6. Discussione
6.2 Prey choice negli adulti
L'adulto di H. axyridis non ha mostrato una significativa capacità di discriminare una preda rispetto ad un'altra, poiché quando messo di fronte ad una scelta il coccinellide non ha mostrato preferenze. Questo potrebbe essere dato dal fatto che, in condizioni di laboratorio, al coccinellide mancano delle informazioni base per localizzare la preda, informazioni che normalmente sono presenti in un ambiente
37
naturale. Le condizioni del laboratorio, infatti, non permettono di offrire al predatore informazioni e stimoli essenziali, in particolare quelli derivanti dalla emissioni di volatili da parte delle piante infestate dalla vittima e che costituiscono una sorgente di informazioni importanti, in grado di modificare sia il comportamento degli erbivori sia quello dei loro nemici naturali (Cortesero et al., 2000). La produzione di allomoni da parte della pianta infestata da afidi (Alhemedi et al., 2010) aiuta il coccinellide nella localizzazione della vittima in condizioni naturali, come pure il contrasto con il colore della foglia (Obata, 1997). In generale, poi, si osserva una maggiore capacità discriminatoria dell'adulto legata alla vastità di stimoli a cui l'insetto fa affidamento per la localizzazione della preda, che sono di tipo visivo e chimico (Obata, 1997).
Propylea quatuordecimpunctata: viene attaccata maggiormente dagli adulti di H. axyridis rispetto a
quanto non facciano le larve e mostra anche in questi test poca mobilità se non sottoposta a stimoli. Tuttavia, l’adulto di H. axyridis durante la predazione nei suoi confronti non ha mai portato a termine l'attacco, desistendo dal consumare la vittima. E’ noto in generale che larve, pre-pupe e pupe sono spesso equipaggiate con composti chimici e contengono sostanze simili agli alcaloidi difensivi presenti nelle loro uova. In più, adulti, larve e pre-pupe, se disturbati, possono rilasciare secrezioni (Majerus, 1994) contenenti alcaloidi e pirazine che fungono da deterrenti per molti predatori (Ware e Majerus, 2008). Più in generale, le specie di coccinelle hanno alcaloidi specie-specifici (Pasteels et al., 1973) che possono agire da deterrente nell'intraguild predation da parte di altre specie di coccinellidi. Questo potrebbe essere la spiegazione al comportamento di H. axyridis nei confronti di Propylea quatuordecimpunctata, seppure da dimostrare con ulteriori e più approfondite indagini.
Scymnus marginalis: assieme a P. quatuordecimpunctata, è soggetta ad una maggiore predazione da
parte degli adulti di H. axyridis rispetto alle larve, tale predazione risulta comunque non significativa nonostante le dimensioni ridotte rispetto a H. axyridis e l'assenza di alcaloidi (Pasteels et al., 1973).
6.3 Time choice
Per quanto riguarda la differenza fra i tempi di localizzazione, non si hanno differenze significative quando il predatore si trova a localizzare la preda tipica o una alternativa, in nessuno dei due stadi. Tuttavia, pur non esistendo significative differenze fra i tempi di scelta nei due stadi del predatore è possibile evidenziare una tendenza da parte degli adulti di H. axyridis ad impiegare più tempo nella
38
localizzazione della preda. Questo dato conferma la difficoltà dello stadio adulto di orientarsi nella localizzazione e scelta della preda in assenza delle informazioni derivanti dall'ambiente esterno (Obata, 1997; Alhmedi et al., 2010) mentre nel caso della larva, anche se meno influenzata dalle condizioni di laboratorio rispetto all'adulto, le difficoltà sono legate alla casualità dell'incontro (Kawai, 1976).
In conclusione i risultati hanno mostrato differenze tra i comportamenti nella ricerca della preda da parte degli stadi giovanili rispetto agli stadi adulti.
Le larve hanno dimostrato di utilizzare, principalmente, comportamenti elementari nella scelta della preda manifestando una ricerca caotica e intensa per poter continuare il proprio sviluppo. Gli adulti invece, non hanno manifestato nessuna selettività ma è stato possibile osservare come la ricerca sia basata su stimoli tattili e chimici mediante l'utilizzo delle antenne e una loro continua pulizia.
