Patterns of Host Exploitation by Meteorus communlS (Hymenoptera: )

KENNETH J. WEST AND JEFFREY C. MILLER Department of Entomology, Oregon State University, Corvallis, Oregon 97331

Environ.Entomol.18(3): 537-540 (1989) ABSTRACT The Meteorus communis (Cresson)was reared from 7 of 15 species of noctuids collected in peppermint and . of these species consistently Downloaded from https://academic.oup.com/ee/article/18/3/537/2393821 by guest on 29 September 2021 involved the second through penultimate instars. The most frequently parasitized species were (Hufnagel) (35.3%),Dargida procinc!a (Grote) (15.0%),and saucia (Hubner) (6.9%). Parasitism of P. saucia, A. ipsilon, and Autographa califomica (Speyer) (3.8%)by M. communis are new host records. Alsoparasitized were the lesscommon Pseudaletia unipuncta (Howorth) (20.0%), pyramidoides (Guenee) (10.0%),and Xylena nupara Lintner (5.0%). Percentage parasitism by M. communis was significantly higher in alfalfa than in peppermint. Three or four overlapping generations of M. communis may occur per year in western Oregon. Larvae of M. communis overwinter inside the host larvae.

KEY WORDS Insecta, MeteoTUs communis, hosts, parasitism

TIn: BIOLOGY of only a few species of Meteorus plots to assess the parasitism of noctuid larvae were Haliday has been described. In general, species of established in peppermint and alfalfa near Cor- M('teorus are solitary or gregarious endoparasitoids vallis, Benton Co., Oreg. Four peppermint fields of larval Coleoptera and , and many and three alfalfa fields were sampled during 1984. are polyphagous. Typically, development proceeds The original four plus three additional peppermint through three instars (Simmonds 1947, Balduf 1968, fields and two of the three original alfalfa fields Askari et al. 1977). The third instar exits the host were sampled during 1985. and spins a cocoon which characteristically dangles Sampling was conducted and standardized as fol- by a thread from the vegetation. Larvae of M. lows. A series of 25 sweeps in a 1200 arc with a communis (Cresson) overwinter inside host larvae standard (38 cm diameter) sweep net were taken. (Simmonds 1947, Balduf 1968, Shaw 1981, West Sweeps were started at the edge of a field and taken 1988). 1 m apart progressing into the field. This process One of the most frequently reared was repeated moving laterally 3 m until a total of from noctuid larvae collected on peppermint and 250 sweeps was taken or 250 noctuid larvae had alfalfa in western Oregon is M. communis (J.C.M.), been collected. This pattern was altered in the case unpublished data; Coop 1987). Known hosts for M. of damp foliage (the number of sweeps per transect communis include the noctuids Amphipyra py- reduced to 10) or if the density of noctuid larvae ramidoides Guenee, Dargida procincta (Grote), was extremely low (number of sweeps increased to Eupsilia sedus (Guenee), Lithophane bethunei 500). Weekly sampling was initiated in early July (Grote and Robinson), L. laticineae Grote, Ortho- during 1984 and mid-June during 1985, when lar- sia himsci (Guenee), O. revicta (Morrison), Pseu- vae were present in both crops, and was continued daletia unipuncta (Howorth), the Iymantriid Ma- until peppermint was sprayed or harvested (about lacosoma americanum (F.), and the tortricid 10 August) or until the final cut in alfalfa (about Grapholitha sp. (Krombein et al. 1979). We found 15 September). Samples were taken on a monthly 110 published information on the biology of M. basis during the winter of 1984-1985 on alfalfa communis. Here we report the patterns of host regrowth. None of the peppermint fields had suf- utilization by M. communis on peppermint and ficient winter growth to facilitate sampling. alfalfa in the Willamette Valley of Oregon. Larvae were placed in plastic cups (473 m\) while in the field; they were returned to the laboratory, where individual larvae were placed into 30-ml Materials and Methods cups, provisioned with cubes of artificial diet We monitored the relative abundance of im- (BioServ #928L2, Cabbage Looper Diet; French- mature Lepidoptera by species and age class in town, N.].) and sealed with paper insert lids. Diet alfalfa and peppermint to address some of the fac- was changed at least every 2 d. The species and tors likely to influence the relative abundance of instar of each noctuid were recorded at time of M. communis throughout the growing season. Study collection. Larvae were reared in the laboratory at

0046-225x/89/0537-0540$02.00/0 © 1989 EntomologicalSocietyof America 538 ENVIRONMENTAL ENTOMOLOGY Vol. 18, no. 3

Table 1. Numbers and species of noctuid larvae col- 50 lected in peppermint and alfalfa and parasitism of those species by M. communis Autographa caLifornica

