J Pest Sci (2005) 78: 217–226 DOI 10.1007/s10340-005-0096-3

ORIGINAL ARTICLE

Stanislav Trdan Æ Ljiljana Andjus Emilija Raspudic´Æ Milica Kacˇ Distribution of Aeolothrips intermedius Bagnall (Thysanoptera: ) and its potential prey Thysanoptera on different cultivated host plants

Received: 24 January 2005 / Published online: 13 July 2005 Ó Springer-Verlag 2005

Abstract The results of the monitoring of Thysanoptera Introduction species on cultivated plants in Slovenia (2000–2001), Croatia (1994–1996), and Serbia and Montenegro as crop pests (1988–2003) are presented in this study. The aim of the investigation was to study the host plant distribution of Thrips are mostly known as living in flowers, but the predator Aeolothrips intermedius Bagnall and its many species live only on leaves, a few are predatory, and potential prey. Banded thrips were found on 30 different probably about half of the species feed on fungi (Palmer host plant species belonging to 16 botanical families, et al. 1989). The damage on plants caused by thrips due to always in mixed populations with phytophagous or their feeding or their deposition of eggs is often ascribed to facultative phytophagous insects (including 18 Thysa- other pests, especially if observed and/or evaluated by noptera species). On the vegetative parts of the culti- non-professionals. Of 5,000 thrips species recorded vated plants, banded thrips were found less numerous in worldwide, only 1% are known to be economically spite of the massive population of some harmful thrips harmful (Mound and Teulon 1995); while on different species. This indicates highly important role of pollen as continents, the number of harmful Thysanoptera species alternative food for Aeolothrips intermedius. very rarely approaches the number of 10 (Jenser and Czencz 1988; Chang 1995; Gahukar 2004). In this sense, Keywords Aeolothrips Æ Distribution Æ Hosts Æ western flower thrips (Frankliniella occidentalis [Per- Predator Æ Thysanoptera gande]) (van Dijken et al. 1994; Kirk and Terry 2003) and onion thrips (Thrips tabaci Lindeman) (van Rijn et al. 1995; de Jager et al. 1997; Theunissen and Schelling 1997) are considered to be harmful in Europe, while banded Communicated by W. Schwenke greenhouse thrips (Hercinothrips femoralis [Reuter]), thrips (Thrips simplex [Morison]), pear thrips S. Trdan (&) Chair of Entomology and Phytopathology, (Taeniothrips inconsequens [Uzel]), pea thrips (Kakothrips Biotechnical Faculty, Department of Agronomy, pisivorus [Westwood]), privet thrips (Dendrothrips ornatus University of Ljubljana, Jamnikarjeva 101, [Jablonowski]) and some other species are considered less 1111 Ljubljana, Slovenia important, since their occurrence and the following E-mail: [email protected] damages on cultivated plants are not observed every year Tel.: +386-1-4231161 Fax: +386-1-4231088 in economically important level (Kirk 1985; Jenser and Czencz 1988; Andjus 1997; Trdan 2003). L. Andjus Natural History Museum, Njegosˇ eva 51, 11000 Belgrade, Serbia and Montenegro Thrips as biological control agents E. Raspudic´ Faculty of Agriculture, University of J. J. Strossmayer in Osijek, Though much progress has been made during the last Trg Sv. Trojstva 3, 31000 Osijek, Croatia 15 years in the sense of making the control of insects M. Kacˇ (Insecta), mites (Acarina) and other pests on cultivated Chair of Chemistry, Biotechnical Faculty, crops more environmentally friendly (Wilson et al. 1995; Department of Food Science and Technology, University of Ljubljana, Jamnikarjeva 101, Milevoj 1999), and the number of such investigations is 1111 Ljubljana, Slovenia steadily increasing during the last years (Arthurs et al. 218

