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Efficacy of Piper (Piperaceae) Extracts for Control of Common

Home and Garden Insect Pests

Article  in  Journal of Economic Entomology · September 2004

DOI: 10.1603/0022-0493-97.4.1390 · Source: PubMed

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 HORTICULTURAL ENTOMOLOGY

Efficacy of Piper (Piperaceae) Extracts for Control of Common

Home and Garden Insect Pests
I. M. SCOTT,1 H. JENSEN,1 R. NICOL,1 L. LESAGE,2 R. BRADBURY,3 P. SÁNCHEZ-VINDAS,4
L. POVEDA,4 J. T. ARNASON,1 AND B.J.R. PHILOGÈNE1

J. Econ. Entomol. 97(4): 1390Ð1403 (2004)

ABSTRACT Extracts from three species of the plant family Piperaceae, Piper nigrum [L.], Piper
guineense [Schum & Thonn], and Piper tuberculatum [Jacq.], were tested for efÞcacy against insects
from Þve orders. All three species contain isobutyl amides, plant secondary compounds that act as
neurotoxins in insects. These materials are considered safe to mammals because Piper spp. were used
for centuries for spice and medicinal purposes. When 24-h P. nigrum LC50 values were compared
between common insect pests from eastern Canada and the northeastern United States, the most
sensitive species in order of increasing lethal concentration were eastern tent caterpillar, Malacosoma
americanum (F.) ⬍ European pine sawßy larvae, Neodiprion sertifer (Geoffroy) ⬍ spindle ermine
moth larvae, Yponomeuta cagnagella [Hübner] ⬍ viburnum leaf beetle larvae, Pyrrhalta viburni
[Paykull] ⬍ stripped cucumber beetle adults, Acalymma vittatum (F.) ⬍ Colorado potato beetle
adults, Leptinotarsa decemlineata (Say) ⬍ Japanese beetle adults, Popillia japonica [Newman] ⬍ hairy
chinch bug, Blissus leucopterus hirtis [Montandon]. The life stage tested was the point at which each
species causes the greatest amount of damage to the host plant and the point at which most gardeners
would likely choose to treat with a conventional synthetic insecticide. Greenhouse trials revealed that
the pepper formulations also had a repellent activity, thus protecting plant leaves from 1) herbivory
(lily leaf beetle, Lilioceris lilii [Scopoli], adults and larvae and stripped cucumber beetle adults) and
2) oviposition [European corn borer, Ostrinia nubilalis (Hübner)]. Combinations with other botanical
extracts were additive at best in toxicity and repellent trials. Nontarget toxicity to beneÞcial inver-
tebrates is a possibility because the P. nigrum LC50 for beneÞcial ladybird beetles was 0.2%. P. nigrum
extracts can provide a reasonable level of control against lepidopteran and European pine sawßy larvae
and also will work as a short-term repellent and feeding deterrent. It is recommended that the use of
Piper extracts be restricted to small-scale spot treatments in residential areas where insect pest
outbreaks have occurred.

KEY WORDS Piperaceae, Piper nigrum, piperamides, efÞcacy, repellent effect

BIOPESTICIDES OF PLANT ORIGIN were reviewed recently pepper, Piper nigrum L., and African Guinea pepper,
(Regnault-Roger et al. 2002), and it was concluded Piper guineense Schum & Thonn, but many other spe-
that botanicals have considerable market potential as cies in the family are also insecticidal (Bernard et al.
reduced risk control agents. In addition, the National 1995).
Research Council (2000) in the United States recom- Early investigations with P. nigrum extracts indi-
mended a number of areas where botanicals meet cated that isobutyl amides were responsible for the
current and future requirements for alternative pest toxicity of the extracts to the adzuki bean weevil,
control. Lydon and Duke (1989), Isman (1994), and Callosobruchus chinensis L. (Miyakado et al. 1979,
MacKinnon et al. (1997) surveyed several plant fam- 1980). Three of the isobutyl amides isolated from P.
ilies that show promise as sources of new botanical nigrum, pipercide, pellitorine, and piperine, were
insecticides. Members of the pepper family Piper- toxic at 0.15, 2, and 20 ␮g/male C. chinensis, respec-
aceae produce phytochemicals with insecticidal ac- tively (Dev and Koul 1997). Guineensine, isolated
tivity. The most widely recognized species are black from P. guineense, had similar activity to pipercide
when tested topically on the cowpea weevil, Calloso-
1 Biology Department, University of Ottawa, Ottawa, Ontario, K1N bruchus maculatus F. (0.25 versus 0.84 ␮g/male 48-h
6N5 Canada. LD50 values, respectively). Pepper extracts containing
2 ECORC, Agriculture & Agrifood Canada, Ottawa, Ontario, Can- mixtures of isobutyl amides are also highly effective
ada. (Scott et al. 2002).
3 EcoSafe Natural Products, Saanichton, British Columbia, V8M

2C2 Canada.
Essential oils of P. nigrum were found to effectively
4 Facultad de Ciencias de la Tierra y del Mar, Universidad Nacional, protect stored wheat from the stored grain pests Sito-
Heredia 3000, Costa Rica. philus oryzae L. and Rhyzopertha dominica F. at con-

0022-0493/04/1390Ð1403$04.00/0 䉷 2004 Entomological Society of America

August 2004 SCOTT ET AL.: Piper EXTRACTS FOR CONTROL OF HOME AND GARDEN PESTS 1391

centrations ⬎100 mg/liter for up to 30 d (Sighamony viburni Paykull (Chrysomelidae)]; 2) Dermaptera

