Seasonal Biology of Melanagromyza Albocilia (Diptera: Agromyzidae) and Seasonal Patterns of Field Bindweed Infestation, Under Field Conditions in Slovakia

Seasonal biology of Melanagromyza albocilia (Diptera: Agromyzidae) and seasonal patterns of field bindweed infestation, under field conditions in Slovakia

Peter Téth, Massimo Cristofaro & Ludovit Cagafi

Toth, P., Cristofaro, M. & Cagafi, L. 2005: Seasonal biology ofMelanagromyza albocilia (Diptera: Agromyzidae) and seasonal patterns offield bindweed infestation, under field conditions in Slovakia. — Entomol. Fennica 16: 254—262.

Melanagromyza albocilia Hendel, a stem borer of field bindweed (Convolvulus arvensis) was observed at three sites in Slovakia during 1999 and 2000. The insect overwintered as pupa. The first generation adults emerged in May and those ofsecond during June—August. Percentage ofinfested field bindweed plants varied from 46.7% to 99.2% and number ofinfested stems from 4.1% to 37.2%. In-

fested stems contained an average of 0.84 i 0.40 to 1.34 i 0.50 specimens (larvae, parasitized larvae, pupae). The population ofM. albocilia was the highest in 1999, with mean densities of 1.57 i 1.01 to 3.21 i 2.13 specimens/plant. All parasitoids emerged from larvae were identified as Braconpicticornis Wesmael. P. Totk, Slovak Agricultural University, Department of Plant Protection, A. Hlinku 2, 94976 Nitra, Slovak Republic; E—mail.‘ [email protected] M. Cristofaro, ENEA C. R. Casaccia, BIOTEC, via Anguillarese 301, 00060 Rome, Italy; E—mail.‘ massimo. cristofaro@casaccia. enea. it E. Cagan, Slovak Agricultural University, Department ofPlant Protection, A. Hlinku 2, 94976 Nitra, Slovak Republic; E—mail.‘ [email protected]

Received 21 January 2004, accepted 15 August 2005

1. Introduction 1976). In Mediterranean Europe, 139 species of phytophagous arthropods were collected from Field bindweed (Convolvulus arvensis Linnaeus) C. arvensis and its close relatives (Rosenthal & is considered one ofthe most important weeds in Buckingham 1982). For more details, see Toth & the world. Many scientists have surveyed Cagan (2005). phytophagous insects associated with Convolvu— During 1996, field surveys of phytophagous lus spp., usually to find and evaluate potential insects associated with Convolvulus spp. started biocontrol agents. Mohyuddin (1969) listed 125 at various localities of Slovakia. During the insects and mites known to attack Convolvulus study, high numbers of data have been collected, spp. and Calystegia spp. found by him in Canada and partly already published (Toth et al. 1997, and by others throughout the world. In Pakistan, 1998, 2001, 2003, Toth 2000, Defago et al. 2001, Baloch (1974) found 35 species of insects and 2 Toth & Cagafi 2005). Melanagromyza albocilia species of mites in association with Convolvulus Hendel, 1931 (Diptera: Agromyzidae), a native spp. The insect fauna noted on the field bindweed European stem borer of C. arvensis, was consid- in Egypt consisted of 13 species (Awadallah et al. ered as a very promising candidate for the biolog- ENTOMOL. FENNICA Vol. 16 ° Biology and host infestation ofMelanagromyza albocilia 255

Table 1. Details of localities weekly checked in southwest of Slovakia for the occurrence of M. albocilia (1998— 2000). Coord. = coordinates; Alt. = altitude in metres; Temp. = average temperature in °C; Rad. = global radiation; Crop = covering crop species.

