TEAM NEWSLETTER

No. 6 December 2008

TO OUR READERS

The first meeting of TEAM in Majorca was very combination exerted an additive effect on male sexual successful. Over 140 scientists from various countries signalling, female visitation, and finally male mating participated and numerous papers of high quality were success. Pereira and co-authors suggest that males presented. This success is due to our sponsors, the that have access to an optimal diet would have the TEAM Steering Committee, the Scientific Committee, energetic capacity to exploit the physiological effects the symposia and forums organizers, the efforts of the of additional methoprene. In longevity experiments, local Organizing Committee headed by Miguel methoprene application alone did not affect male Miranda, and above all the active participation of survival. Protein availability seems to be an important TEAM members and friends. factor that increases male longevity. These experiments, regardless of the protein or methoprene I am happy to inform you that the special issue of the effects, reveal detrimental effects of male–male and Journal of Applied Entomology with papers from the male–female interactions in male longevity, with male– meeting (see contents below) was released in male interactions being substantially more costly for December 2008. The release took place well before it males. The importance of these findings for a better was initially planned due to the intensive work of David understanding of the sexual behaviour and associated Nestel, Miguel Miranda and me, as well as Christine costs of male Caribbean fruit as well as of their Denys and Stefan Vidal from the Journal of Applied application for controlling A. suspensa by applying the Entomology. I thank all of you who contributed to the Sterile Release Technique is discussed. reviewing process and of course the authors of the papers. Also, in this newsletter Kirsten Köppler contributes a note on the occurrence of non-diapausing individuals During the meeting in Mallorca the Steering in the cherry fruit in Germany, which eclose during Committee decided that TEAM would make awards to the same vegetation period shortly after pupation. persons with notable contribution to the field of fruit Though this phenomenon has been observed by flies. The first recipient of this award is Jorge others as well, its significance has not yet been fully Hendrichs, who has had an outstanding contribution to elucidated. the field and has also greatly encouraged and supported the establishment of our group. On behalf of the TEAM steering committee I would like to wish to all our members and friends a very happy In Majorca, the TEAM Steering Committee made and prosperous 2009, with lots of personal and several important decisions regarding the future of the professional achievements. group. It was decided to hold our next scientific meeting in 2011 in either Morocco, or Kenya or Greece. We also explored the idea of establishing Nikos Papadopoulos working groups on specific fields of fruit fly research Chairman of the Steering Committee and applications. These groups would review the University of Thessaly, School of Agriculture status of each field and provide the ground for 38442 N. Ionia (Volos) Magnisias, Greece developing collaborative projects. I hope that during [email protected] the next few months we will be able to announce the first working groups. We also discussed issues related to the current newsletter and how we can improve it further and make it more useful to our group and fruit fly workers all over the world.

The current newsletter includes a paper by Rui Pereira, John Sivinski and Peter Teal, on the effects of adult food and juvenile hormone application on the sexual performance and other fitness components of the Caribbean Fruit Fly, Anastrepha suspensa. When offered alone, protein rich diet or methoprene increased male sexual performance. However, their

TEAM Newsletter No. 6 (December 2008) 1 JUVENILE HORMONE ANALOG AND DIETARY PROTEIN IMPROVE MALE CARIBBEAN FRUIT FLY SEXUAL BEHAVIOR

