This review covers aspects of putting essential oil compounds to use either as allelochemicals that manipulate the host selection process of Thysanopteran pest species or as botanical insecticides that kill these pests. Thysanoptera (thrips) make an especially interesting case study in this field, because their increasing economic impact puts some urgency on the development of novel control strategies, especially strategies that incorporate natural compounds. Known facts about the host selection behaviour of thrips are briefly summarized, and methods for the evaluation of thrips responses to volatile and non-volatile plant compounds are outlined. Recent results on the search for attractive and repellent volatiles and for feeding and/or oviposition deterrent essential oil compounds are listed in detail and their potential for use in control strategies against thrips pests is discussed. An overview of plant essential oils used either for insecticidal spraying treatments of crops or for fumigation of crops in greenhouses or fumigation chambers completes the picture of bioactivities. Finally, an outlook on the perspectives for future control strategies against thrips pests is given, including thoughts on the direction of further research needed to fully evaluate the thrips control potential of plant essential oils.
HarrewijnP, MinksAK, MollemaC. (1994) Evolution of plant volatile production in insect-plant relationships. Chemoecology, 5, 55–73.
2.
Regnault-RogerC. (1997) The potential of botanical essential oils for insect pest control. Integrated Pest Management Reviews, 2, 25–34.
3.
IsmanMB. (2000) Plant essential oils for pest and disease management. Crop Protection, 19, 603–608.
4.
MorseJG, HoddleMS. (2006) Invasion biology of thrips. Annual Review of Entomology, 51, 67–89.
5.
LewisT. (1999) Thrips and their control. Pesticide Outlook, 10, 73–77.
6.
KirkWDJ. (2003) The pest and vector from the west: Frankliniella occidentalis. In 7th International Symposium on Thysanoptera: “Thrips, Plants, Tospoviruses: the Millenial Review, 2.-7. July 2001, Reggio Calabria, Italy. MarulloR, MoundL (Eds). CSIRO Entomology, Canberra, Australia, 33–44.
7.
CannonRJC, MatthewsL, CollinsDW. (2007) A review of the pest status and control options for Thrips palmi. Crop Protection, 26, 1089–1098.
8.
ChildersCC. (1997) Feeding and oviposition injuries to plants. In Thrips as crop pests.LewisT (Ed). CAB International, Oxon, UK, 505–537.
9.
Van RijnPCJ, MollemaC, Steenhuis-BroersGM. (1995) Comparative life history studies of Frankliniella occidentalis and Thrips tabaci (Thysanoptera: Thripiadae) on cucumber. Bulletin of Entomological Research, 85, 285–297.
10.
JensenSE. (2000) Insecticide resistance in the western flower thrips Frankliniella occidentalis. Integrated Pest Management Review, 5, 131–146.
11.
BrødsgaardHF. (2004) Biological control of thrips on ornamental crops. In Biocontrol in protected culture.HeinzKM, van DriescheRG, ParrellaMP (Eds). Ball Publishing, Batavia, USA. 253–264.
12.
ShippJL, RamakersPMJ. (2004) Biological control of thrips on vegetable crops. In Biocontrol in protected culture.HeinzKM, van DriescheRG, ParrellaMP (Eds). Ball Publishing, Batavia, USA. 265–276.
13.
GerinC. (1994) Etude par olfactometrie de l'attactivite de la plante hote sur Frankliniella occidentalis. Mededelingen Faculteit Landbouwetenschappen Rijksuniversiteit Gent, 59, 701–707.
14.
ShamshevIV, SelytskayaOG, ChermenskayaTD, BurovVN, RoditakisN. (2003) Behavioural responses of western flower thrips (Frankliniella occidentalis (Pergande)) to extract from meadow-sweet (Filipendula ulmaria Maxim.): laboratory and field bioassays. Archives of Phytopathology and Plant Protection, 36, 111–118.
15.
SmitsPH, van DeventerP, de KogelWJ. (2000) Western flower thrips: reactions to odours and colours. Proceedings of the section Experimental and Applied Entomology of the Netherlands Entomological Society, 11, 175–180.
16.
De KogelWJ, KoschierEH. (2001) Thrips responses to plant odours. In 7th International Symposium on Thysanoptera: “Thrips, Plants, Tospoviruses: the Millenial Review, 2.-7. July 2001, Reggio Calabria, Italy. MarulloR, MoundL (Eds). CSIRO Entomology, Canberra, Australia, 189–190.