Locality Alt.a Coord.a Relief Temp.b Rad.b Crop

Cajkov 206 48°17’51"N Plain 9.0 105—110 Sunflower 18°35’36"E Mala n / 103 47°50’45"N Hilly 10.2 115< Sunflower Hronom 18°41 ’24"E country Tehla 194 48°12’16"N Hilly 9.7 110—115 Sunflower 18°21 ’53"E country a Magellan GPS 315 2 b According to Zat’ko (2002); Global radiation is expressed in kcal/cm . ical control ofbindweed (Toth et al. 1998, 2000). causes the infested branches to dry up (Spencer This was the reason for the intensive study ofthis 1973). species. Because of the cryptic behavior of M. albo— Very little information is available about M. cilia, little or no information exists on its occur- albocilia. It is a greenish-black species with wing rence during the growing season and infestation length from 2.1 mm in male to 2.5 mm in female ofthe host plant. The objectives ofthis study were (Hendel, 1938, Spencer, 1966). It is known from to determine the following: (1) occurrence and Austria and Hungary (Hendel 1938, Soos & Papp abundance of M albocilia in cultivated areas in 1984), Egypt (Spencer 1973, Awadallah et al. Slovakia; (2) the over-wintering survival and 1976), Israel (Spencer 1973, 1974), England, emergence pattern of M albocilia; (3) the sea- Denmark and Sweden (Spencer 1976), Italy sonal biology ofM albocilia; and (4) the infesta- (Rosenthal & Buckingham 1982), Czech Repub- tion rate of field bindweed in the southwest of lic (Vala & Cemy 1997), Slovakia (Toth et al. Slovakia. 1998) and Serbia (Spasic & Smiljanic 1998). Fe- males ofM albocilia insert their eggs under the epidermis of field bindweed leaves. Hatched lar- 2. Material and methods vae feed on the leaftissue and make linear serpen- tine mines extending to the midribs, where each To determine the seasonal occurrence of M al— larva bores downwards towards the base of the bocilia and field bindweed infestation by the stem or even into the root. At this site, pupation agromyzid fly under field conditions in Slovakia, takes place after the larva gnaws an oval exit hole phenologically mature plants from 91 localities through which the emerged adult can find its way were investigated during the growing seasons of out (Awadallah et al. 1976). M. albocilia has 1999 and 2000. Three localities in the southwest been recorded solely from field bindweed in na- of Slovakia (Table 1) were checked weekly, from ture, though Rosenthal and Buckingham (1982) the end of May until the beginning of October. listed also Convolvulus althaeoides L. as an alter- Crops, where the samples were taken, were native host. grown according to local farmers’ practices. A related species M. convolvuli Spencer, Each week thirty plants per site were ran- 1971 was found in C. arvensis stems in Pakistan domly selected for dissection. For each selected (Spencer 1971). Females ofM convolvuli punc- plant, the number of healthy, dried and infested ture C. arvensis leaves with their ovipositors and stems was recorded. The stems were periodically feed on the ensuing exudates (Baloch 1977). Lar- dissected for developmental specimens (larvae, val feeding ofM convolvuli in the stems or lateral pupae and parasitized larvae). The presence ofthe branches considerably weakens the plant and in larvae and pupae could be detected only when the non-irrigated areas of limited rainfall the com- plants were dissected. The stems and roots were bined effect of the drought and larval attack first cut across (a few cm above the root). The in- 256 Trim et at. 9* ENTGMGL. FENNTCA Vet“ t6

Bite thistrihntien

The distrihutien ef MC atheet’tt’a in Stet/atria is shewn in Fig. t. The fty was feund at 9t teeati~ ties. eeniirrning that it is a eemmen inseet in Sievehia and it is eiesety retated tn the distrihu-— Fig. t. titer: tien ef its hest C. never/rat’s. The speeies eeeurred Hendet in dievatda (htaetr dets tndieete identities even at: eetder teeatities with an attitude mere where the species was reeerded). than 4th.) m ant. and the infestatien aehie‘ved sire—-