M. 25 Species Crop· No. com- munisb Autographa californica a, m 1,997 + 0 Peridroma saucia a, m 1,203 + 50 Mamestra configurata a, m 853 til Peridroma saucia Dargida procincta a, m 564 + l:l. II) Agrotis ipsilon a 312 + II) Xestia adda a, m 92 ~ 25 Amphipyra pyramidoides a 48 + U) Leucanla farcta a 44 0 Downloaded from https://academic.oup.com/ee/article/18/3/537/2393821 by guest on 29 September 2021 Trichoplusia nl a, m 40 ....•0 Xylena nupara a,m 23 + 0 M Pseudaletia unlpuncta a 10 + II) 50 Heltothis phloxiphoga a, m 8 l:l. Dargida procincta Hello/his zea a 1 M 1 II) Scotogramma trifoltl a .a 25 Spodoptera praefica a 1 § • a, alfalfa; m, peppermint. Z b +, parasitized; -, not parasitized. 0 50 Meteorus communis a constant temperature of 24 (±2)OC and observed daily for parasitoid emergence. The time it took 25 M. communis to exit from hosts in the laboratory was correlated with the host instar at the time of collection by regression analysis. Cadavers of hosts o from which M. communis had exited were dis- 6/09 7/19 8/28 10/07 sected within 12 h for evidence of superparasitism. Date Fig. 1. Seasonal abundance (per 100 sweeps) of M. Results and Discussion communis and three of its principal hosts in peppermint and alfalfa during 1985; *, peppermint; 0, alfalfa. A total of 5,197 noctuid larvae representing 15 different species were collected (Table 1). Seven species were parasitized by M. communis. Para- M. communis from winter collections of D. pro- sitism of Agrotis ipsilon (Hufnagel) (35.3%), Peri- cincta (10.8%) and A. califomica (6.0%) in alfalfa. drama saucia (Hubner) (6.9%), and Autographa Of those species not parasitized, only Mamestra califomica (Speyer) (3.8%) are new host records. configurata Walker was collected in sufficient Also parasitized were P. unipuncta (20%), D. pro- numbers to suggest that it is not a suitable host cincta (15%), A. pyramidoides (10%), and Xylena species. It is possible that parasitism of some of the nupara Lintner (5%). Additionally, we recovered other species may occur, but insufficient numbers of larvae were collected during this study. For ex- Table 2. M. communis parasitism of field-collected ample, Trichoplusia ni (Hiibner) was readily ac- noctuid larvae collected from peppermint and alfalfa in cepted and suitable for M. communis in laboratory western Oregon during 1984-1985. tests (West 1988). Larvae of three noctuid species were parasitized Alfalfa Peppermint by M. communis in peppermint and alfalfa (Table Host Yr % % 2). Most often, M. communis was associated. with No· Para- No." Para- P. saucia, D. procincta, and A. califomica in alfalfa sitizedb sitizedb and peppermint. The most frequently parasitized Peridroma 1984 25 8 330 2 species, A. ipsilon, was extremely rare in mint. saucia 1985 310 12 167 6 The relative abundance of M. communis and Dargida 1984 104 12 17 0 the three hosts common to both crops is given in procincta 1985 256 18 44 5 Fig. 1. The peak in host abundance during July Agrotis 1984 286 36 1 0 was caused by the presence of P. saucia, A. cali- ipsilon 1985 15 27 1 0 fomica, and D. procincta in alfalfa and P. saucia Autographa 1984 317 3 226 3 and A. califomica in peppermint. The increase in californica 1985 423 6 380 2 abundance of hosts was followed by a similar in- crease in M. communis-parasitized hosts in alfalfa, • Susceptible stages only, second to fifth instars (second to fourth of A. californica). whereas in peppermint no such response was ob- b Determined by rearing. served. Junp 1989 WEST & MILLER: HOST EXPLOITATION BY M. communis 539 Downloaded from https://academic.oup.com/ee/article/18/3/537/2393821 by guest on 29 September 2021

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4 . '[ r ..'ll. . J: . . ...,.:. . .·hl.:. . : ··l· .'.".~'..' .' ~: . o Days :3 4 5 B Instar Collected Fig. 2. Rplationship between the number of days required for M. communis to exit from a host after eo\lpclion and the instar of that host at the time of collection (least-squares regression T' = 0.22, Y = 12.97 - 1.545x, n = 1(6).

