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Insect Pests
                                                                                                                                                                             May 2007
                                                                                                                                                                                 IP-27




                   Using Clovers as Living Mulches

        To Boost Yields, Suppress Pests, and Augment Spiders

                     in a Broccoli Agroecosystem

                                           Cerruti R2 Hooks,a Raju R. Pandey,b and Marshall W. Johnsonc

    a
     CTAHR Department of Plant and Environmental Protection Sciences; bHimalayan College of Agricultural Sciences and Technology, 

                         Kathmandu, Nepal; cDepartment of Entomology, University of California, Riverside




Summary                                                                                            economically important crop with an undersown plant
A field study was conducted to examine the influence of                                            species that has no direct market value but is used to di­
intercropping broccoli (Brassica oleracea L.) with three                                           versify the agroecosystem or influence the main crop.
living mulches on caterpillar pest and spider densities                                                 The impact on insect pest densities of undersowing
and crop yield. Broccoli was grown in bare ground or                                               vegetable crops with living mulches has been examined
intercropped with strawberry clover (Trifolium fra-                                                mainly in Brassica crops (Asman et al. 2001; Hooks
giferum L.), white clover (Trifolium repens L.), or yel­                                           and Johnson 2001, 2002). In most of these studies,
low sweetclover (Melilotus officinalis L.). Lepidopteran                                           fewer insect pests were found on interplanted vegetable
(butterfly and moth) eggs and caterpillar densities were                                           crops (Hooks and Johnson 2003 and references therein).
significantly greater on broccoli in bare-ground plots                                             However, Brassica crops are typically slow growing
compared with broccoli intercropped with clover dur­                                               and do not compete well with background vegetation.
ing the late broccoli growth cycle. More spiders were                                              Therefore, most of these studies reported significant
found on bare-ground broccoli during early crop growth;                                            yield reductions, possibly caused by interspecies com­
however, during the later growth period, spider counts                                             petition.
were significantly higher on broccoli in intercropped                                                  Several strategies may be implemented to limit com­
plots. The number of insect contaminants found in                                                  petition among Brassica crops and their companion
harvested broccoli crowns were significantly less in                                               plants. These strategies may include
intercropped than in bare-ground broccoli plots. The                                               •	 proper fertilization and irrigation of the main crop
weight of harvested crowns was similar in intercropped                                             •		 use of vigorous or rapidly growing crop cultivars
and bare-ground habitats.                                                                          •		 optimal spacing between the main crop and compan­
                                                                                                       ion plants
Introduction                                                                                       •		 use of less competitive background plants (e.g., low
An established cover crop that is interplanted and grown                                               canopy height)
with an annual row crop is known as a living mulch.                                                •	 timely planting of the main crop and companion plant
Living mulches can provide many benefits to a crop­                                                •		 planting the intercrop at a lower seed rate or in nar­
ping habitat, including weed control, reduced erosion,                                                 rower strip
enhanced fertility, and improved soil quality (Lanini et                                           •		 suppression of the intercrop (e.g., mowing, reduced
al. 1989). However, recent studies have shown that when                                                fertilization and irrigation) at critical times
living mulches are undersown with a vegetable crop                                                 •	 use of a self-suppressing companion plant (i.e., one
they can also help reduce injury imposed by insect pests                                               that dies during the critical period of crop growth)
(Costello and Altieri 1995, Hooks et al. 1998, Hooks and                                           •		 use of non-crop borders surrounding the field crop.
Johnson 2001). Undersowing is the intercropping of an

Published by the College of Tropical Agriculture and Human Resources (CTAHR) and issued in furtherance of Cooperative Extension work, Acts of May 8 and June 30, 1914, in coopera­
tion with the U.S. Department of Agriculture. Andrew G. Hashimoto, Director/Dean, Cooperative Extension Service/CTAHR, University of Hawai‘i at Mänoa, Honolulu, Hawai‘i 96822.
An equal opportunity/affirmative action institution providing programs and services to the people of Hawai‘i without regard to race, sex, age, religion, color, national origin, ancestry, dis­
ability, marital status, arrest and court record, sexual orientation, or status as a covered veteran. CTAHR publications can be found on the Web site <http://www.ctahr.hawaii.edu/freepubs>.
UH–CTAHR                                                                                              IP-27 — May 2007