itar tevets te these at tewer situated. and warmer fested stems were eieariy Visihie due tn the hrewn teeaiities. tEtut tarvae were net parasitized at interier er the hete. sneh stems were separated. eeider teeatities at higher attitudes (285—435 in The rest ef stems were eut aeress again. aheut: ten a.s.i.). The highest attitude where tar‘vat parasites eentimeters higher. were ehser‘ved was 245d.) rn a..s.t. The preeedure was repeated. ateng the wheie tength ef the stem te nete infestatien. The stems with ne infestatien were eensidered as heatth; as 5.2a Seasenat hietegy wett ptants witheut infested stems were rnartted as heatth ptants. Based en these resu’its) pereent-u Fig. 2 shews seasenat hieiegy ef M1 attracted in age ef infested. ptants and stems was eateutated. seuthwestern Stet/atria. The insect everwintered Separated (infested) stems were sptit iengthwise. as pupa. and the emergenee efthe first generatien tty fettewing the feeding trash efthe tarvae either adutts started in the beginning efh/iay and eentinw dewn er up the stems. the iar‘vae er pupae nerw ued untit the end efh/tay in an e‘utdeer inseetary. matty eeutd he feund. thrpae ehtained. in this way Fiftymtwe aduits were reeevered in the spring at were transferred. te rearing eentainers Within the stems and. rents ef eaeh ptant. the pesitien and MQNTH VT VET iVTTT mix Xi numher ef and tart/ae t E H tarvae. pupae parasitised 5 E a was were s a reeerded. P‘upae and parasitised tart/as i I I 5 t i transferred in tuhes tn the F a ptastie taheratery. I E i E E E was a F The materiat kept in piastie tuhes with E l E

fer aeratien in an eutdeer I perferated tep inseetary E Z te assess s l the peried efadntt emergenee. itinerged I E i E t a tarvat parasiteids were eet_ieeted., eeunted. and E i suhmitted fer identitieatien. Veueher speeimens Fig. 2. desserts) hteiegy et at. eiheetiie Hendei in efparasiteids have 'heen retained in the eeiteetien seuthwestern Etievatda (eased en the data trern teeaitm ef the first anther. Besides the regutar e’hserw’au ties Tehie. state a l tirenem and defines; Tahte t). % 86’ tiens during t999 and 208%. pupae and parasitm m4... tees —~ an... seen :3 teed tarvae eftt/f. etheettta were eetteeted at the na- asst 39” three teeaiitiies atse during the fatt seasen at t. 5998 K

- ”/1, \ . te determine its everwwintering heha‘vier. as \s as g tn 88 further ieeatities were a. ,' r additien, sampted // we... threugheut Ste'vatria. in ditthrent geegraphie and 20 fl,” “egg-X: “Ha... etimatie the fati seasens ef t998 ------4.’ regiens. during ”a “Emmmfi‘; (/ ’ sate“ ‘tt—ts 20% (Fig. t) te reeerd the presenee ef the t—shisy ay nests-ea negates any in the stems. with a. agremyzid ity Starting sampie Fig. 3. its. atheeiiie adutt emergence in eutdeer ef it) ptants per site, the numher reaehed a maxi-— tnseetary during the spring tdhhmfiddt (the ratte (w) mum ef 3d ptants per site if fty tarvae and/er pun trern the tetai aduits emerged during spring eaeh pae were net feund. within the first t t) ptants. year). ENTOMOL. FENNICA Vol. 16 ° Biology and host infestation ofMelanagromyza albocilia 257

Table 2. Population dynamics of M. albocilia at three localities in Slovakia in 1999. Numbers of specimens are based on totals for 30 plants each date.