The Caribbean fruit fly, Anastrepha suspensa (Loew) mating success. Experiments conducted in laboratory (Figure 1), like many polyphagous tephritids (Diptera: and field cages (Figure 2), show and advantage of the ), adopts a lek polygyny mating system. addition of both factors. Of the total matings, 55% (in The success of sterile insect technique (SIT) directed laboratory) and 59% (in field cages) were performed towards this pest requires the release of males that by M+P+ males, which were significantly higher can compete with wild males and attract wild females compared with males from all the other treatments (Knipling 1955, Hendrichs et al. 2002). Mass-reared (Figure 3). males must form leks, engage in male-male agonistic interactions, court females, and mate to transfer sterile sperm (Shelly and Whittier 1997). Effects of application of a juvenile hormone analog, methoprene, and dietary protein on male Caribbean fruit fly sexual success were evaluated in the four possible combinations of the two factors (methoprene and protein). The manipulation of juvenile hormones titers can contribute to an increase in male pheromone production and under certain conditions, sexual competitiveness. At the same time, it leads to earlier sexual maturation which can reduce the cost of the SIT, particularly in fruit fly species with a long precopulatory periods like A. suspensa, due to space savings at fly handling facilities. However, accelerated maturity may have nutritional consequences since there is less time for flies to acquire reserves. Thus the addition of protein rich adult diet may be particularly important when hormones titers are manipulated. Figure 2. Field cage used in the experiment. Laboratory, field cage, and greenhouse experiments compared male sexual performance and other + + reproductive parameters on a lifetime and daily basis. The earliest mating occurred on day 4 for M P males, on day 5 for M+P- males and on day 6 for males that did not receive methoprene. In addition, higher Major components of sexual success studied include: + + (1) male sexual performance on a daily and lifetime percentages of M P males mated consecutively over 3 or more days than males in other treatments. On a basis; (2) lek tenure, as influenced by male-male + + interactions; (3) male sexual attractiveness to females; daily basis M P males were not only more likely to and (4) male survival. mate, but 10% (14 males) were able to mate 3 times on the same day. Only 1 M+P-, 1 M-P+, and none of the M-P- males were able to mate so often.

The combination of methoprene application with a supply of protein-enriched food resulted in greater male sexual success than when either was provided separately. It seems likely that with an optimal diet would have the energetic capacity to exploit the physiological effects of additional methoprene.

80

70

60 a A

50 Field cage Laboratory 40

30 Matings (%) B b 20 b B

Figure 1. Male Anastrepha suspensa 10 c C 0 M+P+ M+P- M-P+ M-P- Treatment 1. Male sexual performance In this study we evaluate the effects of a juvenile Figure 3. Percentage of matings per treatment of male hormone analog (methoprene) application, of protein Caribbean fruit flies in laboratory and field cage experiments, when treated or not with methoprene (M+/M-) and fed or not in the adult diet and their interactions on the sexual + with protein (P /P) (mean plus standard deviation). Different performance of male A. suspensa, specifically male lowercase and capital letters represent significant differences competitiveness in agonistic interactions with among treatments, for field cage and laboratory tests competing mates, attractiveness to females, and respectively (Tukey’s test, Į=0.05).

TEAM Newsletter No. 6 (December 2008) 2 To measure the impact of both intersexual and 2. Lek tenure intrasexual activities on male survival in A. suspensa Experiments in field cages were conducted with males and the interacting effects of reproductive behavior, treated with different methoprene and protein diet and methoprene levels, three experimental groups regimens. Initiation and participation in leks, male of individually-caged flies were monitored for a period pheromone-calling, male position in the lek, male-male of 35 days: males caged alone with no interactions; and male-female interactions, and sexual success those with daily male-male interactions; and those with were observed. Significant effects of methoprene a daily male-female interaction. In the last two cases application, protein supply, and the interaction of the the “interacting” fly was presented to the “focal” fly for methoprene application and protein supply were found the two hours that coincided with peak time for sexual in lek initiation, lek participation, number of males activity (Landolt and Sivinski 1992). The intensity of calling and matings performed (Figure 4). the sexual interactions was manipulated within each experimental group by the addition of a methoprene application and/or dietary protein to various subgroups

90 of males. 80 Lek initiation Lek participation 70 a a For all combinations of experimental groups and a Males calling 60 a Mating methoprene application there were no significant 50 differences in longevity when protein was provided. 40