Each approach should be viewed with caution and               Arthropod census
weighed against its potential to negate any positive          Caterpillar pest stages and predators on broccoli plants
benefits of pest suppression.                                 were sampled at 7-day intervals until the harvest period.
   This publication describes a field experiment that         Sampling was stratified according to plant structure (5
was part of a continuing effort to refine management of       upper, 10 middle, and 10 lower positioned leaves). Egg,
lepidopteran pests in broccoli by mixed cropping and          caterpillar, and pupa stages of the diamondback moth,
undersowing it with other plant species. The primary pur­     Plutella xylostella L., imported cabbageworm, Pieris
pose of this study was to determine if specific undersown     rapae L, and cabbage looper, Trichoplusia ni Hübner,
living mulches would have the ability to suppress lepi­       were counted separately.
dopteran pest densities without reducing crop yields to
economically unacceptable levels. We focused on cover         Broccoli yield
crops in the genus Trifolium partially because of their       The diameter and weight of 16 broccoli crowns, chosen
low-growing structural similarities to yellow sweetclover.    at random from the interior rows of each plot, were mea­
Additionally, because spiders were recurrently observed       sured at harvest time. The heads were then completely
preying on various lepidopteran caterpillars (Hooks and       dissected and examined for insects, insect parts, and as­
Johnson 2002), their densities were compared among            sociated contaminants (e.g., frass, webbing, cocoon).
undersown and bare-ground broccoli.
                                                              Statistical analysis
                                                              The effects of habitat type on the experimental param­
Materials and methods
                                                              eters were analyzed using analysis of variance (Proc
Experiment layout                                             GLM, SAS Institute, Cary, NC 1990) and predetermined
The field experiment was conducted during the summer          orthogonal comparisons to separate mean differences.
2000 at the University of Hawai‘i at Mänoa’s Poamoho          Within the model, the following predetermined contrasts
Research Station on O‘ahu. The four cropping systems          were conducted: broccoli-clovers vs. monoculture; true
examined were broccoli plants undersown with: (1) yel­        clovers (white and strawberry) vs. yellow sweetclover;
low sweetclover (Melilotus officinalis L.) seeded at 72 g     and strawberry clover vs. white clover. Because eggs of
per row, (2) strawberry clover cv. O’Conners (Trifolium       moths and butterflies were acutely low, all species were
fragiferum L. seeded at 58 g per row, (3) white clover        pooled together by stage (e.g., egg, larvae) prior to final
cv. New Zealand (Trifolium repens L.) seeded at 54 g          analysis. The criteria for significance was P < 0.05.
per row, and (4) broccoli monoculture used as a check
treatment. Experimental plots were 11 m x 11 m with
                                                              Results
each treatment replicated four times and arranged in a
randomized complete block design.                             Clover growth
   The living mulches were sown on 10 July 2000 in all of     The yellow sweetclover canopy expanded over the soil
the undersown plots. Undersown plots contained 10 rows        surface faster than the other undersown living mulches.
of broccoli with 11 rows of the living mulch species and      Subsequently, the amount of exposed soil surface area
monoculture plots contained 10 rows of broccoli. Broccoli     between broccoli rows was significantly less in broc­
seedlings, cv. Liberty (Petoseed Co., Saticoy, California),   coli–yellow sweetclover contrasted with broccoli under­
were grown for 5 weeks in the greenhouse before being         sown with strawberry clover and white clover on each
manually transplanted on September 18 and 19.                 sampling date. Strawberry clover canopy development
                                                              occurred at the slowest rate and was significantly less
Clover growth                                                 than that of white clover on most dates.
The spread of the clover canopies was monitored weekly
beginning 16 days after broccoli planting (DAP) until         Arthropod census
broccoli harvest. Five areas between two adjacent clover      Effect of clovers on leidopteran pest densities. Lepi­
rows in each plot were randomly selected, excluding           dopteran populations were low during the experiment.
border rows, and the distance of exposed soil surface (not    Approximately 85, 10, and 5 percent of the lepidopteran
covered by clover canopy) between rows was measured.          fauna observed were P. rapae, T. ni, and P. xylostella,

2
UH–CTAHR                                                                                                                       IP-27 — May 2007



Figure 1. Mean population densities of lepidopteran (a) eggs and (b) larvae in bare-ground broccoli (BG); broccoli–
strawberry clover intercrop (SC); broccoli–white clover intercrop (WC); and broccoli–yellow sweetclover intercrop (YSC).
                  Figure 1.
*** indicates intercrops significantly less than bare ground; s indicates (SC) significantly less than (WC); w indicates (WC)
significantly less than (SC); y indicates (YSC) significantly less than (SC + WC); and ns means no significant differences
exist (P > 0.05).
                                                        0.7
                                                                    BG          eggs                                   A
                                                        0.6                                   ***
                                                                    SC
                                                        0.5         WC
                 Number of lepidopteran per leaf leaf




