009; Mullen 2009; Battisti et al.In 2011). contrast human to threat serious a pose also which but 1984), &Kumada Kawamoto 1971; considered to be adefense vertebrateagainst predators (Gilmer 1925;Pesce &Delgado Some Lepidoptera, aslarvaeadults, or carry urticating setae that are generally Introduction T. pinivora, Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 The release setaeof by the larvae was explored by Démolin (1963), who showed The urticating setae of the larvae of processionary moths, namely the pine andthe oak processionary moth 2 Comment citer cedocument: ; setae area few hundredmicrometer long, with a sharp proximal end Thaumetopoea pityocampa,

rd larval instar or 4 T.processionea the northern pine processionary moth T. pityocampa rd larval instar, depending on (Lepidoptera, , with a density of 3 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article To obtain of samples urticating setaethe of three and methods Materials domestic mammalians. may have an impact on non-target vertebrates exposedto them, such as humans and could be functional to improve the protection of colonies from predation, and in turn airborne stateaccording theirto size.We hypothesize thatthe different size of setae investigate: i)the variation insize of setae; ii) the dynamical of properties setae in the that canbe useful to clarify their importance as environmental hazards.Specifically we preservationafter and treatment with temperature as high as 110°C. of years 12 after power irritating their retain insects that dried demonstrated setae in the environment even after exposure to ratherextreme conditions: Hase (1939) Meyer 2006; Vega et al. 2011). An aggravating factor is the long persistence of the with airborne setae (Maier etal. 2003; Fuentes Aparicioetal. 2006; Gottschling & setae could occur. This isimportant mostbecause reactions inhumans are associated on temperature and windspeed; even at a distance of 2 kmconsiderable numbers of maximumconcentration ofsetae spread atadistance of 200 tometers, 500 depending showedthatthe Thephysical to prevailing authors properties inrelation wind. 11°11’E, m 477) for natural conditions. Samples were collectedin 2008 from Tregnago(Italy) (45°30’N, larvae in the last instar from rearing in Padova University campus or from colonies in 18°16’E, m 7) in 2009 for in 2009 7) m 18°16’E, 10°47’E, m 245) in 2009 for in 2009 245) m 10°47’E, dispersion model for the oak processionary moth, decreased moving away thefrom infestedforest. Fenk et al. (2007)developed asetal progressively setae of abundance withthe traps, inpollen setae found (1990) Lamy Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 In this paper we address a few traits theof setae of three Comment citer cedocument: T. pityocampa T. pinivora T. processionea

, from, the island Gotlandof (Sweden) (56°56’N, , and, from CaprinoVeronese (Italy) (45°35’N, . Thaumetopoea T. processionea Thaumetopoea species, we collected , based on their on , based species 4 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article processionea of setae is done with the aerodynamic diameter d capacity, weanalyzedtheir aerodynamic main A features. descriptive characterization surrounding): settling velocity (resulting from the force of gravity theand friction by the air molecules under consideration.The measures aerodynamicof diameter allowcalculating the that awater droplet would have in order tosettle with the same velocity as the particle release, c the wind velocity,v Hitachi TM-1000) (n= 108 for stereomicroscope andwere measured using ascanning electron microscope (SEM under mirrors from forceps with extracted randomly were Setae collection. individuals,each one from a different colony, thatwere directly frozen after the In order to explore the length anddiameter of setae,we randomly selected 10 Dynamical properties density density and to the axis: movement parallelthe to setae axismajor theand second for movement perpendicular were usedin the calculations, with the first giving the aerodynamic diameter for equations following The microscopy. SEM by measure setae of length and diameter We obtained the aerodynamic diameter using equations providedthe air. by Cox (1970) and the where d g is the whereacceleration of gravity, setae, reaching atthe ground the distance ofL=hc/v Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 In order to evaluate the behavior of setae in the environment andtheir dispersion Knowing these parameters, we calculated the potential distance spreading of f is the seta diameter, ȕ ). the aspect ratio. Comment citer cedocument: ȡ s