2003; Kishimoto 2003; Nielsen et al. 2003), it is an The great majority of the species of this (about 100 interesting fact that there are not enough reliable data species have been recorded so far) spend most of their lives on the predatory efficiency of the autochthonous repre- in flowers, feeding on small and pollen (Har- sentatives of the Thysanoptera order. per et al. 1989; Patrzich and Klumpp 1991; Mound and Some species of the Haplothrips, Scolothrips, Frankli- Kibby 1998; Yee et al. 2001). Till now, what is known on nothrips, Karnyothrips and Aeolothrips genera as well as the impact of two different types of food for the life and those of the Aleurodothrips, Allelothrips, Androthrips, development of facultative predators is not adequate, al- Apterygothrips, Desmidothrips, Desmothrips, Leptothrips, though some studies in this context were performed in the Erythrothrips, Mymarothrips, Parascolothrips, Podothr- last years (von Zegula et al. 2003). ips, Stictothrips, Stomatothrips, Symphyothrips, Trichinothrips, Veerabahuthrips, Xylaplothrips and Andrewarthaia genera (Palmer et al. 1989; Riudavets Aeolothrips species as predators 1995; zur Strassen 1995; Sabelis and van Rijn 1997) are well-known predators; but in Europe, particularly some More than 20 species of the Aeolothrips genus were re- species of the genus Aeolothrips are considered as poten- corded in Europe (Table 1). The majority of the repre- tially important autochthonous facultative predators. sentatives of the genus are considered to be facultative

Table 1 Check-lists of Aeolothrips species recorded in ten different European countries and Iberian Peninsula