Date Tehla Mala n / Hronom Cajkov

La Pb PL° L P PL L P PL

1—10 June 0.26:0.09 0.19:0,08 0 ——————

11—20 June 0.28:0.27 0.29:0.32 0 ——————

21—30 June —————————

1—10 July 0.10:0.09 0.97:0.11 0 —————— 11—20 July 0.45:0,50 0.69:0.45 0.03:0.05 0.20:0.24 0 0 0.26:0.31 0.24:0.11 0 21—31 July 0.89:0.11 0.61:0.51 0.19:0.09 0.54:0.09 0.97:0.43 0.08:0.09 0.33:0.09 0.61:0.41 0.16:0.11 1—10 Aug. 1.37:0.75 1.00:0,46 0.41:0.31 1.30:0.44 0.78:0.10 0.55:0.44 0.23:0.11 0.63:0.17 0 11—20 Aug. 1.76:0.73 0.96:0.32 0.51:0,21 1.06:0.38 0.74:0.42 0.57:0.22 0.47:0.30 0.52:0.14 0.21 $0.27 21—31 Aug. 2.92:0.24 1.15:0.26 0.64:0.56 1.06:0.15 0.84:0.15 0.41:0.16 0.22:0.01 0.22:0.29 0.18:0.06 1—10 Sept. 2.35:0.78 1.02:0.57 1.58:0.66 0.88:0.41 0.72:0.38 0.30:0.22 0.39:0.22 0.31:0.30 0.22:0.19 11—20 Sept. 1.04:0.11 1.93:0.91 2.10:0.30 0.47:0.14 0.46:0.34 0.60:0.53 0.45:0.28 0.38:0.12 0.18:0.07 21—30 Sept. 1.73:0.24 0.67:0.17 1.64:0.72 ——————

a Mean of larvae per 10 evaluated stems 1r SEM b Mean of pupae per 10 evaluated stems 1r SEM c Mean of parasitized larvae per 10 evaluated stems 1r SEM

1999, 64 in 2000 and 39 in 2001 from the pupae was 1.30 i 0.44 larvae per 10 stems in 1999, but collected in the fall season. Maximum adult only 0.94 i 0.35 at Cajkov in 2000. The occur- emergence was from 6—10 May 1999 and 2000 rence of larvae was constant during the study. (52.0% and 89.1%) but from 16—20 May 2001 The number of larvae decreased sharply during (70.7%) (Fig. 3). The first empty pupal cases September. within the field bindweed stems were found dur- The first pupae were found at the beginning of ing the second half of June. Second-generation June and the number reached a maximum from adults occurred during a relatively long period August through September. The number ofpupae (June—August 1999). The first flies ofthe second per 10 stems reached a maximum of 1.93 i 0.91 generation emerged from the field collected pu- from 11—20 Sept. 1999 and 0.72 i 0.20 from 1—10 pae on 17 June 1999 in an outdoor insectary. The Sept. 2000 at Tehla. Nevertheless, the maximum last adult ofthe second generation emerged on 28 ofpupae was recorded in July at Mala n/Hronom August 1999 during the study. 7.3% of the sec- each year (Tables 2—3). ond-generation adults were collected in June, The number of pupae was markedly reduced 30.9% in July, but the majority (61.8%) within as a result of high larval mortality due to parasit- August 1999. ism. Although the first larval parasitoids were ob- Larvae were found from the middle of May served already during June, the larval para- until the end of growing season. The highest sitization was relatively low. The proportion of numbers were found during August and in the parasitized larvae was 22.6% at Tehla, 19.2% at first half of September. Although the peaks were Mala n/Hronom and 16.7% at Cajkov in June similar at all three localities evaluated, the num- 2000. The number ofparasitized larvae increased bers of larvae (but also pupae) per 10 stems were until the end of vegetation in September. During markedly lower at Mala n/Hronom and Cajkov September 2000, the parasitized larvae consti- (Tables 2—3). From June to the beginning of Sep- tuted 60.4%, 57.5%, and 41.2% of the all devel- tember, the number oflarvae fluctuated from 0. 10 opmental specimens recorded in Tehla, Mala i 0.09 to 2.92 d: 0.24 in 1999 and 0.10 i 0.17 to n/Hronom and Cajkov, respectively. All of the 1.45 i 0.41 (means and standard errors are given larval parasitoids were identified as Bracon here and hereafter) per 10 stems in 2000 at the 10- picticornis Wesmael, 1838 (Hymenoptera: Bra- cality Tehla. At Mala n/Hronom, the maximum conidae). While the larval parasitization was sta-