et al. 1986). Stored beans were protected from the [earwig Forficula auricularia L. (ForÞculidae)]; 3)
bruchid Acanthoscelides obtectus Say by ground black Hemiptera [hairy chinch bug, Blissus leucopterus hir-
pepper for up to 18 wk (Baier and Webster 1992). P. tus Montandon (Lygaeidae)]; 4) Hymenoptera [Eu-
guineense-treated kaolin powder at 150 ␮l/g reduced ropean pine sawßy, Neodiprion sertifer (Geoffroy)
the average adult emergence of C. maculatus by 100% (Diprionidae)]; and 5) Lepidoptera [European corn
after 30-d treatment (Kéṏta et al. 2000). Dust and borer (Pyralidae); spindle ermine moth, Yponomeuta
ether-extract formulations of P. guineense also were cagnagella Hübner (Yponomeutidae); and eastern
effective at controlling C. maculatus, at concentrations tent caterpillar, Malacosoma americanum Fabricius
between 0.5 and 0.75 g/20 g cow pea seed within 36 h (Lasiocampidae)].
after treatment (Mbata et al. 1995). Emergence of Nontarget toxicity also was evaluated using an in-
adults from treated eggs was prevented successfully sect species that would likely encounter the effects of
with dust and oil treatments at 0.25 g/seed. the botanical treatment. Applications in the garden
The efÞcacy of Piper extracts against a few other could affect the ladybird beetle Hippodamia conver-
insect pests also has been demonstrated. The termite gens (Guérin-Méneville) (Coccinellidae) as beneÞ-
Macrotermes nigeriensis Sjostedti was controlled with cials or predators of garden insect pests.
a 5% aqueous solution of P. guineense applied topically The target insect pests and nontarget invertebrates
(Ivbijaro et al. 1993). Javier and Morallo-Rejesus were treated with Piper extract formulations to
(1986) determined that semipuriÞed organic solvent achieve the following objectives: 1) establish concen-
extracts of P. nigrum were more toxic than crude tration levels for the target insects and toxicity values
extracts against the house ßy, Musca domestica L.; to protect the nontarget invertebrate; 2) evaluate
cutworm Spodoptera litura F.; black armyworm, Spo- knockdown, repellent action, and feeding deterrent
doptera exempta (Walker); and diamondback moth, effect under controlled treatment conditions; and 3)
Plutella xylostella (L.). Ewete et al. (1996) showed determine residue levels for Piper active compounds
that P. guineense incorporated into the diet of the on contact surfaces. The overall objective was to pro-
European corn borer, Ostrinia nubilalis (Hübner), at vide an in-depth evaluation of the efÞcacy, toxicity,
300 mg/liter reduced larval growth by 27% and in- and environmental fate of pepper extracts.
creased the time to adult emergence and reduced egg
production at concentrations ⬎10 mg/liter. In terms
of nontarget effects, aqueous mixtures of oven-dried Materials and Methods
and powdered P. guineense at 10 mg/liter were found
Piper Extracts and Commercial Botanical
to be effective at controlling fourth instars of mosquito
Formulations
Aedes aegypti L., but they were not toxic to other
aquatic organisms (Okorie and Ogunro 1992). Seed material for both P. nigrum and P. guineense
Other species of Piper that also contain piperamides were purchased from commercial suppliers in Canada,
include long pepper, Piper longum L. from South Asia the United States, and Togo, West Africa. Leaves of P.
and Piper tuberculatum Jacq. from South and Central tuberculatum were collected in Costa Rica by Pablo
America. The latter is of particular interest because of Sanchez and Luis Poveda near San Carlos. Voucher
high concentrations of piperamides in the leaves specimens have been placed in the University of Ot-
(Scott et al. 2002). Along with P. nigrum and P. tawa and the Universidad Nacional (Heredia, Costa
guineense, P. tuberculatum may offer the most prom- Rica) herbaria. P. nigrum and P. guineense peppercorns
ising and active pepper extract available for the de- and P. tuberculatum leaf material were ground and the
velopment of a commercial insect control product due active constituents extracted following the methods
to its long traditional use, low associated health and described in Scott et al. (2002). Natural solvents and
environmental risk, and the relative abundance of emulsiÞers were incorporated into the formulation to
material. reduce the risk of toxicity to the applicator and the
Most previous evaluations have focused on major environment. The Piper extracts were formulated by
crop and nuisance pests, and little research has been R. Bradbury (EcoSafe Natural Products, Saanichton,
undertaken on insects of the home and garden where British Columbia, Canada) as follows: 20% extract, 70%
a botanical product might gain ready acceptance. We tetrahydrofurfuryl alcohol (THFA; Penn Specialty
developed a practical formulation that was evaluated Chemicals, Memphis, TN), and 10% emulsiÞer (Alka-
against target insect species that are of concern to muls EL-719 ethoxylated castor oil, a gift of Rhodia,
gardeners and horticulturalists in eastern Canada and Cranbury, NJ). The piperamide concentration in the
the northeastern United States and were selected for extracts and formulations was analyzed based on the
use in efÞcacy trials based on their abundance and methods of Scott et al. (2002)). High-performance
availability. These include insects from the orders 1) liquid chromatography (HPLC) analysis was con-
Coleoptera [Colorado potato beetle, Leptinotarsa de- ducted using a Prostar model pump, model 330 UV/
cemlineata (Say) (Chrysomelidae); Japanese beetle, Vis, photodiode array detector and model 410 au-
Popillia japonica Newman (Scarabaeidae); lily leaf tosampler (Varian Chromatography Systems, Walnut
beetle, Lilioceris lilii Scopoli (Chrysomelidae); Creek, CA).
striped cucumber beetle, Acalymma vittatum (F.) Other botanical extracts were obtained to act as
(Chrysomelidae); and viburnum leaf beetle, Pyrrhalta synergists or to compare with the insecticidal and
1392 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 97, no. 4

