Host plants of the European tarnished plant bug Lygus rugulipennis Poppius (Het., Miridae)

J. Appl. Ent. 111 (1991), 484498 0 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0931-2048

Department of Environmental Sciences, University of Kuopio and Department of Agricultural and Forest Zoology, University of Helsinki, Finland

Host plants of the European tarnished plant bug Lygus rugulipennis Poppius (Het., Miridae)

By J. K. HOLOPAINEN and ANNA-LIISA VARIS

Abstract

Literature regarding the host plants of the European tarnished plant bug (Lygus rugulzpennzs Po pius) is surveyed and records of some unpublished host plants are added. The recorded hosts incluie 437 plants from 57 families. Of these 50 host plants are classified to genus and 387 to species or subspecies level. Oviposition of L. rugulipennis o r growth of its nymphs has been observed on 59 plant species. The most important host families are Brassicaceae, Asteraceae and Fabaceae. The highest total numbers of recorded host plants are from Poland, Finland and France. Damage to crop plants caused by L. rugulipennis has been most often recorded on alfalfa, clover, potato, cereals and sugarbeet. L. rugulzpennis probably has the broadest documented host plant range after Philaenus spumarius (L.) (Horn., Cercopidae), which has nearly 500 documented host plant species. The distribution and attractiveness of different hosts is discussed.

1 Introduction

Polyphagous Lygus bugs are pests of several cultivated plants especially in northern America (e.g. 112; 41; 134) and their numbers on crop plants can be reduced by attractive trap crops (e.g. 95). The European tarnished plant bug, Lygus rugulipennis Poppius is the most common pest in this genus in many countries in Europe, causing deformations of plant tissues and growth losses (eg. 124, 21, 27, 53) and it is a potential vector of plant viruses (121). L. rugulipennis is spread all over the Palaearctic region from tundras to the deserts of North Africa (40) but it does not occur in North America, where it is replaced by the tarnished plant bug Lygus lineoluris (Palisot de Beauvois) (34). BECH (9) listed over 80 host plants of L. rugulipennis, but since then numerous new host plant reports have been published.

The genus Lygus belongs to subfamily Miridae of the order Heteroptera and has about 300 species (52). KNIGHT (60) suggested that group of species were confined under the name of Lygusprutensis. WAGNER (130) confirmed the L. pratensis complex and separated a subgenus named Exolygus. KBLTON (58) considered Exolygus a junior synonym of Liocoris and promoted this subgenus to generic status. However, International Commission of Zoological Nomenclature (1963, Opinion 667) designated that Exolygus will always be a junior synonym of Lygus. This has confirmed that economic ‘lygus bugs’ will remain in the genus Lygus (52). L. rugulipennis has been formerly included Lygus pratensis complex raised by KNIGHT (60). REUTER (91) described it with the name L. pratensis pubescens. WAGNER (129), KULLENBERG (70) and LESTON (74) used the name L. pubescens. LIN- NAVUORI (75) reported that L. pubescens is identical with L. rugulipennis, which was described by POPPIUS (89). Since that priority is given to name L. rugulipennis. In reports published before WAGNER’S (129) and KULLENBERG’S (70) publications L. rugukpennis is included in L. pratensis group and it is not possible to evaluate if the reports concern L. pratensis or L. rugulipennis. However, according to SOUTHWOOD (105), the publication of WILSON (133) concerns L. rugul@ennis.

U.S. Copyright Clearance Center Code Statement: 0931-2048/91/1105-0484 $ 02.50/0

Host plants of Lygus rugulipennts 485

In this report we list the known host plants of L. rugulipennis. The objective of the present report is to gather the basic information of the host plant preference of L. rugulipennis in order to find potential trap crop plants o r families in the control of Lygus bugs in conifer nurseries or agricultural crops.

2 Materials and methods

Thc method of literature search was very similar to the rules presented by YOUNG (135). If the bug was reported from a plant species that plant was considered as a host plant, because Lygus bugs do not land on a plant that they will not at least probe (135). The first citation for a particular host plant from each country was included the host plant list. If egg laying or nymphal growth on specific host plant was observed, it was indicated in the host plant list to be evidence of the suitability of the plant for the food plant of the bug. Also, this food plant information from the later reports not included in the host reference list, was included and indicated in the host plant list.

The databases of biological, agricultural and medical literature were searched for host plants of L. rugulipennis. The literature from 1940 to late ~ O ' S , which is not available in databases was also searched. The variety of languages in Europe caused some limitations in the listing of plant species and it is possible that in cases where only the native names were used, and scientific names were not given, some plant species might be excluded. With the help of colleagues, lists of unreported host plants were obtained from several countries.

The plants are classified into families according to HAMET-AHTI et al. (47). The scientific names of the plants are according to MOORE (80) and DONY et al. (36).

3 Results and discussion

3.1 Host plant numbers

The recorded hosts consist total of 437 plants from 57 families. Of these plants, 50 (11 %) were classified to genus and 387 (89 YO) to species or subspecies level. Some of the host plants reported to genus level may concern the same species which is reported to species level. Of the host plants reported to generic level, 15 did not have any other host plant from the same genus and 35 had some species classified to species or subspecies level. Thus, the total number of reported host plant species is at least 402 (387 + 15), but it is probably higher. Oviposition or growth of nymphs is recorded from 59 plant species.

