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North Central Regional

Publication 581

Biological control of insects


and other pests of greenhouse crops

Susan E. Rice Mahr


Raymond A. Cloyd
Daniel L. Mahr
Clifford S. Sadof
acknowledgements
This publication was funded, in part, by a
grant from the National Agricultural
Pesticide Impact Assessment Program of
the United States Department of
Agriculture. University of Wisconsin-
Extension and the College of Agricultural
and Life Sciences, University of
Wisconsin-Madison provided additional
support, as did the National Biological
Control Institute of USDA-APHIS.
We thank Dr. David R. Gillespie,
Agriculture Canada; Carol Glenister, IPM
Laboratories, Inc.; Dr. Kevin Heinz,
University of California, Davis; Dr. Richard
K. Lindquist, The Ohio State University;
Dr. Oscar Minkenberg, University of
Arizona; Dr. Lance Osborne, University of
Florida; Dr. Karen Robb, University of
California Cooperative Extension; Dr.
John A. Sanderson, Cornell University; Dr.
J. Les Shipp, Agriculture Canada; and Dr.
Kirk Smith for reviewing and providing
insightful comments on earlier versions
of the manuscript.
We extend sincere gratitude to those
who provided photographs of natural
enemies, without which this publication
would be much less useful. Their contri-
butions are acknowledged alongside
the individual photographs.
The support of the Cooperative
Extension Publishing Unit of University
of Wisconsin-Extension—especially
Susan Anderson, Linda Deith,
Jody Myer-Lynch, and Tim Steller—is
deeply appreciated.
Biological control of insects
and other pests of greenhouse crops

Susan E. Rice Mahr


Raymond A. Cloyd
Daniel L. Mahr
Clifford S. Sadof

University of Wisconsin-Extension, Cooperative Extension


preface
Why use biological control? Why this publication?
Since the late 1940s, much of the insect Biological control relies on living organ- research, or to “tinker with the system”
control in the United States has been isms that must have food and shelter to get optimum results. Therefore, we
based on the use of synthetic chemical and that interact, often in complex ways, provide background information on bio-
insecticides. Insecticides are relatively with the pests, the crop, and other envi- logical control in general, and on the
easy to use and have generally provided ronmental factors. The pest manager (a specific pests and their natural enemies
effective pest control; it is likely that they grower or crop consultant) should be that occur in greenhouse crops to help
will always be a component of pest man- able to recognize important natural you refine your pest management
agement programs. Unfortunately, insec- enemies, understand their needs, and program. More than 1,000 research,
ticides have some undesirable attributes. know how to maximize their effective- technical, and popular publications have
They usually present some degree of ness. A good understanding of the rela- been carefully reviewed and summa-
hazard to the applicator and other people tionships between pests, their natural rized for this publication. I hope
who may come in contact with them; enemies, and the environment is essen- Biological Control of Insects and Other
they can leave residues that some find tial for the successful implementation of Pests of Greenhouse Crops helps you plan
unacceptable; they can contaminate soil biological control. The need for this type and conduct successful biological
and water and affect wildlife, aquatic life, of knowledge is the rationale for this control.
and other nontarget organisms; they can publication. Daniel L. Mahr, Coordinator
interfere with beneficial organisms, such Most insects that attack greenhouse Extension Biological Control Programs
as pollinating insects and the natural crops are readily controlled by biological Department of Entomology
enemies of pests; and insects can develop means. This publication provides infor- University of Wisconsin-Madison
resistance to insecticides, effectively elim- mation on important natural enemies September 2001
inating those materials as pest manage- and offers practical advice on how to
ment options. For these reasons, there is use beneficial organisms for pest man-
growing interest among farmers, horti- agement in greenhouses. The publica-
culturists, and gardeners to explore and tion emphasizes the use of biological
adopt methods that reduce pesticide use. control, but it also tells how best to use
Biological control represents one alterna- cultural and least-toxic chemical controls
tive to the use of insecticides. Biological if it becomes necessary to supplement
control is the conscious use of living ben- biological control. Therefore, the publi-
eficial organisms, called natural enemies, cation is truly a manual for biologically
for the control of pests. Virtually all pests based integrated pest management.
have natural enemies, and many pests Chemical insecticides are fairly straight-
can be effectively controlled by forward to use—after identifying the
managing these natural enemies. pest problem, you can choose an appro-
Although biological control will not priate material, read the label, and follow
control all pests, it can be the foundation directions to achieve good control.
of an approach called integrated pest Because biological controls are based
management, which combines a variety upon living organisms that have specific
of pest control methods in an ecologi- needs and behaviors, it is not always
cally safe system. Biological control can easy to give a recipe to assure success.
be effective, economical, and safe, and it With commercially available natural
should be used more widely than it is enemies, there will be instructions as to
today. release rates and timing. However, with
other approaches to biological control, it
may be necessary to conduct your own
table of contents
Section 1
Understanding pests and their
biological control . . . . . . . . . . . . . . . . 1
The biology of insects . . . . . . . . . . . . . . 1
The natural enemies of insects . . . . . . . . . 5
The general approaches to
biological control . . . . . . . . . . . . . . 7
The economics of pest control . . . . . . . . . 9
Section 2
Practical approaches to the biological
control of greenhouse pests . . . . . . . . . 11
Aphids . . . . . . . . . . . . . . . . . . . . . . 14
Caterpillars . . . . . . . . . . . . . . . . . . . 26
Fungus gnats and shore flies . . . . . . . . . 30
Leafminers . . . . . . . . . . . . . . . . . . . 34
Mites . . . . . . . . . . . . . . . . . . . . . . . 41
Scales and mealybugs . . . . . . . . . . . . . 49
Thrips . . . . . . . . . . . . . . . . . . . . . . 60
Weevils . . . . . . . . . . . . . . . . . . . . . . 69
Whiteflies . . . . . . . . . . . . . . . . . . . . 72
Miscellaneous pests . . . . . . . . . . . . . . 84
Section 3
Overview of biological control
of greenhouse pests . . . . . . . . . . . . . 89
Contemporary biological control . . . . . . . 89
Integration of biological control of
all pests: Examples . . . . . . . . . . . . . 92
Additional reading . . . . . . . . . . . . . . . . . 94
Glossary . . . . . . . . . . . . . . . . . . . . . . . 96
Index . . . . . . . . . . . . . . . . . . . . . . . . . 98
Disclaimer
The complex interactions between a These references do not constitute
pest, its host crop, its natural enemies, recommendations but are included
various environmental factors, and to indicate potential future develop-
crop management practices make it ments or to point to areas of possible
difficult to provide exact recommen- experimentation by the grower. We
dations for the use of natural have made every attempt to be
enemies in biological control. accurate regarding the commercial
Therefore, the suggestions in this availability of natural enemies and
publication for the release of natural products at the time of publication.
enemies in augmentative biological However, as more natural enemies
control are guidelines rather than are proven to be effective, they will
specific recommendations. Some likely become commercially available.
experimentation on your part will be Check with biological control distrib-
necessary to achieve optimal pest utors to determine the best available
management using natural enemies natural enemies for your needs.
and other control approaches. References to products or companies
Some states have regulations regard- in this publication are for your con-
ing the shipment and release of com- venience and are not an endorse- Measurements
mercial natural enemies, and permits ment over similar products or compa- Many of the beneficial natural enemies
are required in some cases. If you are nies. You are responsible for using described in this publication are very
uncertain about such regulations, pesticides according to the manufac- small. Because of its accuracy, we
contact your state’s department of turer’s current label directions and include metric measurements when
agriculture for more information. federal and state laws. Follow label giving the size of organisms. If you
directions exactly to protect the envi- remember that 1 inch equals approxi-
In several cases we refer to experi-
ronment and people from pesticide mately 25 mm, you will be able to make
mental controls that have not been
exposure. Failure to do so violates the rapid conversions. You can also use the
fully evaluated in the United States,
law. accurate conversion scale below to get a
have not been commercialized, or, in
sense of insect size.
the case of pesticidal products, have
not been registered with the
Environmental Protection Agency. 10 20 30 40
MM

INCH 1

A note about scientific names


Most beneficial natural enemies do not
have common names. Therefore, we use
scientific names throughout the publi-
cation. To gain an understanding of the
nature and use of scientific names, read
the section entitled “Insect
Classification” on page 4.
SECTION

1
Understanding pests
and their biological control

B The biology of
iological control is the use of one or
more types of beneficial organisms,
usually called natural enemies, to
reduce the numbers of another type of
insects
B
iological control is most successful
organism, the pest. Although this publica-
when the pest manager has a fun-
tion focuses on the biological control of
damental understanding of the
pest insects, biological control has also
biology of the pests and their natural
been used to manage other types of
enemies. This section explains some of
pests, including weeds and microorgan-
the principles of insect biology and
isms that cause plant diseases.
defines terms used throughout this pub-
Biological control relies on the interac- lication.
tions of living organisms with the target
pests and the environment. It is therefore Insect structure
more complex than certain traditional The insect body is divided into three
pest control practices, such as the use of regions: the head, thorax, and abdomen
chemical pesticides. This introductory (figure 1).The head contains the brain,
section provides basic information on the mouth and mouthparts, and impor-
the biology of insects, the natural tant sensory organs such as the eyes and
enemies of insects, and the methods antennae. Behind the head is the thorax,
used to implement biological control. to which the legs and wings are attached.
There is also a discussion of the econom- Insects have three pairs of true legs,
ics of pest control as it relates to biologi- although some larval insects, especially
Figure 1. All insects have three body cal control. Terms in boldface are in the caterpillars, possess fleshy auxiliary legs,
regions: head, thorax, and abdomen. glossary at the end of the publication. called prolegs, that are attached to the
abdomen (figure 2).There are usually two
head
pairs of wings, except in true flies, which
have only one pair. In some insects such
as grasshoppers and beetles, the front
pair of wings has become thickened to
serve as a protective covering over the
thorax abdomen abdomen while the insect is not flying.
The abdomen is located behind the
Figure 2. This caterpillar shows the basic body form of a larval insect. Certain thorax. It is usually distinctly segmented
groups of larvae, especially caterpillars and sawfly larvae, have auxiliary legs, and as long as, or longer than, the head
called prolegs, on the abdomen.
and thorax combined. Internally, the
abdomen
thorax abdomen contains most of the digestive
head system, the reproductive system, and
other important organs. It is usually more
flexible than the head and thorax
because it needs to stretch to accommo-
date food, water, air, fat reserves, and
true legs prolegs
eggs.
2 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Insect growth and The two major categories of growth and Most insects undergo a more complex
development development are simple and complete form of juvenile development called
Most insects start life in an egg stage. metamorphosis. Simple metamorphosis complete metamorphosis. These
The act of egg laying is called oviposi- (sometimes called incomplete metamor- insects are worm-like, maggot-like, or
tion. The reproductive adult females of phosis) occurs in those insect species in grub-like in the immature stage. These
many species lay their eggs specifically which the young usually look very similar types of immature insects are called
in the area where the offspring will feed. to the adults, except that wings are larvae (singular—larva). The major
For example, greenhouse whiteflies lay absent and they are not reproductively change in body shape between the
their eggs on the foliage where the mature. In the immature stage, these larval and adult stages requires an inter-
immatures will feed, and fungus gnats insects are called nymphs (figure 4). mediate stage of development, the
oviposit on the surface of the soil or Common insects that undergo simple pupal stage (figure 5). Pupae (singular—
growing medium where the larvae will metamorphosis include dragonflies, pupa) are nonfeeding and inactive. They
develop. Insects have specialized organs mayflies, grasshoppers, crickets, cock- usually change shape (metamorphose)
called ovipositors for depositing the roaches, true bugs, and leafhoppers. in a protected location—such as within
eggs in the appropriate location Greenhouse pests with simple metamor- a cocoon, under tree bark, or in the
(figure 3). Some ovipositors are internal phosis include aphids, whiteflies, thrips, soil—because they cannot fly, walk, or
except during oviposition (as with most mealybugs, and scales. (Whiteflies and otherwise avoid natural enemies and
flies); others are external and very thrips have an inactive nymphal stage environmental extremes. Insects with
obvious (as with crickets and ichneu- that is commonly, but incorrectly, referred complete metamorphosis include all
monid wasps). A few insects, such as to as a pupa, even though these insects beetles, butterflies, moths, bees, wasps,
aphids, give birth to live young. do not have a true pupal stage like ants, flies, and lacewings. Greenhouse
insects which undergo complete meta- pests with complete metamorphosis
morphosis—see below.) Mites, which are include leafminers, fungus gnats, shore
not insects, also develop by simple meta- flies, caterpillars, and weevils.
morphosis.
Figure 3. A cricket (left) and an insect-
parasitic wasp (right), each with an obvious
ovipositor. The cricket inserts its eggs in Figure 4. A plant bug is an example of an insect with simple metamorphosis. After
soil and the parasitic wasp “stings” its host hatching from the egg, the nymph grows, occasionally shedding its skin, until it
insect, laying its egg inside the host. reaches the adult winged and reproductive stage, after which it no longer grows.

simple metamorphosis

ovipositor

complete metamorphosis

Figure 5. The imported cabbageworm is an example of an insect with complete metamorphosis. After hatching from the egg, the
larva grows, occasionally shedding its skin, until it is fully grown. The larva then molts one more time and transforms into the pupa. The
pupa in turn molts and transforms into the adult winged and reproductive stage, after which it no longer grows.
S E C T I O N 1 — U N D E R S T A N D I N G P E S T S & T H E I R B I O L O G I C A L C O N T R O L 3
As both nymphs and larvae grow, they The rate of insect Table 1. Common natural enemies with simple and
periodically have to shed their skins (the growth and develop- complete metamorphosis.
exoskeleton), through a process called ment depends
molting. Most species of insects molt a largely on environ- Simple metamorphosis Complete metamorphosis
(immature stage—nymph) (immature stage—larva)
set number of times before they mental factors (e.g.,
become adults. The distinct immature temperature, bigeyed bugs ground beetles
stages between successive molts are humidity, and avail- damsel bugs hover flies (syrphid flies)
called instars. For example, greenhouse ability of food) and minute pirate bugs lacewings
whiteflies have four instars while veg- the genetic traits of predatory mites lady beetles
etable leafminers have three instars. The the species. Within stink bugs parasitic wasps (such as
first instar is that which hatches from limits, the warmer ichneumonids, braconids,
the egg, the second instar is after the the temperature, the and chalcids)
first molt, and so on. Many natural more rapid the tachinid flies
enemies, especially parasitic wasps, development and
attack only certain instars of the target the shorter the gen-
pest. This can be an important factor for eration time. The length of time required In temperate climates with cold winters,
effective biological control, especially for an insect to complete one genera- insects either die or go into an overwin-
when timing releases of natural tion varies considerably with the type of tering protective state of arrested devel-
enemies. insect, the availability of food, and, to opment called diapause. A given
some degree, the location and climate. species diapauses in a specific stage of
Adult insects are characterized by the
Greenhouse environmental conditions development. Outdoors, aphids over-
presence of wings and by reproductive
usually promote more rapid growth of winter in the egg stage, caterpillar pests
maturity. (With insects it seems that
pest populations than would occur such as the cabbage looper overwinter
there are exceptions to every rule; most
outdoors, although population develop- as pupae, and vine weevils or root
adult aphids and all fleas are wingless.)
ment often slows down under cooler weevils diapause as adults. In the green-
Once an insect reaches the adult stage,
winter conditions. house, many pests do not enter
it doesn’t grow any further and never
diapause; they develop year round. But
molts again. Therefore, small beetles do
certain natural enemies do diapause in
not grow into large beetles; small flies
the greenhouse. The predatory aphid
do not grow into large flies; and so forth
midge Aphidoletes aphidimyza, for
(figure 6).
example, diapauses as pupae, and the
predatory mite Neoseiulus cucumeris as
Figure 6. The upper life cycle shows the common misconception that little insects are
adults, although there are nondiapaus-
smaller versions of adult insects, while the lower life cycle illustrates the correct growth
ing strains of the mite.
pattern of a beetle. When two similar adult insects are substantially different in size,
they are likely different species with different habits.

egg small
beetle
medium
beetle
large
beetle = NO

egg larva pupa adult


beetle = YES
4 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Insect feeding Insect classification We also describe insects using their


Insects with simple metamorphosis Classification is the process of categoriz- family name. Thus, mirid bugs,
often feed as both nymphs and adults in ing organisms into related groups, Deraeocoris brevis and Macrolophus
the same location and on the same based on a standard hierarchy of cate- costalis, are both members of the family
food. This is true of aphids, mites, mealy- gories (see table 2). The most basic Miridae.
bugs, scales, and thrips. The larvae of category in classification is the species,
insects with complete metamorphosis a grouping of animals that is function- A note about mites
often feed in a different location and on ally capable of reproducing and rou- Mites are not insects but are classified
a different food than the adults. For tinely does so, thereby perpetuating the with the insects and several other
example, black vine weevil larvae feed species. The scientific name of a species groups in the phylum Arthropoda.
on roots, but the adults feed on foliage. consists of the genus name (capitalized) Arthropods are categorized by their
Some insects, especially those with plus the species name (not capitalized), hard exoskeleton, or skin, and by their
complete metamorphosis, feed primarily both italicized or underlined. For jointed appendages. Mites are in the
in the immature stages, and adult example, the scientific name of the class Arachnida, which also includes
feeding may be insignificant. For species we call greenhouse whitefly is spiders and scorpions, and the order
example, leafminer maggots feed by Trialeurodes vaporariorum. Using scien- Acari (sometimes called Acarida), which
chewing on plant tissue in their mines, tific names is preferable to using includes all mites. Mites have two body
while the adult flies feed by sucking common or colloquial names because in regions instead of three and usually
plant juices from holes made in the different places the same common have four pairs of legs. No species of
leaves. names sometimes refer to different mite has wings. They develop by simple
species. Also, only very common, very metamorphosis, although in some cases
Insects may feed in two basic ways: by
showy, or very pestiferous species have the immatures are markedly different
sucking fluids or by chewing.
common names. The majority of insects, from the adults in structure and biology.
Mosquitoes, for example, suck blood,
including most beneficial natural Most mites are very small. Those impor-
butterflies suck nectar from a flower,
enemies, are uncommon, small, or non- tant to agriculture are about 1⁄50 inch
and aphids suck plant sap from leaves.
descript, and have never received (1⁄2 mm) in size or less when fully
On the other hand, grubs chew on roots
common names. The Entomological grown. There are many mite pests of
of plants, and caterpillars or grasshop-
Society of America publishes a list of greenhouse crops, as well as several
pers chew holes in leaves. Nymphs and
approved common names that can be predatory mites, especially in the family
adults of insects with simple metamor-
used for individual species. In this publi- Phytoseiidae, that are important natural
phosis usually have the same type of
cation, we will use approved common enemies of small insect pests such as
mouthparts. In all stages of develop-
names in conjunction with the scientific thrips and spider mites.
ment, aphids have sucking mouthparts
and grasshoppers have chewing mouth- names.
parts. The adults of insects with
complete metamorphosis do not neces-
sarily have the same type of mouthparts
Table 2. Example of insect classification, based on the greenhouse
as the larvae. Caterpillars have chewing
whitefly, Trialeurodes vaporariorum.
mouthparts, whereas adult butterflies
have sucking mouthparts. Category Classification of greenhouse whitefly
Many beneficial natural enemies, such as Kingdom Animalia The animal kingdom
lady beetles, praying mantids, and para- Phylum Arthropoda The arthropods
sitic wasps, have chewing mouthparts. Class Hexapoda Insects and their relatives
Other natural enemies, such as the Order Homoptera Aphids, cicadas, leafhoppers,
larvae of lacewings and the nymphs and mealybugs, planthoppers,
adults of the true bugs, have sucking scale insects, treehoppers,
and whiteflies
mouthparts.
Family Aleyrodidae Whiteflies
Genus Trialeurodes A group of several closely
related whitefly species
Species vaporariorum Greenhouse whitefly
S E C T I O N 1 — U N D E R S T A N D I N G P E S T S & T H E I R B I O L O G I C A L C O N T R O L 5

The natural Insects that eat other insects are either


predatory or parasitic. Predatory insects,
fly larvae, lay their eggs in the immedi-
ate vicinity of prey insects, such as
enemies of insects or predators, are usually much larger
than their prey. They eat and kill many
aphids. Some types of predators
undergo simple metamorphosis—the