repellent activity of the Piper extracts. These included plants until bioassays were initiated. P. viburni larvae
Central American Cedrela odorata L. (Meliaceae) ex- were collected from the Kemptville Agricultural Col-
tract; dillapiol, 95% dill oil from Anethum sawa Roxb. lege Campus, Kemptville, Ontario, and The Log Farm,
(Umbelliferae); neem oil Azadiracta indica Juss (Me- Ottawa, Ontario, in May 2002. Larvae were kept in
liaceae) (Ahimsa Alternative Inc., Oklahoma City, containers with fresh European highbush cranberry,
OK); Garlic Barrier concentrated garlic Allium sati- Viburnum opulus L. (Caprifoliaceae), leaves at 10⬚C
vum L. (Alliaceae) (a gift of Garlic Barrier Labs, Glen- and a photoperiod of 16:8 (L:D) h until bioassays were
dale, CA), and Ropel, lemon grass oil Cymbopogon initiated. A. vittatum adults were collected from the
citratus [DC] Stapf. (Poaceae) (Burlington ScientiÞc, Gloucester allotment gardens in August 2002, in Ot-
Farmingdale, NY). All commercial products were ap- tawa and kept at 10⬚C and a photoperiod of 16:8 (L:D)
plied at rates recommended by the manufacturer. h and fed cucumber slices until bioassays were initi-
ated. Wild-type adult L. decemlineata were collected
from organic potato Þelds, Hatley, Québec, and urban
Effect of Sunlight and UV Radiation Exposure to
allotment gardens having pesticide use restrictions,
Piper Extracts
Ottawa, Ontario, in July 2002.
The effect of sunlight and UV radiation on exposed Dermaptera. F. auricularia were collected from a
piperamides either alone or in Piper extracts was as- garden site in Ottawa, Ontario, in July 2001. The ear-
sessed following the methods described in Scott et al. wigs were fed carrot and apple slices and were at held
2003. Degradation of piperamides exposed to full-sun- at 10⬚C and a photoperiod of 16:8 (L:D) h until the test
light conditions was determined by placing 50-␮l ali- was initiated.
quots of 20% extract on glass microscope slides and Hemiptera. B. leucopterus hirtus nymphs and adults
allowing them to dry overnight on the bench top. The were collected in residential turf in Gatineau, Québec,
slides were then exposed to full sunlight for 6 h during in August 2002. The chinch bugs were kept in a mix-
peak daylight hours. Readings of solar radiation were ture of soil and grass mulch at 10⬚C until tests were
taken at 3-h intervals. The glass slides were then rinsed initiated.
with 5 ml of 99% ethanol to wash off the extract Hymenoptera. N. sertifer larvae were collected from
residue. Treated control slides not exposed to full Mugo pine, Pinus mugo variety Mugo (Pinaceae),
sunlight were rinsed using the same method. The trees near Almonte and Pakenham, Ontario, in May
ethanol solutions (1 ml) were then Þltered through a 2002. The larvae were fed freshly cut Mugo pine
0.2-␮m polypropylene Þlter and placed in a 1.5-ml boughs and were held at 10⬚C and a photoperiod of
HPLC vial in preparation for analysis. Analysis was 16:8 (L:D) h until bioassays were initiated.
conducted according to Scott et al. (2002); however, Lepidoptera. M. americanum larvae were collected
a Varian Prostar model pump, model 330 UV/Vis, near Almonte and Pakenham, Ontario, in May 2002.
photodiode array detector and model 410 autosam- Larvae were kept at 10⬚C and fed M. sylvestris leaves
pler, and a Varian reverse-phase C18 column (Varian until trials began. Larvae were separated into instars,
Chromatography Systems), were used. The com- and bioassays were conducted with the different age
pounds were eluted with a binary gradient of aceto- classes where numbers permitted. Y. cagnagella were
nitrile and water, where acetonitrile was increased collected from the Kemptville Agricultural College
from 30 to 90% in 12 min, as described by Scott et al. Campus, Kemptville, Ontario, once in June 2001 and
(2002). Solar radiation readings were taken at each twice in May 2002. The individuals collected in June
time period with a Li-Cor Quantum model LI-192SB 2001 at the prepupal, nonfeeding larval stage, whereas
sensor (Li-Cor, Omaha, NE). The UV light was an the earlier larval stages were fed burning bush, Eu-
Industrial F20T12/BLB blacklight blue lamp that onymus alatus Thunb. Siebold variety Compactus
emits in the near UV, 315Ð 400 nm. The lamp has a deep (Celastraceae). The larvae were kept at 10⬚C and a
blue Þlter to absorb visible light and transmit near UV photoperiod of 16:8 (L:D) h until trials began. O.
at a 3.7-W output. nubilalis larvae and adults were obtained from the
University of Ottawa culture. Second instars were
selected 6 d after eggs hatched and the adults within
Insect Species Collected or Cultured
24 h of emerging from the pupal stage.
Coleoptera. P. japonica adults were collected at one Voucher specimens for all insects collected and
site in Ottawa, Ontario, Canada, in August 2001 and used in efÞcacy bioassays were placed in the Insect
2002. Adults were kept at 10⬚C and a photoperiod 16:8 Biosystematics collection, Eastern Cereal and Oilseed
(L:D) h and fed crab apple, Malus sylvestris L. Miller Research Centre (ECORC), Agriculture and Agri-
(Rosaceae), or mountain ash, Sorbus americana Food Canada, Ottawa, Ontario.
Marsh. (Rosaceae), leaves until bioassays were initi-
ated. L. lilii eggs and adults were collected from sev-
Bioassays for Assessing Insecticidal and Repellent
eral gardens in May and June 2001 and 2002 and at the
Effects
Central Experimental Farm in Ottawa during June and
July 2001 and May 2002. Adults were kept at 10⬚C and Twenty-four-hour LC50 Determination. With the
a photoperiod of 16:8 (L:D) h until bioassays were exception of B. leucopterus hirtus adults, F. auricularia
initiated or kept on Asiatic lily plants (Liliaceae) in the adults, N. sertifer larvae, and Y. cagnagella larvae, all
greenhouse to produce eggs. Larvae were reared on toxicity trials were conducted by spraying to runoff on
August 2004 SCOTT ET AL.: Piper EXTRACTS FOR CONTROL OF HOME AND GARDEN PESTS 1393