Country Species Reference

Austria A. albicinctus Haliday, A. astutus Priesner, A. ericae Bagnall, zur Strassen, personel communication A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. versicolor Uzel, A. vittatus Haliday Croatia A. collaris Priesner, A. ericae Bagnall, A. fasciatus (L.), Raspudic´et al. 2003 A. gloriosus Bagnall, A. intermedius Bagnall, A. melaleucus Haliday, A. propinquus Bagnall, A. versicolor Uzel Germany A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, zur Strassen, personel communication A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. propinquus Bagnall, A. tenuicornis Bagnall, A. versicolor Uzel, A. vittatus Haliday Hungary A. albicinctus Haliday, A. astutus Priesner, A. ericae Bagnall, zur Strassen, personel communication A. intermedius Bagnall, A. melaleucus Haliday, A. priesneri Knechtel, A. propinquus Bagnall, A. versicolor Uzel, A. vittatus Haliday Iberian peninsula A. albicinctus Haliday, A. andalusiacus zur Strassen, A. balati Pelikan, Goldarazena 1999 A. bournieri Lacasa, A. citricinctus Bagnall, A. collaris Priesner, A. cursor Priesner, A. ericae Bagnall, A. fasciatus (L.), A. gloriosus Bagnall, A. guitiani Berzosa y Maroto, A. intermedius Bagnall, A. linarius Priesner, A. melaleucus Haliday, A. melisi Priesner, A. pelikani Titschack, A. propinquus Bagnall, A. pyrenaicus Bagnall, A. quercicola Bournier, A. tauricus Derbeneva, A. tenuicornis Bagnall, A. versicolor Uzel, A. vittatus Haliday Italy A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, zur Strassen, personel communication A. cursor Priesner, A. ericae Bagnall, A. fasciatus (L.), A. gloriosus Bagnall, A. intermedius Bagnall, A. melaleucus Haliday, A. propinquus Bagnall, A. tenuicornis Bagnall, A. versicolor Uzel, A. vittatus Haliday Poland A. astutus Priesner, A. ericae Bagnall, A. fasciatus (L.), Kucharczyk, personel communication A. intermedius Bagnall, A. melaleucus Haliday, A. propinquus Bagnall, A. pulcher Oettingen, A. versicolor Uzel, A. vittatus Haliday Romania A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, Oromulu, personel communication A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. priesneri Knechtel, A. pulcher Oettingen, A. verbasci Knechtel, A. versicolor Uzel, A. vittatus Haliday Serbia and Montenegro A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, Trdan et al. 2003 A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. pulcher Oettingen, A. versicolor Uzel, A. vittatus Haliday Slovenia A. albicinctus Haliday, A. astutus Priesner, A. ericae Bagnall, Trdan et al. 2003 A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. versicolor Uzel The Netherlands A. albicinctus Haliday, A. ericae Bagnall, A. fasciatus (L.), Vierbergen, personel communication A. intermedius Bagnall, A. manteli Titschack, A. melaleucus Haliday, A. propinquus Bagnall, A. tenuicornis Bagnall, A. versicolor Uzel, A. vittatus Haliday 219 predators (Mound and Kibby 1998), though several geographically close countries which exhibit considerably differences originating from different size of the species, different reliefs and climates was undertaken. their preference towards different host plants or ar- thropods, their ecology (especially differences in the periods of high population densities of the prey) as well as other factors are to be considered (Moritz 1982). Materials and methods A. intermedius is the most common species of this genus on the Old Continent (Deligeorgidis et al. 2002) The monitoring of thrips presented in this contribution and it is frequently mentioned in the context of biolog- was carried out in Slovenia during 2000–2001 (Trdan ical control (von Zegula et al. 2003). Adult specimens of 2003), on the territory of Serbia and Montenegro during this species are quite similar to those of Aeolothrips 1988–2003 (Andjus 1997, 1998) and in Croatia during fasciatus L., so the two species can very easily lead to 1994–1996 (Raspudic´2003). Samples were collected confusion (zur Strassen, personel communication). from 26 locations in Slovenia (Fig. 2), from 15 locations Adults and larvae of banded thrips, as is a common in Croatia and from 11 locations in Serbia and Monte- name of A. intermedius, were found as predator of 44 negro (Figs. 1). On some locations, samples were col- species of Thysanoptera order (Riudavets 1995). In lected in different periods. Only adults were sampled Europe, it has usually been mentioned as a predator on because identification of larvae is more difficult (Mound onion thrips (T. tabaci Lindeman) (Bournier et al. and Kibby 1998). 1978; Lacasa et al. 1982; Torres-Vila et al. 1994; All samples showing the presence of A. intermedius Franco et al. 1999), other representatives of Thrips were collected in the open. The sampling was performed genus (Lacasa et al. 1989; Gremo et al. 1997), as well with usual thripsological instruments. Different culti- as together with greenhouse thrips (Heliothrips haem- vated plants were beaten over a small white plastic tray. orrhoidalis Bouche´), Odontothrips confusus Priesner Depending on the host species and growing stage of the (Bournier 1979; Khosbayar 2001) and with the species hosts different aboveground parts of the plants were of Haplothrips genus (Dyadechko et al. 1971). Banded searched more precisely. From the tray surface, the thrips also feed on some mite species, larvae and eggs specimens were removed with a fine brush. Collected of psyllids (Psyllina) and whiteflies (Aleyrodina) as well thrips were kept in plastic 1.5 ml Eppendorf tubes in as on aphids (Aphidina). It was determined by the AGA solution, a mixture of ten parts of 60% ethyl laboratory experiments that A. intermedius can com- alcohol, one part of glycerine and one part of acetic acid. plete its life cycle feeding on Tetranychus urticae Koch, This mixture helps to distend the body of most thrips T. cinnabarina (Boisduval), T. atlanticus McGregor, and keeps the limbs supple. Stored specimens were Panonychus ulmi Koch and Cenopalpus pulcher Cane- transferred to 60% alcohol and kept in the dark at the strinı` & Fanzago, while the life cycle cannot be com- temperature about 0°C to prevent loss of colour (Palmer pleted by feeding on Frankliniella occidentalis et al. 1989; Mound and Kibby 1998; Gonzalez-Zamora (Pergande) (von Zegula et al. 2003), Psylla melanoneura and Garcia-Mari 2003). Fo¨rster, Trioza spp., Trialeurodes vaporariorum (West- Thrips specimens were collected from the end of April wood), Aphis fabae Scopoli, Aphis cracivora Koch, A- until the beginning of September, in different growing cyrthosiphon pisum Harris and Myzus persicae (Sulzer) stages of host plants. The number of the sampling spots (Bournier et al. 1979; Berest 1980). Mostly the prey of on selected parcels and the number of the sampled plant the banded thrips is considered to be pests on culti- parts (organs) of host plants depend on the size of the vated plants in the open, which speaks for the specific parcel, host plant species and population size of thrips. bionomics of the banded thrips. This refers mostly to On larger parcels, where thrips populations were less its susceptibility to different prey species and various abundant, the number of the sampling spots was higher. kinds of pollen. The fact is also in accordance with the Bigger plants (trees, bushes) were the only sampled theory that thrips is not an efficient predator in pro- plants on some parcels, if the thrips populations were tected places. Incapability of thrips rearing in labora- high. Different sizes of samples were taken into consid- tory conditions also confirms this (van Lenteren 1999). eration, depending on the density of the thrips popula- Banded thrips is omnivorous, and can complete its tion on the cultivated plants in question (1–50 specimen development by feeding only on flowers, where it was per plant species per location). found to eat pollen. It does not extract nutritional food In spite of the shortest sampling period in Slovenia, the from leaves and stems, but pierces them to extract number of collected specimens was the highest (67). Few water (Bournier et al. 1979; Lacasa 1980). specimens were collected in Serbia and Montenegro (28) In the European scientific literature, only a few detailed and least of all in Croatia (17). In some cases, specimens and/or abundant enough information on host distribu- on the same host plant and same location were collected in tion of A. intermedius on different cultivated plants different periods, but this was not the rule. (Andjus 2004) and its potential prey species that could be The morphological identification was performed of potential use in the investigation of its predatory role using standard light microscope in the facilities of the (von Zegula et al. 2003), could be found. So, an investi- Institute of Phytomedicine (Biotechnical Faculty, gation of its presence on cultivated crops in three Department of Agronomy, Ljubljana, Slovenia), Insti- 220