A
ll insects and mites have natural
prey as they grow and reproduce. Many various families of predatory bugs are
enemies. Some pests have more—
predators will feed on almost anything examples—whereas others undergo
or more efficient—natural enemies
they can catch, but some specialize in complete metamorphosis—examples
than others. Biological control relies on
consuming certain prey types. Most include the lady beetles and lacewings
effective natural enemies that can be
predators are fairly mobile and can (see table 1). Many important predators
managed by humans.
search rapidly for prey. The adults of rel- are predatory as both immatures and
atively nonmobile species, such as hover adults.
Types of natural enemies
Vertebrate natural enemies of insects Table 3. Major groups (orders) of insects, listed in order of increasing evolutionary
include certain birds, such as flycatchers, complexity.1
woodpeckers, purple martins, starlings,
and chickens; certain mammals, such as Number of
bats, moles, voles, skunks, and hogs; and Type of North American
toads, frogs, and lizards. With a few minor Order Type of insect metamorphosis species2
exceptions, these cannot be managed to Thysanura silverfish simple 18
reduce the populations of pests signifi- Ephemeroptera mayflies simple 610
cantly, and they will not be considered Odonata dragonflies, damselflies simple 400
further in this publication. Plecoptera stoneflies simple 465
Insects and mites that feed on other Phasmatodea walkingsticks simple 30
insects or mites make up the most Orthoptera grasshoppers, crickets, simple 1,080
important group of natural enemies. katydids
This is an extremely large and diverse Dermaptera earwigs simple 20
group. Unfortunately, because beneficial Dictyoptera mantids, cockroaches simple 70
insects are often tiny and nondescript, Isoptera termites simple 45
they are frequently overlooked by even Psocoptera booklice, barklice simple 250
the most dedicated practitioners of bio- Phthiraptera chewing and sucking lice simple 1,000
logical control. Only the large, common, Hemiptera true bugs simple 3,600
or brightly colored species, such as Homoptera aphids, cicadas, leafhoppers, simple 6,300
praying mantids and lady beetles, are mealybugs, planthoppers, (some modified)
commonly recognized. scale insects, treehoppers,
and whiteflies
Thysanoptera thrips simple 700
(but modified)
Neuroptera dobsonflies, lacewings, complete 350
and relatives
Coleoptera beetles (including weevils) complete 23,700
Hymenoptera sawflies, bees, wasps, complete 17,800
ants, parasitic wasps
Trichoptera caddisflies complete 1,260
Lepidoptera moths, butterflies complete 11,300
Diptera flies complete 16,900
Siphonaptera fleas complete 300

Approximate total insect species in the United States and Canada3 86,300
1Additional orders include Grylloblattodea (rock crawlers), Zoraptera (angel insects), and
Mecoptera (scorpionflies).
2An approximation of the number of species in this order found north of Mexico.
3Authorities estimate that 10–25% of all species are still unknown to science.
6 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

The majority of parasites (also called Different parasites attack different Pathogens are microorganisms that
parasitoids) are very small wasps in stages of the host. There are egg para- cause diseases. The most common types
several families of the order sites, nymphal parasites, larval parasites, of insect pathogens are bacteria, fungi,
Hymenoptera. Another important group pupal parasites, and adult parasites. nematodes, viruses, and protozoans.
is tachinid flies (order Diptera, family Parasites are generally host specific, Some insect diseases are highly lethal,
Tachinidae). Parasites are about the attacking only a single insect species or killing a large portion of the insect pop-
same size as their hosts, or smaller. All a group of closely related species. Most ulation. Other diseases are less lethal but
parasites undergo complete metamor- types of insects are attacked by one or may retard insect development, shorten
phosis. In most cases, the adult female more species of parasite. In fact, many the insect’s life, or prevent reproduction.
lays eggs on, within, or near a host parasites themselves are parasitized by Most insect pathogens are pathogenic
insect of the appropriate stage. When other species, called hyperparasites. only to insects, frequently to very small
the egg hatches, the parasite larva Section 2 contains more details on groups of closely related species.
consumes its host. Each parasite larva specific predatory and parasitic insects. Microorganisms that are pathogenic to
can attack only one host. A parasite larva Many of these insects are covered in a specific pest are often harmless to
that feeds and develops within its host even greater detail in the companion other insects, including predators and
is called an endoparasite (figure 7), publication Biological Control of Insects parasites.
whereas one that feeds while attached and Mites (NCR 481), which includes Often, specific conditions are necessary
to the outside of its host is called an descriptions and color photographs of for an insect pathogen to reduce a pest
ectoparasite (figure 8). Endoparasites many types of natural enemies. population effectively. These conditions
are much more common than ectopara- vary with the type of pathogen and the
sites. When a single parasite develops host. Most fungal pathogens are only
within a single host, it is called a solitary effective during periods of relatively high
parasite; when two or more individuals humidity, because it is only under these
of the same species can develop within conditions that the fungi will produce
one host, the species is said to be a spores and the spores will germinate.
gregarious parasite. Viral pathogens are often most effective
when the host population is very high,
which facilitates spread from individual
to individual within the population.
Bacterial pathogens must be ingested to
Figure 7. Endoparasites develop inside be effective, and therefore do not usually
the body of the host insect. kill sucking insects such as aphids and
whiteflies. Nematodes require a thin film
of moisture on which to move, so they
are not effective under dry conditions.

Figure 8. Ectoparasites feed externally,


attached to the outside of the host.
S E C T I O N 1 — U N D E R S T A N D I N G P E S T S & T H E I R B I O L O G I C A L C O N T R O L 7
Because of these characteristics, the
degree of control exerted by naturally The general International, federal, and state agencies,
along with a few land-grant universities,
occurring pathogens is often unpre-
dictable. However, pest managers should
approaches to import natural enemies. An agency
undertaking this process requires strin-
be aware of the symptoms of insect
diseases and should be able to assess
biological control gently maintained quarantine facilities
and highly trained personnel to assure

T
here are three broad approaches to
their impacts on pest populations. the biological control of insects: that the introduction of exotic natural
A few insect pathogens are available importation and permanent enemies does not result in the introduc-
commercially. You can spray these on establishment of new natural enemies; tion of additional pest species.
the crop using conventional pesticide augmentation of existing natural Researchers at several of these facilities
spray equipment. When used in this enemies by releasing predators or para- are currently evaluating exotic natural
fashion, insect pathogens are called sites or by applying microbial insecti- enemies of whiteflies, thrips, and other
microbial insecticides. These can be cides; and conservation of natural pests for possible introduction in sub-
very effective at the time of application, enemies by changing aspects of the tropical areas of the United States. Some
but they usually do not persist in the environment that threaten their survival of these may also be useful for control-
greenhouse environment. Their advan- or effectiveness. ling certain greenhouse pests if com-
tages include wide availability from mercial suppliers of natural enemies
Importation of natural enemies is produce them in the future.
commercial sources, safety to humans, based on the understanding that the
and safety to nontarget organisms, most effective natural enemies may not Augmentation of natural enemies
including beneficial insects. However, occur in the region where the pest is means increasing natural enemy
microbial insecticides have some disad- causing damage. Many pests in the numbers through human involvement.
vantages: they may be more costly to United States are native to other parts of Natural enemies moving into green-
use than traditional pesticides; multiple the world, especially Europe, Australia houses will rarely be abundant enough
applications may be necessary because and northern Asia, and were introduced to maintain the pest population below
residual activity is short; and if other accidentally into this area by such damaging levels. Increasing the natural
types of insect pests are present, you human activities as immigration and enemy population improves the chance
may still need to suppress these with commerce. Although modern federal of gaining economic control over the
other controls. Microbial insecticides and state quarantine practices help pests. This typically requires purchasing
containing the bacterium Bacillus exclude many serious foreign pests, and releasing natural enemies from
thuringiensis are discussed in Section 2 occasionally new alien species are intro- commercial vendors, many of which are
of this publication. More information on duced and become established. For insectaries. An insectary is a facility that
insect pathogens and microbial control example, the southeast Asian thrips grows insects, often mass-producing
can be found in the companion publica- Thrips palmi is now a problem in Florida. natural enemies for augmentation
tion Biological Control of Insects and purposes. The three types of natural
Mites (NCR 481) and in Alternatives in The most efficient natural enemies are enemies commercially available for
Insect Management (NCR 401). those that evolved with the pest and are purchase and release in augmentation
therefore adapted to the pest’s life cycle, programs are generalist natural
behavior, and other characteristics. enemies, which are usually predatory
Scientists search throughout the native insects; specialist natural enemies,
range of the pest for natural enemies usually parasites; and insect pathogens
that have the potential for controlling produced as microbial insecticides.
the insect in its new area. Exotic natural
enemies are carefully screened to be Generalist natural enemies feed on a
certain that they have no undesirable variety of prey and may or may not be
characteristics. Then they are introduced adapted to specific types of prey. For
into the target region, with the goal that example, lacewings and certain lady
they become permanently established beetles prefer to feed on aphids and
and provide ongoing biological control similar insects, but will also consume
of the pest. This process is called foreign small caterpillars, spider mites, insect
exploration for natural enemies. eggs, and other types of prey. Some of
8 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

the species of egg parasites in the genus Inoculation is the release of small meet the natural enemies’ needs and
Trichogramma attack the eggs of many numbers of natural enemies when the allow them to stay where you want
types of moths. Praying mantids are pest population is still at low levels. The them. A simple environment, character-
generalists that are not recommended strategy of inoculation is to encourage ized by few plant and insect species,
for augmentation because they are the released natural enemies to repro- often encourages natural enemies to
indiscriminate in prey choice; they will duce rapidly enough to counter the leave. Once natural enemies leave to
eat beneficial and innocuous insects as build-up of the pest. This is the more seek food or other necessities, they are
well as pests. “natural” of the two approaches because unlikely to return. Unfortunately, it is
Specialist natural enemies attack only a it relies on the reproduction of the ben- often impractical to provide the diver-
specific type of pest or a group of eficial insect and takes advantage of its sity necessary for natural enemy repro-
closely related species. For example, the biological characteristics, such as the duction in greenhouses. Thus, continued
parasite Encarsia formosa attacks only ability to seek out scattered prey. introductions into greenhouses are
whiteflies, and Dacnusa sibirica attacks Inoculative releases are usually less usually required to maintain the natural
only leafminers. Although most com- expensive than inundation, but they enemy population.
mercially available specialist natural require routine monitoring of pest and One of the major deterrents to effective
enemies are parasites, some are preda- natural enemy populations to be certain natural enemy activity is the use of
tors. The mealybug destroyer, that the pest is being controlled. Often, a broad-spectrum pesticides, because
Cryptolaemus montrouzieri, and the sequence of inoculative releases at pre- natural enemies are just as susceptible to
spider mite destroyer, Stethorus scribed intervals is more effective and pesticides as are the target pests.
punctum, are both specialized predatory less expensive than an inundative Frequently one pest will be under good
lady beetles. release. biological control, but another requires a
Microbial insecticides also tend to be Conservation of natural enemies pesticide to prevent economic damage
fairly host specific, but their range of entails preserving those predators, para- from occurring. Pesticide applications
activity can vary. For example, Bacillus sites, and pathogens that occur naturally may eliminate natural enemy popula-
thuringiensis (Bt) can kill mosquitoes, or that have been introduced through tions, leading to a secondary pest
beetles, and caterpillars, but the activity importation or augmentation programs. outbreak. This is the term used when a
depends on the strain of bacterium. The Natural enemies, like all living things, species that was not causing damage
caterpillar-active strains of Bt are fairly have specific requirements for life, becomes a problem because of major,
broad spectrum, affecting almost all growth, and reproduction. These usually rapid, changes in the pest’s envi-
caterpillars. On the other hand, most requirements include sufficient food ronment. A similar phenomenon, pest
insect-pathogenic viruses are highly and water, and shelter which allows resurgence, occurs when the population
selective, often affecting only one or a natural enemies to avoid environmental of the pest that was the target for the
very few species. extremes such as high temperatures pesticide application rebounds rapidly
and flooding. Providing natural enemies after the initial application, often because
There are two general approaches to the
with necessary resources and protecting of the elimination of its own natural
augmentative release of natural
them from major mortality factors, such enemies.The use of chemical pesticides
enemies. Inundation is the mass release
as virulent diseases and toxic chemicals, may provoke both phenomena.
of large numbers of natural enemies to
are the two main approaches to natural No chemical insecticide is completely
gain rapid control of a damaging or
enemy conservation. specific to one target pest species. Even
near-damaging pest population. This
method only works when the natural Natural enemies benefit from living so-called natural products such as
enemy can kill the pest relatively rapidly. among a variety of food sources. In the pyrethrum, a plant extract, can be
If the natural enemy acts slowly, then absence of their preferred prey, general- damaging to some natural enemies. The
the pest population is likely to do con- ist natural enemies can shift to other insecticide’s residual activity may deter-
siderable damage before it is brought types of prey. For optimal reproduction, mine how harmful it is. While pyrethrum,
under control. some adult natural enemies also require rotenone, and many other plant-derived
certain nutrients from such sources as insecticides are highly toxic to natural
plant exudates, flower nectar, or the enemies at the time of application, these
honeydew excrement of sucking insects. products break down very rapidly in the
A diverse environment is more likely to environment. They generally will not
S E C T I O N 1 — U N D E R S T A N D I N G P E S T S & T H E I R B I O L O G I C A L C O N T R O L 9
harm natural enemies released after the The damage caused The costs of managing
insecticide application. However, many by each pest the pests
insecticides have a period of residual Some pests cause much more damage The product used for controlling a pest,
activity lasting several days to several than others. Caterpillars feeding directly any equipment required for application,
weeks. These not only kill natural on the flowers of ornamentals cause and the labor required for the control
enemies at the time of application, but more damage than fungus gnats process are the primary costs of pest
can also kill some individuals introduced breeding in the potting medium of the management. The costs of pest monitor-
later. Many insecticides fall into this same plants. Sweetpotato whiteflies ing also can be significant. These include
category. Those materials safest to remove more sap and produce more the costs of traps and other monitoring
natural enemies include the microbial honeydew than greenhouse whiteflies. equipment, as well as the salary or con-
insecticides (e.g., Bt) and insecticidal Western flower thrips transmit tomato sultant fee of the individual responsible
soaps. spotted wilt virus and impatiens for monitoring.
necrosis virus to ornamental plants, When considering costs, it is important
The economics reducing their marketability, whereas
greenhouse thrips only create feeding
to consider the entire pest complex.
Typically, a particular biological control
of pest control scars on the leaves. Each particular pest
problem requires its own specific
method is effective against only a single

R
eleases of natural enemies can type of pest or a narrow range of pests.
often provide adequate pest solution. For example, some parasitoids only
control and result in a relatively attack certain aphid species. If numerous
undamaged crop. However, the cost of The level of the pest types of pests are present, several bio-
purchasing, monitoring, and managing population logical control agents may be necessary.
natural enemies may be greater than It is impractical, and usually impossible, This could be much more costly than
the cost of using insecticides. to make your greenhouse completely using a broad-spectrum insecticide.
pest free; the costs of control would far
The operational expenses of producing
exceed the benefit of controlling the last Effectiveness of control
the crop, including pest management
few insects. Therefore, growers must methods
costs, must be made up by the revenue
monitor carefully and routinely for pest Label directions for traditional chemical
from marketing the crop, as determined
activity to determine the relative abun- insecticides can usually tell you how to
by the quantity and quality of the crop
dance of each pest throughout the achieve maximum control. Such precise
and the marketing approach used. The
growing season. Growers should also recommendations may not be possible
costs of pest management must be
keep weekly records of pest activity and for many biological controls. Complex
weighed against the potential economic
relate these to actual insect damage. environmental factors can make it diffi-
impact of each type of pest and related
Because of the relatively constant green- cult to predict the ultimate degree of
factors. These may include (1) the
house environment, most greenhouse control that releasing natural enemies
amount of physical damage that each
pests can occur at any time. During will provide. Furthermore, some com-
pest can cause to the crop—that is, the
warm months pests may enter the mercially available natural enemies are
loss in yield or quality, (2) the actual
greenhouse from outside, and those much more effective than others.
number of pests present, (3) the cost of
that are established in the greenhouse Although there are general guidelines
managing the pests either individually
may develop rapidly. In northern states for the release of natural enemies,
or, more appropriately, as an entire pest
with cold winters, pest activity usually growers must gain some experience in
complex, (4) the effectiveness of the
declines when the days are shorter and their own greenhouses to determine the
control methods used, and (5) the end
the temperatures lower, and when there effectiveness of the chosen biological
use and market value of the crop.
is no pest invasion from outside. control agents, and then must decide if
the level of effectiveness is acceptable.
10 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

The end use and market value One of the most frequently asked ques- The costs of different pest control strate-
of the crop tions is,“How many bugs will it take to gies varies, and because crop suscepti-
The best pest control method for your cause economic damage to my crop?” bility and market values vary, the EIL and
crop may depend on the end use and There is no simple answer. However, you the ET are not fixed values. There may be
market value of the crop. Whether a can consider the five factors listed several values for each crop, depending
grower is cultivating chrysanthemums above to develop action thresholds. on the end use, and for each pest.
for cut flowers or for potted plants, for These are actual pest population levels Chrysanthemums grown for cut flowers
example, may determine the amount of at which you will apply controls to avoid can tolerate some leafminers on lower
damage that can be tolerated. Fungus economic injury. There are two types of leaves because only the top half of the
gnats are unacceptable on potted action thresholds. The economic injury plant is marketed. Potted chrysanthe-
chrysanthemums nearing harvest but level (EIL) is that pest population that mums cannot have any significant
are of no importance on cut chrysanthe- will cause economic injury, all other leafminer damage. As a result, the
mums. Market value is determined by factors considered (figure 9). At this thresholds for leafminers are higher on
many factors, such as available supply, point, the cost of applying controls chrysanthemums grown for cut flowers
competition, quality, cultivar, and exactly offsets the economic loss caused than those for potted plants.
consumer demand. Values vary from by the pest. If controls are exerted at The above discussion assumes that the
season to season and even within the population levels below the EIL, then grower will purchase biological control
season. When supply is high and prices the control costs exceed the benefits. agents. Some natural enemies invade
low, quality must often be the highest in Conversely, if controls are not exerted greenhouses, but for the most part
order to sell the product at all. Quality until the pest population surpasses the introductions are necessary to provide
becomes more flexible when supply is EIL, some economic loss to the crop will control. In a few situations, introduced
low and prices are high. Unfortunately, have occurred. It is usually impractical to natural enemies become established so
this market information is not usually apply controls exactly as the pest popu- that biological control is effective and
available at the time pest control deci- lation reaches the EIL and still keep the permanent, requiring little if any
sions are made. However, your market- pest from causing economic damage. ongoing economic input. However,
ing strategies can also influence the The economic threshold (ET) is a popu- regular releases are usually necessary to
gross value of the crop. Biological lation level somewhat below the EIL that sustain biological control in green-
controls can be more expensive than allows sufficient lead time to implement houses.
traditional chemical approaches, but if control. By applying controls at the ET
the additional expense is offset by a you improve the chances of achieving
new marketing strategy—by selling economic pest control.
greenhouse tomatoes to the organic
market, for example—then this may still
be a smart approach. Figure 9. A hypothetical example of an uncontrolled pest population
exceeding the economic injury level (EIL). Action should be taken at the
economic threshold (ET) to avoid this problem.
number of insects

EIL

ET

1 2 3 4 5 6
generation
EIL = economic injury level ET = economic threshold
SECTION