Males (%) Males b bbb The same was true when protein was withheld. 30 bb b b + 20 However, for the P treatment, survival was cc - 10 c significantly higher than P in all interaction groups. No c 0 effect of methoprene application was observed in any M+P+ M+P- M-P+ M-P- of the experimental groups. Treatments

Figure 4. Lek parameters and sexual success of male Longevity in the three interaction groups showed that Caribbean fruit flies presented as percentages of males within each treatment, males maintained alone (mean plus standard deviation), when treated or not with + - + - survived significantly longer than those interacting with methoprene (M /M ) and fed or not with protein (P /P ). Bars males or females. with the same letter for each parameter were not significantly different (Tukey’s test, Į=0.05). Observations based on 41 leks (lek initiation), where 212 males participatd, 178 called, Male-male interaction effects on survival were more and 73 matings occurred. costly than male-female interactions, with the exception of the M+P- treatment where mortality in the context of male-male interactions was not significantly From the males that initiated a lek, 78% went on to different from male-female interactions. copulate, which represents a high probability of success. Lek initiation in various tephritids can be 5. Discussion dependent on the nutritional status of the male (Yuval Male sexual performance, in terms of lek tenure, et al. 1998) or perhaps by the male’s hormonal state attractiveness to females, winning in agonistic (Teal et al. 2000). In the case of the Caribbean fruit fly, encounters and number of copulations obtained, was either methoprene or protein resulted in higher rates of significantly improved through both applications of the lek initiation and ultimately greater sexual success juvenile hormone analog, methoprene and the addition (Pereira et al. in press). of protein to adult diet. Used together they had an additive effect. Protein by itself had a positive impact 3. Male attractiveness to females on male longevity and on balanced nutritional status. Significant effects of methoprene application, protein supply and the interaction of the methoprene The advantages of good diet are well understood in application and protein supply on male calling many insects and the costs and benefits of foraging for duration, female visitation were found both in food have been examined (Yuval et al. 2002). laboratory and greenhouse conditions. Male calling Differences in male sexual success can be in part due frequency, regardless of treatment, was correlated to differences in their diets. In addition to a well- with female visitation. nourished male being capable of greater activity, females may choose males on the basis of their Laboratory and greenhouse experiments, using perceived or inferred nutritional status. By mating with individual and aggregated males respectively, males that are able to maintain their position in the lek, obtained similar results. M+P+ males spent significantly females are mating with males that have been more time calling than males from other treatments effective in acquiring food, and consequently may and attracted more females, and M-P- males called provide genes that affect offspring viability and male significantly less than males from other treatments and fitness (Field et al. 2002). Another advantage of a attracted significantly fewer females. Overall, the protein diet is a clear positive impact on longevity immediate effects of both methoprene and protein which is correlated with a potential increase in were to increase male signaling and enhance sexual offspring production. success, and the interaction of these two factors was additive. Greater pheromone emission could be due to Less well understood is the role of methoprene, i.e., either more males calling or because they produce juvenile hormone, titer in sexual behavior and more pheromone per unit of calling time. selection. For example, given the great sexual advantages available to males that have higher 4. Male survival juvenile hormone titers, the question arises as to why