                                                                    YSC
                                                        0.4
                  Number of lepidopteran per




                                                        0.3                                             s        ***
                                                              ns                   ns
                                                        0.2           ns
                                                                           ns
                                                        0.1
                                                         0

                                                        0.8                                                            B
                                                                                 larvae                           ns
                                                        0.7
                                                        0.6
                                                        0.5
                                                        0.4           w
                                                                                                        ***
                                                        0.3                 y      *** y      ***
                                                        0.2
                                                        0.1   ***
                                                         0

                                                              16      23   30      37         44        51       58
                                                                           Days after planting
                                                                           Days After Planting


respectively. From early to mid-season, no significant                                 mordax L. Koch (Clubionidae) composed approximately
differences were detected in the abundance of eggs                                     77, 13, 7, and 3 percent of the spider fauna, respectively.
among broccoli habitats (Figure 1a). However, during                                   Significantly fewer spiders were encountered on broccoli
the late broccoli growth cycle, more eggs were found                                   plants with clovers compared with bare-ground broccoli
in monoculture than in undersown broccoli. Similarly,                                  during the initial three sampling dates. However, during
more caterpillars were recorded in monoculture com­                                    the later part of the broccoli growth cycle, this trend re­
pared with undersown broccoli from mid- to late season                                 versed, and more spiders were found on broccoli plants
(Figure 1b).                                                                           in clover than in bare-ground plots (Figure 2).
   Impact of clovers on spider abundance. At the ex­                                      Additionally, fewer spiders were found on broccoli
perimental site, four spider species frequently inhabit                                grown with the true clovers (e.g., strawberry clover,
Brassica plants. Nesticodes (= Theridion) rufipes Lucas                                white clover) compared with plants grown with yellow
(Theridiidae), Neoscona oaxacensis Keyserling (Ara­                                    sweetclover during mid-season. Fewer spiders were also
neidae), Oxyopes sp. (Oxyopidae), and Cheiracanthium                                   observed on broccoli plants undersown in strawberry

                                                                                                                                                3
UH–CTAHR                                                                                                          IP-27 — May 2007



Figure 2. Mean population densities of spiders in bare-ground broccoli (BG); broccoli–strawberry clover intercrop (SC);
                Figure 2.
broccoli–white clover intercrop (WC); broccoli–yellow sweetclover intercrop (YSC). * indicates (BG) significantly less
than intercrops; *** indicates intercrops significantly less than (BG); c indicates (SC + WC) significantly less than (YSC);
s indicates (SC) significantly less than (WC); and ns means no significant differences exist (P ≥ 0.05).


                                               1
                Number of spiders per leaf


                                                          BG
                 Number of spiders per leaf




                                              0.8         SC                       *cs
                                                          WC
                                                                    ***
                                                          YSC
                                              0.6
                                                                                                *         s
                                                            ***             c
                                              0.4   ***

                                              0.2

                                               0
                                                    16      23      30     37       44         51        58
                                                                    Days after planting
                                                                  Days After Planting




clover compared with white clover from mid- to late                        reducing crop yields to unacceptable levels. We found
season.                                                                    that the number of insect contaminants per broccoli
                                                                           crown was significantly reduced on plants undersown
Crown contamination                                                        with strawberry clover and white clover compared to
At harvest, broccoli crowns were infested with various                     bare-ground broccoli without causing any yield reduc­
stages of Pieris rapae, Trichoplusia ni, and Plutella                      tions. Additionally, spider densities found on broccoli
xylostela (Table 1). T. ni caterpillars and pupae were the                 plants seemed to be influenced by the amount of clover
most abundant lepidopteran contaminants encountered in                     canopy. As the clover canopies expanded and approached
broccoli heads. However, significantly more individuals                    the broccoli plants, more spiders were found on the
of all three species were found in crowns harvested from                   broccoli foliage.
bare-ground broccoli compared to undersown broccoli.
                                                                           Impact on spider abundance
Crop yield                                                                 During the early part of the season, the living mulches
The largest crowns by diameter and mass were harvested                     may have negatively influenced biological control activ­
from broccoli undersown with strawberry clover or                          ity on broccoli plants by serving as a “sink” for spiders.
white clover (Table 2). These crowns weighed signifi­                      Spiders may have preferred the micro-environment
                                                                                                                                  2
cantly more than those harvested from broccoli–yellow                      and prey selection within the clovers and thus did not
sweetclover plots. Yellow sweetclover plots contained                      colonize neighboring broccoli plants. Similar observa­
the smallest crowns, by weight.                                            tions were made in previous field experiments in which
                                                                           fewer spiders were found on broccoli plants intercropped
Discussion                                                                 with peppers or yellow sweetclover, but as the season
The purpose of this study was to determine if undersown                    progressed these differences diminished.
clovers could reduce lepidopteran pest densities without

4
UH–CTAHR                                                                                                                   IP-27 — May 2007



Table 1. Mean number of lepidopterans per broccoli head in four broccoli habitats during summer 2000 (mean ± SE).