2
Practical approaches
to the biological control
of greenhouse pests
B
iological control relies on living Sanitation is the most important cultural
organisms that interact, often in practice for preventing pest problems.
complex ways, with the pests, the Keep the greenhouse clean. The cleanest
crop, and other environmental factors. greenhouses usually have the fewest
The pest manager needs a different type pest problems. Remove dead foliage,
of knowledge and skill, as well as extra broken pots, and trash, and place it in
effort and patience, to implement bio- areas away from the greenhouse opera-
logical control instead of using chemical tions. Keep areas beneath benches as
control. A good understanding of the dry as possible. Remove plastic, boards,
relationships between pests, their and debris that may serve as hiding
natural enemies, and the environment is places for pests such as slugs or
essential. sowbugs. Eliminate standing water and
algae growth; fungus gnats and shore
Successful implementation of biological
flies breed in these conditions. Remove
control in greenhouses does not
weeds under the benches and “pet
depend on natural enemies alone. To be
plants” that are a refuge for insects.
effective it must be part of an integrated
Aphids, thrips, mites, whiteflies, and
pest management program. Integrated
other pests developing on weeds
pest management (IPM) is the ecologi-
outdoors can enter the greenhouse
cally sound use of all available methods
through ventilators or unscreened
to control pests. One of the goals of IPM
doors. To minimize this, mow any grass
is the reduction of pesticide use and, if
or kill weeds in a 15–30 ft strip around
sprays are necessary, the selection of
the greenhouse.
those that are the least disruptive to
natural enemies. That means incorporat- Sanitation inside the greenhouse can be
ing many cultural practices to prevent or more important than preventing migra-
delay the appearance of pests in green- tion from the outside. Use screens on
house crops. It also means monitoring vents and doors if the pest populations
pests regularly to detect their arrival entering the greenhouse from outside
early and keep track of pest and natural are large enough to justify the expense.
enemy populations throughout the Know where the pests are coming in
growing season. In addition, this before installing screens, so you don’t
approach requires an understanding of install unnecessary ones. Remember
all the alternative pest management that screens reduce air flow when
options, both chemical and nonchemi- placed around vents. The smaller the
cal. In the following sections on specific pore size of the screen, the harder it is to
pests, the methods for using and con- get air through. Screens may place
serving natural enemies in an IPM severe strain on exhaust fans. To coun-
context are described in detail. teract airflow problems, expand the
surface area of the intake vents. Work
with reputable suppliers to develop
proper designs for your needs. They
12 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

should have information on the porosity Plant management practices that It is important to have a knowledgeable
of screens for airflow, the insects meant promote healthy plants also help reduce person, such as a consultant or trained
to be kept out, and screen installation. pest problems. Many pests develop employee, on site for monitoring. Weekly
Screens must be cleaned regularly for better on stressed or lush plants than on monitoring is necessary in most green-
proper airflow. Double door entries, with normal plants. Maintain proper soil houses. Monitoring usually entails
negative pressure to prevent pest entry, moisture and avoid over- or under-fertil- counting pests caught on sticky traps
can provide a high level of pest exclu- izing. High nitrogen levels are often placed over crops in the greenhouse as
sion for special situations, such as virus- associated with severe pest infestations. well as visual counts of pests and bene-
sensitive crops and cuttings. Plant selection may make the difference ficial insects on leaves or other plant
It is important to start with clean plants between success and failure in biologi- parts. When recording the number of
and keep them separate from infested cal control on a particular crop. Many pests and natural enemies, be sure to
plants. Pests can easily move from an pests develop better on some crop culti- note the temperature, date, time of day,
infested crop to an uninfested one. vars than on others. Select plants known numbers caught per sticky trap, and
Avoid starting new plants in a green- to be the least sensitive to the pests that location of infested plants and traps. Use
house with an existing crop. Hanging you are dealing with. Eliminate the most a map of the greenhouse to update the
baskets often contain pests that easily susceptible cultivars if possible, or location and density of pests. Different
fall onto the new crop. Carefully inspect isolate them to reduce contamination of methods for monitoring different pests
all new plant material for pests before other cultivars. and their natural enemies are discussed
placing it in growing areas to prevent in more detail in the following sections
Monitoring pest and natural enemy
contamination of the entire range. Treat on specific pests.
populations on a regular basis is a
or quarantine any shipment that is crucial component of any biological
infested. If only a few plants are infested, control program. A good monitoring
remove or destroy them. Also, start with program will help detect the pests early,
steam-sterilized soil or soilless potting time the release of natural enemies for
medium. Dispose of or pasteurize used maximum effectiveness, and determine
media. the result of natural enemy intro-
If pests are already present, rogue out ductions. Finding pest infestations
severely infested plants. After harvest, when populations are small
remove all crop residue promptly from increases the likelihood of success
the greenhouse. Pest outbreaks in new with biological control. It is also
crops can start from pieces of infested easier to determine which benefi-
leaves or stems from the old crop. You cials to use when you know the
should also institute policies to reduce magnitude of the pest problem.
the accidental spread of pests on
workers’ clothing or implements. Restrict
movement through infested areas and
always visit them last. Consider banning
yellow and blue clothes from the green-
house because they are attractive to
whiteflies and thrips, respectively.
Susan E. Rice Mahr
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S 13
Augmentation is necessary to achieve Decide which delivery service to use for The rest of this section discusses the
effective control with most of the para- your location and provide a delivery major pests of greenhouse crops. It
sites, predators, and pathogens dis- address where the shipment will not be includes an overview of the damage
cussed, although a few invade green- exposed to extreme temperatures and caused by each pest, a brief description
houses naturally and provide some will be cared for as soon as it arrives. Be of the pest and its life cycle, a statement
control. The suggested release rates and prepared to make releases when the on monitoring the pest, and detailed
methods for natural enemies that are shipment arrives, or to provide the information on selected parasites, pred-
produced commercially are only general proper storage conditions until releases ators and pathogens of those pests. The
guidelines. Specific release rates are dif- can be made. Check the shipment to possibilities for effective biological
ficult to provide because of the com- make sure you received the species and control of each pest are reviewed, as are
plexity of biological interactions in the quantities you ordered, that the alternative control methods, most of
greenhouse. Numerous variables affect shipment is not damaged, and that the which are nonchemical. Some of the
the performance of natural enemies and natural enemies are viable. Contact the natural enemies discussed show
therefore the number necessary to supplier immediately if there is any promise for biological control in green-
maintain pest populations below problem. Understand proper release house crops but were not commercially
economic injury levels. These include procedures and make the introductions available at the time of printing. Check
the crop variety and growth stage, size according to your supplier’s recommen- natural enemy suppliers for current
of the pest population, environmental dations. In general, releases should be availability.
conditions, release methods, the quality made during the cooler part of the day,
of the natural enemies released, and the either in the early morning or late after-
presence of other insects. Appropriate noon. Never release natural enemies
release rates often can be determined during the heat of the day. Evaluate the
only through experience. Many suppliers success of the release by monitoring
provide recommendations for their both the natural enemy and pest
products. populations.
Ordering and receiving shipments of Residual broad-spectrum pesticides,
predators and parasites requires some including sulfur-containing fungicides
advance planning. Determine which sup- (which are toxic to predatory mites)
pliers offer the natural enemies you need should not be used within a month of
and contact them to confirm availability. parasite or predator release. There
One source of such information is should be no chemical residues on
Suppliers of Beneficial Organisms in North either the plants or the greenhouse
America, which is listed in the additional structure before starting a biological
reading section at the end of this publi- control program. Determine the pest
cation. If importing natural enemies from control history of incoming plant
outside the country, it may be necessary material since pesticide residues on new
to obtain import permits from the cuttings can also be toxic to natural
Animal and Plant Health Inspection enemies.
Service (USDA-APHIS) well ahead of time.
Also, some states have regulations
regarding the interstate shipment of
commercial natural enemies. If you are
uncertain about such regulations, contact
your state’s agriculture department.
14 Aphids
A
phids are one of the most serious Aphids feeding on flowers make the
pests of greenhouse-grown crops. plants unmarketable. Their presence is a
Available natural enemies and
their potential for control Almost all species of plants grown nuisance on ornamental plants because
Aphids have many in greenhouses are susceptible to some they leave cast skins stuck to the plant
predators and para- aphid species. Of the numerous aphid when they molt, which detracts from the
sites that can be species that infest greenhouse crops, value of the plant.
highly effective. The the most common is the green peach
potential for suc- aphid. It attacks a wide range of hosts, Description and life cycle
cessful biological but especially the Solanaceae (tomato, Most aphids are 1⁄8 inch (2–3 mm) long.
control is high. pepper, eggplant) and chrysanthemums. Their pear-shaped bodies have “exhaust
The cotton, potato, and foxglove aphids pipes” (cornicles) protruding from the
are other common species with a wide back end of the abdomen. Wingless
Order Homoptera: Aphids, host range. The cotton aphid is impor- forms of the green peach aphid are yel-
leafhoppers, and scales tant on cucumbers and chrysanthe- lowish green in summer and pink to red
Family Aphididae: Aphids mums, while the other two are impor- in fall and spring. Winged forms are
tant on Solanaceae. The chrysanthemum brown. Cotton aphids may be green,
Chrysanthemum aphid,
aphid infests chrysanthemums and the yellow, mottled, or blackish.
Macrosiphoniella sanborni
rose aphid infests roses. Many other Chrysanthemum aphids are maroon or
Cotton or melon aphid, aphid species may occur in green- dark brown, stem-feeding aphids. Most
Aphis gossypii houses. other aphids are green, gray, or yellow-
Foxglove aphid, Aulacorthum green and feed on leaves or growing tips.
solani Damage Each female aphid reproduces for a
Green peach aphid, Myzus Aphids feed by inserting their mouth- period of 20–30 days, giving birth to
persicae parts into leaves, buds, or stems and 60–100 live nymphs. The nymphs look
sucking out the sap. Chlorotic spots may like the adults but are smaller. The
Potato aphid, Macrosiphum
develop where cell contents have been nymphs mature and can produce off-
euphorbiae
removed. Bud feeding results in dis- spring within a week at greenhouse
Rose aphid, Macrosiphum rosae torted leaves and, at high densities, can temperatures. Winged females that can
cause stunting and/or wilting. Severe migrate are produced when the food
damage to the top of the plant may supply becomes short or the area
reduce the number of flowers produced. becomes overcrowded. Males and eggs
Aphids also excrete honeydew on which are produced only in northern areas, in
sooty molds grow, and some species response to colder temperatures and
may transmit viruses. shorter days. Aphids overwinter as eggs
outdoors in northern areas, but in
greenhouses will continue to develop
year-round. Oregon State University Extension Service

The hollow remains, or mummy, of an aphid parasitized by a


braconid wasp. The “trapdoor” was created by the adult wasp at
emergence. Aphid mummies in a colony usually indicate that
many of the apparently healthy aphids are also parasitized.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S A P H I D S 15
Monitoring Aphelinus abdominalis. This solitary Aphidius colemani. This braconid wasp
Look for aphids on as many plants as endoparasitoid parasitizes primarily is cosmopolitan—that is, it occurs in
possible. Check the lower leaves, around potato aphid and foxglove aphid. The most parts of the world. It reproduces
the terminal or flower buds, and on the 1⁄ 8-inch (2.5–3 mm) adults are black well on cotton aphid, green peach
top 6 inches of the stems of older plants. with a yellow abdomen. Females prefer aphid, and other species, but not on
Different aphid species prefer different to oviposit in third instar nymphs. Adult potato or foxglove aphid. The appear-
parts of the plant. Cast skins, honeydew wasps emerge from black mummies ance, life cycle, and biology of this
droplets, and sooty mold are indications through a jagged hole. Females host species is similar to A. matricariae (next
of an infestation. Aphid infestations are feed on some species of aphids. It is page), but females lay more eggs—an
often localized. Mark infested plants and commercially available. average of 388—over their 4–5 day life
check the progress of control efforts. Aphelinus flavipes. This is a parasite of span. It is commercially available in the
Yellow sticky traps can be used to detect cotton aphid similar to A. semiflavus United States.
winged aphids moving into the green- (below) in size and development. A. Aphidius ervi. This cosmopolitan
house. However, if winged aphids are flavipes also produces a black mummy. It species parasitizes numerous aphid
detected on yellow sticky traps in a is not available commercially. species in many different crops. In
closed greenhouse, this probably indi- greenhouses, they parasitize larger
Aphelinus semiflavus. This wasp para-
cates a serious infestation is already aphids than does A. colemani, particu-
sitizes several aphid species, including
present. larly potato aphid and foxglove aphid.
the green peach aphid. Often confused
with A. asychis, a European parasite of The small black females can lay about 50
Natural enemies spotted alfalfa aphid, the stocky adults eggs a day during the first 5–7 days
Numerous parasites, predators, and after emergence. Parasitized aphids turn
are very small (1⁄25 inch, 1 mm) with
pathogens attack aphids. Only those into a golden yellow-brown mummy.
elbowed antennae. Females lay single
that have shown promise for use in The adult wasps emerge through a small
eggs in an average of 200 early-instar
greenhouses are discussed here. Most round hole cut in the back of the
aphids. The parasite pupates inside the
wasps have similar developmental mummy. A generation can be com-
mummy, which turns a dull black. The
processes. They are born in the aphid pleted in 12 days at about 75°F. It is
females also kill three to five young
and, as larvae, consume it from the commercially available.
aphids each day by feeding directly on
inside. They pupate inside the mummy
aphid body fluids that exude from holes
and later emerge.
made with the ovipositor. It is not yet
Parasites available commercially.
The parasitic wasps that are important
natural enemies of aphids do not have
common names. Those discussed here
are in the families Aphelinidae and
Braconidae. The Braconidae includes the
subfamily Aphidiinae, which is occasion-
ally treated as a separate family, the
Aphidiidae.
Koppert B.V.

Aphidius colemani, a braconid wasp, parasitizing an aphid.


16 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Aphidius matricariae. This 1⁄16 inch


(2 mm) long, black braconid wasp from
Europe is one of the most common and
effective parasites of the green peach
aphid. A. matricariae is not a good
parasite of cotton aphid or potato aphid.
Females lay 50–150 eggs singly in aphid
nymphs of all sizes. The wasp larvae

Max Badgley
consume the aphids from inside. As the
larvae mature and the aphids are killed,
over about 7–10 days, the aphids turn
into mummies—smooth, shiny, and light
Aphidius matricariae, a braconid wasp, parasitizing an aphid.
brown to silvery-gold. After the larvae
pupate, each adult wasp emerges in
Diaeretiella rapae. This cosmopolitan Lysiphlebus testaceipes. This aphidiid
about 5 days through a round exit hole
aphidiid species, probably native to wasp is an effective parasite of all instars
cut in the mummy. In addition to killing
Europe, is normally a parasite of the of cotton aphid, but not of green peach
aphids directly, mechanical disturbance
cabbage aphid, Brevicoryne brassicae, aphid or potato aphid. The aphid
of aphid colonies by the searching
but it readily attacks green peach aphid mummies are yellowish brown to
activity of the adult wasps causes many
in the greenhouse. Females lay an brown. It is adaptable to a range of
aphids to fall off the plants and die. This
average of 85 eggs during their lifetime climatic conditions. This wasp is com-
wasp is available commercially.
and prefer half-grown nymphs over first mercially available.
instars or adults. Parasitized aphids
Predators
become golden brown mummies. This
Aphidoletes aphidimyza. The larvae of
insect is sold commercially.
this midge feed on over 60 aphid species,
Ephedrus cerasicola. This European including all that occur on greenhouse
aphidiid wasp parasitizes green peach crops.The adults are small—less than 1⁄16
aphid. The aphids become black
J. van Schelt, Koppert B.V.

inch (2 mm) long—black, delicate flies,


mummies about 12 days after the female similar in appearance to fungus gnats,
oviposits in the aphids. The parasitized that live for an average of 10 days by
aphids often move from the plant to the feeding on aphid honeydew.They hide
pot or support structures before beneath the leaves during the day and
becoming mummies. The adult wasps are active at night. Females deposit
emerge about a week after mummies 100–250 tiny (less than 1⁄64 inch, or 0.3
The midge Aphidoletes aphidimyza. appear. E. cerasicola is not yet commer- mm), shiny orange eggs singly or in small
Females can lay up to 250 eggs cially available in North America. groups among aphid colonies.The bright
among aphid colonies. orange, slug-like larvae that hatch in 2–3
days inject a toxin into aphids’ leg joints
to paralyze them, then they suck out the
aphid body contents through a hole
chewed in the thorax.The larvae grow up
to 1⁄8 inch (3 mm) long and each kills
4–65 aphids per day. Larvae can consume
aphids much larger than themselves and
may kill many more aphids than they eat
when aphid populations are high. After
3–7 days, the larvae drop to the ground
Cliff Sadof

Midge larvae Aphidoletes


aphidimyza are highly effective
predators of aphids.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S A P H I D S 17
and burrow 3⁄4–11⁄2 inches into the soil The aphidlions that emerge look like Chrysoperla comanche—Comanche
to pupate. Optimal conditions for their green-gray alligators with mouthparts lacewing. This commercially available
development are 73°–77°F and a relative like ice tongs. An aphidlion seizes its species of green lacewing is closely
humidity of 80–90%. A. aphidimyza enters prey, punctures it with its long jaws, related to C. rufilabris (below). It
diapause under short day conditions.This then injects a paralyzing venom and commonly occurs in orchards and vine-
insect is commercially available. sucks out the body fluids. In addition to yards at low elevations in the southern
Chrysoperla (=Chrysopa) carnea— aphids, aphidlions will feed on a wide parts of Texas, New Mexico, Arizona, and
green lacewing. The larvae of these variety of soft-bodied insects including California. Its biology and development
lacewings are known as aphidlions for a thrips, mealybugs, immature whiteflies, is similar to C. carnea, but it is better
reason—they are voracious feeders that and small caterpillars, as well as insect adapted for development under dry
can consume up to 425 aphids or other eggs and spider mites. They will also conditions.
prey per week. The light-green adult consume each other if no alternative Chrysoperla rufilabris. This is another
lacewings have slender antennae, prey is available. After growing to 3⁄8 species of green lacewing sold for
golden eyes, and large, veined, gauze- inch (1 cm) long during a 2–3 week control of soft-bodied pests, including
like wings that are 1⁄2–3⁄4 inch (1.3–2 period, the larva spins a spherical, white aphids. It is common in tree crops in the
cm) long. They are slow-flying, nocturnal silken cocoon in which it pupates, northeastern United States. Its biology
insects that feed on nectar and pollen, emerging in 5 days as an adult. The and development is similar to C. carnea,
and emit a foul-smelling fluid from adults require pollen in order to repro- but it is better adapted for development
special glands when captured. Female duce. Green lacewings are available from under humid conditions than C. carnea.
green lacewings usually lay up to 300 many commercial suppliers.
eggs in groups on leaves, over a period
of 3–4 weeks. Each egg sits on the end

Richard Lindquist, The Ohio State University


of a slender stalk, about 1⁄3 inch (8 mm)
long, attached to the leaf’s surface.
Freshly laid eggs are green, but they
change to whitish gray as they get close
to hatching.

A green lacewing larva, also called an aphidlion, of


Chrysoperla comanche.
Daniel L. Mahr

The distinctive stalked egg of the green


lacewing, Chrysoperla carnea. The stalk is
about 1⁄3 inch long.
Cliff Sadof

An adult green lacewing, Chrysoperla rufilabris.


18 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Deraeocoris brevis. This mirid bug feeds Hippodamia convergens—convergent Other lady beetles. Scientists have
primarily on aphids and whiteflies, lady beetle. Adults are the common red examined several species of lady beetles
although it will attack thrips and small lady beetles with black spots on each as potential biological control agents of
caterpillars. Both nymphs and adults are wing cover. Although they will attack aphids in greenhouses. Introduced at a
predaceous. The 3⁄16 inch (5 mm) long various soft-bodied insects, convergent ratio of one adult lady beetle to 20
females lay up to 200 eggs in plant lady beetles are sold primarily for aphid aphids, Cycloneda sanguinea larvae con-
tissue. Nymphal development takes control. Each will consume as many as trolled cotton aphid on cucumber. In a
25–30 days, and the adults live about 3 2,000 aphids during its life. Females lay series of experiments on cucumbers in
weeks. Optimum conditions are up to 1,500 orange, bullet-shaped, 1⁄25 small greenhouses, the second and third
64°–85°F with 30–60% relative humidity. inch (1 mm) long eggs in clusters on generations of the beetle eliminated the
D. brevis enters diapause when leaves near the prey. These hatch into cotton aphid. However, the commercially
daylength is less than 10 hours and tem- tiny, black, alligator-shaped larvae with available Coleomegilla maculata was
peratures are below 73°F. This bug is conspicuous orange markings. They are ineffective against cotton aphid on
commercially available. somewhat slender, with the body cucumber because it would not remain
tapering to a point at the rear and on the leaves. Adalia bipunctata and
prominent legs that stick out from the Coccinella septempunctata have been
sides. They feed for 3–4 weeks, consum- used experimentally in Finland for aphid
ing 500–1,000 aphids or similar prey control on chrysanthemums and roses.
during their growth. They generally Lemnia biplagiata, imported from
pupate where they were feeding. Vietnam for use in Russian greenhouses,
Hippodamia convergens is one of the controlled cotton aphid on cucumber
most widely available lady beetles in and green peach aphid on peppers.
North America. Many of these beetles are not suitable
for commercial production and only C.
maculata is currently available.
University of Florida

A convergent lady beetle, Hippodamia convergens. Lady


beetles are voracious feeders, consuming as many as 2,000
University of Florida

aphids in their lifetime.