individual host plant leaves allowed to dry at room 0.5% P. nigrum. Each treatment was replicated three
temperature for 30 min. Test insect larvae were then times with 20 P. japonica per plant. Each plant was
treated by spray to drip, patted dry, and placed on the individually caged with a wire frame and mesh cov-
leaves with the same treatment inside a glass petri ering. The number of leaves on each plant was stan-
plate or 500-ml Mason jar. Each species was treated dardized by removing all but 10 undamaged leaves per
with six P. nigrum concentration levels, including a plant. A leaf was considered damaged if L. decemlin-
formulation blank, based upon the results of a range- eata feeding caused partial or complete surface area
Þnding test. Ten larvae were used per two to three loss compared from the start of the trial. The plants
leaves, and each treatment was replicated at least two were checked visually for damage to leaves after 96 h.
times where numbers permitted. Test containers were Prepupal L. lilii larvae and adults were tested using
kept at room temperature. The LC50 value refers to potted Asiatic lily plants treated with P. nigrum ex-
mortality after 24 h, determined by touching the larvae tracts. Asiatic Lilium (Liliaceae) were purchased in
with a probe to elicit a response. the spring as bulbs, Orange Pixie, Butter Pixie, and
Dermaptera. F. auricularia were sprayed to runoff Latoya varieties, or as plants, Cancun, Orange Pixie,
with solutions of P. nigrum and P. guineense extracts at Butter Pixie, and Lennox varieties, and were treated
concentrations of 0.1, 0.5, and 1% and then placed with formulated P. nigrum extract in the range of 0.125,
individually in plastic cups, 10 replicates per treat- 0.25, 0.5, and 1%. At each concentration level three
ment. Controls consisted of water only and a formu- replicate plants were treated along with three repli-
lation blank consisted of a 5% formulation mixture in cate controls: a formulation blank of equal EC con-
water. Mortality was observed after 24 h. centration. Ten prepupal L. lilii larvae were placed on
Hemiptera. B. leucopterus hirtus adults were treated each plant, which was checked after 24 h to assess
with the spray to drip method but were then placed in larval mortality, movement from the treated leaves,
petri plates with Whatman Þlter paper. Ten chinch and feeding damage to leaves. Adults also were placed
bugs were used per treatment with two and three in three large cages where three plants per cage were
replicates for the Þrst and second trials, respectively. treated either with 1% P. nigrum, neem oil, or Ropel.
Six P. nigrum concentration levels were tested, includ- The choice test was checked after 96 h to assess dam-
ing a formulation blank or emulsiÞable concentrate age to leaves from feeding as described previously.
(EC), based upon the results of a range-Þnding test. A choice test using cucumber plants, Cucumis spp.
After 24 h, the number of dead adults on the Þlter (Cucurbitaceae), variety Bush Pickle, was conducted
paper was determined by touching the chinch bug where plants were sprayed to runoff with either 0.1 or
with a probe to elicit a response. 0.5% P. nigrum extract and placed in a cage with both
Hymenoptera. N. sertifer larvae were separated into a water and formulation blank control. Sixty A. vitta-
instars, and bioassays were conducted with the fourth tum adults were released in each of three replicate
or Þfth instars where numbers permitted. N. sertifer cages. Each plant was surrounded with a plastic collar,
larvae were placed onto the tips of needles of P. mugo which was capped at the end of 96 h to remove each
boughs freshly cut and then were sprayed to runoff plant but not lose the insects feeding on it. The plants
with the same range of P. nigrum concentrations. The were then cooled to 10⬚C so that the number of adult
boughs were then placed inside a 500-ml Mason jar to beetles on each plant could be collected and the num-
allow them to remain upright and covered with a mesh ber of damaged leaves counted.
lid. After 24 h, the number of dead or moribund larvae Lepidoptera. Green pepper plants, Capsicum an-
at the bottom of the jars was determined. num L. (Solanaceae), were grown to the early fruiting
Lepidoptera. Prepupal Y. cagnagella larvae col- stage and then chosen on the basis of damage-free
lected in 2001 were treated with control (water only), fruits. Ten plants per treatment were sprayed with
1% formulation blank, and P. nigrum at 0.01, 0.05, and either C. odorata extract plus dillapiol, 95% dill oil from
0.1%. Larvae were dipped into the solutions and then A. sawa (1:1 for the highest concentration and 0.6:1 for
placed in a covered petri plate. Ten larvae per repli- the lowest, due to availability of extract) at 0.1 and
cate were used, with three replicates per treatment. 0.03% extract or P. tuberculatum extract at 0.1 and
Survival of larvae was noted 48 h after treatment, and 0.05%. The control plants were sprayed with 4:1 95%
then the number of adults that emerged successfully ethanol:distilled water. The following day, the plants
within the following 2-wk period. were hand infested with three second instars of O.
nubilalis larvae plus one egg mass. On the third day,
the plants were resprayed. After 11 d, the fruits and
Repellent, Antifeedant, and Oviposition Deterrent
plants were tallied for surviving larvae.
Effects
O. nubilalis adults were collected within 24 h of
All Piper spp. repellent trials were conducted in the emerging from the pupal stage. Five female and Þve
greenhouse at Carleton University. male adults were aspirated into a 500-ml ßask. Green
Coleoptera. P. japonica adults were caged on Ex- pepper plants at the mature fruiting stage were treated
plorer rose plants (Rosaceae) variety Louis Jolliet to with either water, formulation blank, 0.5% P. guineense
determine the repellent effect of P. nigrum by com- extract, Ropel or Garlic Barrier, both at the recom-
paring damage to leaves caused by feeding. Treated mended application rates. Each plant was sprayed
plants were sprayed either with 100 ml of water only, until runoff (⬇100 ml), with four replicates for the
or water combined with 2.5% formulation blank or control-, EC blank-, and P. guineense-treated plants
1394 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 97, no. 4

Table 1. Range of piperamide concentrations (milligrams per gram) in P. nigrum, P. guineense, and P. tuberculatum extracts used
in efficacy trials 2001 and 2002

Extract Year Dihydropiper longuminine Piperlon guminine Dihydropiperine Piperine Total

P. nigrum 2002 0.4 1.9Ð2.9 17.7Ð22 370Ð427 391Ð451
2001 0.5Ð1.8 0.9Ð5.1 10.4Ð23.7 288Ð514 299Ð538
P. guineense 2002 31.2 28.1 84.4 53.5 197
2001 3.6Ð8 2.7Ð7.7 9.8Ð29.9 33.4Ð68.8 49Ð114
P. tuber 2002 59Ð65.5 2.7Ð7.5 6Ð15.5 4.7Ð13.5 72Ð102

and three for the remaining treatments. Plants were tracts used in 2002 had higher piperamide levels than
left to dry and then caged by enclosing the plants with in 2001, probably due to a difference in the source of
a metal frame and mesh net. The adult O. nubilalis the peppercorns.
were added to the cages and left in the greenhouse for Tests with different formulation components indi-
96 h. The cages were reopened and the surviving cated that 1) the three pepper extracts were soluble in
adults, the number of leaves with egg masses, and the ethanol and THFA, and 2) the two emulsiÞers se-
total number of egg masses were counted. lected, Jeneil Co. biosurfactant JBR-325 and -425, and
Rhone-Poulenc castor oil Alkamus EL-719, rated con-
sistently high in terms of emulsion bloom in water,
Nontarget Invertebrates
emulsion suspensibility, and spreading area. THFA
Ladybird Beetles. Adult H. convergens were pur- was chosen over ethanol based on the lower ßashpoint
chased from Biobest, Leamington, Ontario, through and potential for registration, and Alkamus EL-719, a
Plant Products, Brampton, Ontario. The adults were synthetic ethoxylated castor oil, was preferred over
kept at 4⬚C and were not fed before or during the test. JBR-425, a natural rhamnolipid, because it was less
All H. convergens tests were initiated within 2 wk of toxic in house ßy toxicity trials (unpublished data).
obtaining the adults. The toxicity test procedure fol- The Þnal composition of the pepper formulation for
lowed that previously described for the 24-h LC50 testing was pepper extract 20%, THFA 70%, and emul-
determination. Adults were dipped into P. nigrum so- siÞer 10%. The formulation was diluted in water at the
lutions ranging from 0.01 to 1%, patted dry on Þlter time of spraying to the appropriate concentration.
paper, and then placed into a plastic petri plate with
Whatman #1 Þlter paper. Ten H. convergens were
treated per plate with three replicate plates per con- Effect of Sunlight and UV Radiation on Piper
centration. Extracts
The piperamide content of extracts was found to be
Statistics stable over several months in the laboratory at room
temperature, but the formulations are susceptible to
Probit analysis (Hubert and Carter 1990a) was used photodegradation. Piperine in the extracts degraded
to determine the LC50 values for P. nigrum. Compar- quickly after exposure to sunlight (Fig. 1). Pure pip-
ison of the LC50 values between trials for each species erine also degraded quickly under UV lamp exposure
tested was conducted using a ␹2 or Z-test (Hubert and with a half-life of ⬇39 min, suggesting that the deg-
Carter 1990b). Results of all other bioassays were radation was a direct photolysis, not a photosensitized
tested for normality, and the data were transformed if reaction from some pepper pigment. After 6-h expo-
necessary. A one-way analysis of variance was per- sure to sunlight, all amides in the P. guineense extract,
formed with a post hoc TukeyÕs multiple comparison including piperine (t0.5 ⫽ 49 min), were below de-
of means test (SYSTAT 1999). tection (data not shown). Light peak light levels were
measured between 1,230 and 1,410 ␮Einsteins m⫺2
Results s⫺1.
Pepper Formulation Development
Insecticidal Activity
Black pepper extracts contained large amounts of
piperine and smaller amounts of piperlonguminine, Accurate LC50 values for the P. nigrum formulation
dihydro-piperine, and dihydro-piperlonguminine were obtained with seven selected urban insect pest
(Table 1). The four piperamides from each batch were species and one beneÞcial insect (Tables 2 and 3). In
totalled to provide an estimate of the total amide general, the extracts were more toxic to larval insects
content, although a few minor compounds were not where LC50 values ranged from 0.018 to 0.103% (Table
quantiÞed. The total ranged from 299 to 588 mg/g in 2) than adult insects, where LC50 values ranged from
P. nigrum, from 49 to 197 mg/g in P. guineense, and from 0.103 to 0.746% (Table 3). Lepidoptera and one hy-
72.4 to 102 in P. tuberculatum. Piperamide levels from menopteran species were more susceptible (0.018 Ð
P. nigrum batches prepared for the efÞcacy trials in 0.075%) than Coleoptera or the one Hemiptera
both 2001 and 2002 were found to have an overlapping (0.103Ð 0.746%) to the P. nigrum extracts. No direct
range of concentrations, whereas the P. guineense ex- comparisons were made between life stages for the
August 2004 SCOTT ET AL.: Piper EXTRACTS FOR CONTROL OF HOME AND GARDEN PESTS 1395

