Introduction ORIGINAL ARTICLE. Stanislav Trdan Æ Ljiljana Andjus Emilija Raspudic Æ Milica Kacˇ

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1 J Pest Sci (2005) 78: DOI /s ORIGINAL ARTICLE Stanislav Trdan Æ Ljiljana Andjus Emilija Raspudic Æ Milica Kacˇ Distribution of Aeolothrips intermedius Bagnall (Thysanoptera: Aeolothripidae) and its potential prey Thysanoptera species 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 species on cultivated plants in Slovenia ( ), Croatia ( ), and Serbia and Montenegro ( ) are presented in this study. The aim of the investigation was to study the host plant distribution of the predator Aeolothrips intermedius Bagnall and its potential prey. Banded thrips were found on 30 different host plant species belonging to 16 botanical families, always in mixed populations with phytophagous or facultative phytophagous insects (including 18 Thysanoptera species). On the vegetative parts of the cultivated plants, banded thrips were found less numerous in spite of the massive population of some harmful thrips species. This indicates highly important role of pollen as alternative food for Aeolothrips intermedius. Keywords Aeolothrips Æ Distribution Æ Hosts Æ Predator Æ Thysanoptera Communicated by W. Schwenke S. Trdan (&) Chair of Entomology and Phytopathology, Biotechnical Faculty, Department of Agronomy, University of Ljubljana, Jamnikarjeva 101, 1111 Ljubljana, Slovenia stanislav.trdan@bf.uni-lj.si Tel.: Fax: L. Andjus Natural History Museum, Njegosˇ eva 51, Belgrade, Serbia and Montenegro E. Raspudic Faculty of Agriculture, University of J. J. Strossmayer in Osijek, Trg Sv. Trojstva 3, Osijek, Croatia M. Kacˇ Chair of Chemistry, Biotechnical Faculty, Department of Food Science and Technology, University of Ljubljana, Jamnikarjeva 101, 1111 Ljubljana, Slovenia Introduction Thrips as crop pests Thrips are mostly known as insects living in flowers, but many species live only on leaves, a few are predatory, and probably about half of the species feed on fungi (Palmer et al. 1989). The damage on plants caused by thrips due to their feeding or their deposition of eggs is often ascribed to other pests, especially if observed and/or evaluated by non-professionals. Of 5,000 thrips species recorded worldwide, only 1% are known to be economically harmful (Mound and Teulon 1995); while on different continents, the number of harmful Thysanoptera species very rarely approaches the number of 10 (Jenser and Czencz 1988; Chang 1995; Gahukar 2004). In this sense, western flower thrips (Frankliniella occidentalis [Pergande]) (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 greenhouse thrips (Hercinothrips femoralis [Reuter]), gladiolus thrips (Thrips simplex [Morison]), pear thrips (Taeniothrips inconsequens [Uzel]), pea thrips (Kakothrips pisivorus [Westwood]), privet thrips (Dendrothrips ornatus [Jablonowski]) and some other species are considered less important, since their occurrence and the following damages on cultivated plants are not observed every year in economically important level (Kirk 1985; Jenser and Czencz 1988; Andjus 1997; Trdan 2003). Thrips as biological control agents Though much progress has been made during the last 15 years in the sense of making the control of insects (Insecta), mites (Acarina) and other pests on cultivated crops more environmentally friendly (Wilson et al. 1995; Milevoj 1999), and the number of such investigations is steadily increasing during the last years (Arthurs et al.