TEAM Newsletter No. 6 (December 2008) 3 these levels are not generally higher than they addressed. Alternative sources of less expensive apparently are (i.e., as high as they are after protein need to be identified and suitable means of treatment), or perhaps more heuristically, what limits their incorporation into agar diets developed. male hormone production to levels below those that Additionally, inoculation with bacteria to restore the gut would yield more copulations? In general, there may fauna after irradiation could further enhance optimal be limits on the availability of juvenile hormone diets (Lauzon et al. 2000). The effect of methoprene components, or there may be physiological costs to when provided in a diet rather than being topically juvenile hormone production (in the present study applied needs to be determined, since the additional juvenile hormone analog is “free” to males), physiological effect on A. suspensa could be different. or there may be expensive physiological Should this be the case, perhaps an alternative consequences to higher juvenile hormone titers. technique of topical application such as spraying pupae, would be efficacious. Methoprene also causes accelerated maturation. Not only are such males more sexually successful on a References daily basis, they also begin their reproductive lives Dodson, G. 1982. Mating and territoriality in wild Anastrepha earlier. All other things being equal, males that mature suspensa (Diptera: Tephritidae) in field cages. J. Georgia Ent. Soc. 17: 189-200. sooner have more sexual opportunities. So why then Field, S. A., R. Kaspi, and B. Yuval. 2002. Why do calling do they not mature as quickly as they could (as quickly medflies (Diptera: Tephritidae) cluster? Assessing the as in these experiments) in nature? Tephritids often empirical evidence for models of medfly lek evolution. Fla. have extended, pre-reproductive periods in both males Entomol. 85: 63-72. and females, although these tend to be longer in Hendrichs, J., Robinson, A. S., Cayol, J. P., and W. Enkerlin. females. Male maturation in A. suspensa (Dodson 2002. Medfly areawide sterile insect technique programmes 1982, Teal et al. 2000) and Anastrepha ludens (Loew) for prevention, suppression or eradication: The importance (Robacker et al. 1991) occurs in 5 to 7 days. This is of mating behavior studies. Fla. Entomol. 85: 1-13. Knipling, E. F. 1955. Possibilities of insect control or due to the anauthogeneous nature of tephritids, i.e., eradication through the use of sexual sterile males. J. adults emerge with unmatured gonads (Yuval et al. Econ. Entomol. 48: 459-462. 2002). It is known that the timing of maturation is Landolt, P. J., and J. Sivinski. 1992. Effects of time of day, flexible. Environmental factors, such as presence of adult food, and host fruit on incidence of calling by male the male pheromone, can be used as cues for timing Caribbean fruit flies (Diptera: Tephritidae). Environ. facultative A. suspensa female maturation (Pereira et Entomol. 21: 382-387. al. 2006). However, changes in pre-reproductive Lauzon, C. R., R. E. Sjogren, and R. J. Prokopy. 2000. periods under mass-rearing suggest the parameters of Enzymatic capabilities of bacteria associated with apple maggot flies: A postulated role in attraction. J. Chem. Ecol. maturation are set by nutritional requirements, i.e., that 26: 953-957. flies given a predictable laboratory diet are selected to Liedo, P., E. De Leon, M. I. Barrios, J. F. Valle-Mora, and G. mature sooner. Pereira et al. (2006) saw this clearly Ibarra. 2002. Effect of age on the mating propensity of the when they found that the time under domestication Mediterranean fruit fly (Diptera: Tephritidae). Fla. Entomol. affected the female pre-maturation period (mass- 85: 94-101. reared flies mature earlier than recently domesticated Pereira, R., P. E. A. Teal, J. Sivinski, and B. D. Dueben. flies and wild flies took longer still). The same 2006. Influence of male presence on sexual maturation in phenomenon occurs in male Ceratitis capitata (Wied.) female Caribbean fruit fly, Anastrepha suspensa (Diptera: Tephritidae). J. Insect Behav. 19: 31-43. (Liedo et al. 2002). Pereira, R., J. Sivinski, and P. E. A. Teal. Influence of methoprene and dietary protein on male Anastrepha Regardless of the unanswered evolutionary questions suspensa (Diptera: Tephritidae) mating aggregations. J. posed by these experiments, there are clear pragmatic Insect Physiol. (in press). applications for the SIT. This technique is a principal Robacker, D. C., R. L. Mangan, D. S. Moreno, and A. M. non-chemical component of pest tephritid Tarshis-Moreno. 1991. Mating behavior and male mating management (Hendrichs et al. 2002), and success in wild Anastrepha ludens (Diptera: Tephritidae) on improvements in production costs and insect quality a field caged host tree. J. Insect Behav. 4: 471-487. Robinson, A. S., J. P. Cayol, and J. Hendrichs. 2002. Recent influence the efficacy of the technique and even the findings on medfly sexual behavior: implications for SIT. practicality of its use under certain circumstances Fla. Entomol. 85: 171-181. (Robinson et al. 2002). Economic support for these Shelly, T. E., and T. S. Whittier. 1997. Lek behaviour of area-wide programs is based on ecological, insects, pp. 273-293. In B. Crespi and J. C. Choe (eds.), commercial, and regulatory aspects (Munford 2004). The evolution of mating systems in insects and Arachnids. However, the final objective of the technique Cambridge Press, Cambridge, UK. (eradication, prevention or suppression) should Teal, P. E. A., Y. Gomez-Simuta, and A. T. Proveaux. 2000. influence the strategies adopted. Mating experience and juvenile hormone enhance sexual signaling and mating in male Caribbean fruit flies. Proc. Nat. Acad. Sci. 97: 3708-3712. The delay between adult emergence and sexual Yuval, B., R. Kaspi, S. Sholmit, and M. S. Warburg. 1998. maturity poses a significant problem for SIT programs Nutritional reserves regulates male participation in because males must be held for a long period of time Mediterranean fruit fly leks. Ecol. Entomol. 23: 211-215. prior to release, or have to be released before Yuval, B., R. Kaspi, S. A. Field, S. Blay, and P. Taylor. 2002. becoming sexually mature, resulting in fewer surviving Effects of post-ternal nutrition on reproductive success of to maturity and copulation. This means that early male Mediterranean fruit flies (Diptera: Tephritidae). Fla. sexual maturation results in more flies and in lower Entomol. 85: 165-170. costs, due to space savings at fly handling facilities. Rui Pereira1,2, John Sivinski1 & Peter E. A. Teal1 Incorporation of these experimental results into SIT programs requires that certain technical problems be