Fig. 1. Natural log of piperine concentration after exposure of P. guineense extract (dashed line) to full sunlight and
piperine to UV radiation (solid line) over 1.5 h.

same insect species in this study; however, it was ob- Repellent, Antifeedant, and Oviposition Deterrent
served previously that sensitivity to P. nigrum extracts Effects
was less for Colorado potato beetle late instars and adults
compared with early instars (Scott et al. 2003). Coleoptera. Behavior modiÞcation effects (repel-
Although LC50 data were not available, toxicity as- lent and antifeedant effects) of pepper formulations
sessments at several concentrations were made for were clearly evident with some situations, but not
other insects. Survival of adult earwigs F. auricularia others. In a no-choice experimental design with L. lilii
decreased signiÞcantly at 0.5% and 1% P. nigrum (F ⫽ larvae, fewer larvae remained on treated lily plants
14.7; df ⫽ 7, 16; P ⬍ 0.008) and 1% P. guineense (TukeyÕs (expressed as mobile insects) and fewer damaged
multiple range test, P ⫽ 0.001) after 24 h (Fig. 2). leaves were observed (F ⫽ 11.189; df ⫽ 15, 32; P ⬍
These latter results also show comparable insecticidal 0.001) compared with controls as the P. nigrum dose
activity for both the African and black pepper. P. was increased from 0.125 to 0.5% (Fig. 4). These two
nigrum extracts up to 0.1% did not reduce late instar, concentration-dependent relations were signiÞcant
prepupal Y. cagnagella survival signiÞcantly after 48 h (P ⫽ 0.05) with R2 ⫽ 0.86 for leaves damaged and R2
(F ⫽ 13.712; df ⫽ 3, 8; P ⫽ 0.855), but effects were ⫽ 0.94 for insect mobility. Pepper treatment also re-
manifested later in the life cycle where 0.01 and 0.1% duced the survival of all exposed larval insects (F ⫽
P. nigrum reduced survival to adult emergence signif- 13.223; df ⫽ 3, 8; P ⫽ 0.002) but not the number of dead
icantly (TukeyÕs multiple range test, P ⫽ 0.058 and or moribund larvae after 24 h (F ⫽ 18.073; df ⫽ 3, 8;
0.004, respectively) (Fig. 3). P ⫽ 0.131).

Table 2. P. nigrum LC50 values, 95% confidence intervals, and slope of probit lines for selected insect larvae

Insect Order: Family n P. nigrum LC50 (%) 95% C.I. (%) Slope
Eastern tent caterpillar Lepidoptera: Lasiocampidae 358 0.018 0.015, 0.022 2.84
European pine sawßy Hymenoptera: Diprionidae 354 0.046 0.04, 0.054 2.72
Spindle ermine moth Lepidoptera: Yponomeutidae 357 0.075 0.054, 0.124 1.15
Viburnum leaf beetle Coleoptera: Chrysomelidae 160 0.103 0.071, 0.137 2.03

Table 3. P. nigrum LC50 values, 95% confidence intervals, and slope of probit lines for selected insect adults

Insect Order: Family n P. nigrum LC50 (%) 95% C.I. (%) Slope
Striped cucumber beetle Coleoptera: Chrysomelidae 240 0.103 0.087, 0.13 3.16
Convergent lady beetle Coleoptera: Coccinellidae 540 0.213 0.173, 0.276 2.03
Colorado potato beetle Coleoptera: Chrysomelidae 300 0.498 0.363, 0.652 1.45
Japanese beetle Coleoptera: Scarabidae 539 0.532 0.446, 0.616 2.11
Hairy chinch bug Hemiptera: Lygaeidae 214 0.746 0.518, 1.361 1.49
1396 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 97, no. 4

Fig. 2. Adult F. auricularia percentage of survival ⫾ SE 24 h after water control; formulation control (EC); and 0.01, 0.1,
and 1% P. nigrum (Pn) and P. guineense (Pg) extract treatment. The treatment means with the same letter are not signiÞcantly
different (TukeyÕs test, P ⬎ 0.05).