17.
BerryN, ButlerRC, TeulonDAJ. (2006) Responses of New Zealand flower thrips (Thrips obscuratus) to synthetic and natural stimuli (odour and colour) in a wind tunnel. New Zealand Journal of Crop and Horticultural Science, 34, 121–129.
18.
HoltmannH. (1963) Untersuchungen zur Biologie der Getreide-Thysanopteren, Teil II. Zeitschrift für angewandte Entomologie, 51, 285–299.
19.
DethierVG, Barton BrowneL, SmithCN. (1960) The designation of chemicals in terms of the responses they elicit from insects. Journal of Economic Entomology, 53, 134–136.
20.
GaumWG, GiliomeeJH, PringleKL. (1994) Resistance of some rose cultivars to the western flower thrips, Frankliniella occidentalis (Thysanoptera: Thripidae). Bulletin of Entomological Research, 84, 487–492.
21.
KoschierEH, de KogelWJ, VisserJ.H. (2000) Assessing the attractiveness of volatile plant compounds to western flower thrips Frankliniella occidentalis. Journal of Chemical Ecology, 26, 2643–2655.
22.
VisserJH. (1986) Host odor perception in phytophagous insects. Annual Review of Entomology, 31, 121–144.
23.
PowEM, HooperAM, LuszniakMC, PickettPA, WadhamsLJ. (1998) Novel strategies for improving biological control of western flower thrips an protected ornamentals - attraction of western flower thrips to verbena plants. In Proceedings of the Brighton Crop Protection Conference Pests and Diseases. Vol. 2. British Crop Protection Council, Farnham, UK, 417–422.
24.
ChermenskayaTD, BurovVN, ManiarSP, PowEM, RoditakisN, SelytskayaOG, ShamshevIV, WadhamsLJ, WoodcockCM. (2001) Behavioural responses of western flower thrips, Frankliniella occidentalis (Pergande), to volatiles from three aromatic plants. Insect Science and its Application, 21, 67–72.
BruceTJA, WadhamsLJ, WoodcockCM. (2005) Insect host location: a volatile situation. Trends in Plant Science, 10, 269–274.
27.
PowEM, BennisonJA, BirkettMA, LuszniakMJ, ManjunataM, PickettPA, SegersIS, WadhamsLJ, WardlowLR, WoodcockCM. (1999) Behavioural responses of western flower thrips (Frankliniella occidentalis) to host plant volatiles. In Proceedings of the 6th International Symposium on Thysanoptera, 27 April – 1 May 1998, Antalya, Turkey. VierbergenG, TunçI (Eds). Akdeniz University, Antalya, Turkey, 121–128.
28.
ManjunathaM, PickettJA, WadhamsLJ, NazziF. (1998) Response of western flower thrips, Frankliniella occidentalis and its predator Amblyseius cucumeris to chrysanthemum volatiles in olfactometer and greenhouse trials. Insect Science and its Application, 18, 139–144.
TeulonDAJ, HollisterB, ButlerRC, CameronEA. (1999) Colour and odour responses of flying western flower thrips: wind tunnel and greenhouse experiments. Entomologia Experimentalis et Applicata, 93, 9–19.
31.
FreyJE, CortadaRV, HelblingH. (1994) The potential of flower odours for use in population monitoring of western flower thrips Frankliniella occidentalis Perg. (Thysanoptera: Thripidae). Biocontrol Science and Technology, 4, 177–186.
32.
HollisterB, CameronEA, TeulonDAJ. (1995) Effect of p-anisaldehyde a yellow color on behavior and capture of western flower thrips. In Thrips Biology and Management: Proceedings of the 1993 International Conference on Thysanoptera.ParkerBL, SkinnerM, LewisT. (Eds). Plenum Press, New York, USA. 571–574.
33.
PenmanDR, OsborneGO, WornerSP, ChapmanRB, McLarenGF. (1982) Ethyl nicotinate: a chemical attractant for Thrips obscuratus (Thysanoptera: Thripidae) in stonefruit in New Zealand. Journal of Chemical Ecology, 8, 1299–1302.
34.