the settling velocity). Although this expression is only thisexpression Although velocity). settling the f T. pityocampa the seta density (with the value of 1,100kg/m ȡ water is water density and , n= 120 for ae , which is the hypothetic diameter s (where his the height ofsetae T. pinivora Ș is the viscosity of , and n= 175 for 175 n= , and 3 ), 0 unit T. 5 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Smirnov test was used. To test for normality in the distribution of the length of released setae, the Kolmogorov- relationships between setal length and diameter wereanalyzed by regression analysis. regression, assuming a sum twoof normal distributions in case of bimodality.The The best fit of distribution of aerodynamic diameter was obtained through a nonlinear Statistics specimens at temperature below the zeroavoid to water evaporation. 1000) equipped with a cool stage (MK3 unit model, Deben UK), which maintained the The specimens were observed by scanning electron microscopy (SEM Hitachi TM- under a stereomicroscope and dissectedtransversally to the fields carrying the setae. length classes in the urticating apparatus. Larvae from the freezerwere kept on ice Last instar larvae theof three species were usedto observe the distribution of the setal Scanningelectron microscope analysis pinivora The sampledsetae were collectedfrom last instar larvae ( microscope and the pictures were analyzed for measuring the length theof setae present. fromthe central part theof disc (photograph area x 1.2 mm) 1.8 were taken atthe was removedwith a scalpel andtransferred to amicroscope slide. Digital photographs (Barbaro & Battisti 2011). After the treatment, the adhesive sealtape (diameter 14 mm) simulate amechanical disturbance, such as a typical attack by vertebratea predator Vortex a using vortexed, then was vial sealedwith an adhesive transparent PVC tape (Comet transferredto acylindrical vial (17 x 45 mm)with a flat bottom. The opening was disturbance, wecollected onelarva from each colony, which were individually Length of released setae conditions. validfor laminar flow,it is a good indicator for spreading also in turbulent flow Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 In order to evaluate the length theof setae released by larvae after amechanical n= 9 , T. processionea Comment citer cedocument:

n = 8). “ RX3 mixer at 2,400rpm for 10 seconds, to “ , Tesa AG - Hamburg, D). The T. pityocampa n = 5 , T. 6 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article 20 m of height anda windvelocity of 2 m/s, under laminar flow conditions. standard deviation, andthe parameter a (39.4) is thenumber of setae of the mode. peaks at 12.3 µm (variance 1.59 dispersion distances are 21 and 7.4 km. two modes in this sample. x populations of setae, b where x best fit tothe data is by the sum oftwo Gauss curves,with the equation: 2.4 km for the long setae. at 20m heightof ina day with awind velocity of 2m/s is 6.5 kmfor the short setae and than that ofshortsetae (1.6 mm/s). The horizontal distance traveledfor aseta released this sample.The average settling velocity longof setae (6.1 mm/s) was 5 times higher andthe a twoparameters populations setae,of b where x optimum function optimum is: distribution (Fig. 1). The best fit to the data is by the sum of two Gauss curves, the In Dynamical properties Results Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 1 and x T. pityocampa It mustbe mentionedthat the velocities anddistances given above are for the The settling velocity is 5 mm/s, resulting in a dispersion of 8 km for a release at where In The settling velocities are 1.9 and mm/s,5.4 respectively, and the corresponding The distribution of the aerodynamic diameter of 1 1 2 and x and x , and the two parameters a y = y = T. processionea a a 2 1 1 2 x , respectively and7 13.6µm, are the mean diametertwo of different , respectively 7.6 and 12.9 µm, are the mean diameter of two different · · 0 Comment citer cedocument: exp(-0.5 ·((x x - exp(-0.5 ·((x x - (12.3 µm) is the mean diameter of setae, b (1.6 µm , the distribution of aerodynamic diameter showed a bimodal 1 1 (1 µm (1 (1.7 µm 1 (8.8) and a , the distribution of aerodynamic diameteris andunimodal 2