260 Tdth er al. *9 ENTOMOL. FENNlCA Vol. l6

:3 mean of specimens 3 r3 mean of specimens (13 . 4 per infested stem g: per Infested stem

-- is mean of specimens a: a mean of specimens 3 """"""""""""""""""""""""""""""""""""""""""""" (is per infested plant [g §g§g per infested plant g 3 ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 2— —— a __ U *‘i Q. 555; :3; as ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 9'3 5555 5555 5555 " """"""""""""""""""" 5+... 3:5: :§:§ 3:5: e 2 ______0 u.- gig: igig igig gig: o ______s 1-. = it 5: : gigig gigi igigi igig gigi igigi gigig gigi 1 (B IEIEI IEIE EIEIE EIEI EIEIE E E IEIE_ IEIEI_ IEIE_ “3 t fit it: it gigs at: gags sgsgs gsgsg gag; 93 E . gigig gg gg gg gigig gg l ' E c} 535353 353535 EEEEIE 353535 535353 35353 EEIEI IEIEIE EIEIEI IEIEIE 9 I “June June“I Au I June July Aug SeptiJuiy Aug Sept‘Juiy Aug Sept JulyAug s’ea Ziuiy septiiuneJuly Aug Sept TEHLA MALA N 1 HR. CAJKQV TEHLA MALI-l N 1 HR. CAJKOV 8. Mean number of M. albociiia Fig. 7. Mean number of M. elbocilia specimens (lar— Fig. specimens (lar— vae, infested stems vae, pupae, parasitized larvae) per infested stems pupae, parasitized larvae) per and of field bindweed under the field conditions and plants of field bindweed under the field conditions plants of three localities in Slovakia in 2000. of three weekly sampled localities in Slovakia in 1999. weekly sampled Numbers of are based on totals for 30 Numbers of specimens are based on totals for 30 specimens evaluated each date. plants evaluated for each date. plants

by temperature but also other factors influencing fields by B. picricornis was important in limiting diapause. in the case of M. aiiiocilia we assume ofthe species. B. picricornis showed the ability to that it has two overlapping generations per year in forage effectively for larvae even in a relatively Slovakia. Prolonged emergence of the second lower population. level ofM’. trlbocilitr (for exam— generation adults might indicate that partial third ple, (liajkov locality). It is a generalist larval ecto— generation could occur. Larvae and pupae were parasitoid. lt has been reared for instance from found. side by side throughout the season. Esperanto connexa (F) (Diptera: 'l‘ephritidae) The later attack of the stem borer fly, consid— (Solbreck 2000), leafrollers in the family rfortri— ered as the second generation caused the heaviest cidae (Lepidoptera) (Diaconu & Lozan 2000), infestation offield bindweed plants (up to 1000o) and Portrait/rte proxima (.lsepeletier) (.lrlymenopte— and increased also the number of stems attacked ra: Tenthredinidae) (Al-Saffar & Aldrich 1997). (Figs. 5—6). Similarly, M. COfiVOZl’Z/lt’i infested up M’. ulbociliu is a new host for B. picticomis. to 63% of field. bindweed plants in Pakistan ln addition to parasitism, high larval mortality (Baloch 1977). was observed through August and September. In contrast to M. sojae, which reached maxim Similarly, Wang (1979) noted high larval mortal_ mally 1.35 specimens per plant in. Sumatra (van ity ofM. sojrre, the actual. reason for this mortality den Berg et al. 1995), the densities ofM. tzlbocilitz remains to be investigated. were considerably higher in Slovakia. Peak den— Feeding by M: albocilia larvae in stems ap— sities generally occurred through August and parently did not impede the supply of water and September with 3.95 i 2.03, 2.34 i 1.65, and 1.68 nutrients to the upper parts of the plant. During i l .13 live specimens per plant recorded at Tehla, the larval period infested plants looked healthy Mala n/l-lronom and Clajkov, respectively (Figs. from the outside. The stems started to become 7__g)_ Van der (loot @930) reported that several. weak and dry after pupation, which takes place eggs were deposited per plant but only one or directly within the stems, probably because the sometimes two larvae of 1M. sojae were found. in larva gnaws an oval exit hole cutting off water the same feeding tunnel, suggesting that intra~ and nutrients flow to the upper part. specific competition was important. Similarly, Larvae ofMI albocit’iu bore only in stem and the number of specimens in the stems infested by roots of plant within the genus COI/lt’OZVMluS M. al'bocilz’rr varied from 0.84 :i: 0.40 to 1.34 rt: (Spencer 1973, Awadallah at all. 1976, Rosenthal 0.50. Nevertheless, we sometimes found up to 5 & Buckingham l98'2). The species appears to be specimens per stem what does not indicate strong highly host specific. Spencer (1965) mentioned: intraspecific competition of the species. “When a species from the family Agromyzidae is Parasitism ofM. albociiia larvae in Slovakian feeding within stems or seeds there is a high de— ENTOMOL. FENNICA Vol. 16 ° Biology and host infestation ofMelanagromyza albocilia 261

