BIOLOGICAL CONTROL Laboratory Assessment of the Effects of Bacillus thuringiensis on Native

1 2 3 3 JOHN W. PEACOCK, DALE F. SCHWEITZER, JANEL. CARTER, AND NORMAND R. DUBOIS

Environ. Entomol. 27(2): 450-457 (1998) ABSTRACT The effect of 2 formulations of Bacillus thuringiensis subsp. kurstaki (Foray 48B and Dipel 8AF) was evaluated on 42 species of native Lepidoptera in laboratory bioassays using instars that are present in the field at the time of gypsy suppression applications. Mortality was significant for 27 of the 42 species evaluated against Foray 48B, and 8 of 14 species evaluated against Dipel 8AF. Susceptible species were noted in 5 of 6 families assayed-Papilionidae, Nymphalidae, Geometridae, Lasiocampidae, Satumiidae, and . The 1 species treated in the Lymantriidae family was not susceptible to B. thuringensis. Treated individuals that survived for a week were likely to reach adulthood. Intrageneric differences in susceptibility to B. thuringiensis were recorded among 8 species of and 3 species of assayed. Of the 18 species assayed as 1st or 2nd instars, mortality was significant, usually exceeding 95%. By contrast, 9 of 11 species not susceptible to B. thuringiensis were assayed as penultimate or ultimate instars. However, species susceptible to B. thuringiensis were found in both early and late instars.

KEY WORDS Bacillus thuringiensis, nontarget Lepidoptera, microbial control

Bacillus thuringiensis subsp. kurstaki is one of the in­ ication program. Sample et al. (1996) also reported a secticides most commonly used to control forest significant reduction in species abundance and rich­ pests. B. thuringiensis has been used extensively in ness in both larval and adult nontarget Lepidoptera suppression and eradication programs against gypsy from field studies in eastern West Virginia. In west­ moth, Lymantria dispar (L.). From 1980 to 1993, > 1.9 central Virginia, Wagner et al. (1996) found that the million ha (4.9 million acres) were sprayed with B. abundance of micro- and macrolepidopteran caterpil­ thuringiensis. In 1994, > 151,000 ha were treated with lars was consistently lower in treated plots. However, B. thuringiensis, including >57,000 ha in and the differences were modest, and nearly every lepi­ 20,000 ha in both Pennsylvania and Maryland. In 1994, dopteran species recovered within a year of treat­ B. thuringiensis was used on >55% of the area sprayed ment. James et al. (1993) demonstrated that B. thu­ for gypsy moth suppression (Anonymous 1994). Be­ ringiensis is toxic to late, but not early, instars of the sides suppression activities aimed at the European cinnabar moth, Tyria jacobaeae (L.), an introduced gypsy moth, eradication efforts for both the European biocontrol agent. and Asian strain of the gypsy moth also occur yearly. Significant reductions in numbers of larvae of non­ In 1992, efforts to eradicate the Asian gypsy moth in target lepidopterans following the use of B. thurin­ certain locations in Oregon, Washington, and the Van­ giensis can indirectly affect other that rely on couver area ofBritish Columbia resulted in >67,000 ha lepidopterous larvae as a primary source of food. Ro­ being sprayed with B. thuringiensis (Anonymous denhouse and Holmes (1992) found that a significant 1992). In 1994, >56,000 ha were sprayed near Wil­ reduction in biomass oflarvae following B. thuringien­ mington, NC, in attempts to eradicate the Asian gypsy sis application led to significantly fewer nesting at­ moth (McGovern 1994). tempts by certain birds. Bellocq et al. (1992) showed Although there is a growing concern over the non­ that the use of B. thuringiensis increased emigration target effects of B. thuringiensis, there has been limited rates and caused dietary shifts in shrews in treated research aimed at determining its effects on native areas. North American species of Lepidoptera. In Oregon, Krieg and Langenbruch (1981) presented an exten­ Miller (1990) demonstrated that both richness and sive summary of the toxicity of B. thuringiensis to diversity of native Lepidoptera associated with Garry Lepidoptera, which suggests extremely broad impacts , Quercus garryana Dougl., were reduced following within the order. Navon (1993) reported on bioassay treatment of oak habitat as part of a gypsy moth erad- techniques and results for various agricultural and forest lepidopteran pests. Faust and Bulla (1982) re­ 1 U.S. Forest Service, retired. Current address: 185 Benzler Lust ported on the susceptibility of 57 lepidopteran species, Road, Marion, OH 43302. most of which also were agricultural and forest pests. z The Nature Conservancy, 1761 Main Street, Port Norris, NJ 08349. Again, broad taxonomic effects were evident within 3 U.S. Forest Service, Northeastern Forest Experiment Station, Northeastern Center for Forest Health Research, 51 Mill Pond Road, the order. Most bioassays have been conducted to Hamden, Cf 06514. determine the susceptibility of B. thuringiensis to lepi-

F.hrh F.hrh ~t>rntinn ~t>rntinn hPfnrP hPfnrP hnt hnt wf'm wf'm rt~~r~y rt~~r~y l as as chambers chambers to to an- ~witPhecl ~witPhecl bv bv 16 16 bv bv 24 24 (9 (9 em) em) before before the the add ition ition

Several Several polyphagous polyphagous species species were were reared reared on on maintain maintain Prunus freshness freshness and and turgidity , , then then placed placed in in Plexi-

Celtis Celtis occidentalis, occidentalis, and and Speyeria Speyeria diana diana on on Viola Viola alba alba sp. sp. L. L. Th e e bouquets bouquets were were placed placed in in tubes tubes of of water water to to

Asterocampa Asterocampa clyton clyton (Boisduval (Boisduval L e conte) conte) on on & & ( potted potted Mill) Mill) Koch., Koch., K. K. Junip erus erus virginiana virginiana and and L., L., Quercus Quercus

Hemileucamaia Hemileucamaia (Drury) (Drury) (2nd (2nd only) only) on on potted potted were were Q. Q. alba; alba; used used for for assays assays in in 1991 1991 involving involving Carya ovata

bouquets; bouquets; Malacosoma Malacosoma disstria disstria (2nd (2nd only) only) and and Freshly Freshly collected collected foliage foliage bouquets bouquets from from native native trees trees

Walker, Walker, and and Antheraea Antheraea polphemus polphemus on on Quercus Quercus not not alba alba sprayed sprayed with with innocuous innocuous material. material.

Vaccinium Vaccinium coymbosum coymbosum cultivar; cultivar; resumens autoclaved autoclaved sample, sample, foliage foliage to to be be used used as as controls controls was was

type type capax capax (Grote), (Grote), andAbagrotis andAbagrotis altemata altemata on on a a tality tality potted potted in in any any of of the the species species assayed assayed against against the the

vinulenta, vinulenta, semitaria Franclemont, Franclemont, ternans ternans Xylo­ (Walker). (Walker). Because Because there there was was no no larval larval mor ­

potted potted Aronia Aronia arbutifolia; arbutifolia; sordida sordida C. C. Grote, Grote, ris), ris), Eupsilia Amphipyra pyramidoides pyramidoides Guenee, Guenee, and and Egira Egira al ­

bouquets; bouquets; praeclara praeclara C. C. Grote Grote Robinson Robinson & & on on mostly mostly Hemileuca Hemileuca M. M. maia , , disstria, disstria, Alsophila Alsophila pometaria pometaria (Har­

Strecker, Strecker, and and vidua vidua (J. (J. C. C. E. E. Smith) Smith) on on Carya Carya BIU/ha BIU/ha ovata ovata to to suitab l e e host host foliage foliage and and bioassayed bioassayed against against

ted ted Acer Acer rubrum; rubrum; Actias Actias luna , , Catocala Catocala Foray Foray obscura obscura 48B 48B was was applied applied at at a a rate rate equivalent equivalent to to 89 89

grotei grotei Riley, Riley, and and Sunira Sunira bicolorago bicolorago (Guenee) (Guenee) on on (N.R.D, (N.R.D, pot­ unpublished unpublished data) data) An An autoclaved autoclaved sample sample of of

magnaria magnaria

Guenee, Guenee,

Eutralepa Eutralepa clemataria, clemataria, Lithophane Lithophane ber ber of of drops/ drops/ em genera lly lly ranges ranges from from 60 60 to to 300 300