Fig. 1 Map of former Yugoslavia with sampling locations of Aeolothrips intermedius Bagnall and its potential prey Thysanoptera species for Croatia and Serbia and Montenegro. Symbols in front of locations represent the families which members are the host plants where Thysanoptera species were recorded

tute of Plant Protection (Faculty of Agriculture, Osijek, Tables give the results of the thrips monitoring in Croatia) and that of the Natural History Museum Slovenia during 2000–2001 (Table 2), in Croatia during (Belgrade, Serbia and Montenegro). The standard 1994–1996 (Table 3) and on the territory of Serbia and morphological identification keys by Priesner (1964), Montenegro during 1988–2003 (Table 4). It gives the Schliephake and Klimt (1979), Jenser (1982), Palmer host plants only for banded thrips, A. intermedius, and et al. (1989), Moritz (1994) as well as those by Mound for those thrips species that were present in considerable and Kibby (1998) were used. number (that means more than five individuals per

Fig. 2 Map of Slovenia with sampling locations of Aeolothrips intermedius Bagnall and its potential prey Thysanoptera species. Symbols in front of locations represent the families which members are the host plants where Thysanoptera species were recorded Table 2 Cultivated hosts, locations and dates of records of banded thrips, A. intermedius Bagnall, in Slovenia during 2000–2001. In the last column, the potential preys of banded thrips (they shared the same plant parts of the hosts) from the Thysanoptera order are presented