2 ; Kishimoto 2003; Nielsen et al. 2003), it is an interesting fact that there are not enough reliable data on the predatory efficiency of the autochthonous representatives of the Thysanoptera order. Some species of the Haplothrips, Scolothrips, Franklinothrips, Karnyothrips and Aeolothrips genera as well as those of the Aleurodothrips, Allelothrips, Androthrips, Apterygothrips, Desmidothrips, Desmothrips, Leptothrips, Erythrothrips, Mymarothrips, Parascolothrips, Podothrips, Stictothrips, Stomatothrips, Symphyothrips, Trichinothrips, Veerabahuthrips, Xylaplothrips and Andrewarthaia genera (Palmer et al. 1989; Riudavets 1995; zur Strassen 1995; Sabelis and van Rijn 1997) are well-known predators; but in Europe, particularly some species of the genus Aeolothrips are considered as potentially important autochthonous facultative predators. The great majority of the species of this genus (about 100 species have been recorded so far) spend most of their lives in flowers, feeding on small arthropods and pollen (Harper et al. 1989; Patrzich and Klumpp 1991; Mound and Kibby 1998; Yee et al. 2001). Till now, what is known on the impact of two different types of food for the life and development of facultative predators is not adequate, although some studies in this context were performed in the last years (von Zegula et al. 2003). Aeolothrips species as predators More than 20 species of the Aeolothrips genus were recorded in Europe (Table 1). The majority of the representatives 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 Croatia Germany Hungary Iberian peninsula Italy Poland Romania Serbia and Montenegro Slovenia The Netherlands A. albicinctus Haliday, A. astutus Priesner, A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. versicolor Uzel, A. vittatus Haliday A. collaris Priesner, A. ericae Bagnall, A. fasciatus (L.), A. gloriosus Bagnall, A. intermedius Bagnall, A. melaleucus Haliday, A. propinquus Bagnall, A. versicolor Uzel A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. propinquus Bagnall, A. tenuicornis Bagnall, A. versicolor Uzel, A. vittatus Haliday A. albicinctus Haliday, A. astutus Priesner, A. ericae Bagnall, A. intermedius Bagnall, A. melaleucus Haliday, A. priesneri Knechtel, A. propinquus Bagnall, A. versicolor Uzel, A. vittatus Haliday A. albicinctus Haliday, A. andalusiacus zur Strassen, A. balati Pelikan, 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 A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, 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 A. astutus Priesner, A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. propinquus Bagnall, A. pulcher Oettingen, A. versicolor Uzel, A. vittatus Haliday A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, 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 A. albicinctus Haliday, A. astutus Priesner, A. collaris Priesner, A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. pulcher Oettingen, A. versicolor Uzel, A. vittatus Haliday A. albicinctus Haliday, A. astutus Priesner, A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. melaleucus Haliday, A. versicolor Uzel A. albicinctus Haliday, A. ericae Bagnall, A. fasciatus (L.), A. intermedius Bagnall, A. manteli Titschack, A. melaleucus Haliday, A. propinquus Bagnall, A. tenuicornis Bagnall, A. versicolor Uzel, A. vittatus Haliday zur Strassen, personel communication Raspudic et al zur Strassen, personel communication zur Strassen, personel communication Goldarazena 1999 zur Strassen, personel communication Kucharczyk, personel communication Oromulu, personel communication Trdan et al Trdan et al Vierbergen, personel communication