3.1.1 Host plants of Lygus rugu@ennis

After the name of each family, total number of species, number of plants determined to species or subspecies level and number of plants determined only to genus level are indicated in brackets. After the name of each species the reference number in reference list [or f = field observation, 1 = laboratory observation by the authors or abbreviation indicating personal communication: Au = Dr. B. AUKEMA (The Netherlands), Co = Mr. COUTINOT (France), De = Dr. M. DETHIER (Switzerland), He = Mr. E. HEISS (Austria), Kh = Dr. M. KHANJANI (Iran), Ra = Dr. VERA RACZ (Hungary), Re = Mr. L. REICHLING (Luxemburg)] and the abbreviation of corresponding country is given in brackets. Asterisk indicate plants from which oviposition or nymphal growth of L. rugulipennis is reported. Actinidiaceae (1/1/0)

Alismataceae (l/l/O)

Amaranthaceae (1 /1/0)

Apiaceae (Umbelliferae) (23/21/2)

Actinidiu chinensis Planch (30, I)

Alismu pluntugo-uquuticu, L. (9, DDR; 71, S)

Amaranthus cuudutus L. (Re, L)

"Aegopodium odugruria L. (f, I, SF) - '"Anethum gruveolens L. (f, SF; 8, DDR; 66, PL; 71, S) - Angelica rum& (37, F) - A. sp. (37, F) - Anthriscus silvestris (L.) Hoffm. (9, DDR; 71, S) - '&Apium gruveolens L. (9, DDR) - "Curum curvi L. (1, SF; 9, DDR; 87, BG) - C. verticillutum (L.) Koch (37, F) - CuuculisplutycuTos L. (37, F) - Conium muculutum L. (Co, F) - Coriundrum sutivum L. (9, DDR; 82; H; 87, BG) - 'iDuucus curotu ssp. sutivus (Hoff.) Arc. (He, A; 8, DDR; 37, F; 65, PL; 124, SF; 128, DK) - Eryngium cumpestre L. (9, DDR; 37, F) - E. muritimum L.

486 J . K . Holopairien and Anna-Lirsa Vans

(37, F) - Focnrculum vulgure Miller (9, DDR; 42, H; 114, PL) - Hevacleurn sphondylium L. (9, DDR) - H . sphondylium ssp. sibivicum L. (87, BG) - Lasrrpitium gallicurn L. (37, F) - Levisticum offirinale Koch (Re, L; 9, DDR) - Pastinaca sutiva L. (9, DDR; 37; F) - P. sp. (He, A) - Peroselinum cvispum (Miller) A. W. Hill (9, DDR; 66, PL) - Pimpinella anisum L. (9, DDR)

Ilex aqulfolium L. (37, F)

Hedeva helix L. (37, F)

Aquifoliaceae ( l / l /O)

Araliaceae (l/l/O)

Asclepiadaceae (2/1/1) Asclepias sp. (87, BG) - Vincetoricurn officinale (37, F)

Aspidiaceae ( l / l / O ) Dryopteris filzx-mas (L.) Schott (Re, L)

Asteraceae (Compositae) (60/47/13) Achillea filioendulina Lam. 171. S) - "-A. millefolium L. if, SF; 9, DDR; 37, F; 71. S; 108, CS) - A . ptarmici L: (Re, L; 37, F) -'A.'sb. (3, TR) - 'Anthemis'tznctoriu L. (9; DDR; 71, S) - A . sp; (He; A) - Arctium tomentosum Miller (1, SF; 71, S) - Arctotis sp. (25, GB) - Artemisza absinthium L. (f, 1, SF) - A . cina Berg. et Pol. (87, BG) - A . drucunculus L. (1, SF) - A . kurilensis Walp. (87, BG) - A. muritimu L. v. suhna Koch (87, BG) - ':-A. vulgaris L. (Au, NL; Re, L; 37, F; 86, DDR; 124, SF; 129, D) - A. sp. (123, SF) - Aster sp. (9, DDR; 133, GB) - Bidens trzpartita L. (71, S ) - Calendula officinulis L. (f, SF) - Curduus sp. (He, A; 3, TR; 9, DDR) - Centaurea cyanus L. (3, TR) - C. scubiosu L. (71, S ) - C. montana L. (37, F) - C. nigru L. (37, F) - "Chrysanthemum majus (Desf.) Aschers. (1, SF) - C. cuncifolium (37, F) - C. myconis (71, S ) - ':C. morifolium Ram. (109, GB) - C. segetum L. (71, S) - C. sp. (Au, NL; De, CH; 9, DDR; 94, CS; 133, GB) - Cirsium unglicum (37, F ) - C. umense (L.) Scop. (37, F) - C. eriophorum (L.) Scop. (37, F) - C. sp. (9, DDR; 129, D) - Dahlia pinnatu Cavann. (9, DDR) - D. cultorum L. (128, DK) D. sp. (132, GB) - Erigevon aureus (He, A) - E. canudensis L. (16, PL; 133, GB) - Eupatorium cannabium L. (37, F ) - Gnuphalium luteoalburn L. (37, F) - ':.G, uli inosum L. (37, F ; 54, SF) - "Heliunthus annuus L. (5, F; 9, DDR; 27, YU; 69, H; 97, BG) - InuBd helenium L. (107, SU) - I . sulicinu L. (f, SF) - 'Yeueanthernum vulgure Lam. (9, DDR; 18, N; 20, D; 71; S; 109, GB; 128, DK) - Mutricaria recutitu L. (f, 1, SF; 20, D; 71, S; 108, CS) - ':-M, rnutricurioides (Less) Porter (1, SF; Au, NL; Co, F; 108, CS; 109, GB) - "-M. sp. (Co, F; 94, CS) - Petusites ulbus (L.) Gaertner (37, F) - Pulicuriu dysentricu (L.) Bernh. (37, F) - ;:.Seneno jucobeu L. (Co, F; 109, GB) - ':.S. viscosus L. (20, D) - '"S. vulgaris L. (Co, F; 20, D; 54, SF; 128, DK, 133, GB) - Solidugo cunadensis L. (He, A; 71, S) - Tunucetum vulgure L. (f, SF; Co, F; Re, L) - T. sp. (129, D) - ::.Tripleurospermum inodorurn Schltz Bip. (18, N ; 20, D; 71, S; 124, SF; 128, DK) - ':.T. muritimum (L.) Koch (Au, NL; 9, DDR; 20, D; 71, S) - Venidiurn sp. (133, GB) - Zinnia sp. (133, GB)