of the Irish Irish gree of probability that such species will be ceedings Royal Academy 97, Royal 2197223. strictly monophagous”. Before ovipositing, fe- Academy, Awadallah, K. T., Tawfik, M. F. S. & Shalaby, F. F. 1976: males field bindweed leaves and feed on puncture Insect fauna ofbind—weed, Convolvulus arvensis L., in the ensuing exudates (Awadallah et al. 1976). Giza, Egypt. 7 Bull. Soc. ent. Egypte 60: 15724. Spencer (1973) suggested three reasons for such van den Berg, H., Ankasah, D., Hassan K., Muhammad, behavior: firstly, to confirm that the host is cor- A., Widayanto, H. A., Wirasto, H. B. & Yully, I. 1995: stem rect; second, to ingest proteins from the host Soybean fly, Melanagromyza sojae (Diptera: Agromyzidae), on Sumatra: Seasonal incidence and which are for maturation ofthe ovaries; 7 necessary the role ofparasitism. International Journal of Pest and third, the actual feeding on the available car- Management 41: 1277133. bohydrates. Baloch (1977) found that freshly Baloch, G. M. 1974: Phytophagous organisms associated emerged females ofM. convolvuli that had not fed with bindweed, Convolvulus spp., in Pakistan. 7 Techn. Bull. Commonw. Inst. Biol. Contr. 17: 29736. on leafjuices offield bindweed laid only infertile Baloch, G. M. 1977: Insects as biological control agents of when on sweet eggs feeding potato (Ipomoea field bindweed, Convolvulvs arvensis. 7 PANS 23: batatas Linnaeus). Based on our data (unpubl.), it 58764. is clear that M. albocilia does not attack I. bota— De'fago, G., Ammon, H. U., Cagan, L., Draeger, B., Grae- M. tas. We propose thatM. albocilia may be a prom- ves, P., Guntli, D., Hoeke, D., Klimes, L., Lawrie, J., Moénne—Loccoz, Y., Nicolet, B., Pfirter, H. A., ising biological control agent of field bindweed Tabacchi, R. & Toth, P. 2001: Towards the biocontrol not in this This also be only respect. species might 7 ofbindweeds with a mycoherbicide. Biocontrol 46: a promising candidate for its primary damage: its 1577173. stem-boring feeding after pupation results in Diaconu, A. & Lozan, A. 2000: Ectoparasitoid braconids weakness and dryness ofstems, as shown by Toth (Hym.: Braconidae) of certain species of leafrollers (Lep.: Tortricidae) of fruit trees. 7 Mitt. Schweiz. (2000). In addition, it probably causes secondary Ent. Gesell. 73: 5711. damage e.g. by “opening the door” to several van der Goot, P. 1930: De Agromyza—vliegj es der inlandse such as and which can pathogens, fungi bacteria, katjang—gewassen op Java. Mededeelingen van het in turn increase the total damage and even cause Instituut voor Plantenziekten. 7 Asian Vegetable Re— deaths of the target weed. Mines formed during search and Development Center, Shanhua, Taiwan, China. 78 larval increase the risk of infection pp.