Habitat                          Pieris rapae            Trichoplusia ni           Plutella xylostella             Total

Bare ground                      0.1 + 0.0              0.8 + 0.10                    0.13 + 0.0            0.87 + 0.0

Broccoli-SC                      0.08 + 0.04              0.10 + 0.04                    0.00 + 0.00            0.18 + 0.06

Broccoli-WC                      0.0 + 0.03              0.00 + 0.00                    0.02 + 0.02            0.07 + 0.03

Broccoli-YSC                     0.0 + 0.03              0.12 + 0.0                    0.03 + 0.02            0.20 + 0.06

Contrast1                                                                      P-value

BG vs. LMs                            0.03                     0.01                          0.006                 0.0001
TCs vs. YSC                           0.07                     0.4                          0.4                   0.39
SC vs. WC                             0.0                     0.62                          0.67                   0.20
1
 SC (strawberry clover), WC (white clover), YSC (yellow sweetclover), BG (bare ground, broccoli monoculture); LMs (living mulches) includes
broccoli-SC, broccoli-WC, and broccoli-YSC; TCs (true clovers) includes broccoli-SC and broccoli-WC.




Conclusion
                                                                           Table 2. Mean head size per broccoli plant in four habitats
Using undersown living mulches seems to be promising                       during summer 2000.
in reducing lepidopteran pest densities and increasing
                                                                                                                  Broccoli parameters1
the activity of spiders in broccoli plantings. In this study,                                                         (mean ± SE)
white clover appeared to be more suited for the broccoli
                                                                           Habitat                          Diameter (cm)       Weight (kg)
system. White clover expands over the soil surface faster
than strawberry clover and may therefore be a better                       Bare ground                        13.0 + 0.3       0.3 + 0.01
weed suppressor. For those farmers looking to create
                                                                           Broccoli-strawberry clover         14.7 + 0.3       0.40 + 0.02
more sustainable cropping practices or practicing organic
farming, undersowing may be a valuable addition to their                   Broccoli-white clover              14.4 + 0.24       0.39 + 0.01
crop production practices.                                                 Broccoli-yellow sweetclover 13.7 + 0.28              0.33 + 0.01
   This field trial showed that it is possible to lower insect
pest density while maintaining crop quality and yield.                     Effect2                                         P -Values

However, insect pest management is only one potential                      Planned contrast
                                                                           BG vs. LMs                              0.0002            0.11

benefit of using leguminous living mulches. Other po­                      TCs vs. YSC                             0.03              0.000
tential benefits not examined during this study include                    SC vs. WC                               0.71              0.1
nematode and weed suppression and enhancement of
soil nitrogen. However, before an undersown compan­                        1
                                                                            Multiply cm by 0.394 to obtain inches, and multiply kg by 2.2 to 

                                                                           obtain pounds.

ion plant is chosen for insect suppression purposes, its                   2
                                                                            SC (strawberry clover), WC (white clover), YSC (yellow sweetclover), 

impact on other important organisms associated with the                    BG (bare ground, broccoli monoculture); LMs (living mulches) include 

crop should be considered.                                                 broccoli-SC, broccoli-WC, and broccoli-YSC; TCs (true clovers) 

                                                                           include broccoli-SC and broccoli and broccoli-WC.


Acknowledgements
The authors wish to thank the crew at the Poamoho                          References and further reading
Research Station for assisting in the field and Dr. Raju                   Asman, K., B. Ekbom, and B. Rämert. 2001. Effect of
Pandey for his notable contributions to this study. This                     intercropping on oviposition and emigration behavior
project was funded by the USDA/CSREES Special                                of the leek moth (Lepidoptera, Acrolepiidae) and the
Grant for Tropical and Subtropical Agriculture Research                      diamondback moth (Lepidoptera, Plutellidae). Envi­
(T-STAR).                                                                    ronmental Entomology 30: 288–294.
UH–CTAHR                                                                                        IP-27 — May 2007