Lady beetle larva feeding on aphids. Note that


some aphids have already been parasitized by
braconid wasps.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S A P H I D S 19
Macrolophus costalis. This mirid bug Micromus angulatus. This brown
was investigated in Poland for control- lacewing has recently been investigated

Oregon State University Extension Service


ling aphids on greenhouse crops. It as a beneficial insect for the biological
feeds on aphids and greenhouse control of small, soft-bodied insect pests
whitefly larvae and was able to control in greenhouses. Its prey includes small
aphids at predator-to-prey ratios of 1:2 aphids, immature whiteflies, and insect
and 1:3. It reduced aphid numbers best eggs. It has been successfully mass
when the aphid populations were high. reared in Germany but is not yet com-
It is not commercially available, mercially available in North America.
although a related species, M. Orius spp. Several species of minute
caliginosus, is offered as a predator of pirate bug, which are generalist preda-
whiteflies. tors, feed on aphids, thrips, spider mites,
whiteflies, and caterpillar eggs, as well as
pollen and plant juices. They The slug-like maggot of the syrphid fly can
consume as many as 400 aphids during
are also cannibalistic under
development.
crowded conditions. The
black, 1⁄16–3⁄16 inch (2–5
mm) long adults are ovoid Syrphid or hover flies. The larvae of
and somewhat flattened, many species of syrphids can control
with distinctively patterned aphid populations rapidly. They occa-
black and white wings. They sionally move into greenhouses from
lay their eggs in leaf tissues outside, where they are common and
with one end of each egg important natural enemies of aphids.
sticking out. The tiny Orius The adult flies, which are often seen on
nymphs are pinkish-yellow or hovering near flowers, are small to
Les Shipp

to light brown. Both nymphs medium in size (3⁄8–3⁄4 inch, or 8–20


and adults are very active mm), often with a striped yellow and
and will feed on small black body resembling honey bees or
The minute pirate bug, Orius spp., is a generalist wasps. Females lay eggs near aphid
that feeds on aphids, thrips, spider mites, whiteflies, aphids, although they will
colonies. The slug-like, pale green to
and caterpillar eggs, as well as pollen and plant juices. feed on other prey if avail-
able. Several species of Orius yellow maggots feed on aphids, scales,
are available commercially. and other insects, growing to 3⁄8–5⁄8
inch (10–15 mm) in length. Larvae can
consume as many as 400 aphids during
their development. Some species
pupate on the foliage near the feeding
site, whereas others leave the plant to
pupate in the soil. The pupa is enclosed
within a puparium—the hardened skin
of the last larval instar—which is often
teardrop-shaped, smooth, and tan
colored. The life cycle of most species
lasts 2–4 weeks. Syrphid flies are not
commercially available.
Max Badgley

An Orius nymph.
20 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Pathogens Eventually the entire body cavity is filled Paecilomyces fumosoroseus. This insect
Beauveria bassiana. This is a common with fungal mass. When conditions are pathogenic fungus has a wide host
soil-borne fungus that occurs world- favorable the fungus will grow through range, with several strains infecting
wide. It attacks a wide range of both the softer parts of the insect’s body, pro- insects in over 25 different families,
immature and adult insects, including ducing the characteristic “white bloom” including aphids, whiteflies, thrips, and
some natural enemies. As with all appearance. Relative humidity must be some natural enemies. Fungal spores
insect-pathogenic fungi, Beauveria 92% or more for B. bassiana to grow bore through the insect’s skin. As the
produces spores that are resistant to outside the insect. These external spores grow, they consume the insides
environmental extremes and are the hyphae produce conidia that ripen and of the insect, eventually killing the host.
infective stage of the fungal life cycle. are released into the environment, com- External mycelium on infected insects is
The spores (conidia) infect directly pleting the cycle. This fungus is commer- white at first, then changes to shades of
through the outside of the insect’s skin. cially available. pink. The infected insect eventually
Under favorable temperature and Metarhizium anisopliae. This insect becomes light gray when spores are
moisture conditions, a conidium pathogenic fungus infects over 200 produced on the outside of its body. The
(singular of “conidia”) adhering to the species of insects. It occurs naturally in infection cycle is very rapid, with sporu-
host cuticle will germinate. The fungal soils throughout the world. Green, cylin- lation occurring within 72 hours of
hypha growing from the spore secretes drical spores are produced in chains infection and peaking at 5–7 days. This
enzymes which attack and dissolve the from infected insects. Because this fungus requires humidity over 90% for
cuticle, allowing it to penetrate the skin fungus has a wide host range and infection and germination is poor when
and grow into the insect body. Once infects some beneficial insects, including humidity is below 98%, which limits its
inside the insect it produces a toxin lady beetles, it may not be compatible utility. Virulence varies among strains;
called beauvericin that weakens the with all aphid predators and parasites. the most virulent strains are being com-
host’s immune system. After the insect However, it is not toxic to plants, honey mercialized. This fungus is commercially
dies, an antibiotic (oosporein) is bees, earthworms, fish, or humans. A for- available in Europe, but is not registered
produced that enables the fungus to mulation of this fungus can be applied for use in the United States.
out-compete intestinal bacteria. like a conventional insecticide, but this is Verticillium lecanii. This insect patho-
not yet registered for use in the United genic fungus infects both aphids and
States on any greenhouse crops. whiteflies. The fungal strain with large
spores infects aphids; the strain with
smaller spores is specific to whiteflies.
Relative humidity over 95% must be
maintained for infection to occur. The
fungal spores kill aphids by growing
through the aphid’s skin. The spores
continue growing inside the insect,
eventually consuming the internal
Don McLean, University of Vermont

contents. Aphids killed by this fungus


remain on the plant, covered in spores.
Under the right conditions spores stick
to healthy aphids that touch them and
initiate a new infection cycle. Vertalec
and Mycotal are commercial formula-
tions of the fungus specific to aphids
that are available in Europe. They are not
currently registered for use in North
The fungus Metarhizium anisopliae infecting a green peach America.
aphid.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S A P H I D S 21
Possibilities for effective more useful in a biological control or up to three pupae per 10 ft2, or
biological control program for aphids on greenhouse 2,000–6,000 pupae per acre are success-
Aphids can be effectively controlled by crops than lady beetles or green ful for controlling aphids on most plants.
several commercially available natural lacewings. However, many aphids killed Two to four applications are often nec-
enemies, released singly or in combina- by midge larvae remain attached to the essary to achieve control. Additional
tion. Aphid populations can increase leaves, which may detract from the introductions of the midge will not be
very quickly, so it is best to release value of ornamental plants. necessary if successive generations are
natural enemies while populations are Aphid midges are shipped as pupae produced during the growing season. If
low. If more than 10% of the plants are (inside their cocoons) in moist vermicu- aphids can be tolerated on the lower
heavily infested, you should reduce lite. Sprinkle the vermiculite with the leaves of the plants, the midge popula-
aphid populations before you release cocoons on moist soil or root medium tions will build up quickly on the supply
parasites or slower-acting predators. Use within the plant canopy. Adults will not of aphids.
either nonresidual chemicals (insect emerge from cocoons that fall on dry Biological control of aphids with A.
growth regulators, soaps, oils, etc.), or areas in direct sunlight. The bright aphidimyza is most successful in green-
predators, such as green lacewing orange midge larvae should be visible houses with soil beds or gravel floors,
larvae, that will quickly eat large among the aphids about a week after probably because the midge larvae
numbers of aphids. the adults have emerged. The adults easily find suitable sites for pupation.
Aphidoletes aphidimyza has been used hide beneath the leaves during the day Sprinkle a thin layer of sawdust or peat
successfully for biological control of and are active at night. Optimal condi- moss between the rows on concrete or
aphids on cucumber, green pepper, tions are 68°–81°F and 50–90% relative plastic-covered floors to provide
tomato, chrysanthemum, rose, and many humidity. Recommended release rates suitable pupation sites in these green-
other ornamentals on a commercial vary depending on the plant type, houses. Without proper pupation sites
scale. Unlike other predators, this midge growth stage, aphid species, and infesta- on the floor, fewer midge offspring
reproduces during the growing season, tion level. Lower release rates may be survive, and weekly releases may be
so only a single release may be neces- used before aphids are observed for necessary throughout the season to
sary, although two to four introductions preventative control. Weekly or biweekly provide adequate control.
are often required. A. aphidimyza may be releases of one or two pupae per plant,

Aphids
Enemies egg ————— nymph ————— adult
Parasitoids

Aphidoletes

Green lacewings

Deraeocoris

Lady beetles

Macrolophus

Minute pirate bugs

Syrphid flies Natural enemy attacks the host


Beauveria
Effectiveness depends on
Metarhizium species of enemy

Paecilomyces Width of bar indicates degree


Verticillium of effectiveness
22 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

The midge can also be introduced from introduced in parasitized aphids, Aphidius ervi and Aphelinus abdominalis
open rearing units or banker plants packaged in boxes of rooted cuttings are two other commercially available
placed in the greenhouse. Aphid species (50 aphids per box of 500 cuttings). It is wasps that can be utilized against
that do not infest commercial green- not as effective against cotton aphid as certain aphid species. A. ervi is most
house crops—such as the grain aphids it is against green peach aphid. effective against potato and foxglove
Sitobion avenae, Metopolophium A. matricariae performs best when it is aphids when released before aphid pop-
dirhodum, and Rhopalosiphum padi—are established in the greenhouse early in ulations build up. It should be released
used as prey for A. aphidimyza. Place the growing season. Native A. matri- at a rate of about 0.15 adults per 10 ft2
these aphids on wheat or barley plants cariae entering the greenhouse from every week. If aphids are already
(the banker plants) in open trays or outside sometimes become established present, suggested release rates are 0.5
boxes. The predators develop on these and provide effective control. However, per 10 ft2 and introductions should be
aphids, and the adult predators disperse in the spring and summer these wasps made every 3 days if aphid populations
to lay eggs on infested crop plants. The are frequently attacked by their own are high. A. abdominalis, another good
rearing units must be established early native hyperparasites, which reduce the parasite of potato and foxglove aphids,
in the growing season and maintained natural or introduced A. matricariae pop- should be released in infested areas only
for several weeks so that a large midge ulation. Adult A. matricariae are attracted since this wasp is not very mobile. Make
population has developed by the time to the color yellow, so yellow sticky weekly introductions of two to four
pest aphids appear. This is not a practical cards should be removed before adult wasps per 10 ft2 depending on
method in large commercial green- releases are made. A release rate of two aphid population density until 80–90%
houses, but boxes containing growing per 20–800 ft2 is recommended. It is of the aphids have become black
barley with grain aphids can be pur- effective when aphid populations are mummies. After that time small quanti-
chased from some suppliers for this use. low, but should be used in combination ties can be released.
One drawback of A. aphidimyza is that it with other natural enemies when popu- Many general predators, such as lady
enters into diapause under cool, short- lations are high. This wasp is not com- beetles, lacewings and hover flies, also
day conditions. Diapause can be pre- patible with insect pathogenic fungi provide effective and rapid control of
vented by leaving on a few incandes- such as V. lecanii which kill the parasite large aphid populations. Adults lay eggs
cent walkway lights all night during larvae inside the aphid. only near large numbers of aphids.
winter months (through late February). Diaeretiella rapae can be as effective as However, most species will not repro-
Larvae are very sensitive to light, so a A. matricariae. It has eradicated green duce in the greenhouse. In order to
single, 100-watt bulb will prevent over peach aphid on chrysanthemum in 6 obtain control you must release them
half of the midges within a circle with a weeks when released at a rate of one every 2–3 weeks.
diameter of 24 yards from diapausing. parasite per 500 aphids. Release rates The convergent lady beetle can
This will not be effective when plants vary by aphid species and stage of infes- suppress high aphid populations
are so large that light does not pene- tation and should be determined on an quickly, but establishing a population in
trate between the rows. If supplemen- individual basis. the greenhouse is difficult. Thus, conver-
tary lighting to prevent the midges from gent lady beetles are used most effec-
Aphidius colemani is a very promising
diapausing is not feasible, and aphids tively to reduce high aphid populations
candidate for biological control in
are a problem in the fall or winter, other that can then be kept under control by
greenhouses because of its high repro-
natural enemies, such as the parasitic other predators, parasites, or pathogens.
ductive potential, short development
wasp Aphidius matricariae, can be used Most commercially available convergent
time, and ability to parasitize several
for aphid control. lady beetles are not insectary-reared,
species of aphids, including cotton
Because Aphidius matricariae does not aphid, melon aphid, and green peach but are collected from winter hiberna-
diapause under winter greenhouse con- aphid. It is more efficient at parasitizing tion sites. Only when their stored winter
ditions as readily as A. aphidimyza, it can cotton aphid on cucumber than A. matri- fat reserves have been depleted will
be an important part of a biological cariae is. they become predaceous. Even then,
control program from fall through early after consuming many aphids, lady
spring. On chrysanthemum, green peach beetles often fly to the vents and leave
aphid has been controlled within 2 to lay eggs outside. These beetles may
months of planting when the wasp was also be parasitized. Repeated releases of
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S A P H I D S 23
convergent lady beetles are often neces- these aphidlions do not eat the new eggs to plants or containers. This is
sary. Determine release rates on a case- lacewing eggs or larvae. Where aphid especially helpful in making applications
by-case basis. Make releases in the densities are low (four or fewer per in hanging baskets.
evening, because bright sunlight plant) lacewing larvae are not effective, The lacewings should be released as
encourages their flight. Keep the vents and the parasite Aphidius matricariae, soon as they begin to hatch. Releases
closed or screened to prevent the which is more effective at low aphid are made by sprinkling the contents of
beetles from leaving the greenhouse. densities, should be used. However, if this the container onto infested plants. The
Green lacewing larvae have controlled wasp is released while lacewing larvae newly hatched larvae will be very tiny
aphids effectively on green pepper, are still feeding, the lacewings may (about the same size as the eggs) so you
parsley, chrysanthemum, snapdragon, consume parasitized aphids, thereby may have difficulty seeing them. The
and many other kinds of ornamentals. In reducing the wasp population. released aphidlions will travel a consid-
an experiment on snapdragon, the The number of lacewings needed for erable distance, up to 100 feet, in search
flower quality of plants with aphids con- effective control depends on the crop, of prey. Making releases early in the
trolled by green lacewings equalled the the growth stage, the aphid species, and morning or late in the day when it is
quality of flowers produced with the degree of infestation. Releases cooler, or on a cloudy day, increases the
chemical applications. Lacewings are should be initiated early, when aphids chances the lacewings will survive.
most effective at high aphid densities, are discovered on a few plants. You can Larger larvae, which consume aphids at
eliminating large aphid populations or achieve control at higher aphid densi- a faster rate than newly hatched larvae,
reducing them to such low levels that ties, but the number of lacewing eggs are available from some suppliers.
additional control is often not necessary. needed is very high. Suppliers usually Because they are cannibalistic, lacewings
The effect of a release will be apparent in make recommendations based on purchased as large larvae must be
1–2 weeks. If additional control is neces- specific situations. For control of shipped in individual containers which
sary, the next release should be delayed moderate aphid infestations, 5–10 increases the cost of the product.
until the larvae from the previous release lacewing eggs per plant or 1,000 eggs Lacewings released as pre-fed adults
have finished their development so per 200 ft2 are recommended. Release that are ready to lay eggs can fly away
recommendations for larger upon opening the shipping container, so
areas start at 5,000 per acre greater care must be taken when releas-
for each application. These ing lacewings at this stage to ensure
insects are extremely effec- their establishment in the infested area.
tive, but cannot multiply in The commercially available mirid bug
greenhouses and must be Deraeocoris brevis can also be used to
released at regular inter- control aphids. Since this insect can
vals—about every 2 weeks. survive on pollen if prey are scarce,
Green lacewings are usually including some pollen-producing plants,
sold as eggs but also may be such as pepper, may help retain the
sent as larvae or adults. Eggs insect in the greenhouse. The bugs are
are sent in a packing material shipped as adults with some nymphs
to cushion and separate the present. Release rates should be
emerging larvae during provided by the supplier.
shipment. The material—rice Several natural enemies of aphids that
hulls, wheat bran, or corn are not commercially available in the
grits, along with moth eggs United States have been investigated in
for food so the larvae will be Europe. The parasites Aphelinus flavipes,
less likely to eat each other— Ephedrus cerasicola, and Lysiphlebus tes-
also makes it easier to distrib- taceipes, and the fungus Verticillium
Ray Cloyd

ute the very tiny eggs evenly. lecanii show promise as effective biolog-
Lacewing eggs can also be ical control agents against aphids in
Lady beetles can be good predators of aphids. mixed with Biocarrier, a greenhouses. Aphelinus flavipes is most
material that helps to glue effective at cooler temperatures, so the
24 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

parasite can overtake a slow aphid pop- Several brands and formulations of the V. lecanii is applied as spores suspended
ulation increase. A high parasite-to-host fungus Beauveria bassiana are available in water for spraying. High humidity is
ratio and early introduction are neces- for use in greenhouses. Because it takes necessary for spore production and
sary for reliable control. A. flavipes was 3–7 days to kill an insect with B. bassiana, infection. When humidity is low, the per-
only able to control cotton aphid on it will take some time to suppress the formance of the fungus is unpredictable.
cucumber when the parasites were pest population when using these Humidity can be increased by dampen-
introduced before or at the time of products. Thorough spray coverage is ing the plants with water sprays. Late
aphid infestation. essential because fungal spores must afternoon applications reduce spore
Ephedrus cerasicola was considered contact the insect for infection to occur. injury by ultraviolet light and desicca-
better than A. aphidimyza in Norway, This fungus is more tolerant of lower tion, since the greenhouse is more
although control by both species was humidity than many other insect patho- humid at night. Alternating two nights
similar, because (1) it does not diapause genic fungi, so high relative humidity is of fogging—to produce the elevated
and therefore provides control through- not necessary for infection to occur. This humidity necessary for infection—with
out the season, (2) it can be introduced fungus is susceptible to some fungi- two nights of ambient conditions may
at lower aphid densities, and (3) fewer cides, so chemical fungicide applications provide an adequate environment for
individuals need to be released. When should not be made within 48 hours of control. However, this fungus may be
E. cerasicola was released early in the B. bassiana applications. Three to five practical for use only in humid areas
season, it kept green peach aphid popu- applications may be necessary to with moderate temperatures, such as
lations on peppers below damage achieve control. rooting benches and shade-cloth
thresholds and eliminated the aphids Although Verticillium lecanii is not avail- covered areas used to induce inflores-
within 2 months. able for use in the United States yet, this cence in chrysanthemums. Repeated
fungus may eventually be registered applications of V. lecanii will be neces-
These wasps are released as pupae
and provide effective control of some sary if humidity is not high enough to
inside aphid mummies. Mummies can
species of aphids here. Under the appro- allow continuous infection. This fungus
be placed among the plants or intro-
priate conditions, it has the potential to may be susceptible to some fungicides.
duced on banker plants in a cage in the
center of the greenhouse. Parasite-to- eliminate the need for insecticide appli- If aphids are numerous, other means of
host ratios of between 1:5 and 1:10 are cations for aphid control. V. lecanii effec- reducing their numbers (such as releas-
probably sufficient. Two introductions at tively controls most aphids, including ing lacewing larvae or chemical control)
10-day intervals of four mummies per green peach aphid, but does not control must be used before treatment with
plant should be effective when there is chrysanthemum aphid. A single spray of V. lecanii. Two to 3 weeks are required to
less than one aphid per plant. At higher V. lecanii was sufficient to control green control an aphid population. If after 3
aphid populations, a single introduction peach aphid on chrysanthemum, but weeks of treatment less than 90% of the
of one mummy per 10 aphids should the fungus did not control chrysanthe- aphids are infected by the fungus, you
suffice. New introductions may not be mum aphid on the same plants. The should employ other means of control.
necessary because this wasp will cotton aphid often escapes contact with Other fungi, such as Metarhizium aniso-
multiply in the greenhouse. However, if the spores because it does not move pliae and Paecilomyces fumosoroseus,
the aphids begin to increase or the around much. Only 80–90% control of may also provide effective control of
parasite dies out, additional releases cotton aphid can be expected. Plants aphids if these products become avail-
should be made at the same ratio. should be treated when small to keep able in the United States. In lab experi-
the aphid population under control. ments, Russian wheat aphid was more
The aphidiid Lysiphlebus testaceipes has
Otherwise, the fluffy white aphid bodies susceptible to P. fumosoroseus than to
been released outdoors in
stuck to the mature foliage will make B. bassiana.
Mediterranean areas for control of
the plants commercially unacceptable.
several aphid species and is being inves-
The fungus also kills A. matricariae and
tigated for control of cotton aphid in
whitefly parasites inside the pest body,
greenhouses. It has considerable poten-
but it seems to be compatible with para-
tial for biological control, but because it
sites of other pests, such as the eulophid
does not parasitize green peach aphid
wasp, Diglyphus begini, that attacks
well, an additional parasite might be
leafminers.
needed to assist in control.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S A P H I D S 25