Berberis thunbergii DC. 'Atropurpurea' (I, SF)

Alnus incuna (L.) Moench (He, A) -A. glutinosu (L.) Gaertner (Au, NL) -A . viridis (Chaix) DC. (He; A) - "Betulu pendulu Roth. (54, SF) - B. sp. (110, D) - Corylus uvellunu L. (71, S)

Berberidaceae ( l / l / O )

Betulacea (6/5/1)

Boraginaceae (3/2/1) Myosotis scorpioides L. (71, S) - M . amensis (L.) Hill (1, SF) - M . sp. (He; A)

Brassicaceae (Cruciferae) (123/122/1) Alliuriupetiolutu (Bieb.) Cavara & Grande (76, PL) -Alyssum ulyssoides (L.) (76, PL) -A . urduini Fritsch. 76, PL) -A . urgenteum All. (76, PL) A. montunum L. (76, PL) -A . suxutile spp. orientule (Ard.) (76, PL) -A . suxutile spp. suxutile L. (113, PL) - Arubidopsis thuliunu (L.) Heyn (113, PL) - Arubis a1 inu L (113, PL) - Armoruciu rusticunu R. Gaert. B. Meyer & Scherb. (113, PL) - Aubrietiu Ll toidea (L.) DC. (76, PL) - Barbureu arcuutu Rchb. (76, PL) - B. intermedid Boreau (76, PL) - B. strictu Andrz. (76, PL) - B. vernu (Mill.) Aschers. (76, PL) - B. vulgaris R.Br. (1 10, D; 113, PL) - Berterou incunu (L.) DC. (113, PL) - B i ~ u t e l l u uuriculutu L. (76, PL) - B. luevigutu L. (113, PL) - Brussicu chinensis L. (76, PL) - B. junceu (L.) Czern. (1, SF; 113, PL) - B. nupus L. (He, A; 113, PL - ':.B. nupus ssp. nupus v. nupobrussicu (L.) Reich. (18, N; 71, S; 76, PL; 122 SF) - ':B. nupus ssp. oleiferu DC. (2, D; 9, DDR) - B. nigru (L.) Koch (1, SF; 113, PL) - B. oleruceu L. v. ucephulu (2, D) - B. oleruceu L. v. botrytis Alef (50, S; 76, PL) - '$B. oleruceu L. v. cupitutu (f, SF; 9, DDR; 18, N; 76, PL) - B. oleruceu v. gernrniferu DC. (76, PL) - B. oleruceu L. v. medullosu The11 (2, D; 122, SF) - B. oleruceu L. v. gongylodes (76, PL; 122, SF) - B. oleruceu v. itulicu Plenck (76, PL) - B. oleruceu v. subuudu L. (76, PL) - B. oleruceu v. subellicu L. (76, PL; 122, SF) - B. oleruceu v. viridis L. (76, PL) - ':B. rupu L. (Co, F; 113, PL) - +B. rupu ssp. rupu L. (9, DDR; 102, S; 122, SF) - '$B. rupu ssp. oleiferu DC. (Co, F; 123, SF) - "B. rupu ssp. sylvestris (L.) Janchen (1, SF) - B. rupu ssp. typzcu (76, PL) - Bunius erucugo L. (113, PL) - B. orientalis L. (113, PL) -