Complessivamente comunque la tendenza all’intraguild predation nei confronti delle specie testate sembra essere maggiormente spostata verso gli adulti rispetto alle larve, anche se ulteriori studi si rendono necessari.
39
BIBLIOGRAFIA
Adriaens T., Branquart E., Maes D. (2003) - The multicoloured Asian ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae), a threat for native aphid predators in Belgium? - Belgian Journal of
Zoology, 133 (2): 195-196.
Agarwala B. K. (1991) - Why do ladybirds (Coleoptera: Coccinellidae) cannibalize? - J. Biosci, Vol.16,
Number 3, September 1991, pp 103-109.
Agarwala B. K., Yasuda H. (2001) - Larval interactions in aphidophagous predators: effectiveness of wax cover as defence shield of Scymnus larvae against predation from syrphids - Entomol. Exp. Appl., 100: 101-107.
Agarwala B. K., Dixon A. F. G. (1992) - Laboratory study of cannibalism and interspecific predation in ladybirds - Ecological Entomology, 17 (4): 303-309.
Alhmedi A., Haubruge E., Francis F. (2010) - Identification of limonene as a potential kairomone of the harlequin ladybird Harmonia axyridis (Coleoptera: Coccinellidae) - European Journal of
Entomology, 107 (4): 541-548.
Allan J. D., Flecker A. S., McClintock N. L. (1987) - Prey preference of stoneflies: Sedentary vs mobile prey - Oikos 49: 323-331.
Bezzerides A. L., McGraw K. J., Parker R. S., Jad Husseini J. (2007) - Elytra color as a signal of chemical defence in the Asian ladybird beetle Harmonia axyridis - Behavior Ecology and Sociobiology, 61: 1401-1408.
Brown P. M. J., Adriaens T., Bathon H., Cuppen J., Goldarazena A., Hägg T., Kenis M., Klausnitzer B. E. M., Kovar I., Loomans A. J. M., Majerus M. E. N., Nedved J., Pedersen J., Rabitsch W., Roy H. E., Ternois V., Zakharov A., Roy D. B. (2008) - Harmonia axyridis in Europe: spread and distribution of a non-native coccinellid - BioControl, 53 (1): 5-21.
40
Burgio G., Santi F., Lanzoni A., Masetti A., De Luigi V., Melandri M., Reggiani A., Ricci C., Loomans A. J. M., Maini S. (2008) - Harmonia axyridis recordings in northern Italy - Bulletin of
Insectology, 61 (2): 361-364.
Burgio G., Santi F., Maini S. (2002) - On intra-guild predation and cannibalism in Harmonia axyridis (Pallas) and Adalia bipunctata L. (Coleoptera: Coccinellidae) - Biological Control, 24 (2): 110-116.
Canepari C. (1983) - Le specie italiane del gruppo dello Scymnus frontalis Fab. con descrizione di due nuove specie (Coleoptera Coccinellidae) - Giornale Italiano di Entomologia, 1: 179-204
Canovai R., Lucchi A. (2011) - La coccinella arlecchino in vigneti della Toscana - L'informatore
agrario, N. 11/2011, p.65.
Canovai R., Loni A., Gandini L., Lucchi A. (2014) - Ladybirds in Tuscan vineyard (Coleoptera: Coccinellidae) - Integrated Protection and Production in Viticulture IOBC-WPRS Bulletin, 105: 241- 244.
Celli G. (1985) - Lotta biologica, produzione agricola, conservazione ambientale, qualche considerazione di ordine generale - in Metodi alternativi alla lotta biologica (I convegno internazionale,
Cesena Agricoltura, Cesena, 1985), Edd. Goidanich G., Baronio: 115-130.
Chapin J. B., Brou V. A. (1991) - Harmonia axyridis (Pallas), the third species of the genus to be found in the United States (Coleoptera: Coccinellidae) - Proceedings of the Entomological Society Washington, 93: 630-635.
Cortesero A. M., Stapel J. O., Lewis W. J. (2000) - Understanding and manipulating plant attributes to biological control - Biol. Control, 17: 35-49.