Alternative control methods


Sanitation Horticultural oils, both refined petro-
Weeds may harbor aphids both inside leum distillate products and those
and outside the greenhouse. Pull made from vegetable oils, can kill
weeds or use commercially available aphids and other insects. These horti-
herbicides; use barriers to prevent cultural oils may be toxic to some
weed growth. plants. Many brands are registered for
use on vegetables and ornamentals in
Insect screens
greenhouses to control many pests,
Screens on vents and doors will help
including aphids.
prevent aphids from getting into the
greenhouse, but often the most effec- Pyrethrins are insecticides derived
tive screens reduce air flow. The from the flowers of the pyrethrum
maximum hole size to exclude cotton daisy, Chrysanthemum cinerariaefolium.
aphid is 341 µ (micrometers). Be sure to They are extremely fast–acting and
screen the greenhouse before the crop cause immediate knockdown, but most
growing season, when the potential for insects can recover after a brief period
aphid problems is still low. of paralysis. Other ingredients can be
added to prevent this recovery. Most
Host plant cultivar pyrethrin products are sold commer-
Aphids feed preferentially or repro- cially as a premix with rotenone, with
duce at different rates on different cul- piperonyl butoxide, or with piperonyl
tivars of chrysanthemum. In one exper- butoxide and diatomaceous earth.
iment, for example, 40 times more
Kinoprene is an insect growth regula-
green peach aphids were found on the
tor registered for use on ornamentals
cultivar Tuneful than on Portrait. Green
for control of aphids and other insects.
peach aphids were found to reproduce
It can be used when aphid mummies
much faster on Tuneful than on Golden
or predators are present. It has minimal
Princess Anne.
activity on natural enemies but may
Chemical control damage some plants. Some rose vari-
Localized infestations and high popu- eties show delayed damage.
lations that need to be reduced before Azadirachtin is another insect growth
predator or parasite introduction can regulator that can be used in rotation
be spot-sprayed with selective chemi- with kinoprene, although it generally
cals. Insecticidal soap can effectively acts as a feeding deterrent for aphids.
control aphids, but should be used This is a commercial formulation of an
only if no predators, including preda- extract from the seeds of the neem
tory mites, or adult parasites are tree (Azadirachta indica). Several for-
present because it will affect exposed mulations are registered for use on
natural enemies. Once the soap is dry, greenhouse ornamentals and
however, it is nontoxic. Thorough vegetables.
coverage of infested surfaces is essen-
tial, and more than one application
may be required.
26 Caterpillars
T
he larval, or caterpillar, stages of Description and life cycle
Available natural enemies and several moths are occasional pests The female moths, which are nocturnal,
their potential for control of many greenhouse crops, espe- lay eggs on plants or other surfaces in
Bacillus thuringien- cially in summer when lights inside the the greenhouse. The caterpillars that
sis (Bt) can be very greenhouse attract moths at night from hatch from these eggs feed on the
effective against outside through open vents. Beet army- underside of the leaf for several days,
young caterpillars, worms are common on carnation, often leaving the upper leaf surface
but other natural chrysanthemum, cyclamen, geranium, intact. As they grow larger they eat
enemies are of snapdragon, and other crops. Cabbage through the whole leaf and may feed on
limited value. loopers are particularly damaging on fruits or flowers. Caterpillars often feed
carnation, chrysanthemum, geranium, at night or on cloudy days and hide
Order Lepidoptera: Butterflies,
nasturtium, and crucifer sets. Corn during the day at the base of the plant
moths, and skippers
earworms prefer the buds of chrysan- or in camouflaged positions along mid-
Family Noctuidae: Owlet moths and themum, gladiolus, and rose, as well as ribs, leaf edges, or petioles. Caterpillars
underwings small tomato fruits. European corn of noctuid moths can reach 2 inches
Beet armyworm, Spodoptera borers tunnel into the stems of plants, (5 cm) in length. The different species
exigua particularly chrysanthemum and dahlia. vary in color. They all pupate on or just
Black cutworm, Agrotis ipsilon Variegated cutworm, a climbing species, under the soil surface.
is one of the most common species on The grayish brown beet armyworm
Variegated cutworm, ornamentals and often causes damage
Peridroma saucia moths are about 3⁄4 inch (2 cm) long.
to tomatoes, carnations, and chrysanthe- Females lay 100–150 eggs in small piles
Cabbage looper, Trichoplusia ni mums. Black cutworms damage on the undersides of lower leaves. The
Corn earworm or tomato seedlings, vegetables, and flower crops. smooth, green caterpillars have a wide
fruitworm, Helicoverpa zea Many other species of cutworms, black stripe down each side and grow to
loopers, or other caterpillars may occa- 11⁄8 inches (3 cm) long. After about 2
Family Pyralidae: Pyralid, grass,
sionally infest greenhouse crops. weeks they pupate in a loose cell on the
and wax moths
medium surface.
European corn borer, Damage
Ostrinia nubilalis Black cutworm adults are dark red to
Caterpillars chew off pieces of foliage
blackish-brown in color, with indistinct
and leave plants ragged in appearance.
dark markings on the wings. The females
Some species also feed directly on
lay eggs singly or in small groups on
flowers or fruit, which is often when
leaves or stems. The greasy-appearing
people first notice the infestation.
larvae are gray to black along their back
Foliage losses usually matter less than
and pale underneath. They grow to
damage to buds. Damaged buds don’t
11⁄4–13⁄4 inches (3.0–4.5 cm) long. They
produce flowers but axillary shoots
pupate under debris on the medium
instead. Shoots tunneled into by
surface.
European corn borers wilt and do not
produce flowers. The adult variegated cutworm has a
wingspan of about 11⁄2 inches (4 cm).
The wings are a mottled dark gray with
a purplish tint and two indistinct spots
in the upper middle of each wing. The
eggs are laid in clusters of 200–500 on
the undersides of leaves. The caterpillars
are gray or brown with blotchy gray and
black markings and a row of pale, small,
almost diamond-shaped spots along the
sides. They grow up to 13⁄4 inches
(4.5 cm) long. They pupate in the soil.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S C A T E R P I L L A R S 27
The 1-inch (2.5-cm) cabbage looper Monitoring Parasites
moths are mottled dark-brown and have Moths are often attracted to a green- Trichogramma spp. Trichogramma
a silvery figure 8 on each forewing. They house’s production or security lights, wasps attack the eggs of over 200
lay round, white eggs singly or in small then enter through openings and species of moths and butterflies. These
groups on the undersides of leaves. The proceed to lay eggs. Adult populations almost microscopic wasps (1⁄64 inch,
caterpillars are light-green with faint can be detected—and low numbers 0.5 mm) are very important in prevent-
white stripes down each side, and they controlled—with blacklights or, for some ing crop damage because they kill their
taper from the tail toward the head, so species, pheromone traps. Small green hosts before the insects can cause plant
the rear is fatter than the front. They fecal pellets near chewed foliage often damage. The female Trichogramma lays
grow up to 11⁄2 inches (4 cm) long. The indicate a caterpillar attack. Search an egg into a recently laid host egg. As
caterpillars move with a characteristic plants for egg masses or young caterpil- the wasp larva develops, the host egg is
looping motion produced by holding on lars when moths are caught in traps. killed and turns black. Each female para-
with the front legs and arching the These pests may also be brought into a sitizes about 100 eggs. The wasp’s short
middle portion of the body to bring the greenhouse as eggs or young caterpil- life cycle of 8–10 days allows their popu-
prolegs (hind legs) forward, then lars on cuttings, so new shipments lation to increase rapidly. These wasps
extending the front of the body while should be carefully inspected. are harmless to people, animals, and
holding on with the prolegs. Pupation plants. There are many commercially
occurs in a loosely woven cocoon Natural enemies available species, such as T. pretiosum,
attached by one side to a plant leaf. Numerous parasites and many predators T. minutum, and T. platneri.
Corn earworm moths vary considerably attack caterpillars, but few are suitable Predators
in color and markings, from tawny with for control in greenhouses. Of the many
Many general predators will feed on
faint markings to reddish-brown with nematodes, viruses, and bacteria that are
caterpillars or their eggs. Larvae of the
heavier markings. The 3⁄4-inch (2 cm) used commercially for control of cater-
commercially available green lacewings
females lay 500–1000 eggs singly on pillars, only one bacterium is useful on
Chrysoperla (=Chrysopa) carnea and
foliage. The caterpillars grow to 11⁄3 inch greenhouse crops. Chrysoperla rufilabris are voracious pred-
(3.5 cm) long and are highly variable in ators that will feed on moth eggs and
color—red, maroon, orange, yellow, very small caterpillars. (See “Aphids” for
green, and nearly black—with a yellow more information.) Lady beetles and
head and alternating dark and light minute pirate bugs may also feed on
stripes along the length of the body. eggs or small caterpillars if other prey is
They bore into the soil to pupate. not available.
Female European corn borer moths are
about 3⁄4 inch (2 cm) long, vary in color
from yellow to light-brown, and have
two zigzag lines across the outer parts
of their wings. Males are slightly smaller
Richard Gagne, Minnesota Department of
Agriculture, Biological Control Program

and light-brown with yellow zigzag


lines. Females lay 400–500 eggs in
masses of 15–20, with the flat eggs over-
lapping each other like fish scales. The
eggs are white when first laid but turn
darker just before hatching. The caterpil-
lars grow to 1 inch (2.5 cm) long and
range in color from light brown to dark
brown to pink or gray, always with a
dark head and an indistinct stripe on the
back. They pupate within flimsy cocoons Trichogramma minutum wasps have a short life cycle
inside tunnels in stems, or in other pro- allowing their population to increase rapidly.
tected locations.
28 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Pathogens During sporulation Bt forms a parasporal Possibilities for effective


Bacillus thuringiensis. This pathogen, body, or crystal, which is the source of its biological control
commonly known as Bt, is a naturally insecticidal properties.The crystals Bt can provide good control of the most
occurring, soil-inhabiting bacterium that dissolve in the gut juices of susceptible common greenhouse caterpillar pests.
is highly selective and active mainly insects and release toxic compounds. However, Bt does not provide persistent
against caterpillars. There are many Later the spores germinate, and bacteria control under greenhouse conditions
types of Bt and they differ both in speci- grow in the insect body.The first observ- because it does not disperse well
ficity to and potency against a range of able reaction after a caterpillar ingests Bt enough and breaks down rapidly in
target insect species. Only two varieties is paralysis of the gut and mouthparts, ultraviolet light. It must be applied
are toxic to insects other than caterpil- followed by cessation of feeding. Paralysis whenever pest populations develop and
lars: one affects the larvae of mosqui- may happen in a matter of minutes, but in a manner similar to conventional
toes, blackflies, and some other types of death usually takes 30 minutes to 3 days insecticides. Applications should not be
flies; the other affects certain beetles. following ingestion. Bt is available in made on a regular schedule, but should
The varieties kurstaki and aizawi are several commercial formulations. be timed based on the occurrence of
available in commercial formulations for eggs or caterpillars on the plants.
control of caterpillars. This bacterium is Monitor moth flights to determine when
nonpolluting and safe to humans. to begin scouting for eggs. Because Bt
must be eaten to be effective, apply it
where the target insects are feeding. If
infestations are localized, make spot
applications. Bt is not effective against
caterpillars feeding within plant
tissues—for instance, inside stems, buds
or nests of webbed leaves. Bt is more
effective against earlier instars than later
instars.

Caterpillars
Enemies egg —— caterpillar —— pupa adult

Trichogramma spp.

Predators

Bacillus thuringiensis

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S C A T E R P I L L A R S 29
Trichogramma wasps provide good
control of certain species of caterpillars
in outdoor crop systems. In most green-
Alternative control methods
houses, caterpillar infestations are Sanitation Baits
sporadic and minimal. Under these con- Numerous plants can serve as reser- Cutworms are attracted to bran baits.
ditions egg parasites are not very effec- voirs for beet armyworms or other One recipe from the 1930s suggested
tive, and spot applications of Bt would caterpillars. Remove weeds that can moistening 12 pounds of bran with a
be more efficient. However, if large be alternate hosts for these caterpil- mixture of 1 pint molasses, 1 ounce
caterpillar infestations occur regularly, lars from inside and outside the banana oil (amyl acetate), and an
Trichogramma released at the first sign greenhouse. Sterilize new soil that insecticide in about 1 gallon of water.
of moth flight through peak egg laying may contain cutworm eggs, larvae, or Spread the poisoned bait late in the
can provide control. Regular scouting to pupae before bringing it into the day so it will be fresh when the
detect the appearance of caterpillar greenhouse. Check new plant cutworms come out to feed.
eggs is necessary to determine when material entering the greenhouse for
wasps should be released so the appro- Chemical control
eggs or larvae and treat or quaran-
priate stage will be available to Spinosad is an insecticide derived
tine any that are infested.
maximize Trichogramma effectiveness. from natural metabolites produced
Most suppliers of Trichogramma can Insect screens under fermentation conditions by the
make recommendations about the Screens on vents and side walls will actinomycete Saccharopolyspora
species and number of wasps to release. prevent moths from getting into the spinosa. It has a high level of contact
Rates should be determined on an indi- greenhouse. and oral activity and rapid speed of
vidual basis. action. It is especially effective
Hand picking
against caterpillars, leafminers, and
The wasps are shipped as immatures Frequent plant inspections and
thrips, but has low to moderate
inside moth eggs glued to small cards removal of all visible caterpillars may
impact on beneficials. It is registered
that can be attached by hand to be impractical for large plantings, but
for control of many pests on land-
infested plants. Keep the cards in a is an option for small or spotty infes-
scape ornamentals; check with your
warm, humid place out of direct sunlight tations. Inspect plants thoroughly to
chemical supplier on availability for
until the emerging adults can be seen as catch eggs and newly hatched cater-
use on greenhouse ornamentals.
small dots moving around in the closed pillars or those that may have
container. A few tiny worms may also be escaped earlier detection.
found in the container because it is very
difficult to obtain 100% parasitization of
the moth eggs, but these worms are
harmless to plants. When most of the
adults have emerged, place the opened
containers in a shaded spot in areas Green lacewing larvae can provide quick
where you suspect moths are laying control of small, localized infestations.
eggs. The adult wasps will fly onto the They prefer to feed on aphids, so their
plants in search of new host eggs to effectiveness will be limited if aphids are
attack. Do not put the cards out before present. Where caterpillar eggs are
the wasps start to emerge from the present, releases should be made as
moth eggs because ants and other described in the “Aphids” section.
predators may eat them. The best time
to make releases is early morning or
evening when direct sunlight will not hit
the cards. Avoid making releases when
the temperature is above 85°F.
30 Fungus gnats & shore flies
F
ungus gnats and shore flies, Description and life cycle
Available although only distantly related, both Adult fungus gnats are 3⁄16 inch
natural enemies occur in the same environmental (4–5 mm), dark brown to black flies that
and their poten- conditions. Both are nuisances, but only rest on the potting medium or plants.
tial for control fungus gnat larvae injure plants directly, The delicate adults are thin with long
Nematodes, feeding on a variety of plants and on legs and antennae. They run rapidly and
Bacillus thuringiensis (Bt), and preda- cultivated mushrooms. are weak fliers. The wings have a distinct
tory mites can provide excellent Y shape in the middle. Females lay
control of fungus gnats. Damage clusters of 2–30 eggs on moist medium
Fungus gnat larvae usually feed on soil surfaces, producing a total of 100–300
Order Diptera: Flies
fungi, algae, and decaying organic small white eggs during a 7–10 day life
Family Sciaridae: Darkwinged matter. However, when populations are span. The larvae are slender maggots
fungus gnats high, they may feed on root hairs and that have a distinct black head and a
Family Ephydridae: Shore flies tunnel into roots and stems just under transparent white body that is 1⁄4 inch
Fungus gnats, Bradysia impatiens the soil surface. Affected plants may turn (6–7 mm) long by the third and final
and B. coprophila yellow, appear stunted, or wilt during instar. They live on or near the surface of
the day under severe attacks. Cuttings, the medium. The larvae feed for 5–14
Shore flies, Scatella stagnalis prior to setting callus tissue, and tender days, then pupate in the soil and
root systems of young seedlings are emerge as adults about a week later.
most susceptible to feeding injury by Fungus gnats are most prevalent in wet
fungus gnat larvae. Plants may eventu- soil mixes that are high in organic
ally die. In addition, adult fungus gnats matter and new media that are micro-
deposit fecal droppings on plants that bially active.
may reduce the aesthetic quality of the Shore fly adults are about the same size
crop. The adults often get trapped in as fungus gnats, but are darker and
surface moisture on leaves, making more robust, with short legs and
plants less salable because they look antennae. They breed in algae and
“buggy.” standing water or water-soaked areas in
Shore flies are algae feeders and do not the greenhouse. Females scatter eggs
normally damage plant material directly, on moist soil or algae. The larvae that
although larvae will feed on roots hatch in 2–3 days live just below the soil
infected with fungi. Black drops of excre- surface, feeding on bacteria, yeasts, and
ment deposited by the flies on leaves algae for 3–6 days. The adults emerge
can make the plant less attractive. after a 4–5 day pupal period and feed
Both shore flies and fungus gnats can on algae.
transmit some fungal diseases, including
Botrytis, Fusarium, Phoma, Pythium, and
Thielaviopsis basicola when they are
larvae, and Pythium, Fusarium, and
Verticillium when they are adults.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S F U N G U S G N A T S & S H O R E F L I E S 31
Monitoring Predators Pathogens
Yellow sticky cards or stakes are very Hypoaspis (=Geolaelaps) miles. This Bacillus thuringiensis var. israelensis.
effective in monitoring adult fungus laelapid mite is a native American soil There are many types of the naturally
gnat populations. Blue sticky traps may dweller that feeds on many soil-inhabit- occurring Bacillus thuringiensis, a soil-
be more attractive to shore fly adults ing arthropods, including fungus gnat inhabiting bacterium commonly called
than yellow cards. See “Whiteflies” for larvae. It is an aggressive predator that Bt. The types differ both in specificity
more detailed instructions. attacks individuals many times its size. and potency against a range of target
The brown adult females are about insect species. Most are highly selective
Potato wedges cut into 1⁄4-inch sections
1⁄ 25 inch (1 mm) long; males are much and active against caterpillars (the
and placed into the growing medium
smaller. Adults consume one to five larvae of butterflies and moths), but the
surface can be used to monitor fungus
fungus gnat larvae daily, killing more variety israelensis (Bti) is toxic to the
gnat larval populations. Leave the
small prey than large prey. When insects larvae of fungus gnats, mosquitoes,
potato pieces on the medium for 72–96
are scarce, H. miles can survive by scav- blackflies, and some other types of flies.
hours. Fungus gnat larvae feeding on
enging on algae and plant debris. Fungus gnat larvae stop feeding and
the exposed potato surface can be
Females deposit one to three eggs per become limp within 24 hours of ingest-
counted.
day. Nonfeeding, six-legged larvae hatch ing Bti. After 2 days the larvae disinte-
in 2–3 days; they molt into eight-legged grate. Bti is more effective against earlier
Natural enemies
nymphs within about a day. This stage instars than later instars. This bacterium
Several natural enemies attack the
lasts 4–5 days, during which the nymphs is commercially available as a microbial
larvae of fungus gnats and shore flies. A
consume 16–33 young fungus gnat insecticide. See “Caterpillars” for more
naturally occurring diapriid wasp para-
larvae. All stages of the mite are found in information on Bt.
sitoid attacks fungus gnats in British
the top inch of the soil or potting
Columbia, but little is known about it or
medium, and prefer moist soil with an
any other parasitoids. A few predators
open structure. They become inactive
and many pathogens, including
below 59°F, but do not diapause in the
bacteria, fungi, and nematodes, attack
winter. H. miles is offered commercially.
fungus gnat or shore fly larvae in the
soil. Only a few have been examined and
are useful for control of these pests in
greenhouses.