2 2

Benjamin) Benjamin) on on Juniperus Juniperus virginian virginian a a bouquets; bouquets; Ennomos tribution. tribution. For For this this spray spray procedure, procedure, the the average average num­

Grote Grote on on potted potted Prunus Prunus serotina; serotina; L. L. lemmeri lemmeri seedling seedling (B (B or or bouquet bouquet & & to to monitor monitor droplet droplet size size and and dis­

(Grote), (Grote), Lithophane Lithophane unirrwda unirrwda (Lint) , , and and L. L. petulca petulca cards cards (Mcac.l. (Mcac.l. Dayton , , OH) OH) were were placed placed next next to to each each

Euchlaena obtusaria obtusaria (Hubner), (Hubner), Eupsilia Eupsilia vinulenta vinulenta times times during during the the ]-min ]-min spray spray period . . Kromecote Kromecote spray spray

are are as as follows: follows: Papilio Papilio glaucus, glaucus, Limenitis Limenitis arthemis arthemis lings lings (F.), (F.), were were placed placed on on a a rotating rotating table table that that revo l ved ved 4 4

oaks (usually (usually Quercus Quercus rubra) rubra) and and the the food food plants plants terial terial used used to to be be treated. treated. Foliage Foliage bouquets bouquets or or potted potted seed­

gypsy gypsy moth moth control. control. Species Species not not assayed assayed on on potted potted red red the the tower tower a t t a a h eight eight of of = 3.5 3.5 m m above above the the plant plant ma­

often often dominant dominant or or codominant codominant in in forests forests sprayed sprayed for for 4 m m 4 high, high, with with th e e nozzle nozzle positioned positioned in in the the center center of of

oaks oaks were were usually usually assayed assayed on on oaks oaks because because they they are are Lewis Lewis 19 7 3) . . The The tower tower was was =2 .5 .5 min min diameter diameter and and

in in the the field. field. Polyphagous Polyphagous species species that that commonly commonly units units use use VMD VMD in in a a cy lindri ca l l spray spray tower tower (Hubbard (Hubbard and and

genus in in th e e case case of of oak oak feeders) feeders) that that is is a a known known PA) PA) host host rotated rotated at at 12 , 000 000 HPM HPM to to generate generate drops drops at at 75-125 75-125

Larvae Larvae were were assayed assayed on on a a plant plant species species (or (or at at l eas t t with with a a mini-Beecomist mini-Beecomist nozzle nozzle (Beecomist, (Beecomist, Telford, Telford,

obtained obtained from from D.F.S. D.F.S. The The pesticide pesticide was was app li ed ed using using a a pro-mist pro-mist system system

in stars. stars. Details Details of!arval of!arval collections collections and and rearing rearing can can be be cidal cidal activity activity of of the the Foray Foray and and Dipel Dipel preparations. preparations.

soma soma disstria disstria Hubner Hubner were were collected collected as as wild wild third third foliage foliage on on eac h h bioassay bioassay date date confirmed confirmed the the insecti ­

were were collected collected as as wild wild l ate ate instars; instars; the the 1993 1993 Malaco­ and and 4th-instar 4th-instar gypsy gypsy moth moth on on treated treated and and untreated untreated

treatment treatment lots . . Most Most of of the the Abagrotis Abagrotis alternata alternata with with (Grote) (Grote) different different batches batches of of Foray. Foray. Bioassays Bioassays with with 2nd­

offspring offspring were were simi larly larly represented represented in in the the control control says says and and for for a a species species were were conducted conducted in in different different years years

of of a a sing le le female If If . . progeny progeny of of 2 2 females females were were species species used, used, also also were were assayed assayed against against Dipel. Dipel. Multiple Multiple as ­

New New Jersey. Jersey. In In most most assays, assays, larva e e were were th e e progeny progeny species species were were assayed assayed against against Foray, Foray, and and 14 14 of of the the 42 42

that that were were t aken aken at at bait bait or or light light most most often often in in southern southern (Foray) (Foray) or or 99 99 (Dipcl) (Dipcl) BIU/ha BIU/ha in in field field programs. programs. All All

collected collected ova ova or or from from ova ova obtained obtained from from wild wild quets quets females females at at rates rates equiva l ent ent to to the the aerial aerial application application of of 89 89

the the species species that that we we evaluated evaluated were were reared reared from from sis sis field­ subsp. subsp. kurstaki kurstaki t o o potted potted seedlings seedlings or or foliage foliage bou­

Larval Larval Collections Collections and and Rearing. Rearing. Larvae Larvae of of most most of of Foray Foray 48B 48B or or Dipcl8AF Dipcl8AF formulations formulations of of B. B. thuringien­

escape escape impact impact from from B. B. thuringiensis thuringiensis app lication s. s. treatments treatments co nsisted nsisted of of undiluted undiluted applications applications of of

larva e e of of these these species species usually usually hatch hatch late late enough enough to to Treatment Treatment of of Foliage, Foliage, Bioassay Bioassay Procedure. Procedure. Foliage Foliage

add add taxonomic taxonomic diversity, diversity, a lthou gh gh some some spring spring says brood brood . .

Smith), Smith), and and Papilio Papilio glaucus glaucus (L.) (L.) also also were were assayed assayed thetic thetic to to diet diet (BioScrv, (BioScrv, Frenchtown, Frenchtown, NJ) NJ) before before the the as­

polyphemus polyphemus (Cramer), (Cramer), Eutrapela Eutrapela clemataria clemataria iment iment E. E. (J. (J. Station Station at at Hamden, Hamden, CT, CT, were were reared reared on on syn­

to to control control gypsy gypsy moth. moth. Actias Actias luna luna (L.), (L.), Antheraea Antheraea cu lture lture maintained maintained at at th e e Northeastern Northeastern Forest Forest Exper­

present present as as larva e e at at the the time time B. B. thuringiensis thuringiensis is is applied applied Gypsy Gypsy moth moth larva e e from from the the continuous continuous l aboratory aboratory

patric patric with with gypsy gypsy moths and, and, more more importantly, importantly, were were are are observed observed at at the the rearing rearing site. site.

and and 26 26 noctuids. noctuids. Most Most of of these these are are forest forest species species s l eeves. eeves. sym­ For For most most species, species, wild wild larvae larvae of of similar similar age age

geometrids, geometrids, 1 1 lasiocampid, lasiocampid, 3 3 saturniids , , 1 1 l ymantriid, ymantriid, 2nd 2nd instars , , l arvae arvae wer e e reared reared outdoors outdoors in in muslin muslin

These These included included 1 1 papillionid papillionid species, species, 3 3 nymphalids mal mal eclosion eclosion , , 7 7 times. times. Except Except for for species species assayed assayed as as 1st 1st or or

Lepidoptera Lepidoptera were were evaluated evaluated in in the the laboratory laboratory ris, ris, assays. assays. Cumberland Cumberland County, County, New New Jersey, Jersey, to to ensure ensure nor­

Selection Selection of of Species. Species. Forty-two Forty-two species species of of native native outdoors outdoors in in a a screened, screened, shaded shaded enclosure enclosure at at Port Port Nor­

priate priate instars instars for for assay, assay, all all adults adults and and eggs eggs were were h e

ld ld

Materials Materials

and and Methods Methods

applications applications typically typically are are scheduled. scheduled. To To obtain obtain appro­

occur occur in in the the field field at at the the time time when when B. B. thuringiensis thuringiensis

target target Lepidoptera. Lepidoptera. Larvae Larvae were were bioassayed bioassayed in in an an instar instar in in which which they they

th e e ef fect fect of of B. B. thuringiensis thuringiensis on on larva e e of of nativ e, e, non­ species. species.