In any analysis of host-parasitoid relationship, it year. We did not detect superparasitism in field- is important to determine the host stages used by collected hosts (based on host dissections, n = 110). the parasitoid for development. From a combi- No hyperparasitoids were reared from M. com- nation of field and laboratory data (Fig. 2), it is munis emerging in the lab (n > 500), but two field- possible to predict the instar in which M. com- collected M. communis pupae yielded a species of /IIullis oviposits in the field. All but the first and Mesochorinae (), indicating that last instars of each noctuid species received eggs hyperparasitism occurs in the pupal stagt'. from M. communis. The length of time required A larger percentage of susceptible hosts (second for M. communis to exit hosts (D. procincta, A. through penultimate instars) were parasitized in califomica, and P. saucia) in the laboratory was alfalfa than in peppermint (all species pooled, Ta- longt'f for those exiting younger hosts, presumably ble 2). Overall parasitism was higher in alfalfa. Ten because they had more recently received M. com- hosts parasitized by M. communis were collected TlItmis eggs. The number of days required for M. per 1,000 sweeps in alfalfa versus only two per C011l/ll1mis to exit (at 24 ± 2°C) after field collection 1,000 sweeps in peppermint. This difference was \Vas(i ± SD) 10.2 ± 1.8, n = 19 for second instars; consistent both years and may be a result of du- H.l ± 2.2, n = 42 for third instars; and 6.8 ± 2.4, rational stability of the habitat, host availability in It = 44 for fourth instars (M. communis exited from the agroecosystems, nutritional effects on devel- 2 of 105 fifth instars, which required 4 and 8 d). opment and allelochemical effects on behavior This compares to the 10-d developmental time we caused by the host plants, or differences in the use observed in nur laboratory culture (oviposition to of . exit, reared on third instars of P. saucia). These The influence of each of these variables requires data suggest that hosts are typically parasitized as experimental examination but may be considered second instars in the field. as follows. Peppermint foliage is present only part Neither host species nor instar had an effect on of the year (June to early August), whereas ~Ifalfa the sex ratio of M. communis (1:1) throughout the foliage is present throughout the year. Although A. 540 ENVIRONMENT AL ENTOMOLOGY Vol. 18, no. 3 ipsilon, A. pyramidoides, and P. unipuncta did References Cited not occur in peppermint, proportionately more lar- Askari, A., J. W. Mertins & H. C. Coppel. 1977. De- vae of D. procincta and P. saucia were parasitized velopmental biology and immature stages of Meteo- when they occurred in alfalfa than when they oc- rus pulchricornis in the laboratory. Ann. Entomo!. curred in peppermint (Table 2). There was no dif- Soc. Am. 70: 655-659. ference in the ability of M. communis to develop Baldur, W. V. 1968. Bionomic notes on the hexapo- in hosts fed peppermint versus alfalfa in the lab- dous parasites of Acrobasis rubrifasciella. Ann. Ento- oratory (West 1988). Also, development times for mo!. Soc. Am. 61: 463-476. M. communis reared from P. saucia fed mint fo- Berry, R. E., S. J. Yu & L. C. Terriere. 1980. Influ- liage (Coop & Berry 1986) are not significantly ence of host plants on metabolism and management of variegated cutworm. Econ. Ento- different from times observed with an alfalfa or J. mo!. 73: 771-774. artificial diet (West 1988). Insecticides were used

Coop, L. B. 1987. Management of variegated cut- Downloaded from https://academic.oup.com/ee/article/18/3/537/2393821 by guest on 29 September 2021 on all but one peppermint field, whereas no alfalfa worm in peppermint. Ph.D. dissertation, Oregon State fields were treated. University, Corvallis. The results of our investigation demonstrated Coop, L. B. & R. E. Berry. 1986. Reduction in var- that M. communis is abundant throughout the iegated cutworm (Lepidoptera: ) injury to growing season; is more common in alfalfa than in peppermint by larval parasitoids. J. Econ. Entomo!. peppermint; and is capable of exploiting a large 79: 1244-1248. number of noctuid hosts, some of economic im- Krombein, K. V., P. D. Hurd, Jr., D. R. Smith & B. portance. Further investigations should address the D. Burks. 1979. Catalog of Hymenoptera in Amer- ica north of Mexico, vo!. 1. Smithsonian Institution role of cultural practices and the impact of insec- Press, Washington, D.C. ticide treatments to determine further what factors Shaw, M. R. 1981. Parasitism by Hymenoptera of influence parasitism by M. communis in these hab- larvae of the white admiral butterfly, Ladoga camilla itats. Data from these studies will be useful in in- (L.), in England. Eco!' Entomo!. 6: 333-335. vestigations on the structure and dynamics of para- Simmonds, F. J. 1947. The biology of the parasites sitoid guilds. of Loxostege sticticalis L. in North America-Me- teorus loxostegei Vier. (Braconidae: Meteorinae). Bull. Entomo!. Res. 38: 373-379. Acknowledgment West, K. J. 1988. The biology and host relations of Meteorus communis (Cresson). M.S. thesis, Oregon Paul Marsh identified M. communis and Paul Ham- State University, Corvallis. mond identified the noctuid . We thank V. Brookes and R. Berry for their comments, suggestions, and critical Received for publication 29 March 1988; accepted 12 reviews. This is Oregon State Agricultural Experiment September 1988. Station Technical Paper 8682.