Daloze D., Braekman J. C., Pasteels J. M. (1994/1995) - Ladybird defence alkaloids: structural, chemotaxonomic and biosynthetic aspects (Col.: Coccinellidae) - Chemoecology 5/6, 3/4: 173-183.
41
against bean aphid Aphis craccivora Koch (Homoptera: Aphididae) - Journal of Pest Science, 81 (3): 153-159.
de Almeida L. M., da Silva V. B. (2002) - First record of Harmonia axyridis (Pallas) (Coleoptera, Coccinellidae): a lady beetle native to the Palaearctic region - Revista Brasileira de Zoologia, 19: 941- 944.
De Bach P. (1964) - The scope of biological control - In: De Bach P (ed) Biological control of insect
pests and weeds. Chapman and Hall, London: 3-20.
Dixon A. F. G. (1985) - Structure of aphid populations - Annu. Rev. Entomol., 30: 155-174.
Dixon A. F. G. (2000) - Insect Predator-Prey Dynamics: Ladybird beetles and Biological Control -
Cambridge, United Kingdom: Cambridge University Press.
Dutto M. (2010) - Harmonia axyridis (Pallas, 1773) (Coleoptera, Coccinellidae): un'infestante stagionale emergente in ambito domestico - Biologia ambientale, 24 (2): 5-8.
Eastop V. F. (1966) - A taxonomic study of Australian Aphidoidea (Homoptera) - Aust. J. Zool. 14: 399- 592.
Ejbich K. (2003) - Producers in Ontario and northern U.S. bugged by bad odors in wine - Wine Spectator, 15 May: 16.
El-Sebaey I. I. A., El-Gantiry A. M. (1999) - Biological aspects and description of different stages of
Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) - Bulletin of the Faculty of Agriculture, Cairo University, 50 (1): 87-97.
Ettifouri M., Ferran A. (1993) - Influence of larval rearing diet on the intensive searching behavior of
Harmonia axyridis [Col.: Coccinellidae] larvae - Entomophaga 38: 51-59.
42
(Coleptera, Coccinellidae) larvae - Journal of Insect behavior, 5 (2): 147-159.
Ferran A., El-Arnaouty S., Beyssat-Arnaouty V., Galal H. (2000) - Introduction and release of the coccinellid Harmonia axyridis Pallas for controlling Aphis craccivora Koch on faba beans in Egypt -
Egypt. J. Biol. Pest Control 10: 129-136.
Fox L. R. (1975) - Cannibalism in natural populations - Annu. Rev. Ecol. Syst., 6: 87-106.
Gordon R. D. (1985) - The Coleoptera (Coccinellidae) of America north of Mexico - Journal of the New
York Entomological Society, 93: 1-912.
Gourreau J. M. (1947) - Systematique de la tribu des Scymnini (Coccinellidae) - Ann. De Zoo., Eco.
Ani., Institut National de la Recherche Agronomique, Paris: 122 pp.
Grez A. A., Zaviezo T., Roy H. E., Brown P. M., Bizaman G. (2016) - Rapid spread of Harmonia
axyridis in Chile and its effects on local coccinellid biodiversity - Diversity and Distributions: 22 (9):
982-994.
Grill C. P., Moore A. J. (1998) - Effects of a larval antipredator response and larval diet on adult phenotype in an aposematic ladybird beetle - Oecologia, 114: 274-282.
Gutierrez A. P., Morgan D. J., Havenstein D. E. (1971) - The ecology of Aphis craccivora Koch and SCSV. I. The phenology of aphid populations and the epidemiology of virus in pastures - J. Appl. Ecol., 8: 699-721.
Hamilton W. O. (1964) - The genetical evolution of social behaviour - I. J. Theor. Biol., 7: 1-16.
Han B. Y., Chen Z. M. (2000) - Behavior response of four Leis axyridis varieties to volatiles from tea and Toxoptera aurantii - Chinese Journal of Applied Ecology, 11: 413-416.
Han B. Y., Chen Z. M. (2002) - Composition of the volatiles from intact and tea aphid-damaged tea shoots and their allurement to several natural enemies of the tea aphid - Journal of Applied Entomology,
43
126: 497-500.