KirkWDJ. (1985) Effect of some floral scents on host finding thrips (Insecta: Thysanoptera). Journal of Chemical Ecology, 11, 35–43.
35.
TeulonDAJ, PenmanDR, RamakersPMJ. (1993) Volatile chemicals for thrips (Thysanoptera: Thripidae) host finding and applications for thrips pest management. Journal of Economic Entomology, 86, 1405–1415.
36.
MuraiT, ImaiT, MaekawaM. (2000) Methyl anthranilate as an attractant for two thrips species and the thrips parasitoid Ceranisus menes. Journal of Chemical Ecology, 26, 2557–2565.
37.
TeulonDAJ, ButlerRC, JamesDE, DavidsonMM. (2007) Odour-baited traps influence thrips capture in proximal unbaited traps in the field. Entomologia Experimentalis et Applicata, 123, 253–262.
38.
TeulonDAJ, DavidsonMM, HedderleyDI, JamesDE, FletcherCD, LarsenL, GreenVC, PerryNB. (2007) 4-Pyridyl carbonyl and related compounds as thrips lures: effectiveness for onion thrips and New Zealand flower thrips in field experiments. Journal of Agricultural and Food Chemistry, 55, 6198–6205.
39.
HooperAM, BennisonJA, LuszniakMC, PickettJA, PowEM, WadhamsLJ. (1999) Verbena x hybrida flower volatiles attractive to western flower thrips, Frankliniella occidentalis. Pesticide Science, 55, 660–662.
40.
ImaiT, MaekawaM, MuraiT. (2001) Attractiveness of methyl anthranilate and its related compounds to the flower thrips, Thrips hawaiiensis (Morgan), T. coloratus Schmutz, T. flavus Schrank and Megalurothrips distalis (Karny) (Thysanoptera: Thripidae). Applied Entomology and Zoology, 36, 475–478.
41.
KnudsenJT, TollstenL, BergstromLG. (1993) Floral scents – a checklist of volatile compounds isolated by headspace techniques. Phytochemistry, 33, 253–280.
42.
WilsonCW, ShawPE, KnightRJ, NagyS, KlimM. (1985) Volatile constituents of carambola (Averrhoa carambola L.). Journal of Agricultural and Food Chemistry, 33, 199–201.
43.
TeulonDAJ, RamakersPMJ. (1990) A review of attractants for trapping thrips with particular reference to glasshouses. IOBC/WPRS Bulletin, 13 (5), 212–214.
44.
DavidsonMM, ButlerRC, WinklerS, TeulonDAJ. (2007) Pyridine compounds increase trap capture of Frankliniella occidentalis (Pergande) in a covered crop. New Zealand Plant Protection, 60, 56–60.
45.
Van TolRWHM, JamesDE, de KogelWJ, TeulonDAJ. (2007) Plant odours with potential for a push-pull strategy to control the onion thrips, Thrips tabaci. Entomologia Experimentalis et Applicata, 122, 69–76.
46.
HowlettFM. (1914) A trap for thrips. Journal of Economic Biology, 9, 21–23.
47.
BrødsgaardHF. (1990) The effect of anisaldehyde as a scent attractant for Frankliniella occidentalis (Thysanoptera: Thripidae) and the response mechanism involved. IOBC/WPRS Bulletin, 13 (5), 36–38.
48.
BaranowskiT, GorskiR. (1993) Chemical attractants in monitoring glasshouse pests. MaterialyXXXIIISesij Naukowej IOR, Poznan, Poland, 94–97.
49.
TeulonDAJ., HollisterB, CameronEA. (1993) Behavioural responses of western flower thrips to anisaldehyde, and implications for trapping in greenhouses. IOBC/WPRS Bulletin, 16 (2), 177–180.
50.
RoditakisNE, LykouressisDP. (1996) Prospects for the use of volatile chemicals and a new pyrrole in integrated pest management of western flower thrips. Acta Horticultura, 431, 513–520.
51.
DavidsonMM, ButlerRC, TeulonDAJ. (2006) Starvation period and age affect the response of female Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) to odor and visual cues. Journal of Insect Physiology, 52, 729–736
52.
TeulonDAJ, NielsenMC, JamesDE, WinklerS, McLachlanARG, PerryNB. (2007) Combination of two odour chemical lures does not increase thrips capture in field bioassays. New Zealand Plant Protection, 60, 61–66.