results results of of labor atory atory bioassays bioassays aimed aimed at at determining determining gous gous species species were were reared reared and and assayed assayed on on the the same same

nontarget nontarget species species in in the the laboratory . . We We report report the the phipyra, phipyra, and and Alsophila. Alsophila. Oligophagous Oligophagous and and monopha­

evaluate evaluate the the toxicity toxicity of of B. B. thuringiensis thuringiensis against against all all native, native, , Sericaglaea, , Chaetaglaea, Eupsilia, Eupsilia, , Orthosia, Am ­

dopterous dopterous pests, pests, but but there there have have been been few few attempts attempts other other to to host host (usually (usually oak) oak) for for the the assays . . These These included included

Aprill998 Aprill998 PEACOCK PEACOCK ET ET B. B. AL.: AL.: thuringiensis thuringiensis AssESSMENT AssESSMENT OF OF ON ON NATIVE NATIVE 451 451 LEPIDOPTERA LEPIDOPTERA

this this that that indicates indicates This This replicates. replicates. ll ll a from from some some ing ing other other in in rates rates Mortality Mortality species. species. butterfly butterfly 4 4 and and umiid umiid

includ­ s, s, lt adu as as eclose eclose did did arvae arvae l treated treated (54.5%) (54.5%) 22 22 sat­ the the for for 100% 100% - 90 was was e e larva treated treated of of Mortality Mortality

of of 12 12 However, However, analysis. analysis. l l statistica formal formal precluding precluding 1). 1). (Table (Table Lymantriidae Lymantriidae lone lone the the not not but but Noctuidae Noctuidae 26 26

replicates, replicates, treatment treatment between between differed differed species species this this for for of of 14 14 and and Satumiidae, Satumiidae, all3 all3 Lasiocampidae, Lasiocampidae, single single the the

Mortality Mortality . . ]. ]. virginiana on on instars instars second second as as lemmeri lemmeri Geometridae, Geometridae, 7 7 of of 5 5 Nymphalidae, Nymphalidae, 3 3 all all Papilionidae, Papilionidae,

L. L. assayed assayed also also We We L. L. unaffected. unaffected. were were unimoda unimoda of of the the included included This This species. species. (64.3%) (64.3%) 42 42 of of 27 27 for for corded corded

stars stars in penultimate penultimate However, However, 7). 7). day day (by (by petulca petulca of of L. L. re­ was was mortality mortality Significant Significant Assays. Assays. 48B 48B Foray Foray

stars stars in in 2nd 2nd for for and and grotei grotei of of L. L. s s instar antepenultimate antepenultimate

Results Results

for for significant significant was was Mortality Mortality 4). 4). (Table (Table evaluated evaluated cies cies

spe­ 3 3 Lithophane Lithophane the the for for recorded recorded were were differences differences

intrageneric intrageneric Catocala, Catocala, the the with with case case the the was was As As . . pupae treatment treatment for for 75% 75% least least at at was was success success sion sion

Foray. Foray. to to susceptible susceptible not not eclo­ (3) (3) and and feeding, feeding, completed completed survivors survivors treatment treatment

e e wer apparently apparently stars) stars) in in ate ate l as as (evaluated (evaluated Edwards Edwards these these of of 90% 90% east east l at at (2) (2) 7, 7, day day on on alive alive were were arvae arvae l

H. H. W. W. similis similis and and midinstars) midinstars) as as C. C. (evaluated (evaluated sordida sordida control control and and treatment treatment the the of of 90% 90% east east l At At 1) 1) ( ( ria: ria:

affected. affected. ly ly C. C. significant not not were were instars instars st st la whereas whereas crite­ following following the the met met species species Insensitive Insensitive pupation. pupation.

7), 7), day day (by (by Foray Foray to to e e usceptibl s were were instars instars ultimate ultimate by by or or 7 7 or or 5 5 days days by by controls controls in in mortality mortality 5% 5% than than

pen­ praeclara, praeclara, In In instars. instars. y y l ear in in C. C. greatest greatest mortality mortality more more no no and and arvae arvae l treated treated in in mortality mortality 95% 95% least least at at

with with , , foliage thuringiensis-treated thuringiensis-treated B. B. consuming consuming after after was was there there criteria: criteria: following following the the met met sensitive sensitive highly highly

mortality mortality significant significant showed showed groups groups age age both both in in e e larva as as considered considered Species Species thuringiensis. thuringiensis. to to B. B. insensitive insensitive

ilia ilia . . thuringiensis C. C. B. B. of of applications applications field field during during or or sensitive sensitive highly highly as as species species categorized categorized also also We We

only only instar instar ate ate l as as present present be be would would it it though though < < even even 0.05. 0.05. if if P P thuringiensis) thuringiensis) to to B. B. e e l susceptib

instars, instars, (3rd/4th) (3rd/4th) mid mid and and (1st/2nd) (1st/2nd) ly ly ear both both as as ated ated thus thus (and (and different different significantly significantly considered considered was was vae vae

evalu­ was was ilia ilia praeclara. praeclara. and and C. C. Grote, Grote, coccinata coccinata ilia, ilia, ­ lar l l contro and and treatment treatment between between Mortality Mortality foliage. foliage.

instars­ ate ate l as as assayed assayed were were that that others others 3 3 and and - Grate control control versus versus treated treated on on species) species) some some for for pupation pupation

la la eel lin and and (Cramer), (Cramer), ilia ilia vidua, vidua, Strecker, Strecker, obscura obscura or or 7 7 (day (day 5 5 day day at at mortality mortality the the compare compare to to used used

instars­ 2nd 2nd and and 1st 1st as as assayed assayed species species 4 4 ll ll a included included was was 1958) 1958) (Fisher (Fisher test test exact exact 's 's Fisher . . Analysis Data Data

These These 4). 4). (Table (Table evaluated evaluated were were that that Catocala Catocala genus genus species. species. the the of of subset subset

noctuid noctuid e e th of of species species 8 8 of of 6 6 in in significant significant was was tality tality a a for for recorded recorded were were lso lso a weights weights dry dry lt lt adu or or weights weights

Mor­ Mortality. Mortality. Larval Larval in in Difierences Difierences lntrageneric lntrageneric pupal pupal live live and and mortality, mortality, pupal pupal pupation, pupation, to to Days Days

. . 4) and and 2 2 (Table (Table results results similar similar recorded. recorded. were were eclosion eclosion adult adult of of Dates Dates NJ. NJ. Norris, Norris, Port Port

very very giving giving always always instar instar same same the the in in once once than than more more in in location location sheltered sheltered a a in in ground ground the the on on outdoors outdoors kept kept

assayed assayed were were species species Eight Eight adulthood. adulthood. reached reached and and were were pupae pupae overwintering overwintering with with Containers Containers gence. gence.

recovered recovered that that e e larva some some had had species species affected affected cantly cantly emer­ lt lt adu until until conditions conditions ambient ambient under under held held were were

signifi­ the the of of 14 14 larvae, larvae, treated treated the the among among mortality mortality Pupae Pupae sleeves. sleeves. rearing rearing their their in in cocoons cocoons spin spin to to allowed allowed

substantial substantial Despite Despite 3). 3). (Table (Table stages stages pupal pupal and and pupal pupal were were or or cocoons, cocoons, spin spin to to which which in in leaves leaves dead dead and and

pre­ the the during during occurred occurred astyanax astyanax arthemis arthemis Limenitis Limenitis towel towel paper paper crumpled crumpled given given were were ground ground above above pate pate

in in mortality mortality and and pupation, pupation, through through 6 6 day day until until nificant nificant pu­ that that Species Species container. container. cardboard cardboard wax wax or or plastic plastic

sig­ not not was was mortality mortality a a in in others, others, 5 5 For For medium medium pupation pupation treatment. treatment. a a as as after after moss moss peat peat damp damp , , sterilized

d d 5-7 5-7 significant significant was was mortality mortality species, species, affected affected 27 27 heat­ of of em em 10 10 = with with supplied supplied were were soil soil the the enter enter

the the of of For21 For21 2). 2). (Table (Table e e larva treated treated among among mortality mortality would would normally normally that that ae ae larv aestivating aestivating or or Pupating Pupating

100% 100% had had which which species species with with along along mortality mortality no no fered fered weekly. weekly. least least at at observed observed was was

suf­ which which species species had had e e larva Noctuidae Noctuidae these these for for and and Mortality Mortality Geometridae Geometridae assay. assay. before before used used species species

the the both both example, example, For For variable. variable. highly highly were were families families host host the the of of plants plants unsprayed unsprayed wild-growing, wild-growing, on on eeves eeves l s

in in confined confined and and foliage foliage e e th from from removed removed were were vae vae

lar­ surviving surviving monitoring, monitoring, long-term long-term for for selected selected If If d. d.