Host plants of Lygus rugulzpennis 487

:‘-Cakile murztimu Scop (37, F; 113, PL) - Curnelina mzcrocurpa Andrz. (1 13, PL) - C. sativu (L.) Cram ( I 13, PL) - ‘“Cupsellu bursa-pustoris (L.) Medic. (I , SF; 9, DDR; 20, D, 71, S; 113, PL) - C. grundiflora Boiss. (76, PL) - Cardaria draba ( L . ) Dew. (76, PL.) - Cardumznopsis arenosu (L.) Hayek (113, PL) - Cheirunthus cheiri L. (113, PL) - Cocbleariu officinalis L. (113, PL) - Conringia orientalis (L.) Andrz. (76, PL) - Coronopus didymus (L.) Sm. (76, PL) - C. squamurus (Forskil) Acsch. (76, PL) ~ Crambe abyssinica Hochst. (38, DDR; 76, PL) - C. cordifoliu Stev. (76, PL) - C. maritimu L. (113, PL) - Descurainia sophia Prantl(ll3, PL) - Diplotaxis murulis (L.) DC. (113, PL) - D. tenuzyolia (L.) DC. (113, PL) - Erucu vesicuriu (L.) Cav. ssp. sutivu (Miller) Thell. (113, PL) - Emcastrum gullicum (Willd.) 0. E. Schultz (113, PL) - Erysimum cheiruntoides L. (1 13, PL) - E . crepzd2folzum Rchb. (76, PL) - E . diffusum Ehrh. (76, PL) - E . hieraciifolium L. (76, PL) - E . hungaricum Zap (113, PL) - E . pannonicum (L.) CR. (76, PL) - E . perofskiunum Fish. et Mey (76, PL) - E. pzeninicum (Zap.) Pawl. (113, PL) - E. pulchellum (Willd.) Boiss. (76, PL) - E . repundurn L. (113, PL) - E . wahlenbergii (Asch. et Engl.) Borb. (76, PL) - Fibzgiu clipeuta (L.) Medicus (76, PL) - Hesperis matronulis L. (113, PL) - Hirschfeldia incanu (L.) Lagreze-Fossat (76, PL) lberis amara L. (76, PL) - Iberzs umbelluta L. (113, PL) - Isatis tinctoria L. (He, A; 113, PL) - Lepidium campestre (L.) R. Br. (113, PL) - L . densiflorum Schrad. (76, PL) - L. raminifolium L. (76, PL) - L. Iatifolium L. (113, PL) - L. neglectum Thell. (76, PL) - L. perfotatum L. (76, PL) - L. ruderale L. (76, PL) - L. sativum L. (113, PL) - L . virginicum L. (1 13, PL) - Lobuluriu maritimu (L.) Desv. (113, PL) - Lunariu unnua L. (76, PL) - L . redzvivu L. (76, PL) - Matthiolu bicornis (Sibth. et Sm.) DC. (76, PL) - M . incuna (L.) R. Br. (113, PL) - M. sinuutu (L.) R. Br. (37, F) - Moricundiu arvensis (L.) DC. (76, PL) - Myugrumperfoliatum L. (76, PL) -Nasturtium officinule (L.) R. Br. (76, PL) - Nesliupuniculutu (L.) Dew. (1 13, PL) - Pelturiu ulliucea Jacq. (76, PL) - Raphunus ruphunistrum L. (109, GB; 113, PL) - R. sativus L. (113, PL) - R. sativus v. niger L. (76, PL) - R. sativur v. oleiformis Pers. (76, PL) - R. sativus v. rudiculu Pers. (76, PL) - Rapistrum perenne (L.) All. (76, PL) - R. rugosum (L.) All. (3, TR; 76, PL) - Rorippa amphibia (L.) Beser (113, PL) - R. urmorucioides (Tausch.) Fuss. (76, PL) - R. austriucu (Cr.) Bess. (76, PL) - R. palustris (Leyss.) Bess. (76, PL) - R. sylvestris (L.) Bess. (1, SF; 113, PL) - Sinupis ulbu L. (1, SF; 113, PL) - ‘:.S. urvensis L. (Co, F; 76, PL) - S. sp. (Co, F; 2, D) - Sisymbrium altissimum L. (76, PL) - S. uustrzucum Jacq. (76, PL) - S. irio L. (1 13, PL) - S. loeselii L. (113, PL) -S. officinule (L.) Scop. (113, PL) - S. orientale L. (76, PL) -S. strictissimum L. (113, PL) - Succowzu bulearica (L.) Med. (76, PL) - “Thluspz uruense L. (1, SF; 9, DDR; 20, D; 71, S)

Buxus sempervirens L. (Au, NL)

Cumpanula medium L. (87, BG) - Lobeliu wens L. (37, F)

Cannabis sutzvu L. (86, DDR) - Humulus lupulus L. (110, D; 132, GB)

Loniceru morrowii A.Gray (117, N) - Syrnphoricurpos rivularis Suksd. (117, N) - Viburnum tinus L. (37, F)

‘“Stelluriu media (L.) Vill. (9, DDR; 54, SF; 109, GB) - Cerustiumfontunum Baumg. (1, SF)

Atriplex hulimus L. (37, F) - A . hostutu D.C. (37, F) - putulu L. (20, D) - A . sp. (9, DDR) - ‘+Beta vulgaris L. v. altzssimu Doell (9, DDR; 15, PL; 49 D; 56, H, 108, CS; 123 SF) - B. vulgaris L. v. conditivu Alef (126, SF) - B. vulgaris L. v. muritimu (L.) Arcangeli (37, F) - B. vulgaris L. v. v u v , (122, SF) - B. s!. (133, GB) - “Chenopodium album L. (1, SF; Re, L; 9, DDR; 37, F) - C. am roszotdes L (37, F) C. bonus-henricus L. (25, GB; 37, F; 87, BG) - C. botryoides (37, F) - C. hybridum L. (37, F) - C. polyspermum L. (He, A) - “.C. sp. (Co, F; 3, TR; 132, GB) - Corrigiolu littoralis L. (37, F) - Spinuciu oleruceu L. (9, DDR)

Cichorium intybus L. (1, SF) - Crepis turuxifoliu Thuill. (37, F) - Hypochoeris sp. (37, F) - Tuvaxucum Koksughyz Rodin (110, D) - T. sp. (9, DDR)

Hypericum tertupterum Fries (37, F) - H. sp. (3, TR)

Sedum sediforme (Jacq.) Pan. (37, F)

Cucumis sutivus L. (68, PL; 125, SF)

Chumuecypuris luwsoniunu (A. Murr.) Parl. (Au, NL) - Juniperus communis L. (71, S) - Thuju occzdentulis L. (Au, NL)

Buxaceae (l/l/O)

Campanulaceae (2/2/0)

Cannabaceae (2/2/0)

Caprifoliaceae (3/3/0)

Caryophyllaceae (2/2/0)

Chenopodiaceae (17/15/2)

Cichoriaceae (5/3/2)

Clusiaceae (2/1/1)

Crassulaceae (1/1/0)

Cucurbitaceae (l/I/O)

Cupressaceae (3/3/0)

488 J . K Holopuinen und Annu-Liisu Vuris

Dipsacacea (1/0/1)

Ericacea (3/3/0) Scabiosu sp. (He, A)

Cullunu vulguris (L.) Hull (Au, NL; 37, F; 71, S; 109, GB, 111, PL; 124, SF) - Ericu scopuriu (37, F) - Vuccinium myrtillus L. (20, D; 71, S)