3 219 predators (Mound and Kibby 1998), though several differences originating from different size of the species, their preference towards different host plants or arthropods, 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). A. intermedius is the most common species of this genus on the Old Continent (Deligeorgidis et al. 2002) and it is frequently mentioned in the context of biological control (von Zegula et al. 2003). Adult specimens of this species are quite similar to those of Aeolothrips fasciatus L., so the two species can very easily lead to confusion (zur Strassen, personel communication). Adults and larvae of banded thrips, as is a common name of A. intermedius, were found as predator of 44 species of Thysanoptera order (Riudavets 1995). In Europe, it has usually been mentioned as a predator on onion thrips (T. tabaci Lindeman) (Bournier et al. 1978; Lacasa et al. 1982; Torres-Vila et al. 1994; Franco et al. 1999), other representatives of Thrips genus (Lacasa et al. 1989; Gremo et al. 1997), as well as together with greenhouse thrips (Heliothrips haemorrhoidalis Bouche ), Odontothrips confusus Priesner (Bournier 1979; Khosbayar 2001) and with the species of Haplothrips genus (Dyadechko et al. 1971). Banded thrips also feed on some mite species, larvae and eggs of psyllids (Psyllina) and whiteflies (Aleyrodina) as well as on aphids (Aphidina). It was determined by the laboratory experiments that A. intermedius can complete its life cycle feeding on Tetranychus urticae Koch, T. cinnabarina (Boisduval), T. atlanticus McGregor, Panonychus ulmi Koch and Cenopalpus pulcher Canestrinı` & Fanzago, while the life cycle cannot be completed by feeding on Frankliniella occidentalis (Pergande) (von Zegula et al. 2003), Psylla melanoneura Fo rster, Trioza spp., Trialeurodes vaporariorum (Westwood), Aphis fabae Scopoli, Aphis cracivora Koch, A- cyrthosiphon pisum Harris and Myzus persicae (Sulzer) (Bournier et al. 1979; Berest 1980). Mostly the prey of the banded thrips is considered to be pests on cultivated plants in the open, which speaks for the specific bionomics of the banded thrips. This refers mostly to its susceptibility to different prey species and various kinds of pollen. The fact is also in accordance with the theory that thrips is not an efficient predator in protected places. Incapability of thrips rearing in laboratory conditions also confirms this (van Lenteren 1999). Banded thrips is omnivorous, and can complete its development by feeding only on flowers, where it was found to eat pollen. It does not extract nutritional food from leaves and stems, but pierces them to extract water (Bournier et al. 1979; Lacasa 1980). In the European scientific literature, only a few detailed and/or abundant enough information on host distribution of A. intermedius on different cultivated plants (Andjus 2004) and its potential prey species that could be of potential use in the investigation of its predatory role (von Zegula et al. 2003), could be found. So, an investigation of its presence on cultivated crops in three geographically close countries which exhibit considerably different reliefs and climates was undertaken. Materials and methods The monitoring of thrips presented in this contribution was carried out in Slovenia during (Trdan 2003), on the territory of Serbia and Montenegro during (Andjus 1997, 1998) and in Croatia during (Raspudic 2003). Samples were collected from 26 locations in Slovenia (Fig. 2), from 15 locations in Croatia and from 11 locations in Serbia and Montenegro (Figs. 1). On some locations, samples were collected in different periods. Only adults were sampled because identification of larvae is more difficult (Mound and Kibby 1998). All samples showing the presence of A. intermedius were collected in the open. The sampling was performed with usual thripsological instruments. Different cultivated plants were beaten over a small white plastic tray. Depending on the host species and growing stage of the hosts different aboveground parts of the plants were searched more precisely. From the tray surface, the specimens were removed with a fine brush. Collected thrips were kept in plastic 1.5 ml Eppendorf tubes in AGA solution, a mixture of ten parts of 60% ethyl alcohol, one part of glycerine and one part of acetic acid. This mixture helps to distend the body of most thrips and keeps the limbs supple. Stored specimens were transferred to 60% alcohol and kept in the dark at the temperature about 0 C to prevent loss of colour (Palmer et al. 1989; Mound and Kibby 1998; Gonzalez-Zamora and Garcia-Mari 2003). Thrips specimens were collected from the end of April until the beginning of September, in different growing stages of host plants. The number of the sampling spots on selected parcels and the number of the sampled plant parts (organs) of host plants depend on the size of the parcel, host plant species and population size of thrips. On larger parcels, where thrips populations were less abundant, the number of the sampling spots was higher. Bigger plants (trees, bushes) were the only sampled plants on some parcels, if the thrips populations were high. Different sizes of samples were taken into consideration, depending on the density of the thrips population on the cultivated plants in question (1 50 specimen per plant species per location). In spite of the shortest sampling period in Slovenia, the number of collected specimens was the highest (67). Few specimens were collected in Serbia and Montenegro (28) and least of all in Croatia (17). In some cases, specimens on the same host plant and same location were collected in different periods, but this was not the rule. The morphological identification was performed using standard light microscope in the facilities of the Institute of Phytomedicine (Biotechnical Faculty, Department of Agronomy, Ljubljana, Slovenia), Insti-

4 220 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, Croatia) and that of the Natural History Museum (Belgrade, Serbia and Montenegro). The standard morphological identification keys by Priesner (1964), Schliephake and Klimt (1979), Jenser (1982), Palmer et al. (1989), Moritz (1994) as well as those by Mound and Kibby (1998) were used. Tables give the results of the thrips monitoring in Slovenia during (Table 2), in Croatia during (Table 3) and on the territory of Serbia and Montenegro during (Table 4). It gives the host plants only for banded thrips, A. intermedius, and for those thrips species that were present in considerable 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