Family Host species Location Date Potential preys from the Thysanoptera order

Alliaceae Allium cepa L., aLjubljana 6 Jun 2000, 18 Jul 2001 Thrips tabaci Lindeman Allium fistulosum L. Asparagaceae Asparagus officinalis L. aLjubljana 18 Jul 2001 Chirothrips manicatus Haliday bSˇ empeter pri Novi Gorici 22 Aug 2001 Asteraceae Aster spp. Ljubljana 29 Aug 2001 Frankliniella intonsa (Trybom) Chamomilla spp. Spodnje Sˇ kofije 7 Jun 2001 Thrips vulgatissimus Haliday Dahlia pinnata Cav. Sˇ empeter pri Novi Gorici 22 Aug 2001, 5 Sep 2001 Microcephalothrips abdominalis (Crawford) Helianthus annuus L. Ljubljana 12 Aug 2001 Thrips physapus Linnaeus, Thrips tabaci Lindeman, Frankliniella intonsa (Trybom) Brassicaceae Brassica spp. Ljubljana 2 May 2001 Frankliniella intonsa (Trybom) Pacinje 22 Apr 2000, 27 Apr 2001 Strocˇ ja vas 21 Apr 2000 Sˇ ikole 5 Jun 2000 Caryophyllaceae Dianthus spp. Ljubljana 12 Aug 2001 Thrips simplex (Morison), Thrips atratus Haliday, Sˇ empeter pri Novi Gorici 22 Aug 2001 Thrips tabaci Lindeman Cichoriaceae Lactuca sativa L. Ljubljana 12 Aug 2001 Thrips spp. Fabaceae Medicago sativa L. Dolenja vas 18 Aug 2001 Odontothrips confusus Priesner, Frankliniella intonsa Ljubljana 22 Aug 2000, 12 Aug 2001 (Trybom), Thrips tabaci Lindeman Mala Slevica 18 Aug 2001 Podnanos 22 Aug 2001 Phacelia tanacetifolia L. Ljubljana 12 Aug 2001 Frankliniella intonsa (Trybom) Trifolium pratense L., Preserje pri Radomljah 12 May 2000, 29 Jun 2000, Frankliniella intonsa (Trybom), T. repens L. 15 Jun 2001, 17 Aug 2001 Haplothrips spp., Thrips tabaci Lindeman Ljubljana 6 Jun 2000, 22 Aug 2000, 12 Jun 2001, 22 Jun 2001, 12 Aug 2001 Iridaceae Gladiolus spp. Ljubljana 12 Aug 2001 Frankliniella intonsa (Trybom), Thrips simplex (Morison), Pacinje 25 Jul 2001 Thrips fuscipennis Haliday, Thrips tabaci Lindeman Postojna 27 Aug 2001, 3 Sep 2001 Zˇ elimlje 23 Aug 2001 Oleaceae Olea europaea L. bOsp 7 Jun 2001 Thrips spp. Poaceae Avena sativa L. cLjubljana 19 Jun 2001, 22 Jun 2001, Haplothrips aculeatus (Fabricius), Limothrips denticornis Haliday, 30 Jul 2001 Frankliniella tenuicornis Hood dRadomlje 18 Jun 2000 dZˇ upelevec 2 Jul 2000 Hordeum vulgare L. cSpodnje Sˇ kofije 28 Apr 2000 Limothrips denticornis Haliday, Haplothrips aculeatus (Fabricius), dSpodnji Brnik 4 Jun 2001 Frankliniella tenuicornis Hood cPtujska gora 2 May 2001, 5 May 2001 bSpodnji Velovlek 28 Apr 2001 cStojnci 5 May 2001 Triticum aestivum L. bGorisˇ nica 13 May 2000, 14 May 2000, Haplothrips aculeatus (Fabricius), Haplothrips tritici Kurdjumov, 27 Apr 2001, 19 May 2001 Limothrips denticornis Haliday, Limothrips cerealium Haliday, bPragersko 5 May 2001 Frankliniella tenuicornis Hood, Frankliniella intonsa (Trybom) bStojnci 5 May 2001 Zea mays L. bBolehnecˇ ici 6 Jul 2000 Haplothrips aculeatus (Fabricius), Haplothrips spp., Cephalothrips monilicornis (O.M. Reuter), Chirothrips manicatus Haliday 221 222

sample) on the same plants and so present a potential prey for the facultative predator in question. The highest share of specimens were collected on flowers (62.5%), less specimens were collected on leaves (19.6%) and in

Thrips the rest three groups were placed specimens from ears of cereals or different aboveground host plant parts. The number of the specimens in none of these groups did not exceed 9% of the total number of specimens (112). (Trybom), Haliday

Haliday Sampling locations and the host plants of recorded Thysanoptera species are to be found in Figs. 1, 2.