Ray Cloyd

The mites Hypoaspis miles are applied by shaking the container onto
the soil or potting medium.
Max Badgley

The soil-dwelling laelapid mite


Hypoaspis miles.
32 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Nematodes. Several entomopatho- Several species of entomopathogenic as a soil drench once per week for 3
genic (insect-pathogenic) nematodes nematodes, including Steinernema weeks are recommended. Potting
occur naturally in the soil and parasitize feltiae (=Neoaplectana bibionis), S. car- medium, soluble salts, soil temperature,
a variety of soil-inhabiting insects, pocapsae, S. riobravis, and Heterorhabditis moisture, and pH, can all influence the
including fungus gnat larvae. spp. are sold commercially for biological effectiveness of Bt in greenhouses.
Nematodes are small, long, slender control of fungus gnats. H. megadis is Nematodes can also be applied to crops
roundworms. They are about 1⁄64 inch sold in Europe for fungus gnat control. in large quantities as a biological insecti-
(0.5 mm) long, transparent, and practi- The mermithid Tetradonema plicans can cide. Several species and isolates of
cally invisible to the naked eye. They control fungus gnats, but it is not avail- nematodes are marketed under various
require moist soil to survive. When they able commercially. These nematodes are trade names, but not all are equally
find an insect, they enter it through harmless to people, animals, beneficial effective against fungus gnat larvae.
natural openings and release a bac- above-ground insects, and earthworms. Check the label or contact the supplier
terium that kills the host within 48 to confirm that the product is effective
hours. The bacterium itself serves as a Possibilities for effective against these pests. They should be
partial food source for the nematodes, biological control applied to moist, but not saturated, soil.
which complete their development You can control fungus gnats easily by The area to be treated should be
inside the dead insect. The next genera- using biological methods and good san- watered before and after application.
tion of nematodes leaves the insect in itation practices. Shore flies are more dif- The soil temperature must be above
search of new hosts. Unlike plant para- ficult to control with biological 50°F for nematodes to be effective.
sitic nematodes, these nematodes have methods. Application in the early evening or the
no mouthparts and do no damage to morning is recommended to avoid
The biological insecticide Bti controls
plants.
fungus gnats but is not effective against exposing the nematodes to extreme
shore flies. A commercial formulation of heat and sunlight. Nematodes are
this bacterium, Gnatrol, is registered for applied as a spray or a drench on the
the control of fungus gnats on orna- soil surface. Since they can withstand
mentals. Because Bt does not persist in pressures up to 300 psi, nematodes can
the soil, it must be applied whenever be applied with the same equipment
pest populations develop. Applications used for the application of chemical

Fungus gnats and shore flies


Enemies egg ——— larva ——— pupa adult

Hypoaspis

Bt var. israelensis

Nematodes

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S F U N G U S G N A T S & S H O R E F L I E S 33
pesticides and through irrigation or
injector systems. Always check that the
nematodes are alive before applying
Alternative control methods
them. Examine a drop of the dilute spray Sanitation bark, while the most came from media
with a magnifying glass. The nematodes The elimination of breeding areas will containing some hardwood bark.
should be squirming. Inactive nema- reduce fungus gnat and shore fly Potting media may also influence
todes bent at sharp angles are usually problems quickly. Drain wet areas nematode effectiveness.
dead. Nematodes are most effective at and repair drip irrigation systems to Avoid over-fertilizing plants. Excess
soil temperatures of 65°–85°F. They remove standing water and algae fertilizer promotes algae growth
should be applied weekly for 4–8 weeks. where flies can breed. Dispose of or where fungus gnats and shore flies
Although both Bti and nematodes can sterilize infested growing media so can breed.
be effective against fungus gnats, their that emerging flies will not reinfest
control may not always be satisfactory new plantings. Remove all used Chemical control
to the grower. potting medium and plant debris Several insect growth regulators, such
from the greenhouse as this is a place as diflubenzuron, cyromazine and
For long-term control, the soil-inhabiting
where fungus gnats can pupate. fenoxycarb, are registered for use on
predatory mite Hypoaspis miles, intro-
ornamentals to control fungus gnat
duced early in the crop cycle, can Cultural control and shore fly.These insecticides
maintain fungus gnat populations at Avoid overwatering plants as this should be applied as a drench when
acceptable levels.These mites are most allows growing media to remain sat- fungus gnats or shore flies are first
effective when released before fungus urated and promotes the growth of observed.The drenches will not harm
gnat populations are established. H. miles algae which provides an ideal envi- natural enemies but may damage
seems to thrive in the top 1–2 inches of ronment for fungus gnat and shore some types of plants. Read the label
the growing medium and in moist areas fly growth and development. carefully for phytotoxicity information.
where fungus gnats breed. Excellent
Choose the correct planting medium. Horticultural oils are registered for
control of fungus gnats has been
Although soilless root media are con- use against fungus gnats on some
achieved with experimental releases on
sidered excellent for plant growth, they greenhouse crops. Several other syn-
poinsettia and bedding plants.This mite
are also ideal for the growth and thetic insecticides are available for
also works well in combination with
development of fungus gnats and the control of fungus gnats or shore
nematodes or Bti. Commercially
shore flies. In a test of several different flies, but most are very toxic to bene-
produced mites are shipped in vermicu-
media, the fewest adult pests emerged ficial insects.
lite, bran, peat, or a similar carrier. Sprinkle
from media containing pine bark or no
the mite mixture over flats, trays, capillary
mats, and floors. Shake the container
occasionally to keep the mites evenly dis-
tributed throughout the container. For
potted plants, 1⁄4 teaspoon of the mite
mixture sprinkled around the roots of
every second pot, is recommended.
Release rates vary by crop and pest pop-
ulations, from 10,000–20,000 per acre for
greenhouse vegetables to 10–50 per ft2
for bedding plants. Additional releases
will be necessary to maintain control if
pots are removed and replaced regularly.
Researchers are investigating the possi-
bility of inundating propagation beds
with excessive numbers of mites so that
they will be distributed to individual pots.
34 Leafminers
S
everal different species of leafminers Description and life cycle
Available natural enemies and attack greenhouse crops. The most Adult leafminers are shiny black flies
their potential for control common are Liriomyza trifolii and with yellow markings and are slightly
A few parasites the vegetable leafminer. L. trifolii is a cos- smaller than fruit flies, at 1⁄16 inch
are effective mopolitan species, originally from the (1.3–2.5 mm) long. Most species appear
against the Caribbean, that attacks numerous plant very similar, except chrysanthemum
species of species. It is a problem on vegetable leafminer, which is slightly larger and
leafminers crops, including tomato, cucumber, dark gray. The female punctures leaves
commonly lettuce, and pepper, and ornamentals with the ovipositor, leaving a small white
found in greenhouses. The potential such as alstroemeria, cineraria, chrysan- spot at the point of entry. Both female
for successful biological control is high themum, gerbera, gypsophila, and snap- and male flies feed on the sap that
on vegetable crops and moderate on dragon. The vegetable leafminer is a oozes from the punctures. Pale, oval
ornamentals. problem on a wide range of vegetables. eggs are inserted into about 15% of
Other important species include the these punctures. Each female produces
Order Diptera: Flies
chrysanthemum leafminer which is a an average of 60 eggs in her 2- to 3-
Family Agromyzidae: Leafminer flies pest on chrysanthemum and cineraria, week life span. The area of the plant pre-
Liriomyza trifolii and the pea leafminer, a pest of aster, ferred for oviposition depends on the
Vegetable leafminer, carnation, gypsophila, and some veg- leafminer species and the host plant.
Liriomyza sativae etable crops, especially in the coastal The white or yellow maggots hatch in a
valleys of California, Mexico, and South few days and tunnel through the leaf
Chrysanthemum leafminer, America. Liriomyza bryoniae, a European tissue for up to 2 weeks. This produces
Chromatomyia syngenesiae species that attacks vegetable crops, the narrow, winding, white mines visible
Pea leafminer, mainly tomatoes, is not known in North on leaves. The width of the mines
Liriomyza huidobrensis America. increases as the larvae grow. Once larval
development is complete, the maggots
Damage drop out of the leaf and pupate inside
Leafminer larvae tunnel within leaves their hardened skins, or puparia, in
between the upper and lower surfaces, cracks in the soil, on benches, or within
making unsightly white blotches or cupped leaf surfaces. The barrel-shaped
twisting lines. Different species produce pupae change from bright yellow to
different types of mines that may vary brown. Adults emerge in about 10 days.
with the host plant. The adult female
flies also make numerous punctures in Monitoring
the leaves that show up as white spots Begin weekly monitoring of susceptible
on the upper surface. The mines and greenhouse crops after transplanting
punctures interfere with photosynthesis them. Monitor leafminers in two ways.
and, if numerous, can reduce crop yields. Use yellow sticky traps to detect
Young seedlings may be destroyed. leafminers moving into the greenhouse
Infestations on edible portions of veg- and to keep track of adult populations.
etable crops and on ornamentals make Examine the plants visually for both
the plants unmarketable. Although adult and larval damage. In fast-growing
tomato plants can tolerate high levels of crops the damage of the first generation
damage with little effect on yield, may be hidden by new foliage. You may
leafminer feeding accelerates leaf drop not observe the infestation until the
above the developing tomato fruits, second generation attacks the upper
making them vulnerable to sunburn. In leaves of mature plants.
addition, punctured and mined tissue is Yellow sticky traps are easier to handle
more susceptible to diseases, such as and assess than other sampling tech-
bacterial leaf spot on chrysanthemum. niques, such as sweep nets or pupal
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S L E A F M I N E R S 35
trays. The traps are commercially avail- into trays placed beneath the plants. Natural enemies
able or can be made by the grower. Make sure trays have small drain holes Numerous wasp species parasitize
Place the sticky traps just above the so they won’t fill with water. Place the leafminers, but few predators or
crop canopy and adjust the height of pupae in clear plastic containers and pathogens attack leafminers.
the traps as the crop grows. Also place keep the samples at room temperature,
traps near doors, air-intake vents, among out of direct sunlight, until parasites or Parasites
newly arriving plants, and near more flies emerge. All of the parasites of leafminers para-
susceptible varieties to detect new sitize the larval and/or pupal stages.
A sequential sampling plan for L. trifolii in
infestations. The number of traps Only those that have been studied as
chrysanthemum or gerbera crops has
needed depends on the crop, but on possible biological control agents in
been developed in California. An estimate
average they should be spaced 18–23 greenhouses are mentioned below.
of adult leafminer populations is deter-
meters (20–25 yards) apart. Count the These wasps are in the families
mined from the number of flies caught
leafminer adults on the traps once or Braconidae, Eulophidae, and Eucoilidae,
on yellow sticky traps spaced uniformly
twice per week. for which there are no common names.
throughout a uniform crop. Larval densi-
Even when sticky traps are used it is still ties are estimated from leaf samples Dacnusa sibirica. This European
important to check the crop visually. taken during crop growth or at harvest. braconid wasp is a solitary endoparasite
Inspect plants regularly for flies and for Treatments should be applied when of all instars of L. bryoniae, L. trifolii, and
white spots on the upper surfaces of larval and adult populations reach certain the chrysanthemum leafminer. The
leaves, which are the oviposition and thresholds established in the plan. adults are black and 1⁄8 inch (2–3 mm)
feeding punctures of the adults. Sample The methods that have been developed long, with extended, flexible antennae.
leaves weekly to estimate larval Using their antennae and ovipositors,
for sampling leafminers provide fairly
leafminer densities. Select leaves the females locate leafminer larvae
accurate assessments of leafminer pop-
randomly from plants throughout the within the leaf and deposit single eggs
ulations, but the infestation level does
greenhouse or mark plants with colored not always correlate well with plant in the larvae. They are best at finding
tape for continual monitoring of the late-instar larvae and prefer these for
damage or aesthetic injury, especially on
same plants through the season. Record ornamentals. Although it is usually oviposition. Each female lays up to 90
the number of mines per leaf on three eggs during a 2-week life span. The eggs
acceptable to have a maximum of 10%
leaves from either the middle or bottom mined leaves, this maximum varies with hatch in 4 days, and the larvae complete
of each plant, wherever most of the their development in about 2 weeks in
the crop, the time of year, and the
leafminers are found. Sample a the living leafminer larvae. Adult wasps
market for the product. For example, cut
minimum of 10 plants per greenhouse. emerge from the leafminer puparium.
chrysanthemums can have mining
The adults do not host feed. This wasp
Leaf samples can also provide informa- damage on the lower leaves (which are
prefers cool conditions (65°–75°F). It is
tion on parasite activity, although many removed at harvest), while the same
commercially available.
growers do not have the time or expert- plants grown for sale as potted
ise to do this. Learn to recognize adult plants cannot have any mines
parasites and mines containing para- on the mature foliage.
sites. Carefully cut open the mines on Vegetable crops, such as
sampled leaves and examine them for tomatoes, can tolerate fairly
the presence of parasites or leafminer heavy leafminer damage
larvae that have been killed by adult without any effect on yield.
parasite feeding. A 10X–15X magnifier or Information on the population
R. Buhl, Stuttgart, Germany

hand lens, or a dissecting microscope levels present will provide a


will help you observe the insects in basis for making pest manage-
small mines. Parasite pupae and ment decisions, especially for
prepupae can easily be seen through determining how many para-
the leaf by holding it up to a light. sites to introduce.
Record parasitism and mortality rates
weekly. Parasitism can also be checked
by collecting leafminer pupae that fall The European braconid wasp Dacnusa sibirica.
36 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Diglyphus isaea. This Eurasian eulophid Other Diglyphus species. Several other Ganaspidium utilis. This eucoilid wasp,
wasp parasitizes L. trifolii, L. bryoniae, and species of Diglyphus have been studied native to subtropical areas of North
chrysanthemum leafminer. The 1⁄16-inch as possible biological control agents for America, is a larval-pupal parasite of
(1–2 mm) adults are black with short leafminers in greenhouses. The appear- vegetable leafminer and L. trifolii. The
antennae. Females detect leafminer ance, life cycle, and biology of these robust 1⁄20-inch (1.2-mm) adults are
larvae by drumming their antennae species are similar to that of D. isaea. black with slender antennae. Females
along the mines. When a larva is found, None of these wasps are commercially oviposit in leafminer larvae. The eggs
the female drills through the leaf and available in the United States: hatch after the leafminer pupates, and
into the leafminer body with the ovipos- ■ D. begini, a facultative—or environ- the wasp larvae feed on the pupa within
itor, paralyzing the larvae. The female mentally adaptable—gregarious the host puparium. It is not commer-
then either feeds on the body fluids of wasp, attacks all leafminer species. It cially available.
the leafminer larva or lays one to five develops much faster than L. trifolii.
eggs on the body of the host, depend-
■ D. intermedius host feeds on all
ing on its size. The wasp larvae feed
instars of L. trifolii but prefers the
inside the leafminer body for 3–5 days,
third instar for oviposition. It also
then pupate inside the mine. Feeding by
attacks the vegetable leafminer and
the wasps stops leafminer and mine
chrysanthemum leafminer.
development. The adults emerge about
a week later. The females live for 3–4 ■ D. pulchripes parasitizes the veg-
weeks and lay approximately 60 eggs. etable leafminer.
Over half of the leafminer mortality is
caused by adult host-feeding, which is
necessary for egg production. This wasp
prefers warm conditions (75°–90°F). It is
commercially available.

Richard Lindquist
The adult wasp Diglyphus begini attacks all
leafminer species. It is not yet commercially available
in the United States.
R. Buhl, Stuttgart, Germany

The Eurasian eulophid wasp Diglyphus isaea. Adult females


feed on leafminer larvae or lay eggs in them.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S L E A F M I N E R S 37
Opius dimidiatus. This American Oscinidius (=Chrysocharis) parksi. This Pathogens
braconid wasp is one of the most American eulophid wasp is a solitary, Steinernema (=Neoaplectana)
abundant larval-pupal parasites of the larval-pupal endoparasite of L. trifolii and carpocapsae. This nematode is a
vegetable leafminer and L. trifolii, both the vegetable leafminer. The 1⁄25-inch common species with considerable
outdoors and in greenhouses in the (1-mm) adults are metallic black with potential as a biological control agent
United States and in southern Ontario, white legs. Females lay eggs in third- because of its rapid infectious action and
Canada. It will also parasitize L. bryoniae. instar leafminers. The wasp larvae hatch environmental safety. (See “Fungus Gnats
It is not commercially available. and begin development in the leafminer and Shore Flies” for a description and life
Opius dissitus. This braconid wasp is a larvae. Leafminers complete their larval cycle information.) The nematodes enter
native parasite of the vegetable development, then drop to the ground. the mines through small tears in the leaf
leafminer in Florida that will also para- They die when the wasp larvae start to or punctures made by the adult
sitize L. trifolii. The 1⁄16-inch (1.5-mm) pupate. The adult wasps emerge from leafminer, then infect second- and third-
females oviposit in leafminer larvae of the host puparium. By host-feeding, the instar leafminer larvae through body
any instar. After eggs hatch the wasp females kill more leafminers than they openings. Nematodes may also be able
larvae consume the leafminers within need for reproduction. This wasp is not to infect leafminer pupae in or on the
their puparium. Adult wasps emerge commercially available in the United growing medium. Foliar applications of
about 2 weeks after the eggs are laid. States. nematodes may be effective against
This wasp is not commercially available. leafminer larvae in leaves, but very high
Predators
relative humidity is required. The need
Opius pallipes. This European braconid A few general predators, such as spiders
for constant moisture is a major limita-
wasp is a solitary endoparasite of all and ants, will feed on adult or larval
tion in their effectiveness.
instars of L. bryoniae and chrysanthe- leafminers, but none are specific to
mum leafminer larvae. It will lay eggs in leafminers. The tomato bug, Crytopeltis
Possibilities for effective
L. trifolii, but the leafminer kills the eggs. modestus, a facultative mirid predator of
Females locate larvae by drumming Liriomyza spp., has been investigated in
biological control
Biological control of leafminers is
their antennae and inserting the ovipos- California for leafminer control on
possible on many crops. Natural popula-
itor into the mine. They lay single eggs chrysanthemum and tomato. Older
tions of parasites that often enter
into larvae. After the wasp completes nymphs and adults suck out the body
unscreened greenhouses from outdoors
development in the leafminer, the host contents of leafminer larvae within the
in early summer may provide some
dies and the wasp emerges from the mine and larvae that have emerged to
control. Releases of both larval and
mine to pupate. Females do not host pupate. However, because C. modestus
larval-pupal parasites can maintain
feed. This wasp is commercially available feeds on tomato stems and only the
leafminer populations at acceptable
in Europe. older stages eat leafminers, this bug is
levels. However, the high cost of these
considered a pest. Much more research
parasites and uncertain availability may
is necessary before
detract from their usefulness. Also,
it can be recom-
adequate biological control of leafmin-
mended as a bio-
ers on floral or foliage crops is difficult,
logical control
because even a low population of
agent.
leafminers may produce cosmetic
damage.
Richard Lindquist

Diglyphus begini larva developing externally on a leafminer


larva.
38 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Dacnusa sibirica is often used to control Although your best guides are personal some wasps have already emerged from
the first generation of leafminers, while experience and the supplier’s instruc- the puparia. Distribute the parasites
Diglyphus spp. is used against later gen- tions, results of recent research using evenly throughout the greenhouse, but
erations, but appropriate wasp selection leafminer parasites on specific crops can place extra wasps in leafminer hot spots
depends on your specific circumstances. help in selecting the proper natural or near especially susceptible cultivars.
D. sibirica prefers cool temperatures, enemies. Dacnusa sibirica, Diglyphus Proper release rates depend on the
while Diglyphus isaea favors warmer begini, D. pulchripes, and Oscinidius parksi leafminer species, the crop, the growth
conditions. Diglyphus may control subse- have all been shown to be successful as stage of the crop, and the time of year. A
quent generations more effectively if biological control agents on greenhouse single inoculative release of parasites
temperatures are around 80°F. Because tomatoes in various countries. D. pul- may succeed if made early in the
D. sibirica attacks late-instar leafminers chripes and Opius dimidiatus nearly erad- growing season. However, multiple
when mines are already well developed, icated leafminers on tomato in an exper- releases are more likely to provide effec-
this wasp is best for use on crops with imental greenhouse in Ohio within 8 tive control. Suppliers should provide
little aesthetic value, such as cut flowers weeks. Dacnusa sibirica moved from detailed instructions on the use of
where the foliage is not harvested, or on open rearing units to a lettuce crop to natural enemies. Introduce the parasites
young plants that will outgrow any early control nearly 100% of pea leafminer in as soon as the first leafminers are
damage before harvest. Diglyphus the Netherlands. D. intermedius effec- detected on yellow sticky traps or when
attacks young larvae before mining tively controlled L. trifolii on cut chrysan- feeding punctures are observed. It is diffi-
damage is extensive, and therefore is themums in experimental greenhouses cult to control high populations of
particularly suitable for use on flower in California. When D. begini was intro- leafminers even with very high release
crops. Diglyphus can parasitize leafmin- duced early in the crop cycle it virtually rates if the releases are not made early.
ers already parasitized by D. sibirica, so it eliminated leafminers on marigolds Provide the optimal environmental con-
usually becomes the dominant parasite within 2 months and maintained control ditions of 60°–85°F and 50–90% relative
under warm conditions. for another 2 months, even at high humidity. Do not use residual insecticides
leafminer densities. The lower, older within a month of parasite release. If the
leaves were damaged, but the upper leafminer population is high, you can use
portions were free of injury. a nonresidual spray, such as horticultural
Leafminer parasites are shipped either oil, azadirachtin, or natural pyrethrins (not
as pupae in small cardboard boxes or as pyrethroids), to reduce the number of
adults in plastic tubes. Open the parasite leafminers before releasing the wasps.
shipment in the greenhouse in case

Leafminers
Enemies egg ——— larva ——— pupa adult

Dacnusa spp.

Diglyphus

Ganaspidium

Opius spp.

Oscinidius

Nematodes

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S L E A F M I N E R S 39
Tomato crops can tolerate relatively Scientists at the University of California-
high leafminer populations without Davis have developed grower-friendly
yield loss. Weekly releases of one software to help simplify biological
parasite for every 10 new mines on the control efforts against L. trifolii infesting
crop are suggested for the first 6 weeks cut chrysanthemums. The program
of an infestation on tomato. On chrysan- determines the number of Diglyphus to
themum, the recommendation is to release each week for the
release three adult Dacnusa sibirica for duration of the crop to
every 1,000 plants (500 per acre) in the eradicate L. trifolii before
week after planting. Make a second marketable foliage
parasite release at the same rate in the develops. A salable cut
sixth week after planting to control the crop was produced
progeny of any surviving leafminers. without using any pesti-
Diglyphus isaea is often preferred for the cides following the
second introduction. Releases of 500 D. program’s recommended
sibirica adults per acre every 2 weeks release rate.
can be made instead. Either parasite Currently parasites are the
should control an infestation of one best option for biological
mine per six plants. control of leafminers.
Monitor carefully after the releases to be More research must be
sure that parasitism is high. Even if 90% conducted on predators
of the first leafminer generation is para- and pathogens before
sitized, the population can still increase they can be used to
from one mine per 100 plants to about control leafminers.