Harmon J. P., Losey J. E., Ives A. R. (1998) - The role of vision and color in the close proximity foraging behavior of four coccinellid species - Oecologia, 115: 287-292.
He J. L., Ma E. P., Shen Y. C., Chen W. L., Sun X. Q. (1994) - Observations of the biological characteristics of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) - Journal of the Shanghai
Agricultural College 12: 119-124.
Hemptinne J. L. (1988) - Ecological requirements for hibernating Propylea quatuordecimpunctata (L.) and Coccinella septempunctata [Col.: Coccinellidae] - Entomophaga Volume 33, Issue 4, pp 505-515.
Hironori Y., Katsuhiro S. (1997) - Cannibalism and interspecific predation in two predatory ladybirds in relation to prey abundance in the field - Entomophaga, 42: 153-163.
Hodek I. (1973) - Life history and biological properties - In: Hodek I, editor. Biology of Coccinellidae,
70-76. The Hauge, Holland: Dr. W. Junk N. V., Publishers.
Hodek I. (1996) - Food relationships - In: Hodek I, Honek A., editors. Ecology of Coccinellidae.
Dordrecht, The Netherlands: Kluwer Academic Publishers.
Honek A., Evans E. W. (2012) - Food relationships - in Hodek I, Van Emden HF, Honek A., Ecology
and Behaviour of the Ladybird Beetles (Coccinellidae) Wiley - Blackwell ed.: 141-274.
Hoogendoorn M., Heimpel G. E. (2003, in press) - PCR-based gut content analysis of insect predators: use in field-caught predators - Proceedings of the 1st International Symposium on Biological Control of
Arthropods, Honolulu, Hawaii.
Hu Y. S., Wang Z. M., Ning C. L., Pi Z. Q., Gao G. Q. (1989) - The functional response of Harmonia
(Leis) axyridis to their prey of Cinara sp - Natural Enemies of Insects 11: 164-168.
44
Bulletin of the Faculty of Agriculture, Gifu University 39: 31-37.
Hukusima S., Kamei M. (1970) - Effects of various species of aphids as food on development, fecundity and longevity of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) - Research Bulletin of the Faculty
of Agriculture, Gifu University 29: 53-66.
Hukusima S., Ohwaki T. (1972) - Further notes on feeding biology of Harmonia axyridis (Coleoptera: Coccinellidae) - Research Bulletin of the Faculty of Agriculture, Gifu University 33: 75-82.
Iablokoff-Khnzorian S. M. (1982) - Les Coccinelles. Coléopètres Coccinellidae - Boubèe Ed. 568 pp.
Ingels B., De Clercq P. (2011) - Effect of size, extraguild prey and habitat complexity on intraguid interactions: a case study with the invasive ladybird Harmonia axyridis and the hoverfly Episyrphus
balteatus - BioControl., 56: 871-882.
Jamal E., Brown G. C. (2001) - Orientation of Hippodamia convergens (Coleoptera: Coccinellidae) larvae to volatile chemicals associated with Myzus nicotianae (Homoptera: Aphididae) - Environmental
Entomology, 30: 1012-1016.
Johnson B. (1965) - Wing polymorphism in aphids. II. Interaction between aphids - Entomologia Exp.
Appl., 8: 49-64.
Joseph S. B., Snyder W. E., Moore A. J. (1999) - Cannibalizing Harmonia axyridis (Coleoptera: Coccinellidae) larvae use endogenous cues to avoid eating relatives - Journal of Evolutionary Biology, 12 (4): 792-797.
Kajita Y., Obrycki J. J., Slogget J. J., Haynes K. F. (2010) - Intraspecific alkaloid variation in ladybird eggs and its effects on con- and heterospecific intraguild predators - Oecologia, 163: 313-322.
Kalaskar A., Evans E. W. (2001) - Larval responses of aphidophagous lady beetles (Coleoptera: Coccinellidae) to weevil larvae versus aphids as prey - Annals of the Entomological Society of America, 94: 76-81.