In a choice experiment with L. lilii adults, P. nigrum plants (TukeyÕs multiple range test, P ⫽ 0.243) (Fig.
at 1% did not reduce adult feeding damage after 96 h 6). More importantly, there was no difference in the
signiÞcantly compared with the formulation blank number of leaves damaged by the beetles between
(F ⫽ 3.542; df ⫽ 3, 11; P ⫽ 0.074) (Fig. 5), and P. nigrum treatments (TukeyÕs multiple range test, P ⬎ 0.953).
at 1% was not signiÞcantly more effective than either Both 0.5 and 0.1% P. nigrum extracts reduced the
Ropel or neem oil (TukeyÕs multiple range test, P ⬎ number of A. vittatum adults found on cucumber
0.108). The 1% pepper spray caused phytotoxicity: plants after a 96-h choice test (F ⫽ 8.563; df ⫽ 7, 16;
burning of lily leaf tips, which was not observed at P ⬍ 0.007) (Fig. 7A and B, respectively). The A. vit-
lower concentrations that effectively repelled larval L. tatum adults not found on plants were considered
lilii. mobile and factored into the analyses but were not
Rose plants sprayed with 0.5% P. nigrum had fewer shown in Fig. 7A and B. In this situation, no phyto-
adult P. japonica present after a 96-h choice test com- toxicity to cucumber leaves was observed with 0.1% P.
pared with water-treated (control) plants (F ⫽ 7.57; nigrum formulation.
df ⫽ 5, 12; P ⫽ 0.001), but there was no signiÞcant Lepidoptera. The mixture of C. odorata and dillapiol
difference compared with formulation-treated (EC) at 0.1% had a signiÞcant impact on O. nubilalis larval

Fig. 3. Prepupal Y. cagnagella larvae percentage of survival and adult emergence ⫾ SE after 0.01 and 0.1% P. nigrum
treatment. The treatment means with the same letter are not signiÞcantly different (TukeyÕs test, P ⬎ 0.05).
August 2004 SCOTT ET AL.: Piper EXTRACTS FOR CONTROL OF HOME AND GARDEN PESTS 1397

Fig. 4. Mean number of live, dead, and mobile L. lilii larvae ⫾ SE on P. nigrum-treated Asiatic Lilium plants along with
the mean number of damaged leaves ⫾ SE per plant. The treatment means for survival (clear bar), mortality (dark bar), mobile
L. lilii larvae (hatched bar), and number of damaged leaves (shaded bar) with the same letter are not signiÞcantly different
(TukeyÕs test, P ⬎ 0.05).

survival (P ⬍ 0.01), but P. tuberculatum at 0.05 and tions ⬎0.1% were knocked down, but some had re-
0.1% had no signiÞcant effect (Fig. 8). When higher covered by 24 h in the 0.1% treatment.
concentrations of P. nigrum and P. guineense alone
were tested, the results were similar (unpublished
Discussion
data). However P. guineense at 0.5% reduced oviposi-
tion compared with control (F ⫽ 3.182; df ⫽ 4, 13; P ⫽ Botanical insecticides have been used for centuries
0.043) but not the formulation blank, Ropel, or Garlic for crop protection. Only with the development of
Barrier (P ⬎ 0.113) (Fig. 9). synthetic insecticides in the mid-1900s did their use
drop as more effective products took their place.
Within a relatively short time, problems arose with the
Nontarget Toxicity
synthetic products: environmental contamination,
The 24-h LC50 for H. convergens adults was 0.21% poisonings of nontarget species, and resistance. This
(Table 3). Within 1 h, adults treated with concentra- led many to reconsider botanical formulations as nat-

Fig. 5. Percentage of leaf damage ⫾ SE on Asiatic Lilium plants by L. lilii adults during 96-h choice test: plants treated
with either formulation control (EC), 1% P. nigrum (1% Pn), C. citratus oil (Ropel), or Azadirachta indica oil (neem). The
treatment means with the same letter are not signiÞcantly different (TukeyÕs test, P ⬎ 0.05).
1398 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 97, no. 4

Fig. 6. Mean number of P. japonica adults and leaves damaged ⫾ SE on rose plants treated with water (control),
formulation control (EC), and 0.5% P. nigrum. The treatment means for adults/plant (clear bar) and leaves damaged (dark
bar) were compared separately where the same letter indicates no signiÞcant difference (TukeyÕs test, P ⬎ 0.05).

ural alternatives because they are less toxic. However, damage by larvae becomes apparent. A spray mixed
these have always had varying degrees of success, and between 0.05 and 0.1% P. nigrum would initially knock
recently even their continued safe use has been ques- the larvae off leaves and branches and the neurotoxic
tioned. Rotenone and pyrethrum, two of the most activity would prevent them from returning to the
commonly applied by home gardeners and organic plant. Similarly, gardeners could repel L. lilii larvae
farmers, are being reevaluated by the U.S. EPA-based and Y. cagnagella larvae from plants with 0.1% extract
on concerns regarding health effects from long-term formulations. However, larger hard-bodied coleopter-
exposure (Khera et al. 1982, Betarbet et al. 2000). ans, such as L. decemlineata and P. japonica adults,
Plants with an established record for culinary or require higher doses, probably due to their relatively
medicinal use that therefore offer a safer starting ma- greater body mass and the difÞculty of penetrating the
terial have been evaluated in terms of their potential thicker cuticle. A post hoc hypothesis worthy of future
application as insecticides. For the current study, examination is larger larval insects are more sensitive
these plant compounds were considered not as leads to pepper treatments than smaller ones, regardless of
for synthetic insecticides, but for extract-based for- insect order. This is based on the observation that the
mulations that combine all of the co-occurring sec- size of the larval insects (Table 4) was inversely cor-
ondary plant compounds. The advantage of whole related with the toxicity (R2 ⫽ 0.913, Y ⫽ ⫺0.002X ⫹
extracts over single active ingredients was demon- 0.091): the smaller P. viburni were less sensitive than
strated by Feng and Isman (1995), who showed that the much larger M. americanum and N. sertifer. Al-
resistance development occurred with pure azadirac- though concentrations of 0.5% P. nigrum were re-
tin alone but not neem seed extract containing nu- quired to knockdown both P. japonica and L. decem-
merous compounds besides azadirachtin. From the lineata adults, this is still a practical concentration for
regulatory perspective, this complicates the chemistry botanical insecticides.
but simpliÞes the processing and allows for a unique A more suitable explanation of the greater sensitiv-
mixture of actives. The results of the current study ity of lepidopterans and the European pine sawßy
agree with many of the previous studies (Miyakado et larvae may be greater absorption through the cuticle
al. 1979, 1980; Sighamony et al. 1986; Baier and Web- compared with the coleopteran larvae. Based upon
ster 1992; Ivbijaro et al. 1993; Mbata et al. 1995; Kéṏta visual observations the coleopteran, P. viburni, had a
et al. 2000) in terms of the promising potential efÞcacy thicker, tougher cuticle. Structural difference in insect
found in P. nigrum and P. guineense extracts. cuticle between different species has been docu-
P. nigrum extracts were found to provide excellent mented as the reason for different rates of penetration
knockdown of the lepidopteran species and European (Smagghe et al. 1997, Teal et al. 1999). This was ob-
pine sawßy tested. The larvae were sensitive to P. served with cuticle preparations from the adult to-
nigrum and P. guineense treatments ⬍0.1% if applied as bacco budworm, Heliothis virescens (F.), and the adult
a contact insecticide, although some consumption of American cockroach, Periplaneta americana (L.). The
treated plant material cannot be discounted. Both M. obviously thicker cockroach cuticle had a slower pen-
americanum and N. sertifer are pests of ornamental etration rate compared with the moth (Teal et al.
trees and shrubs (Johnson and Lyon 1994) and thus 1999). When the toxicity between the adult coleopter-
could be controlled quickly by the homeowner when ans are compared, cuticle thickness may not play as
August 2004 SCOTT ET AL.: Piper EXTRACTS FOR CONTROL OF HOME AND GARDEN PESTS 1399