Costello, M.J., and M.A. Altieri. 1995. Abundance,          flora diversification. Annals of Applied Biology 138:
  growth rate and parasitism of Brevicoryne brassicae       269–280.
  and Myzus persicae (Homoptera, Aphididae) on broc­      Hooks, C.R.R., and M.W. Johnson. 2002. Lepidopteran
  coli grown in living mulches. Agriculture, Ecosystems     pest populations and crop yields in row intercropped
  and Environment 52: 187–196.                              broccoli. Agriculture and Forest Entomology 4:
Hooks, C.R.R., H.R. Valenzuela, and J. Defrank. 1998.       117–126.
  Incidence of pests and arthropod natural enemies in     Lanini, W.T., D.R. Pittenger, W.L. Graves, F. Munoz, and
  zucchini grown with living mulches. Agriculture,          H.S. Agamalian. 1989. Subclovers as living mulches
  Ecosystems and Environment 69: 217–231.                   for managing weeds in vegetables. California Agri­
Hooks, C.R.R., and M.W. Johnson. 2001. Broccoli             culture. Nov.–Dec., p. 25–27.
  growth parameters and level of head infestations        SAS Institute. 1990. SAS User’s Guide: Statistics. Cary,
  in simple and mixed plantings: Impact of increased        NC.




6

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Using Clovers as Living Mulches to Boost Yields, Suppress Pests, and Augment Spiders in a Broccoli Agroecosystem