TEAM Newsletter No. 6 (December 2008) 4 1Center for Medical, Agricultural and Veterinary e-mail addresses: Entomology, USDA-ARS, Gainesville, Florida, USA. [email protected] [email protected] 2Present address: Insect Pest Control Section, Joint [email protected] FAO/IAEA Programme of Nuclear Techniques in Food and Agriculture, Wagramer Strasse 5, P.O. Box 100, A-1400 Vienna, Austria.

PHOTOS FROM THE FIRST MEETING OF TEAM

For more photos see photo gallery of the event in www.uib.es/depart/dba/Zoology/esp/miranda/TEAM/index.html

JOURNAL OF APPLIED ENTOMOLOGY SPECIAL ISSUE VOLUME 132 ISSUE 9-10 (DECEMBER 2008) - CONTENTS -

EDITORIAL NOTE Bactrocera oleae: a single large population in Northern Current advances in the study of the ecology of fruit flies from Mediterranean basin (p 706-713) , Africa and the Middle East (p 681-681) M. D. Segura, C. Callejas, M. D. Ochando D. Nestel, N. T. Papadopoulos, M. A. Miranda Chueca Seasonality in the occurrence of the Mediterranean fruit fly ORIGINAL ARTICLES [Ceratitis capitata (Wied.)] in the north-east of Spain (p 714- Relationships between the olive fly and bacteria (p 682-689) 721) P. Sacchetti, A. Granchietti, S. Landini, C. Viti, L. Giovannetti, L. A. Escudero-Colomar, M. Vilajeliu, L. Batllori A. Belcari Effect of elevation on spatio-temporal patterns of olive fly Bacteria–diet interactions affect longevity in the medfly – (Bactrocera oleae) populations in northern Greece (p 722- Ceratitis capitata (p 690-694) 733) M. Ben-Yosef, A. Behar, E. Jurkevitch, B. Yuval I. Kounatidis, N. T. Papadopoulos, P. Mavragani-Tsipidou, Y. Cohen, K. Tertivanidis, M. Nomikou, D. Nestel Life-history evolution of an invasive tephritid (p 695-705) A. D. Diamantidis, J. R. Carey, N. T. Papadopoulos Performance and acceptance evaluation of a knowledge- based SDSS for medfly area-wide control (p 734-745)