5-7 5-7 of of period period instars. instars. a a ate ate l over over and and ly ly ear recorded recorded both both in in was was assayed assayed foliage foliage species species e e Som untreated untreated on on d d

< < arvae. arvae. l treated treated for for 0.05) 0.05) ty ty li (P (P morta Significant Significant c c and and treated treated on on mortality mortality Larval Larval Larvae. Larvae. Monitoring Monitoring

instar. instar. last last to to Fourth Fourth h h

species. species. certain certain for for e e larva of of availability availability

. . instar 3rd 3rd to to First First " "

limited limited the the of of because because varied varied replications replications of of number number

the the and and replication replication per per larvae larvae of of number number The The species. species. 10 10 8 8 1 25 25 18 18 42 42 Totald Totald

each each for for foliage foliage treated treated on on luna) luna) for for (10 (10 cations cations A. A.

18 18 6 6 8 8 8 8 ae ae id 26 26

Noctu

repli­ 2-6 2-6 and and (controls) (controls) foliage foliage untreated untreated on on tions tions

0 0 I I 1 1 e e Lymantriida

replica­ 1-4 1-4 were were There There sleeve. sleeve. or or chamber chamber Plexiglas Plexiglas 3 3 3 3 3 3 Satumiidae Satumiidae

1 1 1 1 1 1 1 1 I I Lasiocampidae Lasiocampidae h h eac in in foliage foliage untreated untreated or or treated treated to to confined confined e e larva

3 3 7 7 I I 4 4 4 4 Geometridae Geometridae

5-15 5-15 of of consisted consisted replication replication a a sleeve, sleeve,

a a in in placed placed were were

1 1 1 1 3 3

2 2 2 2 e e Nymphalida

which which of of 50-100 50-100 maia, maia, . . H gregarious gregarious the the for for Except Except

1 1 1 1 I I Papilionidae Papilionidae

foliage. foliage. treated treated to to thuringiensis thuringiensis of of B. B. application application after after

susceptiblec susceptiblec c c susceptible

assayed assayed

h h 1 1 within within

foliage foliage untreated untreated and and treated treated on on placed placed n n n n

No. No.

No. No.

species species Family Family

were were Larvae Larvae instars. instars. 1st 1st some some for for used used were were sleeves sleeves

No. No.

instar" instar" Early Early instarb instarb e e Lat

Muslin Muslin nylon. nylon. fine-mesh fine-mesh of of constructed constructed bags bags in in lings lings

seed­ potted potted of of foliage foliage on on confined confined were were larvae larvae assays, assays,

Lepidoptera Lepidoptera nontarget nontarget of of

most most For For foliage. foliage. ed ed untreat or or eated eated tr to to e e larva of of opecieo opecieo 42 42 for for fwnily, fwnily, by by 48B, 48B, Foray Foray to to Suoceptihility Suoceptihility Table Table 1. 1.

2 2 no. no. 27, 27, Vol. Vol. 452 452 ENTOMOLOGY ENTOMOLOGY ENVIRONMENTAL ENVIRONMENTAL

Prochoerodes Prochoerodes transversata transversata (Drury) (Drury) and and E. E. altemans ultimates ultimates , , survived survived to to adults adults but but none none of of 38 38 antepen-

succumbed succumbed within within the the first first 2-3 2-3 don don treated treated foliage; foliage; for for species. species. In In Lithophane Lithophane grotei, grotei, 35% 35% of of the the treated treated pen­

1st 1st to to 3rd 3rd instar instar with with Foray Foray (Table (Table 1). 1). Most Most ences ences species species in in instar instar sensitivity sensitivity were were noted noted in in 2 2 of of these these

ing ing treatment treatment for for all all of of the the 18 18 species species evaluated evaluated in in the the assayed assayed separately separately in in more more than than 1 1 instar, instar, and and differ­

Instar Instar Difference. Difference. Mortality Mortality was was significant significant follow­ Besides Besides C. C. ilia ilia C. C. and and praeclara, praeclara, 6 6 other other species species were were

apparently apparently

normal normal adults. adults.

1) 1) 0 0

larvae larvae alive alive on on day day 5 5 completed completed feeding feeding and and produced produced with with Foray, Foray, only only 10 10 were were significantly significantly affected affected (Table (Table

B. B. to to thuringiensis. thuringiensis. For For all all Lithophane Lithophane species, species, most most 3 ) . . Of Of the the 25 25 species species eva luated luated in in 4th 4th to to st st la instars instars

rare rare

species species is is ins ensi tiv e e or or only only moderately moderately sensitive sensitive mortality mortality was was not not significant significant until until after after day day 5 5 (Table (Table

Although Although this this species species did did e e not not meet meet our our hi gh ly ly sensitive sensitive cri t eria, eria, >75% >75% of of day day 5 5 survivors survivors died died before before pupation. pupation.

Significant Significant mortality mortality betw d d een een day day 6 6 through through pupation, pupation, but but not not by by day day 5. 5.

c c Highly Highly sensitive sensitive to to Foray Foray 48B. 48B.

No No P P value value reported reported for for species species b b for for which which no no mortality mortality occurred. occurred.

n, n, last last instar. instar. a a

not not

occur occur until until after after day day 6, 6, than than th e e counts counts given given are are a t t the the time time (usually (usually day day 7 7 or or pupation) pupation) si~nificanl si~nificanl mortality mortality occurred occurred (see (see footnoted). footnoted).

See See

Table Table 4 4 for for Catocala Catocala and and Lithophane Lithophane results. results. Counts Counts of of alive alive and and dead dead larvae larvae arc arc al al day day 5 5 or or th e e assays assays unless unless significant significant mortality mortality did did

Abagrotis Abagrotis altemata altemata n/n - 1 1 18 18 0 0 0 0 13 13

Abagrotis Abagrotis altemata altemata n/n - 1 1 29 29 0 0 50 50 0 0

Ortlwsia Ortlwsia hibisci hibisci n - 1 1 20 20 0 0 0 0 39 39

Orthosia Orthosia alurina alurina n - 1 1 18 18 0 0 30 30 7 7 0 . 082 3 3

Ortlwsia Ortlwsia alurina alurina n -2 -2 19 19 I I 20 20 0 0 0.9999 0.9999

Xylotype copax copax n - 1 1 19 19 I I 0 0 48 48 0.2941 0.2941

Sunira Sunira bicolorago bicolorago n n 20 20 0 0 0 0 29 29

Sunira Sunira bicolorago bicolorago n/n - 1 1 20 20 0 0 3 3 0.5498 0.5498 4.'; 4.';

Chaetaglaea sericea n - 1 1 19 19 0 0 4 8 8 l l 0.9999 0.9999

Chaetaglaea Chaetaglaea sericea sericea n - 1 1 20 20 0 0 0 0 20 20

Metaxaglaea Metaxaglaea semitaria semitaria n n 20 20 0 0 51 51 l l 0 . 9999 9999