  • 1. Insect Pests May 2007 IP-27 Using Clovers as Living Mulches To Boost Yields, Suppress Pests, and Augment Spiders in a Broccoli Agroecosystem Cerruti R2 Hooks,a Raju R. Pandey,b and Marshall W. Johnsonc a CTAHR Department of Plant and Environmental Protection Sciences; bHimalayan College of Agricultural Sciences and Technology, Kathmandu, Nepal; cDepartment of Entomology, University of California, Riverside Summary economically important crop with an undersown plant A field study was conducted to examine the influence of species that has no direct market value but is used to di­ intercropping broccoli (Brassica oleracea L.) with three versify the agroecosystem or influence the main crop. living mulches on caterpillar pest and spider densities The impact on insect pest densities of undersowing and crop yield. Broccoli was grown in bare ground or vegetable crops with living mulches has been examined intercropped with strawberry clover (Trifolium fra- mainly in Brassica crops (Asman et al. 2001; Hooks giferum L.), white clover (Trifolium repens L.), or yel­ and Johnson 2001, 2002). In most of these studies, low sweetclover (Melilotus officinalis L.). Lepidopteran fewer insect pests were found on interplanted vegetable (butterfly and moth) eggs and caterpillar densities were crops (Hooks and Johnson 2003 and references therein). significantly greater on broccoli in bare-ground plots However, Brassica crops are typically slow growing compared with broccoli intercropped with clover dur­ and do not compete well with background vegetation. ing the late broccoli growth cycle. More spiders were Therefore, most of these studies reported significant found on bare-ground broccoli during early crop growth; yield reductions, possibly caused by interspecies com­ however, during the later growth period, spider counts petition. were significantly higher on broccoli in intercropped Several strategies may be implemented to limit com­ plots. The number of insect contaminants found in petition among Brassica crops and their companion harvested broccoli crowns were significantly less in plants. These strategies may include intercropped than in bare-ground broccoli plots. The • proper fertilization and irrigation of the main crop weight of harvested crowns was similar in intercropped • use of vigorous or rapidly growing crop cultivars and bare-ground habitats. • optimal spacing between the main crop and compan­ ion plants Introduction • use of less competitive background plants (e.g., low An established cover crop that is interplanted and grown canopy height) with an annual row crop is known as a living mulch. • timely planting of the main crop and companion plant Living mulches can provide many benefits to a crop­ • planting the intercrop at a lower seed rate or in nar­ ping habitat, including weed control, reduced erosion, rower strip enhanced fertility, and improved soil quality (Lanini et • suppression of the intercrop (e.g., mowing, reduced al. 1989). However, recent studies have shown that when fertilization and irrigation) at critical times living mulches are undersown with a vegetable crop • use of a self-suppressing companion plant (i.e., one they can also help reduce injury imposed by insect pests that dies during the critical period of crop growth) (Costello and Altieri 1995, Hooks et al. 1998, Hooks and • use of non-crop borders surrounding the field crop. Johnson 2001). Undersowing is the intercropping of an Published by the College of Tropical Agriculture and Human Resources (CTAHR) and issued in furtherance of Cooperative Extension work, Acts of May 8 and June 30, 1914, in coopera­ tion with the U.S. Department of Agriculture. Andrew G. Hashimoto, Director/Dean, Cooperative Extension Service/CTAHR, University of Hawai‘i at Mänoa, Honolulu, Hawai‘i 96822. An equal opportunity/affirmative action institution providing programs and services to the people of Hawai‘i without regard to race, sex, age, religion, color, national origin, ancestry, dis­ ability, marital status, arrest and court record, sexual orientation, or status as a covered veteran. CTAHR publications can be found on the Web site <http://www.ctahr.hawaii.edu/freepubs>.
  • 2. UH–CTAHR IP-27 — May 2007 Each approach should be viewed with caution and Arthropod census weighed against its potential to negate any positive Caterpillar pest stages and predators on broccoli plants benefits of pest suppression. were sampled at 7-day intervals until the harvest period. This publication describes a field experiment that Sampling was stratified according to plant structure (5 was part of a continuing effort to refine management of upper, 10 middle, and 10 lower positioned leaves). Egg, lepidopteran pests in broccoli by mixed cropping and caterpillar, and pupa stages of the diamondback moth, undersowing it with other plant species. The primary pur­ Plutella xylostella L., imported cabbageworm, Pieris pose of this study was to determine if specific undersown rapae L, and cabbage looper, Trichoplusia ni Hübner, living mulches would have the ability to suppress lepi­ were counted separately. dopteran pest densities without reducing crop yields to economically unacceptable levels. We focused on cover Broccoli yield crops in the genus Trifolium partially because of their The diameter and weight of 16 broccoli crowns, chosen low-growing structural similarities to yellow sweetclover. at random from the interior rows of each plot, were mea­ Additionally, because spiders were recurrently observed sured at harvest time. The heads were then completely preying on various lepidopteran caterpillars (Hooks and dissected and examined for insects, insect parts, and as­ Johnson 2002), their densities were compared among sociated contaminants (e.g., frass, webbing, cocoon). undersown and bare-ground broccoli. Statistical analysis The effects of habitat type on the experimental param­ Materials and methods eters were analyzed using analysis of variance (Proc Experiment layout GLM, SAS Institute, Cary, NC 1990) and predetermined The field experiment was conducted during the summer orthogonal comparisons to separate mean differences. 