Fabaceae (Leguminosae, Papillionaceae) (32/25/7) C~irisus scopurius (L.) Link spp. scopurius (109, GB) - Glycine ma2 (L.) Merr. (136, I) - Luthyrus sp. (37, F) - Lotus sp. (37, F) - Lupinus ulbus L. (23, F) - L. sp. (1, SF; 9, DDR; 14, PL) - "Medicugo suttvu L. (Kh, IR; 1, D; 3, TR; 6, YU; 10, H; 17, CS; 21, F; 22, GR; 33, A; 44, DDR; 77, AL; 79, PL; 88, BG; 123, SF; 127, SU; 133, GB) - M . sp. (37, F) - Melilotus officinulis (L.) Pallas (I, SF) - Onobrychis viciifoliu Scop. (35, H, 59, SU) - Ononis cumpestris L. (37, F) - 0. natrix L. (37, F) - 0. repens spp. repens L. (37, F) - Ornithopus persupillus L. (14, PL) - "Phuseolus vulgaris L. (9, DDR; 61, PL; 124, SF) - Pisum urvense L. (9, DDR; 14, PL; 110, D; 127, SU) - "-P. sutivum I.. (109, GB; 110, D) - Trifoliurn alexundrinum L. (64, H) - T. urvense L. (74, GB) - '". hybridum L. (109, GB) - T. incumutum L. (9, DDR) - '!-T. medium L. (71, S; 128 DK) - '>T. pratense L. (9, DDR; 20, D; 37, F; 33, A; 64, H; 71, S, 79, PL; 100, CS; 123, SF; 127, SU; 133, GB) - T. resupinaturn L. (f, SF) - 'cT. sp. (Co, F; Re, L; 24, R; 94, CS) - Trigonellu coeruleu L. (71, S) - Ulex europueus L. (37, F; 109, GB) - U. sp. (37, F) - Vicia eruccu L. (1, SF) - ::.V.fubu L. (9, DDR, 110, D; 133, GB) - V. suttva L. (9, DDR; 20 D; 127, SU; 133, GB) - '+V. sp. (Co, F)

Facaceae (1/1/0)

Geraniaceae (1/1/0)

Gesneriaceae (1/1/0)

Ginkgoaceae (1/1/0)

Grossulariaceae (5/4/1)

Quercus robur L. (109, GB)

Erodium cicuturium L'Her. (Au, NL)

:>Suintpuuliu ionunthu (H. Wendl) (1, SF)

Ginkgo bzlobu L. (I, SF)

Ribes alpinurn L. (71, S) - R. nigrum L. (f, SF; 51, S; 92, SU; 110, D) - R. rubrum L. (92, SU) - R. uva-erispu L. (133, GB) - R. sp. (133, GB)

/ u g h regia L. (Au, NL)

/uncus gerardi Loisel (71, S)

Glechoma hederacea L. (I, SF) - Lamium album L. (I, SF; 109, GB) - L. purpureum L. (1, SF) - L. sp. (He, A) - Lycopus europaeus L. (71, S) - Mentha aquatica L. (37, F) - M . arvensis L. (1, SF) - M . sp. (86, DDR) - Nepeta sp. (3, TR) - Salvia splendes Sello ex Roem. & Schult (87, BG) - Thymus vulgaris L. (1, SF) - T. sp. (3, TR)

Allium cepa L. (67, PL)

'"Linum usitutissimum L. (f, SF; 102, S)

Alcea rosea L. (87, BG) - Althaea officinalis L. (f, SF; 87, BG) - Hibiscus rosa-sinensis L. (1, SF) - Malva moschata L. (37, F)

Fraxinus excelsior L. (Re, L; 9, DDR)

Juglandaceae ( l / l / O )

Juncaceae (1/1/0)

Lamiaceae (Labiatae) (12/8/4)

Liliaceae (1/1/0)

Linaceae (1/1/0)

Malvaceae (4/4/0)

Oleacea (1/1/0)

Onagraceae (Oenotheraceae) (3/2/1) Chamerion angustifolium (L). J.Holub (f, SF; 9, DDR, 110, D) - Epilobium sp. (4, DDR) - Oenothera biennis L. (37, F)

Papaveraceae (3/3/0)

Pedaliaceae (1/1/0)

Peperomiaceae (1/1/0)

Pinaceae (5/5/0)

Chelidonium majus L. (9, DDR) - Papaver rhoeas L. (f, SF) - Pupuver somniferum L. (1, SF)

Sesamum indicum L. (De, CH; 106, GB)

Peperomia subtrinervis hort. (1, SF)

Larix decidua Miller (Au, NL) - L. leptolepis (Sieb. & Zucc.) (109, GB) - "Piceu abies (L.) (He, A;

Host plunts of Lygus rugulipennis 489

71, S; 124, SF) - "Pznus sylvestris L. (Au, NL; Re, L; 71, S; 109, GB; 111, PL; 124, SF) - Pseudotsugu menziesii (Mirbel) (Au, NL)

Pluntugo lunceolutu L. (71, S ) - '"P. mujor L. (105, GB; 110, D) P. sp. (37, F)