45
Katsanis A., Babendreier D., Nentwig W., Kenis M. (2013) - Intraguild predation between the invasive ladybird Harmonia Axyridis and non-target European coccinellid species - BioControl 58 (1): 73-83.
Katsoyannos P., Kontodimas D. C., Stathas G. J., Tsartsalis C. T. (1997) - Establishment of
Harmonia axyridis on citrus and some data on its phenology in Greece - Phytoparasitica, 25 (3): 183-
191.
Kawai A. (1976) - Analysis of the aggregation behavior in the larve of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) to prey colony - Researches on Population Ecology, 18 (1): 123-134.
Kawauchi S. (1979) - Effects of temperatures on the aphidophagous Coccinellids - Kurume University
Journal 28: 47-52.
Kidd K. A., Nalepa C. A., Day E. R., Waldvogel M. G. (1995) - Distribution of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in North Carolina and Virginia - Proceedings of the Entomological
Society of Washington, 97: 729-731.
King A. G., Meinwald J. (1996) – Review of the Defensive Chemistry of Coccinellids – Chem. Rev., 96 (3), pp 1105-1122.
Koch R. L. (2003) - The multicolored Asian lady beetle, Harmonia axyridis: A review of its biology, uses in biological control, and non-target impacts - Journal of Insects Science, 3, 32: 1-16 (insectscience.org/3.32)
Koch R. L., Galvan T. L. (2008) - Bad side of a good beetle: the North American experience with
Harmonia axyridis - BioControl 53 (1): 23-35.
Koch R. L., Hutchison W. D., Venette R. C., Heimpel G. E. (2003) - Susceptibility of immature monarch butterfly, Danaus plexippus (Lepidoptera: Nymphalidae: Danainae), to predation by Harmonia
46
Komai T. (1956) - Genetics of ladybeetles - Advances in Genetics, 8: 155-188.
Korschefsky R. (1932) - Coccinellidae - Pages In: Schenkling S., editor. Coleopterorum Catalogus, 439-447. Berlin.
Kovach J. (2004) - Impact of multicolored Asian lady beetles as pest of fruit and people - Ann. Entomol., 50: 159-161.
Kuznetsov V. N. (1997) - Lady beetles of Russian Far East - Gainesville, FL: Memoir Seis Editor, CSE: 248 pp
LaMana M. L., Miller J. C. (1996) - Field observations on Harmonia axyridis Pallas (Coleoptera: Coccinellidae) in Oregon - Biological Control 6: 232-237.
Lou H. H. (1987) - Functional response of Harmonia axyridis to the density of Rhopalosiphum
prunifoiae - Natural Enemies of Insects 9: 84-87.
Lucas E., Coderre D., Vincent C. (1997) - Voracity and feeding preferences of two aphidophagous coccinellids on Aphis citricola and Tetranychus urticae - Entomologia Experimentalis et Applicata 85: 151-159.
Lucas E., Coderre D., Brodeur J. (1998) - Intraguild predation among aphid predators: characterization and influence of extraguild prey density - Ecology, 79: 1084-1092.
Mandour N. S., El-Basha N. A. S., Liu T. X. (2006) - Functional response of the ladybird, Cydonia
vicina nilotica to cowpea aphid, Aphis craccivora in the laboratory - Insect Science, 13 (1): 49-54.
Majerus M. E. N. (1994) - Ladybirds - Harper Collins, London.
Masutti L., Zangheri S. (2001) - Entomologia generale e applicata - Casa Editrice Cedam: 938 pp.
47
matsumarae (Homoptera: Margarodidae) in Japan - Environmental Entomology, 15: 976-983.
Meisner M. H., Harmon J. P., Ives A. R. (2011) - Response of Coccinellid Larvae to Conspecific and Heterospecific Larval Tracks: A Mechanism That Reduces Cannibalism and Intraguil Predation -
Environmental Entomology, 40 (1): 104-110.
Michaud J. P. (2001b) - Numerical response of Olla V-Nigrum (Coleoptera: Coccinellidae) to infestations of Asian Citrus Psyllid, (Hemiptera: Psyllidae) in Florida - Florida Entomologist 84: 608- 612.