Fig. 7. Mean number of A. vittatum adults ⫹ SE found on cucumber plants after two 96-h choice test. (A) Plants treated
with water (control), formulation control (EC), and 0.5% P. nigrum. (B) Plants treated with water (control), formulation
blank (EC), and 0.1% P. nigrum. The treatment means with the same letter are not signiÞcantly different (TukeyÕs test, P ⬎
0.05).

great a factor as size (R2 ⫽ 0.967, Y ⫽ 0.021X ⫹ 0.03) ered safe for repeated daily applications because no
because the larger P. japonica and L. decemlineata phytotoxicity was noted at concentrations ⬍0.5%.
required a 5 times higher dose compared with the In the current study, A. vittatum adults were de-
proportionally smaller A. vittatum (Table 4). terred from cucumber plants with a 0.1% P. nigrum
The use of Piper extracts for both antifeedant and spray for a 4-d period (Fig. 7A and B). In practical
oviposition deterrent activity has been well docu- application, this may prevent or reduce the infestation
mented for stored grain insects (Ivbijaro 1990, Ekesi of the Cucurbitaceae roots and fruit by the larvae as
2000, Lale and Alaga 2001). In the current study, re-
well as the spread of the cucumber mosaic virus. An-
pellent activity by Piper extracts alone or in combi-
nation with other botanicals was observed against tifeedant activity also was noted for both P. nigrum and
some common garden insect pests. The most promis- P. guineense as low as 0.01% in a F. auricularia feeding
ing results were seen when 0.125% P. nigrum treat- trial, but this was attributed as much to the formulation
ments protected lily plants. The majority of L. lilii as to the Piper constituents (unpublished data). The
larvae moved or dropped off plants and their feeding repellent effect was conÞrmed when 50 mg of P.
was reduced (Fig. 4). This treatment level is consid- guineense per 30 cm2 repelled the red ßour beetle,
1400 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 97, no. 4

Fig. 8. Mean number of O. nubilalis larvae ⫾ SE found on bell pepper plants 11 d after treatment with water (control),
combined C. odorata and 0.03 or 0.1% Dillapiol (Ced ⫹ Dill), and 0.1 or 0.05% P. tuberculatum (P. tub). The treatment means
with the same letter are not signiÞcantly different (TukeyÕs test, P ⬎ 0.05).

Tribolium castaneum (Herbst), from treated paper The repellent effect of several nonhost volatiles was
discs (Lale and Alaga 2001). tested (Held et al. 2003) to determine whether rose
In the current study, oviposition by O. nubilalis on plants could be protected from P. japonica. None of
green pepper plants was reduced compared with con- the treatments designed to mask the host plant vola-
trols by a 0.5% P. guineense spray for a 4-d period under tiles [red cedar, Juniperus virginiana L.; Osage orange,
greenhouse conditions (Fig. 9). Similarly, studies with Maclura pomifera (Raf.) Schneid; ginko, Ginkgo biloba
P. guineense and A. sativum were equally effective at L.; red pepper, Capsicum frutescens L.; fennel seeds,
reducing the egg hatch of the legume pod borer, Ma-
Foeniculm vulgare Miller; and spearmint, Mentha spi-
ruca vitrata F. (Ekesi 2000), and oviposition of C.
maculatus was reduced by 2 and 3 ml P. guineense/kg cata L.] were effective at repelling P. japonica. Simi-
cowpea seeds compared with control and neem seed larly, the current study determined that P. japonica
oil (Ivbijaro 1990). This suggests that not only would given a choice between P. nigrum-treated roses and
P. guineense extracts reduce lepidopteran pest ovipo- controls would still feed on the treated plants (Fig. 6).
sition but also the hatching success of any eggs that Based upon these results P. japonica does not seem to
were placed on treated leaves. be a pest insect that Piper extracts would affect greatly.

Fig. 9. Mean number of O. nubilalis egg masses found on bell pepper plants 96 h after treatment with either water
(control), formulation blank (EC), 0.5% P. guineense (0.5% Pg), C. citratus (Ropel), or A. sativum (Garlic Barrier). The
treatment means with the same letter are not signiÞcantly different (TukeyÕs test, P ⬎ 0.05).
August 2004 SCOTT ET AL.: Piper EXTRACTS FOR CONTROL OF HOME AND GARDEN PESTS 1401