Haliday According to the thrips monitoring on the area in question, it can be concluded that banded thrips can be found on various plant species of at least 16 different botanical families, which all have representatives among fuscipennis

Thrips fuscipennis Frankliniella intonsa Thrips vulgatissimus economically important cultivated plants. Adults of this facultative predator occur during the last ten days in April, when oil rape is in flower and winter cereals (winter wheat and winter barley) have either completed the phase of tillering (Zadoks’ growth stages 25–29) or have entered the phase of stem elongation (Zadoks’ growth stages 30–32 (Zadoks 1974). When numerous other plant species enter the flowering phase—especially papilionaceous plants (members of the Fabaceae family) and cultivated grasses (members of the Poaceae fam- ily)—the banded thrips inhabit these host plants as well. The density of the thrips population is strongly de- pended on duration of the host plant flowering and probably also on the quality of pollen. The last adults of the banded thrips were found at the end of August or in the first days of September, i.e. in the time when the flowering of the host plants is coming to its end. On 30 plant species where banded thrips was re- corded, populations of 18 other thrips species (mostly of the Terebrantia suborder) were also present—they were mostly phytophagous. Due to its massive populations kofije 7 Jun 2001

ˇ and extreme polyphagy (Deligeorgidis et al. 2002), the flower thrips (Frankliniella intonsa [Trybom]) was stated an 6 Jul 2000 ˇ to be the dominant species. It is a well-known facultative empeter pri Novi Gorici 22 Aug 2001 ikole 15 Aug 2001

PondorPacinjeRakic 5 Jul 2000 5 Jul 2000 phytophagous , pollen being its important food Spodnje S ˇ ˇ LjubljanaRakitnicaS 22 Jun 2001 26 Jul 2001, 18 Aug 2001 S b b b c source (Murai and Loomans 2001). A. intermedius were not found in the flowers of fruit bearing plants and or- namentals of the Rosaceae family (Prunus spp., Malus domestica Borkh., Cydonia oblonga Mill., Pyrus com- munis L.). In the area where the monitoring of thrips was performed, these flower mostly in March and April, when the banded thrips is not active yet. In this case, one L. Brje pri Komnu 9 Jun 2001 cannot describe the Rosaceae family as non-susceptible for the banded thrips, since this same species exhibits a

spp. Postojnamassive 27 Aug occurrence in the flowers of roses (Rosa spp.) in summer. Obviously, growing stages of the fruit bearing Moench Vitis vinifera Fagopyrum esculentum Rosa and ornamental host plants of the Rosaceae family which are attractive to A. intermedius are not timely harmonized with colonisation of the insect in question. It is an interesting fact, that the number of banded thrips in vegetative parts of the plants attacked by

(Contd.) massive populations of more harmful thrips spe- cies—e.g. onion thrips (T. tabaci Lindeman) on the leaves of leek, onion as well as cabbage—is surprisingly Specimens recorded on the leaves Specimens recorded on the ears/panicles Specimens recorded on the flowers and on the leaves Specimens recorded on the leaves and ears/panicles (in the case of grapevine, a bunch) Vitaceae Polygonaceae Rosaceae Specimens recorded on the flowers Table 2 Family Host speciesa b c Locationd low, compared Date to its number in Potential preys from the the Thysanoptera order flowers of some Table 3 Cultivated hosts, locations and dates of records of banded thrips, A. intermedius Bagnall, in Croatia during 1994–1996. In the last column, the potential preys of banded thrips (they shared the same plant parts of the hosts) from the Thysanoptera order are presented