Michaud J. P. (2002a) - Biological control of Asian citrus psyllid, Diaphorina citri (Hemiptera: Psyllidae) in Florida: a preliminary report - Entomological News, 113: 216-222.
Michaud J. P. (2003a) - A comparative study of larval cannibalism in three species of ladybird -
Ecological Entomology, 28: 92-101.
Michaud J. P., Grant A. K. (2004) - The adaptive significance of sibling egg cannibalism in the Coccinellidae: comparative evidence from three species - Ann. Entomol. Soc. Am. 97: 710-719.
Mishra G., Srivastava S., Gupta A. K., Singh S. K. (2005) - Reproductive performance of four aphidiphagous ladybirds on cowpea aphid, Aphis craccivora Koch - Journal of Applied Entomology, 129 (4): 217-220.
Miura T., Nishimura S. (1980) - The larval period and predacious activity of an aphidophagous coccinellid, Harmonia axyridis Pallas - Bulletin of the Faculty of Agriculture, Shimane University 14: 144-148.
Moiser S. E., Haynes K. F., Obrycki J. J. (2010) - Behavioral response to larval tracks and the influence of tracks on intraguild scavenging by coccinellid larvae - Journal of Insect Behavior, 23 (1): 45-58.
Nordlund D. A., Morrison R. K. (1990) - Handling time, prey preference, and functional response for
48
Obrycki J. J., Kring T. J. (1998) – Predaceous coccinellidae in biological control – Annual Review of
Entomology, Vol. 43: 295-321.
Obata S. (1986a) - Determination of hibernation site in the ladybird beetle, Harmonia axyridis Pallas (Coleoptera, Coccinellidae) - Kontyu, 54: 218-223.
Obata S. (1986b) - Mechanisms of prey finding in the aphidiphagous ladybird beetle, Harmonia axyridis (Coleoptera: Coccinellidae) - Entomophaga, 31 (3): 303-311.
Obata S. (1997) - The influence of aphids on the behaviour of adults of the ladybird beetle, Harmonia
axyridis (Col.: Coccinellidae) - Entomophaga, 24: 103-106.
Obopile M., Ositile B. (2010) - Life table and population parameters of cowpea aphid, Aphis craccivora Koch (Homoptera: Aphididae) on five cowpea Vigna unguiculata (L. Walp.) varietes - J. Pest. Sci., 89: 9-14.
Ofuya T. I. (1997) - Control of the cowpea aphid, Aphis craccivora Koch (Homoptera: Aphididae) in cowpea, Vigna unguiculata (L.) Walp - Integr. Pest Manage. Rev. 2: 199-207.
Orlandini G., Martellucci R. (1997) - Melone: lotta biologica all'Aphis gossypii - Colture protette, 6: 33-36.
Osawa N. (1989) - Sibling and non-sibling cannibalism by larvae of a lady beetle Harmonia axyridis Pallas (Coleoptera: Coccinellidae) in the field - Researches on Population Ecology, 31: 153-160.
Osawa N. (1991) - Ecological Studies on the Ladybird Beetle Harmonia axyridis Pallas in a Natural Population - 184 pp. Ph. D. Thesis, Kyoto University. (In Japanese.)
Osawa N. (1992) - Sibling cannibalism in the ladybird beetle Harmonia axyridis Pallas: fitness consequences for mother and offspring - Res. Popul. Ecol. 34: 45-55.
49
Osawa N. (1992a) - Effect of pupation site on pupal cannibalism and parasitism in the ladybird beetle
Harmonia axyridis Pallas (Coleoptera, Coccinellidae) - Japanese Journal of Entomology, 60: 131-135.
Osawa N. (1992b) - A life table of the ladybird beetle Harmonia axyridis Pallas (Coleoptera, Coccinellidae) in relation to the aphid abundance - Japanese Journal of Entomology, 60: 575-579.
Osawa N. (1993) - Population field studies of the aphidophagous ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae): life tables and key factor analysis - Researches on Population Ecology, 35: 335-348.
Osawa N. (2000) - Population field studies on the aphidophagous ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae): resource tracking and population characteristics - Popululation Ecology 42: 115-127.