Table 4. Relationship between P. nigrum toxicity and insect by P. nigrum concentrations in the same range as those
mean dry weight required to knockdown other phytophagous insect
species (Tables 2 and 3). Thus, ladybird beetles, con-
Insect Order LC50 (%) Mean wt. (mg)
sidered beneÞcial and popular biocontrol species,
M. americanum Lepidoptera 0.018 35.9 could be affected when pests are being targeted. Cer-
larvae
N. sertifer larvae Hymenoptera 0.046 17.8
tainly, if H. convergens are sprayed directly with P.
Y. cagnagella larvae Lepidoptera 0.075 2.3 nigrum concentration between 0.1 and 0.5%, they will
P. viburni larvae Coleoptera 0.103 0.4 be susceptible. However, the risk to these predators
A. vittatum adults Coleoptera 0.103 4.1 exposed to homopterans is probably lessened consid-
L. decemlineata adult Coleoptera 0.498 20.1
P. japonica adults Coleoptera 0.532 25.7
ering that the extracts degrade quickly and ladybird
beetles do not feed on treated foliage. Therefore, in an
integrated pest management context, gardeners could
According to Torto et al. (1992), the most potent apply the P. nigrum extracts to knock down the pest
amide in P. nigrum tested against sorghum stem borer insect population, wait several hours for the actives to
Chilo partellus Swinhoe is piperine. It was suggested degrade under full sunlight, and then release H. con-
that the methylenedioxybenzene (MDP) group com- vergens.
mon to the piperamide molecules is an important
factor in antifeedant activity. In both P. nigrum and P.
Recommendations for Successful Insect Control by
guineense extracts, piperine is the major amide present
Using Piper Extracts
(Scott et al. 2002), although several other active
amides are present with the MDP group. In terms of a beneÞcial insect control alternative for
Keeping larvae off the plants is an environmental organic growers and home gardeners, this study found
and practical beneÞt of the Piper extracts especially that Lepidoptera and phytophagous Hymenoptera
because the larvicidal effects of these compounds on could be controlled with P. nigrum extracts at ⬍0.1%.
lepidopteran larvae feeding inside the plant are low. Other important garden vegetable and ornamental
As was observed in this study, P. tuberculatum at 0.1% pests such as L. lilii and P. viburni larvae, and A.
did not repel or effect O. nubilalis larvae placed on vittatum adults were controlled with a concentration
treated green peppers 1 d after spraying (Fig. 9). range between 0.1 and 0.2%. P. nigrum extracts can
However, C. odorata was more effective as a result of knock down adult L. decemlineata at 0.5%; however,
the systemic activity recognized for extracts contain- gardeners would need to scout the Þeld and spot spray
ing liminoids such as azadirachtin from A. indica (Gag- plants when the larvae and adults become active.
non 1992). When the same trial was repeated with Because the half-life of piperine and other piper-
double the concentration of P. nigrum and P. amides is ⬍1 h under full sunlight, residual activity on
guineense, no signiÞcant effect on O. nubilalis survival plant surfaces is very short. Repellent effects were
was noted (unpublished data). observed, but due to the short half-life of active com-
Both a disadvantage and advantage of using a Piper ponents (Scott et al. 2003), daily applications would be
extract is the short residual activity, especially under necessary for plants exposed to full sunlight. The for-
full sunlight. This was further documented under UV mulations prepared and tested indicate that knock
radiation as evidenced in Fig. 1. It was shown that down of insects requires only a short residual effect
under Þeld conditions when P. nigrum was applied to but care should be taken to apply them during periods
deter L. decemlineata adults from feeding on potato of less intense sunlight. Repeated sprays at concen-
plants, there was a rapid loss in residual antifeedant trations below 0.1% would not harm the plant but
activity within 3 h after application, corresponding to would not be practical for large scale agriculture. Sim-
an increase in leaf damage (Scott et al. 2003). Thus, ilarly, concentrations of ⱖ1% have been found to be
Piper formulations, unless prepared with a sunscreen phytotoxic under greenhouse conditions; however,
or sunblock to extend the life of the active ingredients, turfgrass was not harmed by 4% P. nigrum well watered
will not provide an adequate deterrent unless the after application (unpublished data). Treatments to
plants are in shaded areas. Applications could be timed protect shaded plants or when used indoors or in
for later in the day to work as repellents for night- greenhouses would likely extend the residual activity
ßying or -feeding insects or possibly slugs (Mollusca: and thus the antifeedant and oviposition deterrent
Gastropoda). This may explain why Piper extracts effect observed.
have been tested mainly against stored-product in- Piper extracts, like other insecticides, can be haz-
sects, because sunlight is not a factor in the degrada- ardous unless the applicator takes precautions: use of
tion, and residual activity has been observed for up to safety glasses, nose and mouth mask, and gloves. Piper
1 mo (Sighamony et al. 1986, Kéṏta et al. 2000, Ashamo actives are known irritants (Sigma Aldrich 2002), but
and Odeyemi 2001). However, the use of Piper ex- the risk to human health is much reduced because the
tracts, primarily for their knockdown and acute tox- active components have had a safe history as food
icity, does not require the actives to remain on the additives and spices, and the odor of these extracts is
plant surface for longer than the time exposed insects familiar to most people. This, therefore, indicates that
need to absorb them. P. nigrum extracts will not only be beneÞcial as bo-
Another potential disadvantage is the effect upon tanical insect control products but also will provide a
nontarget insect species. H. convergens were affected safe alternative to conventional synthetic insecticides.
1402 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 97, no. 4

Acknowledgments Hubert, J. J., and E. M. Carter. 1990b. COMLC50: a program

in BASIC which tests for the equality of 2 or more LC50s.
We thank R. Wheeler and B. Douglas (Agriculture & Department of Mathematics and Statistics, University of
Agrifood Canada, Central Experimental Farm, Ottawa, On- Guelph, Guelph, Ontario, Canada.
tario) for L. lilli; G. Bushell (Gloucester Allotment Garden, Isman, M. B. 1994. Botanical insecticides. Pestic. Outlook 5:
Ottawa) for L. decemlineata and A. vittatum; D. McCord and 26 Ð30.
R. Jansen (Rideau Hall, Ottawa) for P. japonica; H. Pearson Ivbijaro, M. F. 1990. The efÞcacy of seed oils of Azadirachta
(Kemptville Agricultural College, Kemptville, Ontario) for Y.
indica A. Juss and Piper guineense Schum and Thonn on
cagnagella and P. viburni; D. Bruce and S. Darling (Ottawa)
the control of Callosobruchus maculates F. Insect Sci.
for L. lilii; The Log Farm (Ottawa) for P. viburni; Oblate
Appl. 11: 149 Ð152.
Mission (Arnprior, Ontario) for R. majalis; A. Carpentier
Ivbijaro, M. F., V. C. Umeh, and H.J.W. Mutsars. 1993. Lab-
(Gatineau, Québec) for B. leucopterus hirtus; I. McGregor, B.
oratory toxicity of the crude extracts of Piper guineense
Innouye, and Ottawa Community Garden Networks for as-
Schum and Thonn, Azadirachta indica A. Juss and Parkia
sistance in locating other insects; E. Bruggink (Carleton Uni-
clappertoniana (Jacq) to the termites Macrotermes ni-
versity greenhouses) for horticultural assistance; A. Rao, J.
Livesey, and F. Duval (Biology Department, University of geriensis (Sjostedt) (Isoptera: Termitidae). Insect Sci.
Ottawa) for technical assistance; and N. Cappuccinno for Appl. 14: 229 Ð233.
reviewing the manuscript. Funding was provided by Ontario Javier, P. A., and B. Morallo-Rejesus. 1986. Insecticidal ac-
Ministry of Science and Technology, Canadian Organic tivity of black pepper (Piper nigrum L.) extracts. Philipp.
Growers, and Ontario Graduate Scholarship. Entomol. 6: 517Ð525.
Johnson, W. T., and H. H. Lyon. 1994. Insects that feed on
trees and shrubs. Comstock Publishing Associates, Cor-
nell University Press, Ithaca, NY.
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