) and b ) and y = 2 P ) and b m a · exp(-0.5 · ((x-x a ·exp(-0.5 2 1 1 1 ). The best fit is: fit best The ). (9.6 ) and a 2 )/b )/b 2 (12) are the number of setae ofthe twomodes in (1.8 µm 2 (1 µm 1 1 ) ) 2 2 ) +a ) +a 2 2 ) the of the square standard deviation, ) the square of the standard deviation of 2 2 2 (26.2) are the number of setae of the · exp(-0.5 · ((x · exp(-0.5 - x · exp(-0.5 · ((x · exp(-0.5 - x 0 T )/b) . pinivora 2 ) is andbimodal the 2 2 2 ) the square of its )/b )/b 2 2 ) ) 2 2 ) ) 7 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article processionea cohabit on the whole surface of the mirror, whereas this was not the case for 351 (skewness index = 0.32), whereas in the in whereas 0.32), = index (skewness skewedtothe left in processionea pityocampa The picture theof setae in the showedmirrors differences among the three species. In Distribution ofsetae in the urticating apparatus (skewness index = -0.01). than in the range ofvariation setaof length,as this was wider in T. pinivora pityocampa peak in the class 50-100of species (Fig.3). The shape was bimodal in = 0.12, p<0.01; normal one (Kolmogorov-Smirnov test: (Kolmogorov-Smirnov one normal The analysis ofsetae released by thelarvae showed a distribution different fromthe Length of released setae for (Fig. 2),with a higher slope for shortsetae. The R length were highly correlatedin the threespecies according to a logarithmic function aerodynamic diameter,the smallersetae will muchspread Setal further. diameter and mean aerodynamic diameter. Since the velocity is inversely proportional theto square in all the cases, and were higher for higher were and cases, in all the Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 T. processionea P m) shorterthan the one of the othertwo species. Moreover,the distribution was T. pinivora and , the first peak (22.6% setae)of washigherthan the second peak whereas in the peaks were similar (both about 25%).There was further difference for (Fig. 4). , which showed a single peak in the class of 200-250 T. processionea, T. pinivora Comment citer cedocument: (47-492 (0.62). T. pityocampa P we observedtwo well definedlevels of setae length, that P m anda second peak in the class of200-250 m). A completely different situation occurred A m). in completely occurred situation different

D = 0.07, p<0.01) but with differences shapes among (skewness index = 0.99)and in T. pityocampa T. pityocampa, T. processionea T. T. pityocampa T. 2 (0.92) and for values were statistically significant D = 0.12, p<0.01; D = 0.12, the distribution was symmetric was the distribution T. pityocampa and T. pinivora T.pinivora P T. pinivora m, and a range (56- (46-681 , with, a first P T. pinivora, m. In m. (0.90) than T. T. P T. m) T. D 8 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article , however,such as the brown tail moth the same,the different size may affect bothdispersionskin and penetration. In other size isgiven. As in the studied species of (Notodontidae) (Lamy et al. but 1984), no interpretation about the importance ofsetal examples of variation in setal size,such as in the adult female of demonstrated by our observations atthe microscope. SEM There arefew published classes. Actually the shortand long setae are intermixedthroughout the mirror, as can be explained by the overlapping of two normal distributions for each of the length P considered as bimodal (Sokal & Rohlf 1995), with a first peak in the class of 50-100 measures obtainedfor the same species. Bimodality in processionea where the distribution of setal length has been studied,the oak processionary moth differences may be ascribed to differences insetal extraction methods and sample sizes. processionea 150-250 lower: for lower: to 10 and 6times, respectively.Previously measurespublished of the range were much T. pinivora the longest (680 setae. the ones thatoccur in thetwo of three species areable tospread farther away than long setal size have implications on the dynamical properties. In particular, short setae like from thatexpected by previously and weshowed data, published thatthe differencesin three processionary moth species. We found amarkedly different pattern of setae length remained Here, unknown. wepresentfirst the detailed description ofthesetae size of in individual in an length setal of thedistribution 1982), Lamy et al. 1963; Démolin of studies, for both morphological andfunctional traits (Fabre 1900; Scheidter 1934; Although the setal system ofprocessionary moth has been the subject of a large number Discussion Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 m, and asecond in the class of 200-250 In The results showed awide variation in setal length. In the case of P T. pityocampa T. m (Lamy 1990); for T.pityocampa (47-492 , the distribution appears unimodal (Fenk et al. 2007),confirming our : 150-250 P Comment citer cedocument: m) were about 14 times longerthan the shortest (50 P m) and in P and , 80-160 m (Lamy 1990) and 85-290 T. pinivora