Sericaglaea signata 4th 4th 18 18 0 0 48 48 0 0

Eupsilia Eupsilia vinulenta vinulenta n - l/n - 2 2 20 20 0 0 l l 4:3 4:3 0.9999 0.9999

Eupsilia Eupsilia vinulenta vinulenta n -l/n-2 -l/n-2 20 20 0 0 l l 19 19 0 . 9999 9999

Zale aeruginosa aeruginosa Hatchling Hatchling 1 2 2 0 0 1 9 9 ll ll 0 . 0175 0175

Egira Egira altemans altemans 2nd - 3rd 3rd 18 18 0 0 3 . 5 5 2 2 1.0000 1.0000

Egira Egira altemans altemans

lst lst 20 20 5 5 27 27 22 22 0.0059d 0.0059d

Psaphida Psaphida

resum

ens

1st - 2nd 2nd 20 20

0 0 41 41 :5 0.00001 0.00001 9'' 9''

0 0

Psaphida rolandi n - 1 1 19 19 I I 22 22 0.0001 0.0001 1 8' ' '

Xystopeplus Xystopeplus

rufago rufago 1st-2nd 1st-2nd 28 28 0 0 1 2 2 21 21 :5 0.00001 0.00001

Amphipyra Amphipyra pyramidoides pyramidoides n -1 -1 20 20 0 0 37 37 II II 0.0001 0.0001

Amphipyra Amphipyra

pyramidoides pyramidoides n - 1 1 19 19 2 2 6" 6" 24 24 :5 0.00001 0.00001

Noctuidae Noctuidae

Dasychira obliquata obliquata 4th 4th 20 20 0 0 Z7 Z7 0.9999 0.9999

Lymantriidae Lymantriidae

Actias Actias

luna luna lst lst 26 26 14 14 0 0 96 96 :5 0.00001 0.00001

Antheraea Antheraea

polyphemus polyphemus

lst lst 16 16 4 4 57 57 :5 0.00001 0.00001 .1' ' '

Hentileuca Hentileuca 2nd 2nd maia c c 109 109 I I 0 0 Ill Ill :5 0.00001 0.00001

Hemileuca Hemileuca ma-ia lst lst 20 20

0 0 0 0 51 51 s O.OOOOl O.OOOOl

0 0

Hemileuca Hemileuca

maia maia lst lst 70 70 I I 48 48 312 312 :5 0.001 0.001

Hemileuca Hemileuca 1naia Hatchling Hatchling c c 47 47 0 0 5 5 53 53 :5 0 . 00001 00001

Saturniidae Saturniidae

Malacosoma Malacosoma disstriac disstriac n n 20 20 0 0 44 44 :5 0.00001 0.00001

Malacosoma Malacosoma

disstria disstria 2nd 2nd 23 23 4 4 4" 4" 26 26 :5 0.00001 0.00001

Lasiocampidae Lasiocampidae

Prochoerodes Prochoerodes transversata transversata

2nd 2nd 19 19 I I 28 28 13 13 0.023 7d 7d

Eutrapela Eutrapela c c clemataria Hat ch lin g g 20 20 0 0 4 4 31 31 :5 0.00001 0.00001

Lambdina Lambdina fervidaritf fervidaritf lst lst l7 l7 l l 10 10 26 26 :5 0.00001 0.00001

Ennomos Ennomos ma g naria naria lst lst l7 l7 14 14 0 0 27 27 :5 0.00001 0.00001

Ennomos Ennomos

magnaritf magnaritf lst lst 22 22 l l 0 0 66 66 :5 0.00001 0.00001

Euchlaena Euchlaena obtusaria obtusaria n - 1 1 1 2 2 0 0 1 8 8 0 0

Phiglia Phiglia titea titea n /n-1 /n-1 20 20 0 0 43 43 7 7 0.1801 0.1801

Alsophila Alsophila pom e taria taria

n n

19 19 l l 7 7 0.0164 0.0164 ]] ]]

Geometridae Geometridae

Astercampa Astercampa

clyton" clyton" 4th - 5th 5th 21 21 l l l l 40 40 :5 0.00001 0.00001

Limenitis Limenitis arthemis arthemis

astyanax" astyanax" n /n-1 /n-1 10 10 0 0 0 0 2 0 0 < O.OOOOld O.OOOOld

Speyeria Speyeria

diana 2nd - 3rd 3rd 10 10

0 0 l l 15 15 :5 0.00001 0.00001

0 0

Nymphalidae Nymphalidae

Papilio Papilio

glaucus lst - 3rd 3rd 10 10

0 0 0 0 29 29 :5 0.00001 0.00001

0 0

Papilionidae Papilionidae

No . . a No. No. live live dead dead

No. No. alive alive No. No. dead dead

Species Species Ins Ins tar

a a

ph ph

Control Control Foray Foray

Table Table 2. 2. Mortality Mortality species species sub jected jected in in to to foliage foliage treated treated with with Foray Foray 488 488

Aprill998 Aprill998 PEACOCK PEACOCK ET ET B. B. AL.: AL.: thuringiensis thuringiensis AssESSMENT AssESSMENT oF oF ON ON NATIVE NATIVE LEPIDOPI'ERA LEPIDOPI'ERA 453 453

Significant Significant mortality mortality c c occurred occurred by by day day 7 7 but but not not by by day day 5. 5.

Highly Highly sensitive sensitive to to Foray Foray h h 48B. 48B.

l ast ast instar. instar. an , ,

are are at at the the tim e e (usually (usually day day 7 7 or or pupation) pupation) significant significant mortality mortality occurred occurred (see (see footnote c ). ).

Counts Counts

of of alive alive and and dead dead larvae larvae are are at at day day 5 5 of of the the assays assays unless unless significant significant mortality mortality did did not not occur occur until until after after day day 6, 6, then then the the coun ts ts given given

L. L. unimoda unimoda

n - 1 1 20 20 1 1 35 35 5 5 0.65 31 31

L. L. grotei grotei n - 1 1 20 20 0 0 9 9 ll ll 0 .0001 .0001

L. L. groteib groteib n - 2 2 20 20 0 0 0 0 38 38 :s: 0.00001 0.00001

Lithophone Lithophone petulca petulca

2nd 2nd

9 9 3 3 3 3 10 10 0.0169 c c

C. C.

lineella lineella

1st/2nd 1st/2nd 21 21 6 6 8 8 32 32 :s: O .OOOOl .OOOOl

C. C. simi/is simi/is

n-1/n - 2 2 15 15 3 3 46 46 2 2 0 . 1203 1203

C. C. praeclara praeclara

n n 18 18 0 0 16 16 3 3 0.2 299 299

C. C. praeclara praeclara

n - 1 1 16 16 0 0 5 5 ll ll :s: o.oooo1 c c

C. C. coccinatah coccinatah n - 1 1 18 18 0 0 5 5 45 45 :s: 0.00001 0.00001

C. C. sordida sordida

3rd/4th 3rd/4th 15 15 0 0 26 26 3 3 0.5401 0.5401

C. C. 3rd/4th 3rd/4th iliah iliah 20 20 0 0 2 2 39 39 :s:0.00001 :s:0.00001

C. C. ilia ilia

1st/2nd 1st/2nd 10 10 0 0 0 0 40 40 :s: 0.00001 0.00001

C. C. vidua vidua

2nd/3 rd rd 0 0 16 16 24 24 27 27 :s: O.OOOOl O.OOOOl

C. C.

vidua vidua

1st 1st 17 17 2 2 0 0 19 19 :s: 0.00001 0.00001

C. C.

vidua vidua

1st 1st 17 17 2 2 0 0 20 20 :s:0. 00001 00001

Catocala Catocala

obscura obscura 1st 1st 1 7 7 1 1 14 14 33 33 :s:O.OOOOI :s:O.OOOOI

No. No. aliv e e No. No.

dead dead No. No. alive alive No. No. dead dead

Species Species Ins Ins tara tara

p p

Control Control Foray Foray

Table Table 4. 4. lntrageneric lntrageneric comparisons comparisons of of mortality mortality in in response response to to Foray Foray 48B 48B for for Catocala Catocala and and Litllophane Litllophane (Noctuidac) (Noctuidac)

for for being being hi gh ly ly sensitive sensitive to to B. B. th e e thuringiensis thuringiensis late late for- instars instars (C. (C. ilia ilia sensitive sensitive as as early early and and lat e e in in stars). stars).