53.
KaterinopoulosHE, PagonaG, AfratisA, StratigakisN, RoditakisN. (2005) Composition and insect attracting activity of the essential oil of Rosmarinus officinalis. Journal of Chemical Ecology, 31, 111–122.
54.
MorganAC, CrumbSE. (1928) Notes on the chemotrophic responses of certain insects. Journal of Economic Entomology, 21, 913–920.
55.
KirkWDJ. (1987) Effects of trap size and scent on catches of Thrips imaginis Bagnall (Thysanoptera: Thripidae). Journal of the Australian Entomological Society, 26, 299–302.
56.
DemirelN. (2007) Attraction of color cups and plant compounds to thrips species on organic Napa cabbage. Journal of Entomology, 4, 263–266.
57.
VernonRS, GillespieDR. (1990) Spectral responsiveness of Frankliniella occidentalis (Thysanoptera, Thripidae) determined by trap catches in greenhouses. Environmental Entomology, 19, 1229–1241.
58.
NottinghamSF, HardieJ, DawsonGW, HickAJ, PickettJA, WadhamsLJ, WoodcockCM. (1991) Behavioural and electrophysiological responses of aphids to host and non-host plant volatiles. Journal of Chemical Ecology, 17, 1231–1242.
59.
ThieryD, VisserJH. (1986) Masking of host plant odour in the olfactory orientation of the Colorado potato beetle. Entomologia Experimentalis et Applicata, 41, 165–172.
60.
YamasakiT, SatoM, SakoguchiH. (1997) (-)-Germacrene D: masking substance of attractants for the cerambycid beetle, Monochamus alternatus (Hope). Applied Entomology and Zoology, 32, 423–429.
61.
HarrewijnP, van OostenAM, PironPGM. (2001) Natural terpenoids as messengers. A multidisciplinary study of their production, biological functions, and practical applications. Kluwer Academic Publishers, Dordrecht, 1–440.
62.
RoditakisNE, GolfinipoupouNG, ShamshevIV, RoditakisE. (2002) The perspective in use of attractants in IPM programmes of western flower thrips Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) on greenhouse cucumbers. In Congress abstracts of the 7th European Congress of Entomology, October 7 –13, 2002. Savopoulou-SoultaniM. (Ed). Hellenic Entomological Society, Thessaloniki, Greece. 108.
63.
BennisonJ, MauldenK, DewhirstS, PowE, SlatterP, WadhamsLJ. (2003) Towards the development of a push-pull strategy for improving biological control of western flower thrips on chrysanthemum. In 7th International Symposium on Thysanoptera: “Thrips, Plants, Tospoviruses: the Millenial Review”, 2.-7. July 2001, Reggio Calabria, Italy. MarulloR, MoundL. (Eds). CSIRO Entomology, Canberra, Australia. 199–206.
64.
CookSM, KhanZR, PickettJA. (2007) The use of push-pull strategies in integrated pest management. Annual Review of Entomology, 52, 375–400.
65.
RenwickJAA. (1999) Phytochemical modification of taste: an insect model. In Biologically active natural products: Agrochemicals.CutlerHC, CutlerSJ. (Eds). CRC Press, Boca Raton, USA. 221–229.
66.
RenwickJAA. (1990) Oviposition stimulants and deterrents. In CRC Handbook of natural pesticides, Vol. VI, Insect Attractants and Repellents. MorganED, MandavaNB. (Eds). CRC Press, Boca Raton, USA. 151–153.
67.
IsmanMB. (1999) Pesticides based on plant essential oils. Pesticide Outlook, 9, 68–72.
68.
MollemaC, SteenhuisG, InggamerH. (1995) Genotypic effects of cucumber responses to infestation by western flower thrips. In Thrips Biology and Management: Proceedings of the 1993 International Conference on Thysanoptera.ParkerBL, SkinnerM, LewisT. (Eds). Plenum Press, New York, USA. 397–401.
69.
TsaoR, MarvinCH, BroadbentAB, FriesenM, AllenWR, McGarveyBD. (2005) Evidence for an isobutylamide associated with host-plant resistance to western flower thrips, Frankliniella occidentalis, in chrysanthemum. Journal of Chemical Ecology, 31, 103–109.
70.