T. pinivora T. processionea P m (Scheidter 1934), 93-415 the distribution of setal length can be P Thaumetopoea : 60-260 m. In the only other species sameof genus (56-351 Euproctis chrysorrhoea P m (Démolin 1963); and for P m (Fenk et al. 2007). These 2007). al. et m (Fenk T. pityocampa P the general of shape the seta is m)the same ratios were equal P m (Hase 1939), and Anaphe venata and P m),whereas in T. pityocampa T. pinivora (Linnaeus) T. Butler T. 9 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article explore what happens with setae of different size.The variation in seta size may also be detect setae in human skin severalfor days after the contact, it could beinteresting to penetrate the skin of target organisms. As Fagrell et al. (2008) developed a method to hypothesize that the variation of setal size may involve adifferent capacity setaeof to size was not consideredas a possible factor affecting the penetration efficiency. We variation in seta size,the range identifiedwas smaller than that of our study setaland sharp basal endand it is helped by barbs on the distal end. Although this study explored firstly studied by Hase (1939) on mammalians, such as petsandlivestock. The penetration of setae into the skin was extended protection are higher the than costspossibly imposed by competition. made by these species of processionary moths indicates that thebenefits an from be elucidatedwith appropriate experiments, the large investment urticatinginto setae competition among herbivores should increase.Although the mechanism needs to other preys of vertebrate predators could indirectly benefit of the protection, andthus much farther,with a direct benefit for the colony.In this case,one could speculate that stronger is the protection; in this case the diversity of seta size may extendsuch a barrier the function tokeepaway predators. In this case, thelarger and denseris the the cloud, of shortsetae up to 6km (during the day) and 12 km(during thenight), they could have setae disperse as a cloud around the colony (Fenk et al. 2007), with high concentration humans,it is unlikely that birds or mammalian predators would respond differently.As Although these conclusions have been taken based theon reaction development in as the symptoms appear with a delay of time, whenthe larva has been already killed. urticating setae arean efficient defense system for the not colony fortheindividual,but what the about is known defense mechanism.Battisti thatet al.(2011) out pointed the spp.) over all (GroenenEurope & Meurisse 2012). ( oaks on feeding the genus in one only the this species, of history evolutionary interval ofvariation, remains unexplainedand it is possibly related toa different (Battisti et al.2011). The absence of bimodality in display a high variation of shape that could be potentially linkedto a different function (Kemper 1955) and South American tarantula spiders (Cooke etal. 1972), the setae Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 The seta size canbealsoimportant for impacts on humans andon domestic The role setaof size in protection from predation can be discussedin relation to Comment citer cedocument:

T. pityocampa , showing that the seta enters with the T. processionea , despite alarge Quercus 10 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Cox RG (1970). The motion of long slender bodies in a viscous fluid 1: General theory. Cooke JAL, Roth VD, Miller FH The(1972). urticating hairs oftheraphosidspiders. Battisti A, Holm G, Fagrell B, Larsson S (2011). Urticating hairs in arthropods their– Barbaro L, Battisti A (2011). Birds as predators of pine processionary moth References operations of these forestand urban tree pests. addition, it may usefulbe to pest managers and decision makers in planning the control from predation and for risk assessment toward non-target vertebrates and humans. In this paper may offer an opportunity to explore the importance ofsetal size for protection the soil fromor tents, because theirof high persistence.The information providedin and forest areas (EFSA 2009). This is emphasized by a continuous release ofsetae from moth, such a risk is very high due to the wide distribution of host plants in both urban which humans and domestic mammalians are exposed. In the case of processionary open the way a to better modeling both of the ecology the systemsof and the risks to referencethe to organism that may have causedthem (Hossler 2009). exposurefor and diagnostics, as symptoms are often reported without a specific size for dispersion. This assumption is relevant for the prediction of areas of risk Meyer 2006)are associatedwith airborne material,suggesting the importance of setal and the oak processionary moth (Vega et al. 2011),the northern pine processionary moth people living nearby trees infested by the pine processionary moths humans anddomestic mammalians. Most records of serious symptoms observed in point is the setae dispersion in the environment and the possible consequences for that setae of more diverse than previously considered, as Rodriguez-Mahilloet al. (in press) showed relatedto potential differences in their allergenic content, which toappears muchbe Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 Journal of Fluid Mechanics 44, 791–810. American Museum Novitates 2498, 1–43. 203-220. 56, Entomology of Review Annual significance. medical and nature (Lepidoptera: Notodontidae). Biological Control 56, 107-14. In conclusion, the awareness of a great variation in size of urticating setae may T. pityocampa Comment citer cedocument: contains at least sevendifferent allergens. Another important