5 221 Table 2 Cultivated hosts, locations and dates of records of banded thrips, A. intermedius Bagnall, in Slovenia during 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., Allium fistulosum L. a Ljubljana 6 Jun 2000, 18 Jul 2001 Thrips tabaci Lindeman Asparagaceae Asparagus officinalis L. a Ljubljana 18 Jul 2001 Chirothrips manicatus Haliday b Sˇ 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 Stročja 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., T. repens L. Preserje pri Radomljah 12 May 2000, 29 Jun 2000, 15 Jun 2001, 17 Aug 2001 Frankliniella intonsa (Trybom), 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 Želimlje 23 Aug 2001 Oleaceae Olea europaea L. b Osp 7 Jun 2001 Thrips spp. Poaceae Avena sativa L. Hordeum vulgare L. Triticum aestivum L. Zea mays L. c Ljubljana 19 Jun 2001, 22 Jun 2001, 30 Jul 2001 d Radomlje 18 Jun 2000 d Župelevec 2 Jul 2000 Haplothrips aculeatus (Fabricius), Limothrips denticornis Haliday, Frankliniella tenuicornis Hood c Spodnje Sˇ kofije 28 Apr 2000 Limothrips denticornis Haliday, Haplothrips aculeatus (Fabricius), d Spodnji Brnik 4 Jun 2001 Frankliniella tenuicornis Hood c Ptujska gora 2 May 2001, 5 May 2001 b Spodnji Velovlek 28 Apr 2001 c Stojnci 5 May 2001 b Gorisˇ nica 13 May 2000, 14 May 2000, 27 Apr 2001, 19 May 2001 b Pragersko 5 May 2001 b Stojnci 5 May 2001 Haplothrips aculeatus (Fabricius), Haplothrips tritici Kurdjumov, Limothrips denticornis Haliday, Limothrips cerealium Haliday, Frankliniella tenuicornis Hood, Frankliniella intonsa (Trybom) b Bolehnečici 6 Jul 2000 Haplothrips aculeatus (Fabricius), Haplothrips spp., Cephalothrips monilicornis (O.M. Reuter), Chirothrips manicatus Haliday

6 222 Table 2 (Contd.) Family Host species Location Date Potential preys from the Thysanoptera order b Pondor 5 Jul 2000 b Pacinje 5 Jul 2000 b Rakicˇ an 6 Jul 2000 Ljubljana 22 Jun 2001 Thrips fuscipennis Haliday Sˇ ikole 15 Aug 2001 Polygonaceae Fagopyrum esculentum Moench Rosaceae Rosa spp. Postojna 27 Aug Frankliniella intonsa (Trybom), Thrips Rakitnica 26 Jul 2001, 18 Aug 2001 fuscipennis Haliday Sˇ empeter pri Novi Gorici 22 Aug 2001 Vitaceae Vitis vinifera L. Brje pri Komnu 9 Jun 2001 Thrips vulgatissimus Haliday c Spodnje Sˇ kofije 7 Jun 2001 Specimens recorded on the flowers a Specimens recorded on the flowers and on the leaves b Specimens recorded on the leaves Specimens recorded on the leaves and ears/panicles (in the case of grapevine, a bunch) c d Specimens recorded on the ears/panicles 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 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). Sampling locations and the host plants of recorded Thysanoptera species are to be found in Figs. 1, 2. 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 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 (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 family) the banded thrips inhabit these host plants as well. The density of the thrips population is strongly depended 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 recorded, populations of 18 other thrips species (mostly of the Terebrantia suborder) were also present they were mostly phytophagous. Due to its massive populations and extreme polyphagy (Deligeorgidis et al. 2002), the flower thrips (Frankliniella intonsa [Trybom]) was stated to be the dominant species. It is a well-known facultative phytophagous insect, pollen being its important food source (Murai and Loomans 2001). A. intermedius were not found in the flowers of fruit bearing plants and ornamentals of the Rosaceae family (Prunus spp., Malus domestica Borkh., Cydonia oblonga Mill., Pyrus communis 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 cannot describe the Rosaceae family as non-susceptible for the banded thrips, since this same species exhibits a massive occurrence in the flowers of roses (Rosa spp.) in summer. Obviously, growing stages of the fruit bearing 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 massive populations of more harmful thrips species e.g. onion thrips (T. tabaci Lindeman) on the leaves of leek, onion as well as cabbage is surprisingly low, compared to its number in the flowers of some

7 223 Table 3 Cultivated hosts, locations and dates of records of banded thrips, A. intermedius Bagnall, in Croatia during 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. saccharifera Lange Krndija 5 Jun 1995 Thrips tabaci Lindeman, Frankliniella intonsa (Trybom) 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. b Kopanica 20 Jul 1994 Haplothrips aculeatus (Fabricius), Haplothrips spp., b Berava 20 Jul 1994 Cephalothrips monilicornis (O.M. Reuter), Chirothrips manicatus Haliday b Cˇ epin 24 Jul 1995 b Maslenica 30 Jul 1995 b Marijanci 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 Specimens recorded on the leaves and ears/panicles (in the case of grapevine, a bunch) c d Specimens recorded on the ears/panicles