Osawa N., Nishida T. (1992) - Seasonal variation in elytral colour polymorphism in Harmonia axyridis (the ladybird beetle): the role of non-random mating - Heredity 69: 297-307.
Pasqualini E., Pradolesi G., Melandri M., Scannavini M., Franceschelli F., Cavazza F. (2010) – I prodotti efficaci contro la cocciniglia farinosa della vite – L’Informatore agrario, 46: 75-80.
Pasteels J. M., Deroe C., Tursch B., Brackman J. C., Daloze D., Hootele C. (1973) - Distribution et activites des alcaloides defensifs des Coccinellidae - J. Insect Physiol 19: 1771-1784.
Pell J. K., Baverstock J., Roy H. E., Ware R. L., Majerus M. E. N. (2008) - Intraguild predation involving Harmonia axyridis: a review of current knowledge and future persectives - Biocontrol, 53: 147-168.
Pennacchio F. (2014) - Gli insetti e il loro controllo - Liguori Editore, pp 737.
Phoofolo M. W., Obrycki J. J. (1998) - Potential for intraguild predation and competition among predatory Coccinellidae and Chrysopidae - Entomologia Experimentalis et Applicata, 89: 47-55.
50
Pollini A. (2013) - Entomologia applicata - Edagricole, 2° edizione, 1642 pp.
Polis G. A. (1981) - The evolution and dynamics of intraspecific predation - Annu. Rev. Ecol. Syst. 12: 225-251.
Polis G. A., Myers C. A., Holt R. D. (1989) - The ecology and the evolution of intra-guild predation: potential competitor that eat each other - Ann. Rev. Entomol., 20: 297-330.
Pope R. D. (1953) - Coleoptera Coccinellidae & Sphindidae in Handbooks for the Identification of British Insects - Royal Entomological Society of London Vol. V Part 7., 12 pp.
Porta A. (1929) - Fauna Coleopterorum Italica - Vol III Diversicornia 242-277 pp.
Provost C., Lucas E., Coderre D., Chouinard G. (2006) - Prey selection by the lady beetle Harmonia
axyridis: the influence of prey mobility and prey species. - Journal of Insect Behavior, 19 (2): 265-277.
Raak-van den Berg C. L., De Lange H. J., Van Lenteren J. C. (2012) - Intraguild predation behaviour of ladybirds in semi-field experiments explains invasion success of Harmonia axyridis - PloS one, 7 (7): 1-10.
Rakhshani E., Talebi A. A., Kavallieratos N. G., Rezwani A., Manzari S., Tomanovic Ž. (2005) - Parasitoid complex (Hymenoptera, Braconidae, Aphidiinae) of Aphis craccivora Koch (Hemiptera: Aphidoidea) in Iran - Journal of Pest Science, 78 (4): 193-198.
Ratcliffe S. (2002) - National pest alert: Multicolored Asian lady beetle - USDA CSREES Regional
Integrated Pest Management Program and the Pest Management Centers.
Sakai T., Uehara Y., Matsuka M. (1974) - The effect of temperature and other factors on the expression of elytral pattern in ladybeetle, Harmonia axyridis Pallas - Bullettin of the Faculty of Agriculture,
Tamagawa University, 14: 33-39.
51
Insectology, 60 (1): 89-91.
Sarma K. K., Dutta S. K., Borah B. K. (1996) - Interaction between Aphis craccivora Koch and its predators, Coccinella transversalis Fabricius (Coleptera: Coccinellidae) and Ischiodon scutellaris (Fabricius) (Diptera: Syrphidae) - Journal of biological control, 10 (1/2): 125-128.
Sasaji H. (1971) - Fauna Japonica, Coccinellidae (Insecta: Coleoptera) - Tokyo: Academic Press Japan.
Sasaji H. (1977) - Larval characters of Asian Species of the Genus Harmonia Mulsant - Memoir of the
Faculty of Education Fukui University Series II Natural Science 27: 1-17.
Sato S., Yasuda H., Evans E. W. (2005) - Dropping behaviour of larvae of aphidophagous ladybirds and its effects on incidence of intraguild predation: interactions between the intraguild prey, Adalia
bipunctata and Coccinella septempunctata and the intraguild predation, Harmonia axyridis Pallas - Ecol. Entomol., 30: 220-224.