2000 at the University of Hawai‘i at Mänoa’s Poamoho Within the model, the following predetermined contrasts Research Station on O‘ahu. The four cropping systems were conducted: broccoli-clovers vs. monoculture; true examined were broccoli plants undersown with: (1) yel­ clovers (white and strawberry) vs. yellow sweetclover; low sweetclover (Melilotus officinalis L.) seeded at 72 g and strawberry clover vs. white clover. Because eggs of per row, (2) strawberry clover cv. O’Conners (Trifolium moths and butterflies were acutely low, all species were fragiferum L. seeded at 58 g per row, (3) white clover pooled together by stage (e.g., egg, larvae) prior to final cv. New Zealand (Trifolium repens L.) seeded at 54 g analysis. The criteria for significance was P < 0.05. per row, and (4) broccoli monoculture used as a check treatment. Experimental plots were 11 m x 11 m with Results each treatment replicated four times and arranged in a randomized complete block design. Clover growth The living mulches were sown on 10 July 2000 in all of The yellow sweetclover canopy expanded over the soil the undersown plots. Undersown plots contained 10 rows surface faster than the other undersown living mulches. of broccoli with 11 rows of the living mulch species and Subsequently, the amount of exposed soil surface area monoculture plots contained 10 rows of broccoli. Broccoli between broccoli rows was significantly less in broc­ seedlings, cv. Liberty (Petoseed Co., Saticoy, California), coli–yellow sweetclover contrasted with broccoli under­ were grown for 5 weeks in the greenhouse before being sown with strawberry clover and white clover on each manually transplanted on September 18 and 19. sampling date. Strawberry clover canopy development occurred at the slowest rate and was significantly less Clover growth than that of white clover on most dates. The spread of the clover canopies was monitored weekly beginning 16 days after broccoli planting (DAP) until Arthropod census broccoli harvest. Five areas between two adjacent clover Effect of clovers on leidopteran pest densities. Lepi­ rows in each plot were randomly selected, excluding dopteran populations were low during the experiment. border rows, and the distance of exposed soil surface (not Approximately 85, 10, and 5 percent of the lepidopteran covered by clover canopy) between rows was measured. fauna observed were P. rapae, T. ni, and P. xylostella, 2
  • 3. UH–CTAHR IP-27 — May 2007 Figure 1. Mean population densities of lepidopteran (a) eggs and (b) larvae in bare-ground broccoli (BG); broccoli– strawberry clover intercrop (SC); broccoli–white clover intercrop (WC); and broccoli–yellow sweetclover intercrop (YSC). Figure 1. *** indicates intercrops significantly less than bare ground; s indicates (SC) significantly less than (WC); w indicates (WC) significantly less than (SC); y indicates (YSC) significantly less than (SC + WC); and ns means no significant differences exist (P > 0.05). 0.7 BG eggs A 0.6 *** SC 0.5 WC Number of lepidopteran per leaf leaf YSC 0.4 Number of lepidopteran per 0.3 s *** ns ns 0.2 ns ns 0.1 0 0.8 B larvae ns 0.7 0.6 0.5 0.4 w *** 0.3 y *** y *** 0.2 0.1 *** 0 16 23 30 37 44 51 58 Days after planting Days After Planting respectively. From early to mid-season, no significant mordax L. Koch (Clubionidae) composed approximately differences were detected in the abundance of eggs 77, 13, 7, and 3 percent of the spider fauna, respectively. among broccoli habitats (Figure 1a). However, during Significantly fewer spiders were encountered on broccoli the late broccoli growth cycle, more eggs were found plants with clovers compared with bare-ground broccoli in monoculture than in undersown broccoli. Similarly, during the initial three sampling dates. However, during more caterpillars were recorded in monoculture com­ the later part of the broccoli growth cycle, this trend re­ pared with undersown broccoli from mid- to late season versed, and more spiders were found on broccoli plants (Figure 1b). in clover than in bare-ground plots (Figure 2). Impact of clovers on spider abundance. At the ex­ Additionally, fewer spiders were found on broccoli perimental site, four spider species frequently inhabit grown with the true clovers (e.g., strawberry clover, Brassica plants. Nesticodes (= Theridion) rufipes Lucas white clover) compared with plants grown with yellow (Theridiidae), Neoscona oaxacensis Keyserling (Ara­ sweetclover during mid-season. Fewer spiders were also neidae), Oxyopes sp. (Oxyopidae), and Cheiracanthium observed on broccoli plants undersown in strawberry 3
  • 4. UH–CTAHR IP-27 — May 2007 Figure 2. Mean population densities of spiders in bare-ground broccoli (BG); broccoli–strawberry clover intercrop (SC); Figure 2. broccoli–white clover intercrop (WC); broccoli–yellow sweetclover intercrop (YSC). * indicates (BG) significantly less than intercrops; *** indicates intercrops significantly less than (BG); c indicates (SC + WC) significantly less than (YSC); s indicates (SC) significantly less than (WC); and ns means no significant differences exist (P ≥ 0.05). 