Family Host species Location Date Potential preys from the Thysanoptera order

Chenopodiaceae Beta vulgaris L. var. Krndija 5 Jun 1995 Thrips tabaci Lindeman, Frankliniella intonsa (Trybom) saccharifera Lange Fabaceae Glycine max (L.) Merr. Bizovac 5 Jul 1995, 27 Jun 1996 Frankliniella intonsa (Trybom), Haplothrips spp., Phaseolus vulgaris L. Brela 6 Aug 1995 Thrips tabaci Lindeman, Frankliniella intonsa (Trybom) Pisum sativum L. Brinje 19 Aug 1995 Thrips fuscipennis Haliday, Thrips tabaci Lindeman, Kakothrips pisivorus (Westwood), Thrips atratus Haliday, Frankliniella intonsa (Trybom) Trifolium campestre Schreb. Sˇ tromajerovac 12 Jun 1995 Frankliniella intonsa (Trybom) Trifolium pratense L., Cˇ akovec 15 Jul 1995 Frankliniella intonsa (Trybom), Haplothrips spp., Thrips tabaci Lindeman T. repens L. Iridaceae Gladiolus spp. Visˇ njevac 20 Jul 1996 Frankliniella intonsa (Trybom), Thrips simplex (Morison), Thrips fuscipennis Haliday, Thrips tabaci Lindeman Poaceae Zea mays L. bKopanica 20 Jul 1994 Haplothrips aculeatus (Fabricius), Haplothrips spp., bBerava 20 Jul 1994 Cephalothrips monilicornis (O.M. Reuter), Chirothrips manicatus Haliday bCˇ epin 24 Jul 1995 bMaslenica 30 Jul 1995 bMarijanci 4 Aug 1994 Rosaceae Rosa spp. Ankin dvor 16 May 1995 Frankliniella intonsa (Trybom), Thrips fuscipennis Haliday Sambucaceae Sambucus nigra L. Ankin dvor 16 May 1995 Thrips nigropilosusUzel, Thrips tabaci Lindeman, Haplothrips spp. Solanaceae Nicotiana tabacum L. Pozˇ ega 22 Jul 1995 Frankliniella intonsa (Trybom), Frankliniella tenuicornis (Uzel) Vetovo 22 Jul 1995

Specimens recorded on the flowers a Specimens recorded on the flowers and on the leaves b Specimens recorded on the leaves c Specimens recorded on the leaves and ears/panicles (in the case of grapevine, a bunch) d Specimens recorded on the ears/panicles 223 224

other hosts. This speaks in favour of alternative food importance in the living space of A. intermedius, where

Haliday the insect performs its role as a predator or pollinator. Haliday, (Fabricius), (Trybom) Kurdjumov, (Trybom), Conclusion

spp., It was established that the banded thrips spends much of spp., Chirothrips manicatus its life in the flowers of the host plants, where it feeds on Lindeman, pollen and arthropods. As pollen is an important food source of the banded thrips (Bournier et al. 1979; Lacasa Haplothrips tritici Haplothrips Limothrips cerealium

Frankliniella intonsa 1980), the bionomics and the predatory effectiveness of Haplothrips aculeatus Frankliniella intonsa Haplothrips the species are closely connected with the developmental (O.M. Reuter), Thrips tabaci stages of its hosts. It is well-known that its massive Hood, Hood Haliday, Haliday,

Priesner, occurrence takes place during the flowering of the plants (Fabricius), (Fabricius), (Trybom) (Trybom) (Trybom) (Trybom), which are most attractive for the insect (Khosbayar

Linnaeus, 2001), but it is not sure whether this phenomenon is due Lindeman Lindeman to obviously better accessibility to high quality pollen, because of more arthropods or because of both. Some entomologists ascribe no important role to pollen as far as the development of the thrips is concerned (Milne Limothrips denticornis Frankliniella tenuicornis Frankliniella tenuicornis Cephalothrips monilicornis Frankliniella intonsa Thrips tabaci Thrips tabaci Haplothrips aculeatus Haplothrips aculeatus Limothrips denticornis Thrips physapus Frankliniella intonsa Odontothrips confusus Frankliniella intonsa Frankliniella intonsa et al. 1996; Nakao 1999), while others are convinced that it plays an important part in this game (Tsai et al. 1996). The third group acknowledges that alternative food causes a more numerous offspring and a shorter life cycle (Teulon and Penman 1991), but it most often diminishes the predatory effectiveness of the insect (van Rijn and Sabelis 1993). The host plants do not only provide for alternative Bagnall, on the territory of Serbia and Montenegro during 1988–2003. In the last column, the food but also for a shelter against potential attackers; besides it was established that the plants signalize to the predators the presence of their potential prey (Price et al. 1980; Dicke 1994). The present study merely gives the presence of A. intermedius on various host plants. The A. intermedius economic importance of the predator in the context of its direct influence on the phytophagous thrips is not evaluated. The role of thrips in the environment is complex and