TEAM Newsletter No. 6 (December 2008) 5 A. Cohen, Y. Cohen, D. Broday, D. Timar S. A. Mohamed, S. Ekesi, R. Hanna

Evaluation of a chemosterilization strategy against Ceratitis Parasitism of Bactrocera oleae (Diptera; Tephritidae) by capitata (Diptera: Tephritidae) in Mallorca island (Spain) (p Psyttalia concolor (Hymenoptera; Braconidae) in the Balearic 746-752) Islands (Spain) (p 798-805) A. Alemany, A. González, A. Juan, C. Tur M. A. Miranda, M. Miquel, J. Terrassa, N. Melis, M. Monerris

Female-biased mass trapping vs. bait application techniques Mating competitiveness of the adult oriental fruit fly reared as against the Mediterranean fruit fly, Ceratitis capitata (Dipt., larvae in liquid vs. those raised on standard wheat-based Tephritidae) (p 753-761) diets (p 806-811) M. M. Leza, A. Juan, M. Capllonch, A. Alemany C. L. Chang, D. O. McInnis

Organophosphosphate resistance-related mutations in the Influence of methoprene and protein on survival, maturation acetylcholinesterase gene of Tephritidae (p 762-771) and sexual performance of male Ceratitis capitata (Diptera: E. G. Kakani, K. D. Mathiopoulos Tephritidae) (p 812-819) M. J. Faria, R. Pereira, T. Dellinger, P. E. A. Teal Evaluation of trimedlure dispensers by a method based on thermal desorption coupled with gas chromatography–mass Insect transgenesis applied to tephritid pest control (p 820- spectrometry (p 772-777) 831) C. Alfaro, J. Domínguez, V. Navarro-Llopis, J. Primo F. Scolari, M. F. Schetelig, P. Gabrieli, P. Siciliano, L. M. Gomulski, N. Karam, E. A. Wimmer, A. R. Malacrida, G. Isolation of entomopathogenic fungi from the soil and their Gasperi pathogenicity to two fruit fly species (Diptera: Tephritidae) (p 778-788) Effect of temperature on development and survival of P. Sookar, S. Bhagwant, E. Awuor Ouna immature stages of Bactrocera invadens (Diptera: Tephritidae) (p 832-839) Evaluation of the impact of Diachasmimorpha longicaudata I. Rwomushana, S. Ekesi, C. K. P. O. Ogol, I. Gordon on Bactrocera invadens and five African fruit fly species (p 789-797)

OCCURRENCE OF NON-DIAPAUSING PUPAE IN CERASI L.

Rhagoletis species as temperate insects have a true winter diapause and physiologically it can only be broken, when they have received a certain dose of cold temperatures (Lees, 1955). The lower temperature developmental threshold for e.g. R. pomonella was determined with 6.4 and 8.7°C (Neilson, 1964; Reid and Laing, 1976; Reissig et al., 1979), for R. indifferens with 9°C (AliNiazee, 1975; Stark and Ali Niazee, 1982) and for R. cerasi with 5°C (Boller, 1964). But the diapause regulation of temperate Rhagoletis species is not homogenous and fully understood. Teixeira and Polavarapu (2003; 2005) showed for R. mendax the occurrence of a phenologically distinct late population, which may have been evolved according to different ripening periods of the host fruit. Different populations of R. cerasi differ in post-diapause regulation according to there origin (Boller and Bush, 1974; Haisch and Forster 1975; Baker and Miller, 1978). Additionally, a certain percentage of a R. cerasi population stays in the An adult Rhagoletis cerasi on a cherry. diapause until the vegetation period after next as so- called overlayers (Haisch, 1975).