1 Number of spiders per leaf BG Number of spiders per leaf 0.8 SC *cs WC *** YSC 0.6 * s *** c 0.4 *** 0.2 0 16 23 30 37 44 51 58 Days after planting Days After Planting clover compared with white clover from mid- to late reducing crop yields to unacceptable levels. We found season. that the number of insect contaminants per broccoli crown was significantly reduced on plants undersown Crown contamination with strawberry clover and white clover compared to At harvest, broccoli crowns were infested with various bare-ground broccoli without causing any yield reduc­ stages of Pieris rapae, Trichoplusia ni, and Plutella tions. Additionally, spider densities found on broccoli xylostela (Table 1). T. ni caterpillars and pupae were the plants seemed to be influenced by the amount of clover most abundant lepidopteran contaminants encountered in canopy. As the clover canopies expanded and approached broccoli heads. However, significantly more individuals the broccoli plants, more spiders were found on the of all three species were found in crowns harvested from broccoli foliage. bare-ground broccoli compared to undersown broccoli. Impact on spider abundance Crop yield During the early part of the season, the living mulches The largest crowns by diameter and mass were harvested may have negatively influenced biological control activ­ from broccoli undersown with strawberry clover or ity on broccoli plants by serving as a “sink” for spiders. white clover (Table 2). These crowns weighed signifi­ Spiders may have preferred the micro-environment 2 cantly more than those harvested from broccoli–yellow and prey selection within the clovers and thus did not sweetclover plots. Yellow sweetclover plots contained colonize neighboring broccoli plants. Similar observa­ the smallest crowns, by weight. tions were made in previous field experiments in which fewer spiders were found on broccoli plants intercropped Discussion with peppers or yellow sweetclover, but as the season The purpose of this study was to determine if undersown progressed these differences diminished. clovers could reduce lepidopteran pest densities without 4
  • 5. UH–CTAHR IP-27 — May 2007 Table 1. Mean number of lepidopterans per broccoli head in four broccoli habitats during summer 2000 (mean ± SE). Habitat Pieris rapae Trichoplusia ni Plutella xylostella Total Bare ground 0.1 + 0.0 0.8 + 0.10 0.13 + 0.0 0.87 + 0.0 Broccoli-SC 0.08 + 0.04 0.10 + 0.04 0.00 + 0.00 0.18 + 0.06 Broccoli-WC 0.0 + 0.03 0.00 + 0.00 0.02 + 0.02 0.07 + 0.03 Broccoli-YSC 0.0 + 0.03 0.12 + 0.0 0.03 + 0.02 0.20 + 0.06 Contrast1 P-value BG vs. LMs 0.03 0.01 0.006 0.0001 TCs vs. YSC 0.07 0.4 0.4 0.39 SC vs. WC 0.0 0.62 0.67 0.20 1 SC (strawberry clover), WC (white clover), YSC (yellow sweetclover), BG (bare ground, broccoli monoculture); LMs (living mulches) includes broccoli-SC, broccoli-WC, and broccoli-YSC; TCs (true clovers) includes broccoli-SC and broccoli-WC. Conclusion Table 2. Mean head size per broccoli plant in four habitats Using undersown living mulches seems to be promising during summer 2000. in reducing lepidopteran pest densities and increasing Broccoli parameters1 the activity of spiders in broccoli plantings. In this study, (mean ± SE) white clover appeared to be more suited for the broccoli Habitat Diameter (cm) Weight (kg) system. White clover expands over the soil surface faster than strawberry clover and may therefore be a better Bare ground 13.0 + 0.3 0.3 + 0.01 weed suppressor. For those farmers looking to create Broccoli-strawberry clover 14.7 + 0.3 0.40 + 0.02 more sustainable cropping practices or practicing organic farming, undersowing may be a valuable addition to their Broccoli-white clover 14.4 + 0.24 0.39 + 0.01 crop production practices. Broccoli-yellow sweetclover 13.7 + 0.28 0.33 + 0.01 This field trial showed that it is possible to lower insect pest density while maintaining crop quality and yield. Effect2 P -Values However, insect pest management is only one potential Planned contrast BG vs. LMs 0.0002 0.11 benefit of using leguminous living mulches. Other po­ TCs vs. YSC 0.03 0.000 tential benefits not examined during this study include SC vs. WC 0.71 0.1 nematode and weed suppression and enhancement of soil nitrogen. However, before an undersown compan­ 1 Multiply cm by 0.394 to obtain inches, and multiply kg by 2.2 to obtain pounds. ion plant is chosen for insect suppression purposes, its 2 SC (strawberry clover), WC (white clover), YSC (yellow sweetclover), impact on other important organisms associated with the BG (bare ground, broccoli monoculture); LMs (living mulches) include crop should be considered. broccoli-SC, broccoli-WC, and broccoli-YSC; TCs (true clovers) include broccoli-SC and broccoli and broccoli-WC. Acknowledgements The authors wish to thank the crew at the Poamoho References and further reading Research Station for assisting in the field and Dr. Raju Asman, K., B. Ekbom, and B. Rämert. 2001. Effect of Pandey for his notable contributions to this study. This intercropping on oviposition and emigration behavior project was funded by the USDA/CSREES Special of the leek moth (Lepidoptera, Acrolepiidae) and the Grant for Tropical and Subtropical Agriculture Research diamondback moth (Lepidoptera, Plutellidae). Envi­ (T-STAR). ronmental Entomology 30: 288–294.
  • 6. UH–CTAHR IP-27 — May 2007 Costello, M.J., and M.A. Altieri. 1995. Abundance, flora diversification. Annals of Applied Biology 138: growth rate and parasitism of Brevicoryne brassicae 269–280. and Myzus persicae (Homoptera, Aphididae) on broc­ Hooks, C.R.R., and M.W. Johnson. 2002. Lepidopteran coli grown in living mulches. Agriculture, Ecosystems pest populations and crop yields in row intercropped and Environment 52: 187–196. broccoli. Agriculture and Forest Entomology 4: Hooks, C.R.R., H.R. Valenzuela, and J. Defrank. 1998. 117–126. Incidence of pests and arthropod natural enemies in Lanini, W.T., D.R. Pittenger, W.L. Graves, F. Munoz, and zucchini grown with living mulches. Agriculture, H.S. Agamalian. 1989. Subclovers as living mulches Ecosystems and Environment 69: 217–231. for managing weeds in vegetables. California Agri­ Hooks, C.R.R., and M.W. Johnson. 2001. Broccoli culture. Nov.–Dec., p. 25–27. growth parameters and level of head infestations SAS Institute. 1990. SAS User’s Guide: Statistics. Cary, in simple and mixed plantings: Impact of increased NC. 6