Jim Ducy
one mine per plant in the next genera- Nematodes, however, may
tion. Make weekly introductions of be useful for suppressing
wasps until a well-established popula- leafminer populations. If
tion is present. Weekly introductions constant moisture can be
take into account natural mortality of maintained for at least 24 hours, which
the wasps. Cool, cloudy weather and the may be difficult to achieve in a commer-
use of pesticides will reduce the effec- cial greenhouse, foliar applications of
tiveness of the parasites. Introduce up to the nematode S. carpocapsae are about
400 parasites per acre each week if less as effective as abamectin, but neither
than 90% of the leafminer larvae are treatment eliminates severe leafminer
parasitized. Even when introductions are infestations. Make applications in late
done properly the results may vary, and afternoon or evening because the
extra parasite releases or chemical nematodes may be killed by sunlight or
control may be necessary in some cases. desiccation. Allow several hours of
drying time to give the nematode time
to move to the leafminer larvae. The
optimal temperature for infection is
about 70°F.
40 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Alternative control methods


Sanitation Greenhouse floor Azadirachtin is the active ingredient in
Avoid introducing any new plant The material beneath benches affects commercial formulations of extracts
material that may contain leafminer adult leafminer emergence. Fewer from seeds of the neem tree. This
eggs or larvae into the greenhouses. leafminer pupae survive in gravel natural compound is an insect growth
Rogue out severely infested plants. under the benches than in soil. regulator that has a very low mam-
Promptly remove or destroy all crop Leafminers that tried to pupate on malian toxicity. It is also effective as a
residue containing larval leafminers polyethylene sheeting under tomato repellent or anti-ovipositional treat-
that could continue to develop after plants drowned when the greenhouse ment. Foliar applications kill leafminer
harvest. Use physical barriers, such as was watered. Ants killed many of the larvae in the mines, and reduce
hanging sheets of plastic, to prevent prepupae when the plastic was dry. leafminer pupation and adult emer-
adults from flying to other areas of the However, pupal parasites of leafminers, gence. Solutions of azadirachtin also
greenhouse during harvest. such as Dacnusa sibirica or Opius spp., have controlled leafminers on chrysan-
were also killed. themum for 4 weeks after soaking
Host plant cultivar
rooted cuttings for 2–4 hours. It is reg-
Leafminers prefer and develop better Fertilization
istered for use on greenhouse orna-
on certain crop cultivars. A plant’s Nutrient inputs can affect pests as well
mentals and vegetable crops.
chemical content, nutritional value, as the plant. High levels of nitrogen are
and the distribution and density of its associated with leafminer problems, as Abamectin is an insecticide derived
hairs influence the plant’s attractive- well as spider mite infestations, on from the naturally occurring soil
ness to leafminers. Eliminate the most chrysanthemum. Avoid over-fertilizing microorganism Streptomyces
susceptible cultivars, if possible, or crops. avermitilis. It is not a selective
isolate them to reduce contamination chemical, so it should be used only as
Traps a rescue treatment if biological control
to other cultivars. Chrysanthemum cul-
Yellow sticky traps can be used for fails. It is registered for control of
tivars vary widely in their susceptibility
mass-trapping adult leafminers, as well leafminers and spider mites on orna-
to foliar damage by leafminers. In
as monitoring them. The traps do not mental plants in greenhouses.
general, standard chrysanthemum cul-
catch many parasites of leafminers.
tivars suffer less damage than spray Spinosad is an insecticide derived from
Trapping adults reduced L. trifolii
types. In one experiment, White natural metabolites produced under
damage on chrysanthemum by 50% in
Iceberg and Yellow Iceberg had signifi- fermentation conditions by the actino-
experiments in Canada.
cantly more mines than 15 other culti- mycete Saccharopolyspora spinosa. It is
vars, while Statesman had the fewest Chemical control especially efficacious against leafmin-
mines. In other experiments, foliage of Localized infestations, or high popula- ers, caterpillars, and thrips, but has low
the cultivars Albatross, Bright Yellow tions that need to be reduced before to moderate impact on beneficials. It is
Tuneful, Colonel Comfort, Divinity, Nob predator or parasite introduction, registered for control of many pests on
Hill, Pink Marble, and Sea Foam had should be spot-sprayed with selective landscape ornamentals; check with
less damage than did Iceberg. chemicals. your chemical supplier on availability
Horticultural oils, both refined petro- for use on greenhouse ornamentals.
Insect screens
Leafminers may invade greenhouses leum distillate products and those
during the summer. Screens on vents, made from vegetable oils, can kill
side-walls and doors will help prevent leafminers and other insects. These
leafminers from migrating into the horticultural oils may be toxic to some
greenhouse from outside. However, plants. Various brands are registered
proper screen installation is necessary for use on vegetables and ornamentals
in order to avoid reducing air flow. The in greenhouses to control many pests,
maximum hole size to exclude L. trifolii including leafminers.
is 640 µ.
Mites 41

M
ites are not true insects because Monitoring
Available natural enemies and the adults have only two main Bulb mites are difficult to observe until
their potential for control body regions and eight legs. The their damage becomes apparent. If bulb
There are many highly newly hatched young, called larvae, have mites damage your crop one year, plan
effective predators of only three pairs of legs. After the first control efforts the next year based on
mites. The potential for molt, all subsequent instars, called that experience.
successful biological nymphs, have four pairs of legs. Mites
control is high. are abundant in soil and organic matter, Natural enemies
and many parasitize vertebrates or Bulb mites have few natural enemies,
Order Acari: Mites and
invertebrates. Others are predaceous, and no known parasites or pathogens
ticks
scavengers, or plant eaters. There are control them.
Family Acaridae: Acarid mites numerous families of mites and ticks,
Predators
Bulb mite, Rhizoglyphus echinopus but only those species of economic
Hypoaspis aculeifer. This soil-dwelling,
Family Tarsonemidae: importance for greenhouses are dis-
laelapid mite is a nonspecialist predator
Tarsonemid mites cussed here.
of arthropods, but appears to feed and
Cyclamen mite, Phytonemus reproduce best on bulb mites. The light
pallidus Acarid mites brown adults are less than 1⁄25 inch (1
mm) long and spend all their time on or
Broad mite, Polyphagotarsonemus Bulb mites infest the bulbs of many
latus plant species, including Easter lily, onion, in the soil. Adults consume up to 30 bulb
daffodil, tulip, and hyacinth. mites daily. Females deposit one to three
Family Tetranychidae: Spider mites eggs per day. White, nonfeeding, six-
Twospotted spider mite, Damage legged larvae hatch from the eggs and
Tetranychus urticae Bulb mites feed on the underground become eight-legged nymphs in about a
Carmine spider mite, Tetranychus portion of bulbs, weakening the plant day. The nymphal stage lasts 4–5 days.
cinnabarinus tissue. Feeding damage also promotes During this phase nymphs consume an
the secondary invasion of plant average of 15 bulb mites. Both immature
Lewis mite, Eotetranychus lewisi
pathogens such as Fusarium, Pythium, and adult predators prefer the larval
and Rhizoctonia. On Easter lily (Lilium stage of the bulb mite as prey.
longiflorum), typical symptoms include Development is faster on larval and egg
rosetting, cessation of growth at a stages than on adult prey. Adults may
height of 3–6 inches, chlorosis, and occa- live up to 60 days. They are inactive
sionally death. when temperatures are below 57°F. This
species is available commercially.
Description and life cycle A related species, H. vacua, has similar
The slow-moving adult bulb mites are biology and feeding habits. In small-dish
1⁄ 25 inch (1 mm) long and pearly white trials in laboratory studies, H. vacua
with short, red legs. Females produce consumed 33 bulb mite nymphs during
100–150 eggs, laid singly or in groups development, while adult females ate
on the bulb’s surface, near injured or about 13 mite nymphs per day. H. vacua
decaying tissue or between bulb scales. is not available commercially.
A generation can be completed in
10 days at 81°F.
42 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Possibilities for effective


biological control
Tarsonemid mites Description and life cycle
The adult cyclamen mite is semitrans-
These very small mites infest a wide
Hypoaspis aculeifer or other species may parent and pinkish orange, while the
range of host plants, and because they
have potential as a biological control immature stages are translucent. Broad
are so small their damage can become
agent against bulb mites. In small-scale mite adults are straw-colored with a
extensive before the population is rec-
laboratory experiments H. aculeifer was prominent white stripe down the center
ognized. Preferred hosts of the cyclamen
able to suppress bulb mites to very low of the back. The adults of both species
mite include African violet, azalea,
levels on lily bulbs. The predators did are very small—less than 1⁄100 inch (0.3
cyclamen, fuchsia, geranium, ivy, and
not leave the bulbs until almost all prey mm) long. The life cycles of different tar-
snapdragon. Broad mite is an important
were eaten. In closed plastic bags filled sonemid mites are similar. Their white,
pest of gerbera, and may also be found
with lily bulb scales and vermiculite opaque, ovoid eggs are laid singly.
on other hosts including cyclamen,
(similar to lily bulb propagation condi- Cyclamen mites lay their eggs on upper
geranium, hibiscus, impatiens, ivy, and
tions), Hypoaspis was also able to leaf surfaces, while broad mites usually
peperomia, as well as vegetable
suppress the bulb mite to very low lay eggs on leaf undersides or dark,
bedding plants such as bean, pepper,
numbers. However, no research has moist places on plants. In 2–7 days,
and tomato. These mites cause problems
been conducted on introducing this whitish larvae hatch from the eggs. They
in cool, moist conditions, unlike spider
predator to greenhouses for bulb mite develop for 1–4 days, then pass through
mites, which thrive in a warm and dry
control. The related species H. miles, sold an inactive nymphal stage before
environment.
commercially for fungus gnat control, molting into adults. Broad mites can
died out without noticeable impact on produce a generation in 10 days;
Damage cyclamen mites require 18 days. Relative
the bulb mite population.
Specific symptoms depend on the plant
humidity around 80%, low light, and
species. They range from leaf distortion,
moderate temperatures around 61°F
stunting, and bronzing to plant death.
Alternative Damage often resembles pesticide
favor the increase of both mites.

control methods injury or nutritional problems. However,


unlike those abiotic causes, the distribu-
Monitoring
Frequent visual inspection of sympto-
tion of mite damage within a green-
Sanitation matic plant parts is the best method for
house is patchy. Cyclamen mite infesta-
Avoid introducing infested plants. If detecting infestations. Tarsonemid mites
tions first distort young leaves, which
only a few plants are infested, remove avoid light, prefer high relative humidity,
become rough and wrinkled. Later this
or destroy them to prevent adjacent and tend to be found in the crown of
results in stunted growth and a dwarfed
plants from becoming infested. Avoid their host, between the densely packed
appearance. Damaged gerbera leaves
transporting mites on tools. young leaves of the leaf bud. Cyclamen
have bronzed areas along the midribs.
mite is usually found within buds,
Flowers on infested plants may be dis-
whereas broad mite tends to be on more
torted or may not open. Shoots of plants
exposed surfaces. It may be necessary to
infested with broad mite appear open,
use a low-power microscope to confirm
distorted, shrivelled, and burned. Broad
the presence of these tiny mites.
mites feeding on the underside of
young foliage of gerbera cause the
Natural enemies
leaves to become rigid and curl
No parasites or pathogens provide
downward. Infested tissue on other
effective control of tarsonemid mites. A
plants becomes brown and distorted,
few species of predatory mites feed on
and small leaves and flowers may fall off.
cyclamen or broad mites.
Fruit on tomato, cucumber, peppers, or
eggplant may be russetted or distorted. Predators
Euseius (=Amblyseius) stipulatus. This
phytoseiid mite is a predator of citrus
red mite, Panonychus citri, and feeds on
broad mite in greenhouses as well as
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S M I T E S 43
outdoors on avocado, chili pepper,
citrus, coffee, cotton, mango, tea, and Spider mites Description and life cycle
Spider mites are small, usually only 1⁄50
tomato. It consumes all stages of prey Spider mites are one of the most serious
inch (0.5 mm) long when mature.
except nymphs, and will also feed on pests of ornamental crops, especially in
Twospotted spider mites range in color
pollen and honeydew. This mite does hot, dry conditions. The twospotted
from light yellow or green to dark green
not reproduce at relative humidities spider mite is the most important mite
or brown. All have two dark spots visible
below 50%, nor at very high tempera- species infesting floral crops. It has an
on the abdomen. The carmine spider
tures. It is not available commercially. extremely wide host range, short gener-
mite is plum red. Males tend to be
ation time, and continuous reproduction
Neoseiulus (=Amblyseius) cucumeris. smaller and thinner than females.
when not in diapause.
This commercially available phytoseiid Twospotted spider mite females lay
mite is used mainly for controlling Damage eggs in webbing on the underside of
thrips, but it will also feed on cyclamen Twospotted spider mites feed on the the leaves. Each female produces up to
mite on strawberry. However, it is not underside of leaves, removing leaf cell 120 eggs, depending on the host plant.
recommended for cyclamen mite contents. This results in a chlorotic, On chrysanthemums an average of 14
control in greenhouses. stippled appearance on the leaves of eggs are laid over a 5-day life span,
most plants. However, this stippling may whereas on rose they can lay 112 eggs
Possibilities for effective not be as apparent on thick-leaved over 15 days. The pearly white, perfectly
biological control plants such as chrysanthemum. Leaves spherical eggs hatch into larvae in 4–6
Although some predatory mites will often dry out and fall off. Large popula- days. The larvae are small and white with
feed on broad or cyclamen mite, little tions can severely defoliate or kill plants. only six legs. They molt into eight-
research has been conducted on tech- Webbing produced by spider mites can legged nymphs and look like the adult.
niques for using these predators in cover foliage and flowers, detracting Adults appear after a second nymphal
greenhouses. from the appearance of the plant. stage.
Flower quality and yield is reduced on Lewis mites are slightly smaller than
roses. Northern greenhouses may twospotted mites and have up to three
Alternative receive a respite from spider mites
beginning September and continuing
spots on each side of their body. Their
biology is similar to twospotted mites,
control methods through February, if they go into
diapause.
but they do not diapause. This mite has
been a problem on poinsettia since the
Sanitation mid-1990s, although it will feed on other
Carmine spider mites do not produce
Avoid introducing infested plants. If plants such as cucumber.
the typical speckling of leaves. Instead,
only a few plants are infested, remove
infested leaves become prematurely Spider mites can produce a generation
or destroy them to prevent adjacent
chlorotic with small, transparent lesions. in 7–14 days in warm temperatures. In
plants from becoming infested. Avoid
This type of damage resembles magne- cooler temperatures, generations take
transporting mites on tools.
sium deficiency. Bright yellow patches more than 40 days; below 54°F, develop-
Chemical control develop that eventually turn dark and ment stops. In northern greenhouses
Localized infestations and high popu- spread over the entire leaf. This damage adults may go into diapause for several
lations that need to be reduced may be caused by a toxin injected months, hibernating in cracks and
before predator introduction should during feeding. Even small populations crevices from September through
be spot-sprayed with selective chem- of carmine spider mites can cause February/March. The diapausing mites,
icals. Insecticidal soap controls extensive damage. which are reddish and do not feed, may
exposed tarsonemid mites but also Feeding damage from the Lewis mite be confused with predators or carmine
kills predatory mites and other creates a distinctive stippled appearance spider mites.
natural enemies. However, it is on the leaves. Eventually the entire leaf
nontoxic once it is dry. Thorough becomes bleached, dies, and falls off.
coverage of infested surfaces and
several applications are necessary to
achieve control.
44 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Monitoring Natural enemies Galendromus (=Metaseiulus,


Frequent visual inspection of plant parts Predators are the only effective natural =Typhlodromus) occidentalis—
is the best method for detecting infesta- enemies of spider mites. Numerous western predatory mite. This phyto-
tions of spider mites. Because of their species of predatory mites, some seiid mite is smaller than Phytoseiulus
small size and cryptic behavior, they are beetles, and a few thrips feed on spider persimilis (next page) and develops best
often overlooked until their feeding mites. Mites of the family Phytoseiidae at cooler temperatures, but it tolerates a
damage or webbing becomes apparent. are the most important predators of wide range of relative humidities
They can be found on all areas of the spider mites and the most commonly (40–80%). It is able to regulate spider
plant, but they most often infest the used natural enemies for controlling mite populations at lower densities and
older, middle-age leaves, and the midrib. spider mites in greenhouses. for longer periods of time than P. persim-
ilis, although it will not control spider
Small-scale growers could consider Predatory mites mite populations as quickly. It can also
using plants that spider mites prefer Euseius spp. Several phytoseiids in this survive long periods without prey.
more than the crop plant to detect small genus are commercially available, but Nondiapausing strains have been devel-
populations of mites before damage these predatory mites have not been oped that allow control of spider mites
occurs on the crop—but this is not prac- evaluated for use on greenhouse crops. through the winter, when days are short.
tical in large operations. Highly preferred E. delhiensis (=rubini) will feed on spider Some strains are resistant to a number
plants that can be used this way, as mites, broad mites, thrips, and whitefly of organophosphate insecticides (such
“indicator plants,” include lima beans eggs. E. hibisci feeds on various mites, as as azinphos-methyl, diazinon, and
(tall plants that easily attract blowing well as thrips and sweet potato whitefly phosmet) and a carbamate (carbaryl).
mites) and radishes (much shorter eggs. E. scutalis occurs mainly on trees Several different strains are commer-
plants that show dramatic symptoms of and shrubs in North Africa and the cially available.
leaf distortion). Middle East, where it is associated with
Mesoseiulus (=Phytoseiulus) longipes.
tetranychid mites, whiteflies, and
This predator is similar to P. persimilis in
tenuipalpid mites. Although it is a
activity, but can tolerate warmer tem-
general feeder and will attack thrips,
peratures—up to 70°–90°F—and
spider mites, and whitefly eggs, E.
relative humidity as low as 40%. It is
scutalis prefers pollen and tetranychid
available commercially.
eggs. E. stipulatus will develop on
twospotted spider mite, but it does not Neoseiulus (=Amblyseius) californicus.
lay eggs when feeding on this mite This predatory mite is similar to P. per-
without supplemental food such as similis, but is smaller, pale, and does not
pollen or honeydew. Eggs of this species suppress spider mite populations as
fail to hatch at relative humidities less quickly. However, it is useful for keeping
than 50%. low populations under control because
it can survive longer periods without
prey. It is commercially available.
Some other species of Neoseiulus, such
as the commercially available N. fallacis,
feed on a variety of tetranychid mites
and are commercially available, but little
is known of their utility in greenhouses.
Ray Cloyd

Container of Neoseiulus californicus, a predatory mite.