T. processionea (Maier2003; Gottschling etal. & T. pinivora T. pityocampa (Holm etal. 2009), 11 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Hase A(1939). Über den Pinienprozessionsspinner und über die Gefährlichkeit seiner Groenen F, Meurisse N (2012). Historical distribution of the oak processionary moth GottschlingMeyer S, (2006). S An epidemic airborne disease caused by the oak Gilmer PM(1925). A comparative study of the poison apparatus of certain KawamotoF, Kumada N (1984). Biology andvenoms of Lepidoptera. In: Handbook of Holm G,Sjoberg J, Ekstrand Cet al. (2009). Tallprocessionsspinnarestort – Hossler EW (2009).Caterpillars andmoths. Dermatology andTherapy 22, 353-66. Fagrell B, Jörneskog G, Salomonsson AC, Larsson S, Holm G (2008). Skin reactions Fabre JH (1900). Souvenirs entomologiques, études sur l'instinct et les moeurs des EFSA (2009). Scientific Opinion theof Panel Planton Heath on a pest risk analysis on Fuentes AparicioZapatero V, Remón L,Martínez Molero MIet al. (2006). Allergy to Fenk L, Vogel B, Horvath H (2007). Dispersion of the bio-aerosol produced by the oak Démolin G (1963). Les 'miroirs' urticants la de processionnaire pindu ( Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 Allergologia et Immunopathologia et Allergologia Raupenhaare ( expansion.Agricultural andForest Entomology 14, 147–55. Thaumetopoea processionea processionary caterpillar. Pediatric Dermatology 64-66. 23, 39. lepidopterous larvae. Annals ofthe Entomological Society Americaof 18, 203- Invertebrate Venoms.Dekker, New York, pp. 291-330. Natural Toxins. Vol. In: 2. Tu AT eds, Insect Poisons, Allergens, and other hälsoproblem på södra Gotland. Lakartidningen 106, 1891–94. Schädlingskunde 15, 133-42. pityocampa processionary moth ( induced by experimental exposure to setae from larvae ofthe northern pine insectes. Delagrave, Paris. and extension ofits scope to the EU territory. The EFSA Journal 1195, 1-64. Thaumetopoea processionea pine pine processionary caterpillar ( processionary moth. Aerobiologia Comment citer cedocument: Schiff.).Revue de Zoologie Agricole etAppliquée 10-12, 107-14. Thaumetopoea pityocampa Thaumetopoea pinivora

in Europe suggests recolonization instead of L., the oak processionary moth, prepared by the UK Thaumetopoea pityocampa Thaumetopoea 23, 79-87. 34, 59-63. Schiff.). Anzeiger für ). Contact Dermatitis 59, 290-95. ) inchildren. Thaumetopoea 12 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Pesce H, Delgado QA (1971). Poisoning from adult moths and caterpillars. In Bürchel caterpillars. and moths adult from Poisoning (1971). QA H, Delgado Pesce Novak F, Pelissou V, Lamy M (1987). Comparative morphological, anatomical and Mullen GR (2009). Moths and butterflies (Lepidoptera). In GR Mullen, LA Durden eds. Moneo I,VegaCaballero JM, ML, Vega J,Alday (2003).E Isolation and Matthioli PA (1568). I discorsi di M.Pietro Andrea Matthioli,medico cesareo nelli sei Maier H, Spiegel W, Kinaciyan etT, al. (2003). The oak processionary caterpillaras the Lamy (1990). ContactM dermatitis (erucism) produced by processionary caterpillars Lamy, Pastureaud M, MH,F Novak F, Ducombs G (1984).Papillons urticants Lamy M, Ducombs G, Pastureaud MH, Vincendeau P (1982). Productions Kemper H (1955). Experimentelle Untersuchungen über die durch Afterwolle von Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 Saturniidae). Comparative Biochemistry andPhysiology 88, 141-46. (Lepidoptera, Thaumetopoeidae) and Lepidoptera: biochemical ofthe studies urticating and apparatus urticating hairs of some Medical andVeterinary Entomology,Elsevier, Amsterdam, pp. 353-70. caterpillar characterization Tha p of 1, amajor allergen from the pine processionary libri di PedacioDioscoride Anazarbeo. Valgrisi, Venezia. Dermatology 149, 990-97. ancause of epidemic airborne disease: survey and analysis. British Journal of (genus Thaumetopoea). Journal of AppliedEntomology 110, 425-37. mode de action. Bulletin de la Société Zoologique de France 109, 163-77. microscope électronique à balayage àl´étude leur appareilde urticant et àleur d’Afrique et d’Amérique suddu (g. 515-29. 107, deFrance Entomologique (Lépidoptères). Appareil urticant et appareil deponte.Bulletin la Sociétéde tégumentaires dela processionnaire dupin ( 59. 37- 55, Zoologie Angewandte für Zeitschrift Arthropodenstichen. von Wirkung Euproctis chrysorrhoea Invertebrates Academic, New York, pp. 119–56. pp. York, New Academic, Invertebrates W, Buckley EE eds. Venomous andTheir Venoms, Vol. III: Venomous Comment citer cedocument: Thaumetopoea pityocampa Thaumetopoea pityocampa