Species Species Sensitivity. Sensitivity. Thirteen Thirteen species species met met the the criteria criteria and and the the other other 6 6 were were highly highly sensitive sensitive as as mid-instars mid-instars or or

Eight Eight of of these these species species were were assayed assayed as as ear ly ly instars instars

Discussion Discussion

proved proved to to be be highly highly sensitive sensitive (Table (Table 2). 2).

larvae larvae survived survived to to pupation pupation and and the the treat ed ed last last in in stars stars

were were susceptible susceptible to to Dipel Dipel (Table (Table 5). 5). stria stria to to b e e highly highly sensitive sensitive because because only only 1 1 tr eated eated

treatment. treatment. Five Five of of 11 11 species species test ed ed as as lat er er instars instars tality. tality. However, However, we we would would consider consider 2nd-instar 2nd-instar M. M. dis­

the the ear ly ly instars instars against against Dipel Dipel succumbed succumbed following following sitive sitive b ecause ecause control control larva e e had had grea ter ter than than 5% 5% mor­

species. species. All All treated treated larva e e of of the the 3 3 species species evaluated evaluated in in disstria disstria did did not not meet meet the the criteria criteria of of being being highly highly sen­

mortality mortality was was only only 42% 42% through through adult adult e closion closion for for this this this this species species to to be be highly highly sensitive. sensitive. Second-instar Second-instar M. M.

psilia psilia vinulenta vinulenta was was sensitive sensitive to to Dipel Dipel only; only; although although ment ment replicates replicates with with some some survival, survival, we we s till till consider consider

these these species species showed showed similar similar results results with with Foray. Foray. Eu­ to to pupation pupation and and adult adult e closion . . Despite Despite the the 2 2 tre at­

cant cant during during the the post-treatment post-treatment period period (Table (Table 5); 5); 7 7 of of are are pooled pooled 9.0% 9.0% ( ( 48 48 of of 580) 580) of of tr ea ted ted larva e e survived survived

for for susceptibility susceptibility to to Dipel Dipel 8AF, 8AF, mortality mortality was was signifi­ by by 7 7 day in in 10 10 of of 12 12 replicates, replicates, although although when when all all data data

Dipei8AF Dipei8AF Assays. Assays. In In 8 8 of of 14 14 (57%) (57%) species species eva 2, 2, H. H. luated luated 4). 4). maia maia was was highly highly sensitive sensitive with with 100 % % mortality mortality

(2nd (2nd versus versus last last instar) instar) (Table (Table 2) . . only, only, day day 5), 5), and and Psaphida Psaphida resumens resumens (day (day 7) 7) (Tables (Tables

tepenultimates tepenultimates versus versus pen pen ultimates), ultimates), M. M. or or cocci cocci disstria disstria nata nata (pupation), (pupation), L. L. grotei grotei (antepenultimates (antepenultimates

H. H. maia maia (1st (1st versus versus 2nd 2nd instar), instar), Orthosia Orthosia alurina alurina (an­ instar, instar, day day H. H. 5), 5), maia maia (day (day 7), 7), C. C. ilia ilia (day (day 5), 5), C. C.

3). 3). There There were were no no apparent apparent differences differences in in ins ins tar s s for for (pupation), (pupation), E. E. clemataria clemataria (pupation), (pupation), disstria disstria M. M. (last (last

1st 1st instar instar than than in in the the 2nd 2nd and and 3rd 3rd instar instar (Tables (Tables trat 2 2 e and and d d by by Covell Covell (1984, (1984, f f pl. pl. 54: 54: 15) 15) as as athasaria) athasaria) L. L.

hi gher gher mortality mortality by by pupation pupation for for assayed assayed larvae (Hubner) (Hubner) in in th e e (the (the bivoltine-oak bivoltine-oak feeding feeding species species illu s­

in in 1993 1993 (Table (Table 4). 4). E. E. altemans altemans also also showed showed slightly slightly 5), 5), Ennomos Ennomos magnaria magnaria (day (day 5), 5), Lambdinafervidaria Lambdinafervidaria

All All 2 0 0 larva e e treated treated as as 1st 1st instars instars were were dead dead by by day day 5 5 arthemis arthemis astyanax astyanax ( ( e closion), closion), Astercampa Astercampa clyton clyton (day (day

adults adults compar ed ed with with only only 1 1 of20 of20 first first instars instars in in 1992. 1992. Papilio Papilio g laucus laucus (by (by day day 5), 5), Speyeria Speyeria diana diana (day (day 5), 5), B. B.

instars instars survived survived to to produce produce larg e, e, apparently apparently normal normal trols trols by by days days 5 5 or or 7, 7, or or by by pupation. pupation. Th ese ese includ ed ed

of of 51 51 (19.6%) (19.6%) Foray-treated Foray-treated lat e e 2nd 2nd and and ear ly ly 3rd 3rd tr eated eated larva e e and and no no more more than than 5% 5% mortality mortality in in con­

ultimates ultimates survived survived past past day day 4. 4. Similarly Similarly C. C. in in vidua, vidua, mulation mulation 10 10 foray. foray. There There was was at at l eas t t 95 % % mortality mortality in in

l ast ast insta r. r. an, an,

Egira Egira altemans altemans 1st 1st 7 7 2 2 20 20 55 55

Lithoplwne Lithoplwne petulca petulca 2nd 2nd 0 0 31 31 25 25 77 77

Catocala Catocala praeclara praeclara 0 0 n - 1 1 1 2 2 0 0 68 68

Pro c h eorodes eorodes trans versata versata 2nd 2nd 0 0 5 5 12 12 75 75

Limenitis Limenitis arthemis arthemis astyanax astyanax n/n - 1 1 0 0 10 10 0 0 100 100

Control Control Treated Treated Control Control Treated Treated

Species Species Assayed Assayed instar" instar" 0 - 5 5 d d after after treatment treatment 0 0 d d throu gh gh pupation pupation

%morta lity lity

Table Table 3. 3. Delayed Delayed mortality mortality 5 5 opeeieo opeeieo in in oub jeeted jeeted to to foliage foliage tr ea t ed ed with with Foray Foray 48B 48B

454 454 ENVIRONMENTAL ENVIRONMENTAL ENTOMOLOGY ENTOMOLOGY Vol. Vol. 27, 27, no. no. 2 2

mor- 5 5 dav dav . . However . . 5 day day by by significant significant was was y y lit ta the the of of Some Some . . sensitive highly highly were were species species butterfly butterfly

­ mor , , species the the of of most most For For Mortality. Mortality. Delayed Delayed most most definition, definition, their their by by that, that, found found (1981) (1981) bruch bruch

. . 8AF Dipel Dipel and and 48B 48B Foray Foray both both to to sensitive sensitive Langen­ and and Krieg Krieg varieties, varieties, thuringiensis thuringiensis B. B. several several

ess ess l generally generally e e larva older older and and with with instar, instar, butterflies butterflies of of larval larval to to number number a a related related for for results results tabulating tabulating In In

is is often often thuringiensis thuringiensis B. B. to to sensitivity sensitivity reduced reduced that that species. species. moth moth the the of of species species 38 38 of of 10 10 versus versus sensitive sensitive

belief belief accepted accepted generally generally the the with with consistent consistent is is This This highly highly were were evaluated evaluated we we butterflies butterflies 4 4 the the of of All All

instars. instars. late late or or mid- as as assayed assayed were were species species insensitive insensitive recovery. recovery. for for

the the of of All All Dipel. Dipel. to to insensitive insensitive were were also also Aaltemata Aaltemata necessary necessary be be recolonization recolonization would would Nor Nor thuringiensis. thuringiensis.