Agrostzs stoloniferu L. (63, H) - A . cupilluris L. (I, SF; 71, S) - A. genzculutus L. (1, SF) - Avenu sutzvu L. (9, DDR; 14, PL; 49, D; 71, S; 108, CS; 124, SF) - Culumugrostis epigejos (L.) Roth (I, SF) - Dactylis lomerutu L (I SF) Festucuprutensis Hudson (63, H) - F. rubru L. (32, S; 63, H) - '"Hordeum v u i u r e L. (14,'Pt; 71 s; 108, CS; 122, SF) H . sp. (Re, L) - Lolium perenne L. (1, SF; 2, U) - L. temulentum L. (108, CS) - Moliniu coeruleu (L.) Moench (37, F) - Punicum miliuceum L. (9, DDR; 115, SU) - Phuluris urundinuceu L. (71, S ) - Phleum prutense L. (71, S; 127, SU) - : L P ~ u unnuu L. (54, SF; 108, CS) - P. prutenszs L. (1, SF; 63, H; 137, SU) - Secule cereule L. (14, PL; 49, D; 71, S; 96, SU; 108, CS; 124, SF) - ':.Triticum uestivum L. (9, DDR; 14, PL; 62, H ; 71, S; 83, SF; 96, SU; 108, CS) - Zed mays L. (14, PL; 90, H ; 108, CS)

Fugopyrum esculentum Moench (14, PL) - Polygonum umphibium L. (37, F) - P. uviculure L. (37, F; 108, CS) - P. uviculure ssp. littorule L. (37, F) - P. hydropiper L. (Au, NL; 37, F) - P. luputhifolium L. (37, F) - P. persicaria L. (37, F; 109, GB) - P. polystuchum Meissner (37, F) - P. tuturzcum (37, F) - Rumex ucetosu L. (Co, F; 9, DDR) - R. ucetosellu L. (He, A; 1, SF) - R. crispus L. (Au, NL) - R. dentutus (3, TR) - R. longifolius DC. (Co, F) - ':-R. obtusifolius L. (He, A; 109,

Phntaginaceae (3/2/1)

Poaceae (Graminae) (21 /20/1)

Polygonaceae (15/14/1)

GB) - R. SP. (133, GB) Primulaceae (1/1/0)

Ranunculaceae (1/1/0)

Rosaceae (19/17/2)

Lysimuchiu vulguris L. (1, SF; 71, S)

"Rununculus repens L. (54, SF)

'"Alchemillu subcrenutu Buser (1, SF) - Aphunes urvensis L. (37, F) - Aruncus dioicus (Walt.) Fern (71, S) - Crutuegus luevigutu (Poiret) D.C. (37, F) - C. monogynu Jacq. (37, F; 109, GB) - Filipendulu ulmuriu (L.) Maxim. (Re, L) - Fruguria x ununussu Duch. (f, SF; 43, S; 118, N ; 133, GB) - Mdus x domesticu Borkh. (78, H; 128, DK; 133, GB) - M. sylvestris Miller (9, DDR) - M. sp. (Au, NL) - Physocurpus opulifolius (L.) Max. (117, N) - Prunus domesticu L. (133, GB) - P. pudus L. (71, S ) - Potentillu fruticosu L. (117, N) - Pyrus communis L. (103, N; 133, GB) - Sorbus uucupuriu L. (He, A) - Spiraea x bumuldu 'Froebeli' Bum. (117, N) - Rubus frutzcosus L. (133, GB) - R. sp. (He, A)

Asperulu sp. (3, TR) - Gulium ulbum Miller (1, SF) - G. upurine L. (37, F; 109, GB) - G. palustre L. (71, S) - G. verum L. (32, S) - G. sp. (He, A)

Sulix sp. (Au, NL; Re, L; 71, S)

Astilboides tubuluris (Hemsl.) Engl. (71, S) - Suxifrugu sp. (1, SF)

Digitalis purpureu L. (99, D) - Linuriu vulguris Miller (1, SF) - L. sp. (85, TR) - Pediculuris sylvuticu L. (37, F) - Scrophuluriu nodosu L. (87, BG; 110, D) - Verbuscum lychnitis L. (37, F) - V. nigrum L. (9, DDR) - V. phlomoides L. (37, F) - V. pulverulenturn Vill. (37, F) - V. thupsiforme (37, F) - V. thupsus L. (87, BG) - V. sp. (He, A; 3, TR; 37, F)

Browulliu speciosu Hook (1, SF) - Duturu strumonium L. (87, BG) - Hyoscyurnus niger L. (86 , DDR) - Lycopersicon esculentum Mill. (119, SF) - Nicotiunu tubacum L. (De, CH; 81, DDR; 106, GB; 108, CS; 110, D) - Solunum melongenu L. (110, D) - ' pS . nigrum L. (Co, F; 133, GB) - 'tS. tuberosum L. (7, E; 9, DDR; 14, PL; 49, D; 56, H; 84, S; 94, CS; 121, SU; 124, SF; 133, GB)

"Urticu dioicu L. (Au, NL; He, A; Ra, H; Re, L; 9, DDR; 20, D; 37, F; 71, S; 124, SF; 128, DK; 133, GB) - U. urens L. (Re, L; 9, DDR; 71, S) - U. urens L. ssp. sondenii (f, SF) - U. sp. (De, CH; 94, CS; 129, D)

Vuleriunu off;;n'nulis L. (1, SF; 87, BG)

Rubiaceae (6/4/2)

Salicaceae (1/0/1)

Saxifragaceae (2/1/1)

Scrophulariaceae (12/10/2)

Solanacea (8/8/0)

Urticacea (4/3/1)

Valerianaceae (1/1/0)