erzeugte(Lepidoptera) Dermatitis, vergleichen mit der Anaphe Hylesia metabus . Allergy 34-37.58, Schiff., et g. Thaumetopoea pityocampa Th. processionea Hylesia Cramer (Lepidoptera, ): Contribution du L. Schiff.) 13 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Vega JM, MoneoI, García Ortiz JC, Sánchez Palla P,et al. (2011). Prevalence of Vega JM,MoneoArmentiaI, A,Fernandéz A, et al.(1999). Allergy to processionary Specht A,Corseuil BarretoE, Abella H (2008). Lepidopteros importanciade medica. Sokal RR,Rohlf FJ (1995). Biometry: the principles andpractice ofstatistics in ScheidterF (1934). Auftreten der "Gifthaare" bei den Prozession-spinnerraupen in den Santos-Magadán S, González deOlanoD, Bartolomé-Zavala B, Trujillo-TrujilloM, Rodriguez-Mahillo AI, Gonzalez-Muñoz M, Vega JM et al. in press. Setae from the Werno J,Lamy M (1990). atmospheric pollution: the urticating hairs of the Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 325-31. Lepidoptera). Comptes Rendus l’Académiede des Sciences, Paris,Série III 310, cutaneous reactions to pine processionary moth ( 1418-23. caterpillar ( Principais especies noRioGrande Sul. USEB,do Pelotas, Brasil. biological research. 3rd ed. Freeman,New York. 223-26. 44, Pflanzenschutz und Pflanzenkrankheiten fur Zeitschrift Stadien. einzelnen 109-10. processionary caterpillar: allergicirritant or mechanism? Contact Dermatitis 60, Meléndez-Baltanás A, González-Mancebo E (2009). Adverse reactions theto pine pine processionary caterpillar ( an adult population. Contact Dermatitis 64, 220–28. allergens. Contact Dermatitis. pine processionary moth ( Comment citer cedocument: Thaumetopoea pityocampa

Thaumetopoea pityocampa Thaumetopoea pityocampa Thaumetopoea ). Clinical & Experimental Allergy 29, Thaumetopoeapityocampa ) contain several relevant Schiff.) (Insecta, ) in 14 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Thaumetopoea Figure Distribution1. theof aerodynamic diameter ofthe setaein the three of species Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 . Comment citer cedocument:

15 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article Thaumetopoea Figure Relationships2. between length anddiameter setaeof inthe species three of Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 . Comment citer cedocument:

16 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article test. Figure Length3. of setae released by the threespecies of Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 Comment citer cedocument:

Thaumetopoea in a laboratory ina 17 Version postprint H. (2014). Sizeanddispersion ofurticating setaein threespeciesof processionary moths. Petrucco Toffolo,E.,Zovi, D.,Périn, C. .,Paolucci,P., Roques,A., Battisti,A.,Horvath, Accepted Article the left, and long setaein intermixed Figure 4. SEM images the of urticating apparatus in three Integrative Zoology, 9(3), 320-327. DOI :10.1111/1749-4877.12031 T. pityocampa, T. pinivora, T. processionea Comment citer cedocument: T. pityocampa

and T.pinivora ) showing the occurrence of short of the occurrence ) showing Thaumetopoea . species (from 18