TEAM Newsletter No. 6 (December 2008) 6 In 2008 we observed this phenomenon in three populations of Germany (Dossenheim, Forchheim, Emmerthal). In Dossenheim, we found 0.55 % (N = 196, with 80 male and 116 female flies) out of 35,481 collected pupae, which did not go through a winter diapause. These flies emerged after approximately 22 to 54 days after pupation. The Forchheim population (N = 150) showed an emergence rate of 2.67 % (N = 4 with 3 male and 1 female flies after 31 to 39 days). The Emmerthal population revealed only the emergence of one female out of 1,073 collected pupae after 40 days (0.09 %).

These findings could explain the occurrence of trapped cherry fruit flies (Rebell® traps, Andermatt Biocontrol, Grossdietwil, Switzerland) at the cherry orchard in Dossenheim until the middle of August in A cherry infested by a larva of Rhagoletis cerasi. 2002, the end of August in 2008, as well as until the middle of September in 2004 and 2005, respectively. This early emergence of adults can only be interpreted In contrast to other Rhagoletis species, R. pomonella as a disadvantageous event for these individuals overwinters in a facultative pupal diapause (Prokopy, because no hosts are available any more and possible 1968; Dean and Chapman, 1973; Boller and Prokopy, food limitation as well as the initiation of cold 1976). Different environmental conditions during the temperatures avoids the survival of these flies. larval and pupal stage of host races can cause a pronounced diapause and rapidly initiate adult Nevertheless, further data are needed and research development (Prokopy, 1968). Filchak et al. (2001) must be done to understand the diapause regulation of showed for this species, that photoperiod affects the R. cerasi. genetic regulation of diapause and following, variability in adult eclosion. References On R. cerasi is reported to reveal non-diapausing AliNiazee M. T. (1975): Susceptibility of diapausing pupae of individuals, which eclose in the same vegetation the western cherry fruit fly (Diptera, Tephritidae) and a period shortly after pupation (Boller, Papadopoulos, parasite (Hymenoptera, Diapriidae) to subfreezing pers. comm.). During our own studies, we collected temperatures. Env. Ent. 4: 1011-1013. several thousands of pupae in 2002 to 2007 and we Baker C. R. B. and Miller G. W. (1978): The effect of only found single flies in some of these years, which temperature on the post-diapause development of four emerged without a diapause. Until now, no geographic populations of the European cherry fruit fly quantification of these non-diapausing flies was (Rhagoletis cerasi). Ent. Exp. Appl. 23: 1-13. Boller E. F. (1964): Auftreten der Kirschfruchtfliege possible for several populations. This phenomenon (Rhagoletis cerasi) und Prognose mittels seems not to be a regular event while Boller (pers. Bodentemperaturen im Jahre 1963. Schw. Z. Obst- comm.) could not find non-diapausing pupae during Weinbau 73: 53-58. his work. Boller E. F. and Bush G. L. (1974): Evidence for genetic variations in populations of the European cherry fruit fly Rhagoletis cerasi L. (Diptera: Tephritidae) based on physiological parameters and hybridization experiments. Ent. Exp. Appl. 17: 279-293. Boller E. F. and Prokopy R. J. (1976): Bionomics and management of Rhagoletis. Ann. Rev. Ent. 21: 223-246. Dean R. W. and Chapman P. J. (1973): Bionomics of the apple maggot in eastern New York. Search Agric. Ent. (Geneva, NY) 3: 1-62. Filchak K. E., Roethele J. B. and Feder J. L. (2001): Effects of photoperiod and light intensity on the genetics of diapause in the apple maggot (Diptera: Tephritidae). Ann. Ent. Soc. Am. 94: 902-908. Haisch A. (1975): Zur Puppendiapause der Kirschfruchtfliege, Rhagoletis cerasi L., I Beeinflussung der diapausierenden Puppen durch unterschiedliche Temperaturen und verschieden lange Kälteexposition. Z. angew. Ent. 79: 1-11. Haisch A. and Forster S. (1975): Zur herkunftsspezifischen Diapause-Ausprägung der Europäischen Kirschfruchtfliege, Rhagoletis cerasi (Diptera: Tephritidae). Ent. Germ. 2: 137- 148. Lees A. D. (1955): The physiology of diapause in . Cambridge University Press, Cambridge, 122 pp. Leski R. (1969): Population studies on the cherry fruit fly, Rhagoletis cerasi. pp. 1-7. In: Insect Ecology and the sterile male technique. I.A.E.A., Vienna, 102 pp. Pupae of Rhagoletis cerasi. Neilson W. T. A. (1964): Some effects of relative humidity on development of pupae of the apple maggot, Rhagoletis pomonella (Walsh). Can. Ent. 96: 810-811.