490

Medicago sat iva

Jrifoliurn pratense

Ur i i ca dioica

Solanurn tuberosurn

Trrtrcurn aestivurn

Arternisia vulgaris

Avena sativa

J . K. Holopainen und Anna-Liisu Varis

l I I I I I 1 1

- - I .~ --

~ ~ -- -2

-1

Number of reports

0 2 4 6 8 10 12 14 l f i 18

D Beta vulgaris altissima

Calluna vulgaris

Daucus ca ro ta

Pinus sylvestrrs

1 1 Secale cereale

Fig. 1. The most often reported host plants of L. rugulipennis

3.2 Geographical distribution of host species

L. rugulipennis is found on host plants from 24 European countries and from Iran (table). It occurs also in Belgium (identified by Dr. G. SCHMIDZ from Malaise trap collections, CHRISTIANNE FASSOTTE, pers. comm.), but information of host plants is not available. The highest total number of recorded host plants is from Poland, Finland and France (table). The geographical distribution of the numbers of host plants at least gives information of the areas where this species seems to have economic importance. Damage to crop plants caused by L. rugulipennis has been most often recorded on lucerne, red clover, potato, cereals, and sugarbeet (fig. 1). L. rugulipennis is typically a pest insect in northern and central parts of Europe and only occasionally damages crop plants in southern Europe. According to ERDELYI and BENEDEK (40) L. rugulipennis is most common on lucerne in warm and dry regions, but due to its wide ecological valency, it maintains better than other Mirid pests of lucerne even under cool and wet climate.

3.3 Phylogenetic distribution of host species

The most important host families are the Brassicaceae, Asteraceae and Fabaceae (fig. 2). Of the numbers of L. rugulipennis host species in a plant family, 38 % (n = 53, p < 0.001) was explained by the number of European plant species in that family. However, the importance of different host plant families is affected by the type of host plant studies, which for example in the case of Brassicaceae increased greatly by the Polish studies of the insects occurring on several species of Brassicaceae. The host plants of L. rugulipennis comprise more than 20 % of the known European species in the family Brassicaceae (go), while in the other preferred host families the proportion of host species is less than 10 Yo. Furthermore, in the list of host plants of L. lineoluris, Asteraceae and Fabaceae had clearly more records than Brassicaceae (135).

Host plants of Lygus rugulipennis

Number of reported host plants of L. rugulipennis in European and Asian countries

491

Country Total number Number of economically of host plants important species

Poland (PL) France (F) Finland (SF) German Democratic Republic (DDR) Sweden (S) United Kingdom (GB) Federal Republic of Germany (D) Austria (A) Czechoslovakia (CS) The Netherlands (NL) Bulgaria (BG) Hungary (H) Luxemburg (L) U.S.S.R. (SU) Turkey (TR) Norway (N) Denmark (DK) Switzerland (CH) Yugoslavia (Yu) ’ Italy (I) Albania (AL) Greece (GR) Iran (IR) ’

Romania (R) Spain (E)

137 97 94 62 56 50 41 24 20 19 18 17 15 16 13 11

8 4 2 2 1 1 1 1 1

32 8

43 32 21 27 23

4 12 4 5

11 1

12 1 4 4 3 2 2 1 1 1 1 1

Nevertheless, the Brassicaceae seem to be optimal hosts for L. rugulipennis in many ways. According to TAHVANAINEN (116) an insect herbivore must overcome a hierarchy of environmental barriers to feed on a herbaceous plant. The first barrier is geographical distribution of the plant. Only the plants, introduced or native, occurring in the geographical distribution area of the herbivore are potential host plants. The other barriers are: temporal availability of the plant, habitat type of the plant, microhabitat around the plant and chemical properties of the plant. Most species in the family Brassicaceae are annual or biennial herbs of the early successional phases of field vegetation due to their capability to grow in relatively low temperatures (39). Overwintered adults of L. rugulipennis com- monly occur in fields (124) and other sun-exposed areas ( 5 5 ) in late May and early June. Thus, the temporal availability and plant habitat of Brassicaceae are optimal to L. rugulipennis.

3.4 Attractiveness of the host plants

The attractiveness of a host plant to Lygus bugs depends on many factors such as the age of the plant and the availability of succulent foliage (93), concentration of defensive chemicals (12, 13) availability of nitrogen (31) and the physiological condition of the plant (73). In open-air cage experiments (54) and in the laboratory we observed that many plants listed as a host plant of L. rugulipennis in chap. 3.1.1, were not suitable for feeding and egg laying, if other more attractive plants were available. Among these less attractive plants were, e.g. Chenopodium album, Malus x domestica, Sorbus aucuparia, Humulus lupulus and Chame- rion angustifolium. Plants from the genera that are not very attractive according host plant list, were also unsuitable food plants in laboratory. These were e.g. Allium,Campanula, Convallaria, Ranunculus, Rosa and Viola species.

Most of the favoured host plant species are cultivated plants or they are wild ruderal o r

492 J . K. Holopainen and Anna-Liisa Varis

Number of species

0 300 600 900 1200 1500

Brassicacea - - Asteraceae

Fabaceae

Apiaceae

Po ace a e

Rosaceae

Chenopodiaceae

Polygonaceae

Lamiaceae

Schrophulariaceae

0 European species

6 X l Hosr species So I an ace a e

Betulaceae

Rubiaceae

I I I I

0 30 60 90 120 150

Number of host species Fig. 2. Number of European species and number of L. rugulipennis host plants in different plant families

field plants. Most of the preferred wild plants are nitro- philous (39) such as Urtica dioica, Arternisia vulgaris, and Tripleurospermurn inodorum (124), Matricaria mat- ricarioides, (108) Senecio vulgaris (54). The most often reported cultivated plants Medicago sativa and Trijolium pratense belong to family Fabaceae, in which the plants are fixing nitrogen symbiotically and therefore are more nitrogen-rich than other crop plants (57).