and and bicolorago, bicolorago, S. S. sericea, sericea, obliquata, obliquata, D. D. C. C. species, species, of of B. B. application application 1 1 from from unlikely unlikely be be would would field field the the

these these Of Of A. A. altemata. altemata. and and ( ( Guenee), Guenee), hibisci hibisci Orthosia Orthosia in in ion ion eradicat local local that that so so 0%) 0%) >2 (generally (generally foliage foliage

. . (J Smith), Smith), B. B. alurina alurina Orthosia Orthosia X. X. capax, capax, bicolorago, bicolorago, treated treated on on survivors survivors sufficient sufficient had had criteria criteria sensitive sensitive

Sunira Sunira (Morrison), (Morrison), sericea sericea highly highly Chaetaglaea Chaetaglaea the the meet meet to to failed failed Franclemont, Franclemont, that that assay assay our our in in species species The The

semitaria semitaria Metaxaglaea Metaxaglaea ench), ench), (Fr signata signata Sericaglaea Sericaglaea impossible. impossible. is is recolonization recolonization that that isolated isolated sufficiently sufficiently

E. E. vinulenta, vinulenta, e: e: octuida N N following following the the and and assayed; assayed; riid riid are are colonies colonies ed ed localiz in in species species if if permanent permanent be be could could

­ lymant native native only only the the obliquata, obliquata, Dasychira Dasychira tusaria; tusaria; ications ications erad These These small. small. or or sparse sparse are are populations populations

­ ob Euchlaena Euchlaena geometrid, geometrid, a a included included These These . . both or or if if thuringiensis thuringiensis B. B. with with treated treated areas areas from from eradicated eradicated

Dipel8AF Dipel8AF or or 48B 48B Foray Foray to to insensitive insensitive being being for for criteria criteria be be could could species species Such Such application. application. single single a a even even from from

the the met met assays assays our our in in evaluated evaluated species species 42 42 of of 11 11 Only Only tirpations tirpations ex local local or or crashes crashes population population massive massive incur incur

it. it. within within eradicated eradicated were were they they if if even even area area sprayed sprayed to to expected expected be be can can they they because because thuringiensis thuringiensis B. B. of of

thuringiensis­ B. B. any any quickly quickly ze ze loni reco to to expected expected be be cation cation li app l l aeria the the from from risk risk greatest greatest at at are are probably probably

would would , , therefore and, and, areas, areas, arge arge l over over numbers numbers high high definition definition our our by by sensitive sensitive highly highly are are that that Species Species

relatively relatively in in occur occur assayed assayed species species other other the the of of Most Most well. well. as as Foray Foray to to sensitive sensitive highly highly

adulthood. adulthood. to to survived survived lemmeri lemmeri treated treated 22 22 L. L. of of 12 12 and and were were species species 4 4 other other the the whereas whereas Foray, Foray, to to sensitive sensitive

ensis ensis thuringi B. B. to to insensitive insensitive proved proved capax capax X. X. states. states. highly highly as as not not but but Dipel Dipel to to sensitive sensitive highly highly taria taria was was

several several in in Programs Programs Heritage Heritage Natural Natural by by rare rare as as listed listed pome­ A. A. 5). 5). (Table (Table 7) 7) (day (day pyramidoides pyramidoides A. A. and and 5), 5),

are are capax, capax, Xylotype Xylotype and and lemmeri lemmeri assayed, assayed, L. L. species species (day (day vidua vidua 5), 5), (day (day disstria disstria C. C. (pupation), (pupation), metaria metaria M. M.

noctuid noctuid 2 2 Also, Also, 1995. 1995. in in abandoned abandoned was was list list ­ candidate candidate po A. A. 5), 5), A A day day (by (by clyton clyton product: product: thuringiensis thuringiensis B. B.

the the until until Act Act Species Species Endangered Endangered States States United United the the this this to to sensitive sensitive highly highly being being for for criteria criteria our our met met 5 5 , , 8AF

under under listing listing for for candidate candidate a a was was species species This This . . palachia Dipel Dipel against against evaluated evaluated were were that that species species 14 14 the the Of Of

Ap­ in in suppression suppression moth moth gypsy gypsy for for thuringiensis thuringiensis B. B. adults. adults. of of normal normal apparently apparently duce duce

applications applications forestry forestry in in concern concern particular particular of of is is and and ing ing ­ pro to to survived survived arvae arvae l treated treated the the of of cinata-several cinata-several

­ spray moth moth gypsy gypsy of of time time the the at at stars stars in in y y l ear as as found found coc­ H. H. , , and and maia maia , , feroidaria feroidaria C. C. species-L. species-L. sensitive sensitive

is is studies, our our in in thuringiensis thuringiensis . . B to to sensitive sensitive highly highly be be highly highly the the of of some some for for that that noted noted be be should should . . instar) It It

to to shown shown species species butterfly butterfly the the , , of of one one diana Speyeria Speyeria (antepenultimate (antepenultimate grotei grotei and and L. L. instar), instar), (penultimate (penultimate

studies. studies. other other in in overlooked overlooked coccinata coccinata instar), instar), C. C. penultimate penultimate and and 2nd 2nd ilia ilia (both (both

been been have have might might assay) assay) Limenitis Limenitis our our in in (as (as mortality mortality instar), instar), (last (last disstria disstria C. C. instar), instar), (4th/5th (4th/5th clyton clyton M. M. A A

delayed delayed but but sensitive, sensitive, star), star), in in moderately moderately (last (last only only astyanax astyanax be be to to arthemis arthemis B. B. peared peared were were instars instars late late or or

ap­ Langenbruch Langenbruch and and Krieg Krieg by by listed listed Nymphalidae Nymphalidae mid-instars mid-instars as as sensitive sensitive highly highly were were that that species species 6 6 The The

8AF 8AF Dipel Dipel to to sensitive sensitive Highly Highly c c

arvae. arvae. l control control or or treated treated in in occurred occurred y y lit morta no no which which for for species species for for reported reported e e valu P P No No b b

instar. instar. , , n ast ast l a

0.9999 0.9999 0 0 1 1 31 31 altemata altemata 18 18 Abagrotis Abagrotis 1 1 - n/n

14 14 0.1698 0.1698 1 1 4 4 19 19 2 2 - n alurina alurina Orihosia Orihosia

0 0 0 0 41 41 20 20 n n bicolorago bicolorago Sunira Sunira

0 0 0 0 30 30 20 20 2 2 - 1/n - n sericea sericea Chaetaglaea Chaetaglaea

0.0063 0.0063 0 0 9 9 19 19 lenta lenta vinu vinu 20 20 Eupsilia Eupsilia 2 2 - n

9 9 38 38 0.1423 0.1423 1 1 19 19 unimoda unimoda L. L. 1 1 - n

0 0 28 28 22 22 00001 00001 . 0 $ grotei grotei 20 20 Lithophone Lithophone 2 2 - 1/n - n

3 3 2 2 35 35 19 19 0.00001 0.00001 $ 1 1 - n c c pyramidoides Amphipyra Amphipyra

0 0 0.00001 0.00001 $ 2 2 31 31 17 17 1st 1st c c vidua Catocala Catocala

Noctuidae Noctuidae

0 0 0 0 26 26 20 20 4th 4th obliquata obliquata Daychira Daychira

Lymantriidae Lymantriidae

28 28 0 0 0.00001 0.00001 $ 4 4 23 23 2nd 2nd disstriif disstriif Malacosoma Malacosoma

Lasiocampidae Lasiocampidae

0 0 14 14 47 47 17 17 0.00001 0.00001 $ 1st 1st magnaria magnaria Ennomos Ennomos

11 11 1 1 1 1 2 19 19 0.00001 0.00001 $ n n pometariif pometariif Alsophila Alsophila