TEAM Newsletter No. 6 (December 2008) 7 Prokopy R. J. (1968): The influence of photoperiod, Teixeira L. A. and Polavarapu S. (2005): Diapause temperature and food on the initiation of diapause in the development in the Blueberry maggot Rhagoletis mendax apple maggot. Can. Ent. 100: 318-329. (Diptera: Tephritidae). Env. Ent. 34: 47-53. Reid J. A. K. and Laing J. E. (1976): Developmental threshold and degree-days to adult emergence for overwintering pupae of the apple maggot, Rhagoletis pomonella Walsh, Kirsten Köppler collected in Ontario. Proc. Ent. Soc. Ontario 107: 19-22. Julius-Kühn-Institute (JKI) Reissig W. H., Barnard J., Weires R. W., Glass E. H. and Federal Research Centre for Cultivated Plants Dean R. W. (1979): Prediction of apple maggot emergence Institute for Plant Protection in Fruit Crops and Viticulture, from thermal unit accumulation. Env. Ent. 8: 51-54. Schwabenheimer Str. 101, 69221 Dossenheim, Germany Stark S. B. and AliNiazee M. T. (1982): Model of post- [email protected] diapause development in the western cherry fruit fly. Env. Ent. 11: 471-474. Teixeira L. A. and Polavarapu S. (2003): Evolution of phenologically distinct populations of Rhagoletis mendax (Diptera: Tephritidae) in highbush blueberry fields. Ann. Ent. Soc. Am. 96: 818-827.

FORTHCOMING MEETINGS

IOBC/WPRS Working Group “Induced Resistence in Plants Against Insects and Diseases”. 12-16 May TEAM 2010, Granada, Spain. STEERING COMMITTEE Ecological Impact of Genetically Modified Organisms th Nikos Papadopoulos ([email protected]), Greece (EIGMO), 4 Meeting of the IOBC/WPRS Working Group “GMOs in Integrated Plant Production”. 14-16 Abdeljalil Bakri ([email protected]), Morocco May 2010, Rostock, Germany. Yoav Gazit ([email protected]), Israel 61st International Symposium on Crop Protection. 19 May 2010, Gent, Belgium. Brian Barnes ([email protected]), South Africa Mariangela Bonizzoni ([email protected]), Italy Eight International Symposium on Fruit Flies of Economic Importance (ISFFEI). 12-17 September Massimo Cristofaro ([email protected]), Italy 2010, Valencia, Spain. Slawomir Lux ([email protected]), Poland International Congress on Biological Invasions. 2-6 David Nestel ([email protected]), Israel November 2010, Fuzhou, China. Rui Pereira ([email protected]), Portugal Serge Quilici ([email protected]), France THIS NEWSLETTER NEWSLETTER EDITOR Nikos Kouloussis ([email protected]) This newsletter is intended for the publication of subjects of interest to the members of TEAM. All content is solicited from the membership and should be addressed to: the Chairman of the steering committee, Nikos Papadopoulos, ([email protected]) or to the editor of the newsletter: Nikos Kouloussis Aristotle University of Thessaloniki School of Agriculture 54124 Thessaloniki, Greece [email protected]

TEAM Newsletter No. 6 (December 2008) 8