McKellarRC, McGarveyBD, TsaoR, LuX, KnightKP. (2005) Application of the electronic nose to the classification of resistance to western flower thrips in chrysanthemums. Journal of Chemical Ecology, 31, 2439–2450.
71.
KoschierEH, SedyK, NovakJ. (2002) Influence of plant volatiles on feeding damage caused by the onion thrips Thrips tabaci. Crop Protection, 21, 419–425.
KoschierEH, SedyK. (2003) Labiate essential oils affecting host selection and acceptance of Thrips tabaci Lindeman. Crop Protection, 22, 929–934.
74.
KoschierEH, SedyK. (2002) Die Wirkung pflanzlicher Allomone gegen schädliche Thysanopteren. In 53. Dt. Pflanzenschutztagung, 16.-19.9.2002, Bonn. Biologische Bundesanstalt für Land- und Forstwirtschaft (Ed). Mitteilungen aus der Biologische Bundesanstalt für Land- und Forstwirtschaft Berlin-Dahlem, 390, Parey, Berlin, 159–160.
75.
HiranoC, YasumiK, ItohE, KimCS, HoriikeM. (1994) A feeding deterrent for Thrips palmi (Thysanoptera: Thripidae) found in tomato leaves: isolation and identification. Japanese Journal of Applied Entomology and Zoology, 38, 109–120.
76.
KoschierEH, SedyK. (2003) Effects of plant volatiles on feeding and oviposition of Thrips tabaci. In 7th International Symposium on Thysanoptera: “Thrips, Plants, Tospoviruses: the Millenial Review, 2.-7. July 2001, Reggio Calabria, Italy. MarulloR, MoundL (Eds). CSIRO Entomology, Canberra, Australia. 185–187.
77.
SedyK, KoschierEH. (2003). Bioactivity of carvacrol and thymol against Frankliniella occidentalis and Thrips tabaci. Journal of Applied Entomology, 127, 313–316.
78.
KoschierEH, HoffmannD, RieflerJ. (2007) Influence of salicylaldehyde and methyl salicylate on post-landing behaviour of Frankliniella occidentalis. Journal of Applied Entomology, 131, 362–367.
79.
WiesenbornWD, MorseJG. (1986) Feeding rate of Scirtothrips citri (Moulton) (Thysanoptera: Thripidae) as influenced by life stage and temperature. Environmental Entomology, 15, 763–766.
80.
Van TolRWHM, SwartsHJ, van der LindenA, VisserJH. (2007) Repellence of the red bud borer Resseliella oculiperda from grafted apple trees by impregnation of rubber budding strips with essential oils. Pest Management Science, 63, 483–490.
81.
CowlesRS. (2004) Manipulation of host-finding and acceptance behaviours in insects: Importance to IPM. In Integrated Pest Management: Potential, Constraints and Challenge.KoulO, DhaliwalGS, CuperusGW. (Eds). CABI Publishing, Wallingford UK. 185–204.
82.
KostyukovskyM, RafaeliA, GileadiC, DemchenkoN, ShaayaE. (2002) Activation of octopaminergic receptors by essential oil constituents isolated from aromatic plants: possible mode of action against insect pests. Pest Management Science, 58, 1101–1106.
83.
PriestleyCM, WilliamsonEM, WaffordKA, SattelleDB. (2003) Thymol, a constituent of thyme essential oil, is a positive allosteric modulator of human GABA receptors and a homo-oligomeric GABA receptor from Drosophila melanogaster. British Journal of Pharmacology, 140, 1368–1372.
84.
EnanEE. (2005) Molecular response of Drosophila melanogaster tyramine receptor cascade to plant essential oils. Insect Biochemistry and Molecular Biology, 35, 309–321.
85.
EnanEE. (2005) Molecular and pharmacological analysis of an octopamine receptor from American cockroach and fruit fly in response to plant essential oils. Archives of Insect Biochemistry and Physiology, 59, 161–171.
86.
OparaekeAM, DikeMC, AmatobiCI. (2007) The potential for controlling Megalurothrips sjostedti (Thysanoptera: Thripidae) on cowpea, Vigna unguiculata (L.) Walp flowers using extracts of cashew (Anacardium occidentale L.) products. Archives of Phytopathology and Plant Protection, 40, 252–256.
87.