feeding (Toth 7 Hendel, F. 1938: Agromyzidae. In: Lindner, E. (ed.), et al. Based on all of the aforementioned, 2000). Die Fliegen der Palaearktischen Region: 17570. together with a high infestation of the host plant, Stuttgart, Germany. we conclude thatM. albocilia can be an important Mohyuddin, A. I. 1969: Insect from Calystegia spp. and biological control agent of C. arvensis, especially Convolvulus spp. 17570. 7 Techn. Bull. Commonw. Inst. Biol. Contr. 11: 937104. if it is imported to places where C. arvensis is an Rosenthal, S. S. & Buckingham, R. G. 1982: Natural ene— invasive without associated insect fauna plant mies of Convolvulus arvensis in western Mediterra-

in North There is also the 7 (e. g. America). pre- nean Europe. Hilgardia 5: 1719. sumption that M. albocilia will not experience Solbreck, C. 2000: Ecology and biology of Euphranta

7 high parasitism rates in such areas. connexa (Fabr.) (Diptera: Tephritidae) a seed preda— tor on Vincetoxicum hirundinaria Med. (Asclepia— daceae). 7 Ent. Tidskr. 121: 23730. Acknowledgements. We would like to thank Dr. Josef S06s, A. & Papp, L. 1984: Catalogue of Palaearctic Dip- Lukas (Department of Ecology, Comenius University, tera. Micropezidae7Agromyzidae. Vol. 9. 7Elsevier, Bratislava, Slovakia) for parasitoid identification and Dr. Budapest. Lloyd Knutson (Cornell University, USDA, retired) for Spasic, R. & Smiljanic, D. 1999: Stem-mining flies (Dip- usefiil comments on the manuscript. Part of this work was tera, Agromyzidae) as potential weed biocontrol the Grant No supported by Agency VEGA, project 7 agents. In: Peric, I. & Ivanovic, M. (eds), Proceed— 1/2424/05. ings ofInternational Symposium on Integrated Protec— tion of Field Crops, 25 September 1998, ijacka Banj a, Yugoslavia: 1297135. Plant Protection Society of Serbia, References Belgrade. Spencer, K. A. 1965: The species—host relationship in the Agromyzidae (Diptera) as an aid to taxonomy. 7 Pro- Al Saffar, Z. Y. & Aldrich, J. C. 1997: Factors influencing ceedings ofInternational Congress ofEntomology, 13 the survival ofPontaniaproxima that attack crack wil- July 1965: 1017102. low Salixflagilis. 7 Biology and Environment: Pro- Spencer, K. A. 1966: A revision ofEuropean species ofthe 262 T(W; et al. ' ENTOMOL. FENNICA Vol. 16

genera Melanagromyza Hendel and Hexomyza cal control agent of Field or Lesser Bindweed (Con- Enderlein, with a supplement on the genus Ophiomyia volvulus arvensis L.). 7 Entomologist’s Gazette 54:

Braschnikov. 7 Beitr. Ent. 16: 3760. 2337241. Spencer, K. A. 1971: A new Agromyzid (Diptera) from Toth, P., Vrablova, M. & Cagan, L. 1997: Importance of Pakistan feeding as a stem—borer in Convolvulus noctuid species for biological control of Field Bind- arvensis. 7 Bull. Ent. Res. 61: 3697371. weed (Convolvulus arvensis L.) and Hedge Bindweed Spencer, K. A. 1973: Agromyzidae (Diptera) ofEconomic (Calystegia sepium L.). 7Proceedings ofXIV Slovak Importance. 7Dr. W. Junk B. V., The Hague, Nether— and Czech Plant Protection Conference, Nitra: 3017 lands. 302. K. A. 1974: Some from Spencer, Agromyzidae (Diptera) Toth, P., Vrablova, M. & Cagan, L. 1998: First record of Israel. 7 Israel J. Ent. 9: 1417157. Melanagromyza albocilia (Agromyzidae, Diptera)

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