Polyphagous insects must adapt to variety of defensive plant chemicals. In the plant family Poaceae (Gramineae) defensive chemicals are infre- quent and d o not accumulate in quantity as in many other plant families (48). This family is exploited by the polyphagous members of the family Acridoidea, while they avoid feeding on broad-leaved plants, which means that grasshoppers and locusts are

not adapted to plant toxins (48). Lygus bugs have an opposite tendency. They are rarely found on many grass species, which often are adapted to rather nitrogen poor conditions (39), while they occur on many broad-leaved plants and feed often on flower buds and seeds (120) and apical meristems (124).

The defensive secondary plant compounds have been concentrated in flower buds and seeds of plants and concentration of nitrogen is also highest in these plant organs (11). The wide host plant range of L. rugulipennis suggests that the secondary compounds (e.g. mustard oils in Brassicaceae) are not important feeding barriers to L. rugulipennis and the bugs can somehow operate with numerous compounds that are known to be toxic to herbivores (48).

The preference of i. rugulipennis to feed on flower buds and apical meristems of many plant species is possibly due to higher nutritive value of these plant organs and thus, the nitrogen content of the host plant is the key factor to determine the host plant selection of this species.

3.5 Host plant niches

At the moment L. rugulipennis seems to have the broadest documented host plant niche after the meadow spittlebug, Philaenus spumarius (L.). P. spumarius nymphs have 382 host plants (308 determined to species and 74 to genus level) in USA (131) and according to HALKKA et al. (45, 46) this species has 171 host plants (161 determined to species and 10 to genus level) in Finland. When added to host plant list from USA by WEAVER and KING (131) HALKKA et al. (46) report 114 additional host plants. This means the total of 496 host

Host plants of Lygus rugulipennis 493

plants of which 428 are reported to species level. Furthermore, BUHR (26) reports over 200 plant genus that may act as host plants of P. spurnarius. Polyphagous larvae of the gypsy moth, Lymantriu dispur L., has been estimated to have more than 500 host plant species (72) and the Indian stick insect Curausius morosus (de Sinety) is observed to accept almost all offered plant species as food (29), but comprehensive host plant lists for these species have not yet been published.

Many species in the genus Lygus are polyphagous (110), but for most of the species comprehensive host plant list is lacking. BECH (9) listed over 80 host plants of L. rugulipennis and SCOTT (93) 112 host plants of L. hesperus Knight. According to YOUNG (135) L. limolayis has 328 host plants classified to species and 57 identified only to genus level. In addition to this L. lineolaris is reported from six conifer species (98, 104), which means the total of 391 known host plants.

As YOUNG (135) stated, Lygus bugs might have the widest feeding niche of any known arthropods, when their predator and scavenger capabilities are added to their plant diet. The development from the first-instar nymph to adult exclusively on animal food i.e. insect larvae is possible (20). L. rugulipennis has been also found to predate on eggs of Leptinotursu decemlineuta Say (19) and Pegomya betue Curt. (124). This ability also confirms the attraction of the species to nitrogen rich food. It is known that the nitrogen content in insects is higher than in plant foliage (e.g. 101).

4 Conclusions

The plant species f rom the families Brassicaceae, Asteraceae and Fabaceae seem to be the most suitable trap crops to attract L. rugulipennis from crop plants. The results also suggest that adequate supply of nitrogen is important to keep the attractiveness of trap crop high.

Acknowledgements

We thank B. AUKEMA, M. BIERI, D. COUTINOT, M. DETHIER, C. FASSOTTE, R. C. HEDLUND, E. HEISS, M. KHANJANI, V. RACZ, L. REICHLING and G. SCHMIDZ for providing information of host plants of L. rugulipennis. We are grateful to S. LAUREMA and J. OKSANEN, M. RAATIKAINEN and J. TAHVANAINEN for reading the manuscript and to SILJA MAKELA for the technical assistance. This work was supported financially by the Academy of Finland.

Zusammenfassung

Wirtspflanzen der Triiben Feldwanze, Lygus rugulipennis Poppius (Het., Miridae) Es wird ein Literaturuberblick iiber das Wirtspflanzenspektrum von Lygus rugulipennis gegeben, das durch einige unveroffentlichte Wirtspflanzenangaben erganzt wird. Es konnten 437 Wirtspflanzen aus 57 Familien ermittelt werden. Von diesen sind 50 Wirtspflanzen bis zur Gattung, 387 bis zur Art oder Unterart bestimmt. An 59 Pflanzenarten konnte eine Eiablage oder die Entwicklung der Nymphen beobachtet werden. Die wichtigsten Wirtspflanzen gehoren zu den Familien Brassicaceae, Asteraceae und Fabaceae. Die grofite Wirtspflanzenzahl wird aus Polen, Finnland und Frankreich gemeldet. Schaden an Nutzpflanzen durch L. rugulipennis wurden iiberwiegend an Luzerne, Klee, Kartoffeln, Getreide und Zuckerriiben festgestellt. Neben Philaenus spurnaris (L.) (Hom., Ceropidae), dessen Wirtspflanzenspektrum nahezu 500 Pflanzenarten urnfafit, hat L. rugulipennis das wohl breiteste dokumentierte Wirtspflanzenspektrum. Die Verbreitung und Attraktivitat der verschiedenen Wirte wird diskutiert.

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~

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Authors' addresses: Dr. J. K. HOLOPAINEN, Ecological Laboratory, Department of Environmental Sciences, University of Kuopio, P.O. Box 6, SF-70211 Kuopio; Prof. ANNA- LmA VARIS, Department of Agricultural and Forest Zoology, University of Helsinki, SF-00710 Helsinki, Finland

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Host plants of the European tarnished plant bug Lygus rugulipennis Poppius (Het., Miridae)