SolimanMMM. (2007) Phytochemical and toxicological studies of Artemisia L. (Compositae) essential oil against some insect pests. Archives of Phytopathology and Plant Protection, 40, 128–138.
88.
ChiassonH, VincentC, BostanianNJ. (2004) Insecticidal properties of a Chenopodium-based botanical. Journal of Economic Entomology, 97, 1378–1383.
89.
CloydRA, ChiassonH. (2007) Activity of an essential oil derived from Chenopodium ambrosioides on greenhouse insect pests. Journal of Economic Entomology, 100, 459–466.
90.
GundersonCA, SamuelianJH, EvansCK, BrattstenLB. (1985) Effects of the mint monoterpene pulegone on Spodoptera eridana (Lepidoptera: Noctuidae). Environmental Entomology, 14, 859–863.
91.
GutierrezC, FereresA, ReinaM, CabreraR, Gonzalez-ColomaA. (1997) Behavioral and sublethal effects of structurally related lower terpenes on Myzus persicae. Journal of Chemical Ecology, 23, 1641–1650.
92.
ShaayaE, KostjukovskiM, EilbergJ, SukprakarnC. (1997) Plant oils as fumigants and contact insecticides for the control of stored-product insects. Journal of Stored Products Research, 33, 7–15.
93.
LeeS, PetersonCJ, CoatsJR. (2003) Fumigation toxicity of monoterpenoids to several stored product insects. Journal of Stored Products Research, 39, 77–85.
94.
NegahbanM, MoharramipourS. (2007) Fumigant toxicity of Eucalyptus intertexta, Eucalyptus sargentii and Eucalyptus camaldulensis against stored-product beetles. Journal of Applied Entomology, 131, 256–261.
95.
JanmaatAF, de KogelWJ, WolteringEJ. (2002) Enhanced fumigant toxicity of p-cymene against Frankliniella occidentalis by simultaneous application of elevated levels of carbon dioxide. Pest Management Science, 58, 167–173.
96.
KostyukovskyM, RavidU, ShaayaE. (2002) The potential use of plant volatiles for the control of stored product insects and quarantine pests in cut flowers. Acta Horticulturae, 576, 347–358.
97.
HanJB, AhnKS, LeeCK, KimGH. (2006) Fumigant toxicity of pennyroyal and spearmint oils against western flower thrips, Frankliniella occidentalis. Korean Journal of Applied Entomology, 45, 45–49. (abstract only)
98.
YiCG, ChoiBR, ParkCG, AhnYJ. (2006) Fumigant toxicity of plant essential oils to Thrips palmi (Thysanoptera: Thripidae) and Orius strigicollis (Heteroptera: Anthocoridae). Journal of Economic Entomology, 99, 1733–1738.
99.
ErlerF, TuncI. (2005) Monoterpenoids as fumigants against greenhouse pests: toxic, development and reproduction-inhibiting effects. Journal of Plant Diseases and Protection, 112, 181–192.
100.
WolteringEJ, JanmaatAF, de KogelWJ. (2003) Use of controlled atmospheres to enhance essential oil fumigant toxicity against western flower thrips, Frankliniella occidentalis. Acta Horticulturae, 600, 155–159.
101.
FosterSP, HarrisMO. (1997) Behavioral manipulation methods for insect pest-management. Annual Review of Entomology, 42, 123–146.
102.
KornherrC, BlümelS. (2005) Attraction of the monoterpenoids nerol and carvacrol to the predatory flower bug Orius laevigatus (Fieber). IOBC/WPRS Bulletin, 28 (1), 159–162.
103.
KornherrC, HausdorfH, BlümelS. (2005) Side effects of the monoterpenoids nerol and carvacrol on the predatory flower bug Orius laevigatus (Fieber) in the laboratory. IOBC/WPRS Bulletin, 28 (1), 163–166.
104.
IsmanMB. (2006) Botanical insecticides, deterrents, and repellents in modern agriculture and an increasingly regulated world. Annual Review of Entomology, 51, 45–66.
105.
StewartRR, WeatherstoneI. (2002) Plant volatiles in pest control – regulatory aspects. In Congress abstracts of the 7th European Congress of Entomology, October 7–13, 2002.Savopoulou-SoultaniM. (Ed). Hellenic Entomological Society, Thessaloniki, Greece. 149.