2 2 1 1 20 20 21 21 0.00001 0.00001 $ 4th/5th 4th/5th ° ° clyton Asterocampa Asterocampa

Geometridae Geometridae

dead dead dead dead No. No. No. No. alive alive No. No. alive alive No. No.

star" star" In In Species Species

ph ph

Dipel Dipel l l Contro

8AF 8AF Dipel Dipel to to Lepidoptera Lepidoptera 5. 5. native native Table Table of of species species 14 14 of of Susceptibility Susceptibility

455 455 thuringiensis thuringiensis B. B. April1998 April1998 LEPIDOPTERA LEPIDOPTERA NATNE NATNE ON ON OF OF ASSESSMENT ASSESSMENT AL.: AL.: ET ET PEACOCK PEACOCK 456 ENVJRONMENTAL ENTOMOLOGY Vol. 27, no. 2 tality often will substantially underestimate the true ringiensis had little lasting ill effect on surviving larvae, impact from B. thuringiensis. For 5 species, nearly all even though mortality in treated larvae ranged from 0 of the mortality occurred after day 5 (Table 3). Of to 95% among these same species. particular note was the nymphalid, L. arthemis asty­ On the basis of the available data for the more B. anax, for which no mortality was observed in larvae thuringiensis-sensitive species, we conclude that most that fed on treated foliage, yet all the prepupae and larvae that survived the consumption of treated foliage pupae died before eclosion. All of the L. a. astyanax and reached pupation later produced normal adults. larvae fed untreated foliage completed development For example, 3 H. maia females that survived treat­ and produced normal adults. For 7 highly sensitive ment as larvae laid 2 normal egg rings each. Viable eggs species and 4 other species, 75-100% of the day 5 also were obtained from treated survivors of A pome­ survivors died before pupation, usually during days taria, C. sericea, and D. obliquata, although fecundity 6-10 (Table 2) . For example, the 9 P. resumens that was very low in the former species. Nevertheless, our survived to day 5 were dead by day 7 and only 4 of the data do not allow a conclusion that all survivors of B. 18 P. rolandi surviving to day 5 lived long enough to thuringiensis-sensitive species were completely unaf­ pupate. Our data suggest that one cannot conclude fected. that a species is unaffected or only moderately af­ Taxonomic Patterns. Despite the observed intrafa­ fected by B. thuringiensis solely on the basis of low milial variation in sensitivity to B. thuringiensis, some mortality during the first 5 d or so after consuming taxonomic patterns were apparent from this study. treated foliage. We suggest the B. thuringiensis assays Penultimate and last in stars of at least 7 of 8 species of should be monitored for at least 7 d, and preferably to noctuids were insensitive to B. thuringiensis. pupation. The 8 late instars we assayed included 2 species of Sublethal Effects and Pupal Viability. We at­ Lithophane, and 1 each of Eupsilia, Chaetaglaea, Seri­ tempted to investigate sublethal effects despite having caglaea, Metaxaglaea, Xylotype, and Sunira, but only L. few survivors of B. thuringiensis-sensitive species; in grotei incurred significant mortality. Sensitivity among certain assays this often precluded rigorous statistical early- to mid-instar Xylenini (L. grotei, L. lemmeri, L. analysis. Larvae developed more slowly than their petulca, X. rufago) varied widely, but was significant control counterparts for most species evaluated from for 3 species. day 7 to pupation. This was especially true for L. Both species of the noctuid genus Orthosia were lemmeri treatment larvae that finished feeding =11 d little affected by B. thuringiensis, which agrees with after control larvae. The 4 surviving P. rolandi were reports by Krieg and Langenbruch (1981). These au­ delayed =8 d. For H. maia, treatment larvae took thors also reported the response of additional (prin­ nearly 3 wk longer than controls to complete larval cipally European) species of Alsophila, Ennomos, development, although this delay did not result in Lambdina, Malacosoma, Dasychira, and Papilio to sev­ lower pupal weights (male: 1.1 versus 1.0 g; female: 1.4 eral varieties and formulations of B. thuringiensis; their versus 1.5 g) or reduced survival of pupae. Many other findings largely agree with ours. When our results are species, including all sensitive species and some in­ pooled with those of Krieg and Langenbruch, Catocala sensitive ones, showed some apparent delay in devel­ and Lithophane remain the only genera for which opment to pupation, typically about 5 d. Again, small comparison of effects of B. thuringiensis on more than sample sizes (especially among surviving larvae from 3 congenerics is possible, and in both genera these treated foliage) precluded statistical analysis of this effects (based entirely on our results) varied signifi­ effect. Although the larvae of all these species later cantly among the species tested. produced apparently normal pupae and adults, de­ It is important to note the B. thuringiensis was ap­ layed development could have some biological con­ plied to treated foliage under ideal conditions in this sequences in the field, including increased parasitism study. The amount applied (equivalent of 89 BIU I ha (Ticehurst et al. 1982, Wallner et al. 1983, Weseloh for Foray and 99 BIU/ha for Dipel) is the maximum 1984) and reduced weight and fecundity (Schweitzer allowed in gypsy moth aerial application programs. 1979, Schneider 1980). Further, Foray and Dipel were applied to foliage in a Adult weights of species with significant larval mor­ laboratory spray tower, which resulted in nearly uni­ tality were similar for control and surviving treated form coverage. Such uniformity is seldom achieved Xystopeplus rufago (Hubner) (control versus treated: under field conditions. Thus, the laboratory mortality 0.045 versus 0.046 g), C. vidua (0.273 versus 0.253 g), recorded for certain species under these ideal condi­ C. praeclara (0.142 versus 0.148 g), and C. lineella tions might be greater than that which would occur in (0.035 versus 0.037 g). This also was true for adult the field. weights obtained for some species that were not sus­ It is difficult to generalize when predicting the sus­ ceptible to B. thuringiensis. These included the ceptibility of native Lepidoptera to B. thuringiensis, geometrid E. obtusaria (control versus treated: 0.050 especially for mid- or late instars. For taxa such as versus 0.053 g) and the noctuids S. signata (0.079 butterflies, many species appear to be highly sensitive. versus 0.117 g), C. sericea (0.074 versus 0.087 g), S. Conversely, xylenine noctuids apparently are mostly bicolorago (0.019 versus 0.018 g), X. capax (0.063 ver­ insensitive, especially as late in stars. But for many taxa sus 0.056 g), 0. alurina ( 0.066 versus 0.059 g), 0. hibisci the issue of susceptibility must be dealt with on a (0.07 versus 0.07 g), and A altemata (0.055 versus species-by-species basis, with special attention given 0.058 g). For these species, it is apparent that B. thu- to the phenology of species at the time of application

arthropod species species to to Bacillus Bacillus thuringiensis, thuringiensis, pp. pp. 837 - 896.In 896.In tober tober 1997. 1997.

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west west region. region. Gypsy Gypsy Moth Moth News News 29: 29: 8 - 9. 9. Ticehurst, Ticehurst, M., M., R. R. A. A. Fusco, Fusco, E. E. and and M. M. Blumenthal. Blumenthal. 1982. 1982.

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servation servation concern concern (1 (1 was was highly highly sensitive, sensitive, 1 1 was was in­

crobial crobial pest pest control control agent agent on on nontarget nontarget Lepidoptera . . Am. Am.

our our assays, assays,

though though

we we

did did evaluate evaluate 3 3 species species of of con­

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of of th e e effects effects of of ami­

to to able evaluate evaluate threatened threatened or or endangered endangered species species in in

Service , ,

Forest Forest

Health Health Protection, Protection,

Mor ga ntown , , WV. WV.

or or endangered endangered species species of of Lepidoptera. Lepidoptera. We We were were un­

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April1998 April1998 B. B. thuringiensis thuringiensis PEAcocK PEAcocK ET ET AL.: AL.: 457 457 AssESSMENT AssESSMENT OF OF ON ON NATIVE NATIVE LEPIDOPIERA LEPIDOPIERA