21 Jul 1994, 14 Aug 1994, 30 Apr 1995 the present investigation deals only partly with this issue. It was established that banded thrips reduces the pop- ulation densities of other thrips and insect species in the open (Franco et al. 1999), while its role becomes much less important in greenhouses (in some European in 10 Jul 1994, 21 Jul 1994 inin 11 May 1994, 18 May 1994, 2 Jun 1994 18 May 1993, 11 May 1994, 2 Jun 1994 ˇ ˇ ˇ in 10 Jul 1994, 21 Jul 1994, 28 Jul 1994 inin 30 Apr 1995, 4 May 1995, 8 9 May May 1995, 1993, 12 2 May Jun 1995 1994, 10 Jul 1994, ˇ ˇ ˇ countries species Franklinothrips megalops [Trybom], F. arevac 1 Aug 1998 ˇ MirijevoNovi Beograd 7Resnik May 1993 23Surc Jun 1998 28 Jun 1994 Beli potokSurc 21 MaySurc 1994 vespiformis [Crawford] and Karnyothrips melaleucus c c d b c b b Surc Bela Crkva 26 Apr 1995 JajinciZlatibor 10 25 May Aug 2002, 1997 25 Jul 2002, 9 Jun 2003 Rudnik 26 Aug 1997 Bagnall ’’are used‘‘ in this context (Mulder et al. 1999; EPPO 2003), since it has not been normally recorded in L., L. Poz L. L.

L. Surc greenhouses while the monitoring of the thrips species was performed. L.

L. A. intermedius is very probably an effective predator spp. Surc only in an environment where it does not lack alternative spp. Avala 3 Jun 2003 food; namely, only single specimen of the predator was Triticum aestivum Zea mays Brassica Aster Helianthus annuus Medicago sativa Trifolium pratense T. repens Hordeum vulgare found on vegetative plant parts in spite of massive populations of its potential prey specimens. Future investigation should focus on providing improved life Cultivated hosts, locations and dates of records of banded thrips, conditions of this thrips in the sense of offering a high quality alternative food. As numerous investigations Specimens recorded on the leaves Specimens recorded on the ears/panicles Specimens recorded on the flowers and on the leaves Specimens recorded on the leaves and ears/panicles (in the case of grapevine, a bunch) potential preys of banded thrips (they shared the same plant parts of the hosts) from the Thysanoptera order are presented Brassicaceae Table 4 FamilyAsteraceae Host species Location DateSpecimens recordeda on the flowers b c d Potential preys from the Thysanoptera order Fabaceae Poaceae show, this is of utmost importance for a satisfactory 225 predatory effectiveness of facultative predators. A good EPPO (2003) Commercially used biological control agents (Insecta, example of this is the so-called intercropping, mixed Thysanoptera),http://www.eppo.org/STANDARDS/biocon- trol/bio_list.htm#biolist (27 October 2003) sowing (planting) of different plant species, e.g. cabbage- Franco S, Beignet P, Rat E, Thibout E (1999) The effects of thrips white clover (Theunissen and Schelling 1997), where the on wild and cultivated alliaceous plants in France. Phytoma time of massive occurrence of pests on vegetative parts 514:41–44 of cultivated plants (cabbage) coincides with the flow- Gahukar RT (2004) Bionomics and management of major thrips species on agricultural crops in Africa. Outlook Agric 33:191– ering of the plants which are susceptible for the banded 199 thrips (white clover). Goldarazena A (1999) The kingdom in the Iberian penin- Studying the presence of A. intermedius on various sula and Balearic islands. Order Thysanoptera. Family Aeol- host plants which is presented in this paper constitutes a othripidae Uzel, 1895, http://www.fauna-iberica.mncn.csic.es/ necessary foundation for further investigations on the faunaib/arthropoda/insecta/thysanoptera/aeolothripidae.php. (19 November 2003) predatory efficiency of banded thrips. The data on Gonzalez-Zamora JE, Garcia-Mari F (2003) The efficiency of presence of other thrips species (the majority being several sampling methods for Frankliniella occidentalis (Thy- phytophagous or facultatively phytophagous) on the san., ) in strawberry flowers. J Appl Entomol same plant species are also welcome in the context of 127:516–521 Gremo F, Bogetti C, Scarpelli F (1997) The thrips damaging to knowledge on potential prey of the predator (von Zegula strawberry. L’Inf Agrar 53:85–89 et al. 2003). 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