Savoiskaya G. I., Klausnitzer B. (1973) - Morphology and taxonomy of the larvae with keys for their identification - In: Hodek I., editor. Biology of Coccinellidae, 36-55. The Hauge, Holland: Dr. W. Junk
N. V., Publishers.
Seo M. J., Kim G. H., Youn Y. N. (2008) - Differences in biological and behavioural characteristics of
Harmonia axyridis (Coleoptera: Coccinelidae) according to colour patterns of elytra - J. Appl. Entomol.,
132: 239-247.
Shu C. R., Yu C. Y. (1985) - An investigation on the natural enemies of Hyphantria cunea - Natural
Enemies of Insects 7: 91-94.
Snyder W. E., Joseph S. B., Preziosi R. F., Moore A. J. (2000) - Nutritional benefits of cannibalism for the lady beetle Harmonia axyridis (Coleoptera: Coccinellidae) when prey quality is poor -
Environmental Entomology, 29: 1173-1179.
52
invasion by introduced ladybird beetles - Oecologia, 140 (4): 559-565.
Soares A. O., Coderre D., Schanderl H. (2001) - Fitness of two phenotypes of Harmonia axyridis (Coleoptera: Coccinellidae) - European Journal of Entomology 98: 287-293.
Soares A. O., Coderre D., Schanderl H. (2004) - Dietary self-selection behaviour by the adults of the aphidophagous ladybeetle Harmonia axyridis (Coleoptera: Coccinellidae) - Journal of Animal Ecology, 73 (3): 478-486.
Stuart R. J., Michaud J. P., Olsen L., McCoy C. W. (2002) - Lady beetles as potential predators of the root weevil Diaprepes abbreviatus (Coleoptera: Curculionidae) in Florida citrus - Florida Entomologist 85: 409-416.
Takallozadeh H. M. (2003) - Biology, population dynamics and seasonal parasitism of Aphis craccivora and study on interaction among plant host, aphid and major parassitoid in Kerman - PhD Thesis, College
of Agriculture, Tarbiat Modarres University: 177pp.
Tedders W. L., Schaefer P. W. (1994) - Release and establishment of Harmonia axyridis (Coleoptera: Coccinellidae) in the south-eastern United States - Entomological News, 105: 228-243.
Ueno H., Sato Y., Tsuchida K. (1998) - Colour-associated mating success in a polymorphic ladybird beetle, Harmonia axyridis - Funct. Ecol. 12: 757-761.
Varner M., Cacciatore S., Giallo D., Titone G., Mattedi L., Lucchi A. (2015) – Cocciniglia Farinosa – Millevigne, 2: 12-15
Viggiani G. (1994) - Lotta biologica ed integrata nella difesa fitosanitaria - Vol. 1, ed. Liguori, 250 pp.
Völkl W., Vohland K. (1996) - Wax covers in larvae of two Scymnus species: do they enhance coccinellid larval survival? - Oecologia, 107 (4): 498-503.
53
of cannibalism in Harmonia axyridis - Evolutionary Ecology Research 1: 375-388.
Wang S., Michaud J. P., Tan X. L., Zhang F. (2014) - Comparative suitability of aphids, thrips and mites as prey for the flower bug Orius sauteri (Hemiptera: Anthocoridae) - European Journal of
Entomology, 111 (2): 221.
Wang L. Y. (1986) - Mass rearing and utilization in biological control of the lady beetle Leis axyridis (Pallas) - Acta entomologica Sinica, 29: 104.
Ware R. L., Majerus M. E. N. (2008) - Intraguild predation of immature stages of British and Japanese coccinellids by the invasive ladybird Harmonia axyridis - BioControl, 53: 169-188.
With K. A., Pavuk D. M., Worchuck J. L., Oates R. K., Fisher J. L. (2002) - Threshold effects of landscape structure on biological control in agroecosystems - Ecological Applications 12: 52-65.
Yasuda H., Kikuchi T., Kindlmann P., Sato S. (2001) - Relationships between attack and escape rates,