8 224 Table 4 Cultivated hosts, locations and dates of records of banded thrips, A. intermedius Bagnall, on the territory of Serbia and Montenegro during 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 Asteraceae Aster spp. Avala 3 Jun 2003 Frankliniella intonsa (Trybom) Helianthus annuus L. Požarevac 1 Aug 1998 Thrips physapus Linnaeus, Thrips tabaci Lindeman, Surčin 10 Jul 1994, 21 Jul 1994, 28 Jul 1994 Frankliniella intonsa (Trybom) Brassicaceae Brassica spp. Surčin 30 Apr 1995, 4 May 1995, 8 May 1995, 12 May 1995 Frankliniella intonsa (Trybom) Bela Crkva 26 Apr 1995 Fabaceae Medicago sativa L. Surčin 9 May 1993, 2 Jun 1994, 10 Jul 1994, Odontothrips confusus Priesner, Frankliniella intonsa (Trybom), 21 Jul 1994, 14 Aug 1994, 30 Apr 1995 Thrips tabaci Lindeman Jajinci 10 May 2002, 25 Jul 2002, 9 Jun 2003 Zlatibor 25 Aug 1997 Frankliniella intonsa (Trybom), Haplothrips spp., Rudnik 26 Aug 1997 Thrips tabaci Lindeman Trifolium pratense L., T. repens L. c Beli potok 21 May 1994 Limothrips denticornis Haliday, Haplothrips aculeatus (Fabricius), c Surcˇ in 11 May 1994, 18 May 1994, 2 Jun 1994 Frankliniella tenuicornis Hood Poaceae Hordeum vulgare L. d Mirijevo 23 Jun 1998 Haplothrips aculeatus (Fabricius), Haplothrips tritici Kurdjumov, b Novi Beograd 7 May 1993 Limothrips denticornis Haliday, Limothrips cerealium Haliday, c Surcˇ in 18 May 1993, 11 May 1994, 2 Jun 1994 Frankliniella tenuicornis Hood, Frankliniella intonsa (Trybom) Triticum aestivum L. b Resnik 28 Jun 1994 Haplothrips aculeatus (Fabricius), Haplothrips spp., b Surcˇ in 10 Jul 1994, 21 Jul 1994 Cephalothrips monilicornis (O.M. Reuter), Chirothrips manicatus Haliday Zea mays L. Specimens recorded on the flowers a Specimens recorded on the flowers and on the leaves b Specimens recorded on the leaves Specimens recorded on the leaves and ears/panicles (in the case of grapevine, a bunch) c d Specimens recorded on the ears/panicles other hosts. This speaks in favour of alternative food importance in the living space of A. intermedius, where the insect performs its role as a predator or pollinator. Conclusion It was established that the banded thrips spends much of its life in the flowers of the host plants, where it feeds on pollen and arthropods. As pollen is an important food source of the banded thrips (Bournier et al. 1979; Lacasa 1980), the bionomics and the predatory effectiveness of the species are closely connected with the developmental stages of its hosts. It is well-known that its massive occurrence takes place during the flowering of the plants which are most attractive for the insect (Khosbayar 2001), but it is not sure whether this phenomenon is due 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 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 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 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 the present investigation deals only partly with this issue. It was established that banded thrips reduces the population 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 countries species Franklinothrips megalops [Trybom], F. vespiformis [Crawford] and Karnyothrips melaleucus Bagnall are used in this context (Mulder et al. 1999; EPPO 2003), since it has not been normally recorded in greenhouses while the monitoring of the thrips species was performed. A. intermedius is very probably an effective predator only in an environment where it does not lack alternative food; namely, only single specimen of the predator was found on vegetative plant parts in spite of massive populations of its potential prey specimens. Future investigation should focus on providing improved life conditions of this thrips in the sense of offering a high quality alternative food. As numerous investigations show, this is of utmost importance for a satisfactory

9 225 predatory effectiveness of facultative predators. A good example of this is the so-called intercropping, mixed sowing (planting) of different plant species, e.g. cabbagewhite clover (Theunissen and Schelling 1997), where the time of massive occurrence of pests on vegetative parts of cultivated plants (cabbage) coincides with the flowering of the plants which are susceptible for the banded thrips (white clover). Studying the presence of A. intermedius on various host plants which is presented in this paper constitutes a necessary foundation for further investigations on the predatory efficiency of banded thrips. The data on presence of other thrips species (the majority being phytophagous or facultatively phytophagous) on the same plant species are also welcome in the context of knowledge on potential prey of the predator (von Zegula et al. 2003). 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