S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S M I T E S 45
Phytoseiulus macropilis. This mite has consume 30 eggs or 24 immature spider Macrolophus nubilis. This mirid bug is a
been investigated for greenhouse use mites per day. At favorable temperatures voracious predator of spider mites in
on dieffenbachia, dracena, parlor palm and humidity this predatory mite can Europe. It was a promising candidate in
(Chamaedorea elegans), and schefflera develop twice as fast as its prey. Both trials in greenhouses in Uzbekistan, but
(Brassaia actinophylla). The globose, pesticide-susceptible and organophos- it is not commercially available. A related
light- to deep-red females lay oval phate-resistant strains of this predatory species, M. caliginosus, is offered com-
orange eggs that hatch into six-legged mite are available commercially. mercially as a predator of whiteflies.
larvae. Both larvae and nymphs have a Scolothrips sexmaculatus—sixspotted
Other predators
similar white to light orange color. Males thrips. Both the adult and larval stages
Feltiella acarisuga (=Therodiplosis
are identical to females in shape and of this predatory thrips feed on spider
persicae). This predatory gall midge
color but are smaller. These mites have a mites. The tiny, slender, pale amber
occurs throughout Eurasia and North
strong preference for immature spider adults have three dark spots on each
America. It feeds on all stages of spider
mites over adults. Each predator forewing. Females deposit eggs in soft
mites, but generally prefers the eggs or
consumes four to six spider mite eggs or plant tissue. The larvae that hatch are
larvae. Adult gall midges are small,
larvae during its development and an
delicate, pink-brown flies with long legs. white or yellow, without dark markings.
average of eight eggs per day as an
They do not feed and only live 3–4 days The robust, yellow pupae occur on the
adult. P. macropilis has a very short life leaf surface in the mite colonies. S. sex-
after emergence from the cocoon. Each
cycle in comparison to many spider mite
female lays an average of 30 yellow eggs maculatus prefers spider mite eggs, but
species, allowing it to build up quickly to adult females will consume other mite
near high densities of mites, usually
suppress pest populations. In the stages. The rate of predation depends
where webbing occurs. The yellow- or
absence of spider mites they will prey on temperature. Adult females consume
orange-brown midge larvae grow to
on their own immatures. P. macropilis 50 spider mite eggs per day at 86°F but
1
about ⁄16 inch (2 mm) long. They can
occurs naturally in Florida and is avail- only 21 eggs per day at 68°F. It is com-
consume over 300 mite eggs as they
able commercially. mercially available.
complete their development in about a
Phytoseiulus persimilis. This predatory week in the greenhouse. They then spin
phytoseiid mite was accidentally fluffy white cocoons on the underside of
brought from Chile to Germany on leaves, usually along a leaf vein. This
orchid roots and was subsequently sent midge is commercially available. There
to other parts of the world. The 1⁄25-inch are several closely related species that
(1 mm) adults are a shiny orange-red are also good mite predators, including
and pear-shaped, with long front legs F. minuta, a common predator in British
and no spots. Each female produces Columbia.
about 50 football-shaped eggs that are
twice the size of twospotted spider mite
eggs. The eggs hatch in about 3 days
into nonfeeding, six-legged larvae. A day
later they molt into eight-legged
nymphs that consume 10–12 spider
mite eggs or small mites during their
development. The immatures are pale
salmon in color and oval in shape. The
University of Florida

optimum temperature range for devel-


opment is 70°–81°F. Development
ceases and oviposition and longevity
decline sharply at relative humidities
below 60%. Under warm and humid
conditions, the adults of these predators
Phytoseiulus persimilis is the most commonly released
predatory mite used in greenhouses.
46 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Stethorus spp. These spider mite preda- Possibilities for effective Long-term control may result from a
tors are small black lady beetles. Both biological control single inoculative release of these preda-
the larval and adult stages are preda- In Europe, predatory mites have been tors, especially if nondiapausing, insecti-
ceous on all mite stages, but their use in used successfully for several decades to cide-resistant strains are used. However,
greenhouses has not been evaluated. S. manage spider mites in greenhouse veg- chemical control with selective miticides
punctillum, a predator of European red etable production. Predatory mites have may be required during establishment
mite—a pest of fruit trees—is commer- effectively controlled spider mites on and occasionally thereafter. If there is an
cially available. chrysanthemum, rose, and other orna- average of more than one spider mite
mental crops under experimental condi- per leaf, a chemical spray with low
tions. However, the need to prevent toxicity to predatory mites, such as insec-
cosmetic damage on floral or foliage ticidal soap or horticultural oil, should be
crops may make biological control of applied to reduce the spider mites to
mites difficult, especially when pesticides less than 10% leaf infestation. Do not use
that kill predatory mites are used to residual pesticides within a month of
suppress other pests and/or diseases. releasing predatory mites, including
Your spider mite control strategy may sulfur-containing fungicides, because
depend on the crop you raise and condi- they are highly toxic to mites.
tions in your greenhouse, especially tem- Predatory mites are most effective when
perature and humidity. Phytoseiulus per- introduced while spider mite popula-
similis is the most commonly available tions are low, ideally at or before the first
and most commonly sign of spider mite damage. In green-
released predatory mite in houses with a history of spider mite
greenhouses. However, problems, the first releases should be
Oregon State University Extension Service

there are a few crops on made 1 week after plant emergence. If


which P. persimilis cannot be you are using indicator plants, release
used. For example, P. persim- the predatory mites as soon as those
ilis slips off the stems and plants show the first visible signs of
leaves of carnations. It also damage. Proper timing is essential to
does not do well on achieve economic control, especially for
tomatoes because the mites carmine spider mites, which cause
become trapped on glandu- extensive plant damage at low popula-
lar hairs on the leaf petioles tions. Mite damage and reproduction
and stems, and they are can continue for 1–3 weeks before the
affected by toxic com- predators can destroy the mites. Most
The small black lady beetle Stethorus spp. pounds in the leaf. Other failures of biological control occur when
predatory mite species listed the predator is released too late.
in the “Natural Enemies” section also Species selection and release rates vary
provide good control. Galendromus occi- considerably depending on the plant
dentalis and Neoseiulus californicus, for species and the environmental condi-
example, may be better suited for use on tions such as temperature and humidity
semi-permanent greenhouse crops such which influence the growth rate of both
as rose or gardenia than on short-term predator and prey. P. persimilis is an
vegetable crops. These predatory mites excellent predator of spider mites on
can often be used in conjunction with low-growing plants in humid green-
other houses with moderate temperatures.
predators as well. P. macropilis performs better than P. per-
similis on ornamental plants under
warm, humid conditions. Mesoseiulus
longipes is frequently used to control
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S M I T E S 47
spider mites in hot, dry greenhouses on predators per plant is suggested Inundation of a crop with predators at
taller plants. It tolerates lower humidity because of the potential for rapid plant extremely high rates can also reduce
than does P. persimilis. N. californicus damage at low pest populations. pest populations—effectively and
does well on most potted plants in Predatory mites are usually released quickly—but this method may be pro-
greenhouses with moderate tempera- according to one of three distributional hibitively expensive. Another successful
tures and average humidity. A combina- patterns: uniformly, in infested patches, technique uses “banker” cucumber
tion of predators released at regular or from banker plants. Uniform distribu- plants at either end of the greenhouse.
intervals works best in greenhouses or tion of predators throughout the green- The plants are intentionally infested
interior plantscapes with a variety of house is the most common method of with spider mites and predators are
plants and growing conditions. introduction. It provides predictable released on them when damage begins
Plant density and plant architecture levels of control. Introduction in patches to show. Within a month thousands of
influence the distribution of spider of mite damage will often result in predators are available for release on
mites on a plant species and the ease better control than uniform distribution. the main crop. On tomatoes, predators
with which the predators can find The live mites are usually shipped mixed can be moved to hot spots by workers
patches of prey. For example, P. persimilis in vermiculite, bran, or a similar material during harvesting operations. This may
is very efficient on cucumbers that have to cushion them in transit. The carrier- not be practical in large operations.
large leaves and vines that intermingle, mite mixture can be sprinkled directly The predatory gall midge Feltiella can
but less so on peppers with smaller onto the foliage of infested plants and also be used for spider mite control in
leaves that don’t touch. P. persimilis is the mites will disperse on their own. (Be conjunction with P. persimilis. The preda-
also less effective on cut rose varieties sure to shake the container occasionally tory mite has low dispersal ability, so it
with fewer leaves because the mites to distribute the mites throughout the may fail to find patches of high pest
cannot move around as easily on these container evenly.) However, treating density. Feltiella is good at finding hot
plants. N. californicus is a better choice infested plants only requires regular spots, so the two predators are comple-
for control of spider mites on roses, if inspection of all plants until predators mentary. Feltiella is particularly useful on
introduced early. Arranging plants so are well established, which may be com- hairy-leaved plants (such as tomatoes)
that leaves touch may improve biologi- mercially impractical. If indicator plants that P. persimilis does not work well on,
cal control on some plant species. are used to detect the first infestation, and it can also effectively control
On average, one predator for every five you can release the predators on the carmine spider mite. Even though
twospotted spider mites, or a rate of 10 indicator plants and allow them to move Feltiella larvae eat at least five times as
predators per ft2, should be more than out into the crop. many spider mites per day as does
adequate for control. Other predator-to- Phytoseiulus, the midge alone is usually
pest ratios that have been successful not able to control spider mites.
under specific circumstances are 1:6,
1:10, and 1:25, but specific ratios can only
be determined by experience. In green-
houses a rate of two predators per ft2 (or
two per plant if the plants are small) is
suggested. On vegetables, a predator-to- Spider mites
prey ratio of 1:50 may be sufficient, but
Enemies egg larva ——— nymph ——— adult
on ornamental floral or foliage plants
that cannot sustain any aesthetic injury Predatory mites
the ratio may have to be reduced to 1:10
Feltiella
or 1:5. In the United Kingdom, one
predator for every 10 chrysanthemum Sixspotted thrips
cuttings released 2–3 weeks after Stethorus
planting, before damage is apparent, has
given satisfactory control. For the Natural enemy attacks the host Effectiveness depends on species of enemy
carmine spider mite, a rate of 50 or more Width of bar indicates degree of effectiveness
48 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

The midges are shipped as cocoons on


leaves or the carton. The carton should
be opened and placed on the soil
Alternative control methods
surface of plants in the greenhouse, pro- Sanitation enemies. It has a slight effect on
tected from direct sunlight, for at least 1 Remove any weeds or old plants that spider mite eggs but does not affect
week. Adults emerging from the may harbor spider mites. Avoid predator eggs. Once insecticidal soap
cocoons in the container will fly to spreading mites on workers’ clothing dries, it is nontoxic. Thorough
spider mite colonies. High humidity or implements by restricting move- coverage of infested surfaces and
improves midge emergence. Optimal ment through mite-infested areas. several sequential applications are
conditions for Feltiella are 68–81°F and Always visit mite-infested areas last. necessary to maintain spider mite
relative humidity greater than 60%, populations at low levels.
although larvae can tolerate a wider Irrigation and fertilization
Maintaining proper soil moisture, and Horticultural oils, both refined petro-
range of conditions than can the adults.
misting or hosing off plants helps leum distillate products and those
Weekly releases of one per 10 ft2 of
prevent mite populations from devel- made from vegetable oils, can kill
plant material are recommended, but
oping and dispersing to other plants. mites. These horticultural oils may be
because of the high cost of these insects
High nitrogen levels are often associ- toxic to some plants. Various brands
many growers use only one per 40 ft2
ated with severe mite infestations. are registered for use on vegetables
and allow the population to build up
and ornamentals in greenhouses to
over time. Manipulation of the greenhouse control many pests, including spider
Sixspotted thrips appears to be a good environment mites.
predator for use in greenhouses. Unlike High humidity produced by misting
Abamectin is an insecticide/miticide
predatory mites, thrips can fly away to suppresses spider mite populations
derived from the microorganism
find new prey. Release rates will vary and favors P. persimilis and Feltiella.
Streptomyces avermitilis. It is registered
depending on the crop and mite
Chemical controls for control of spider mites and
density. Contact the supplier for recom-
Localized infestations, or high popula- leafminers on ornamental plants in
mended rates.
tions that need to be reduced before greenhouses.
Spider mite populations should be mon- predator introduction, should be spot- Spinosad is an insecticide based on
itored by observing foliage of suscepti- treated with selective chemicals. natural metabolites produced under
ble plants at least weekly. Additional Integrating chemical control on the fermentation conditions by the actin-
predator releases may be necessary upper portion of the canopy with bio- omycete Saccharopolyspora spinosa.
every 2–4 weeks to achieve good logical control on the lower part of the The product is fast acting and works
control within 6 weeks. plant has worked well for commercial both on contact and when ingested. It
rose production. is very effective against thrips; control
There is good potential for the inte- of spider mites is inconsistent, but is
gration of predatory mites and insecti- improved with the addition of certain
cidal soap applications to control adjuvants. It is registered for control of
spider mites on ornamental foliage many pests on landscape ornamen-
plants. Insecticidal soap controls tals; check with your chemical supplier
spider mites, but when wet it also kills on availability for use on greenhouse
predatory mites and other natural ornamentals.
Scales & mealybugs 49

T
he Superfamily Coccoidea is a large Damage
Available natural enemies and and diverse group, closely related to A heavy infestation of soft scales will
their potential for control aphids and whiteflies.They are cause yellowed leaves, distorted foliage
There are divided into three main groups: soft especially at the growing tips, twig
many differ- scales, armored scales, and mealybugs. dieback, or defoliation. But the main
ent species They all feed in a similar manner, by damage comes from the growth of
of scales sucking plant juices through their sooty mold on the clear, sticky
and mealybugs. For some you can needle-like mouthparts. Many also honeydew excreted by scales.
expect to achieve good biological excrete honeydew, which supports the
control with the available natural growth of sooty molds that are cosmeti- Description and life cycle
enemies; for others the potential for cally damaging. Heavy scale infestations Soft scales are round to oval, dome-
successful biological control is lower. can threaten plant health. shaped on top, and 1⁄16–1⁄4 inch
Order Homoptera: Aphids, The adult males and females appear very (2–6 mm) long when mature. Immature
leafhoppers, planthoppers, different. Females and immatures are scales start out light in color and darken
scales, and mealybugs wingless and often legless and don’t at maturity. Brown soft scale females are
Superfamily Coccoidea: Scale insects look like insects. Adult males look pliable, oval, and somewhat flattened.
somewhat like tiny gnats but lack They have a pale brown, yellow, or gray
Family Coccidae: Soft scales
mouthparts and cannot feed. color that is mottled, shiny and crossed
Brown soft scale, by a dark brown grid-like pattern. They
Coccus hesperidum produce live young, not eggs. The
Hemispherical scale, Soft scales female hemispherical scale is hard,
Saissetia coffeae The soft scales are the more important circular, steeply convex, smooth brown,
of the two groups of scales found in and shiny. Numerous eggs pile up
Black scale, Saissetia oleae
greenhouses. A wide variety of the flow- underneath their cup-like bodies. The
Nigra scale, Parasaissetia nigra ering and foliage ornamentals, from black scale adult female is dark, oval,
Family Diaspididae: Armored scales orchids to ferns, are good hosts for soft globular, and has ridges on the scale
Boisduval scale, Diaspis scales. The brown soft scale attacks a that form an “H” pattern. The nigra scale
boisduvalii broad range of hosts, while the black varies in size, shape, and color depend-
scale prefers woody plants. The hemi- ing on its host. It tends to be more oval
Florida red scale, Chrysomphalus
spherical scale favors ferns, asparagus on leaves and more elongated on
aonidum
fern, schefflera, and many nonwoody petioles or thin stems.
California red scale, Aonidiella evergreen plants. Plants in the family Female soft scales produce 50–2000
aurantii Acanthaceae, such as Crossandra and eggs or live young, depending on the
Fern scale, Pinnaspis aspidistrae the zebra plant (Aphelandra squarrosa), species. Mobile, crawler-stage nymphs
are especially susceptible to the hemi- hatch from eggs after 1–3 weeks. They
Ivy or oleander scale,
spherical scale. Another common green- move to a suitable part of the plant,
Aspidiotus nerii
house species is nigra scale. Woody where they settle for the remainder of
Greedy scale, Hemiberlesia rapax plants such as weeping fig, citrus, ivy, their lives. The nymphs go through three
Latania scale, Hemiberlesia and holly are common hosts. instars. A waxy covering envelops the
lataniae female after she becomes an adult. The
Pineapple scale, Diaspis bromeliae covering adheres tightly to the body of
Dictyospermum scale, the female and cannot be separated
Chrysomphalus dictyospermi from it. All three of these soft scales feed
on the phloem and are often associated
Cactus scale, Diaspis echinocacti
with stems or leaf veins.
Family Pseudococcidae: Mealybugs
Citrus mealybug, Planococcus citri
Longtailed mealybug,
Pseudococcus longispinus
50 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Soft scales are most troublesome in Natural enemies Metaphycus helvolus. This small
greenhouses at temperatures around Numerous parasitic wasps, predators, encyrtid wasp from South Africa attacks
68°F, and development does not usually and several pathogenic fungi attack soft young nymphal stages of several species
occur above 86°F. In greenhouses there scales, but only a few have been investi- of soft scales. The male wasp is dark
may be as many as six generations per gated as candidates for use in green- brown. The females are orange-yellow
year, with a new generation produced houses. and about 1⁄25 inch (1 mm) long. Each
every 40–80 days, depending on tem- female lays up to five eggs per day
perature. All stages may be present Parasites under the bodies of late second and
simultaneously throughout the year. The parasitic wasps that are important early third instar scales and kills up to 20
natural enemies of soft scales are all in more nonparasitized scales of various
Monitoring the family Encyrtidae. ages by feeding on them. The females
Early detection will prevent many pest Coccophagus lycimnia. This cosmopoli- lay an average of 400 eggs over their rel-
management problems. The best way to tan wasp will parasitize over 47 species atively long lives. Wasp larvae develop
detect soft scales is to inspect plants of soft and armored scales, and has been singly inside the scale bodies. After
visually, especially new shipments of shown to be effective against brown about 2 weeks the adult wasp emerges
plants. If scales are present, you will soft scales in citrus orchards and orna- by cutting a small hole in the scale. M.
usually find them on the undersides of mental crops. Females oviposit in late helvolus readily attacks black and hemi-
leaves and on stems, although some first- to third-instar hosts but prefer spherical scales, as well as brown soft
species may occur on upper leaf second instars. The scale insect contin- scale, nigra scale, citricola scale (Coccus
surfaces on some plants. The presence ues to develop after being parasitized pseudomagnoliarum), and European fruit
of ants, wasps, or bees may be a sign but dies before maturity. This wasp is lecanium (Parthenolecanium corni). This
that soft scales are present. They are available commercially at least in parasitic wasp is commercially available.
often attracted to the honeydew Europe. Metaphycus luteolus. This wasp, closely
produced by scale insects. Yellowed related to M. helvolus, has provided
foliage or sooty mold on leaves often effective biological control of brown soft
indicates the presence of scales. scale outdoors in California. Both the
males and females of this species are
lemon-yellow. Females lay 5–10 eggs per
day in host scales over their 1-month life
span. The larvae are gregarious internal
parasites of all stages of the brown soft
scale, but solitary parasites of black
scales. This wasp will also parasitize early
stages of other scale species. Larval
development is very rapid; at 75°F adults
emerge after 11 days. M. luteolus is not
available commercially. Another species,
M. zebratus, is commercially available in
Europe.
Ray Cloyd

The adult wasp Metaphycus helvolus is about 1⁄25 inch


(1 mm) long.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S S C A L E S & M E A LY B U G S 51
Microterys flavus. This encyrtid is Chilocorus infernalis. This beetle from Other lady beetles. Many lady beetles
another internal parasite of brown soft the foothills of the Himalayas is more will feed on soft scales if their normal
scale, and of various other soft scales. It cold tolerant than C. nigritis. It is not prey is not available. The mealybug
is available commercially for release in available commercially, but some closely destroyer, Cryptolaemus montrouzieri,
citrus groves. There are several strains related species (C. baileyii and C. circum- will feed on soft scales in greenhouses if
with preferences for different host datus) are available as armored scale mealybugs are not present. The alter-
scales. For example, the California strain, predators in other countries. nate-host scales allow the mealybug
probably of East Asian origin, prefers the Chilocorus nigritis. This beetle, indige- destroyer to survive between mealybug
brown soft scale and will not attack nous to India and East Asia, is a vora- outbreaks. Several species of Scymnus
black scale. This wasp is solitary in small cious predator with a wide host range are known to attack soft scales. The con-
hosts but often gregarious in larger including both soft and armored scales. vergent lady beetle, Hippodamia conver-
ones. Females readily feed on young The adult is a 5⁄32-inch (4-mm), black gens, may provide limited control of soft
scales. beetle with yellow edges on the thorax. scales if released in large numbers.
Other parasitic wasps. Encyrtus infelix A single female may lay over 350 eggs. Green lacewing larvae. These will feed
and Encyrtus lecaniorum have been used One reason for its success in biological on immature scales, but because they
successfully alone and in combination control programs is that adults and also feed on a wide range of other
with Metaphycus helvolus for scale large larvae completely remove the insects, including many natural enemies,
control in French greenhouses. E. lecan- adult female scale from the plant they are not recommended for use
iorum (commercially available in Europe) surface. Both beetle larvae and adults lift when other natural enemies are already
has parasitized hemispherical and soft scales from the host plant with present.
brown scale more effectively than M. extremely sharp mandibles and then eat
Pathogens
helvolus or C. lycimnia. E. lecaniorum the soft parts below. C. nigritis is avail-
Verticillium lecanii. This fungus has
mimics the ants that tend the scales, able in Europe, but not in North
been reported to infect and kill certain
thereby preventing the other parasites America.
soft scales. Preliminary investigations
from attacking the scales. These three Rhyzobius (=Lindorus) lophanthae. indicate it is a promising biological
wasps, used individually, have controlled This small black lady beetle is mainly a control agent for the scale Philephedra
hemispherical scale on ferns, peperomia, predator of armored scales (see tuberculosa on Annona squamosa in
and dieffenbachia. A combination of M. “Armored Scales” for more information), Florida. However, no research has been
helvolus and either E. lecaniorum or although they will eat some soft scales, conducted on its ability to infect or
another wasp, Diversinervus elegans, mealybugs, and smaller insects. Both control soft scales in greenhouses.
gave satisfactory control of black scale adults and larvae feed on the scales. It is
on Aphelandra and Aralia. commercially available.
Predators
Several species of lady beetles are well-
known predators of soft scales outdoors
in different parts of the world. Adults and
Oregon State University Extension Service

the four larval instars all feed on scales.


Females lay eggs on or under the scale.
The elongated, flattened, spiny larvae
that hatch consume large numbers of
scales as they develop over a period of
2–4 weeks. Pupation normally occurs on
the plant. Two to four generations are
produced each year.

Predatory lady beetle larva feeding on soft scale.


52 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Possibilities for effective establish the natural enemy population


biological control
The natural enemies described above
in the greenhouse.
Release live M. helvolus adults two or
Alternative
can control some soft scales in certain
situations. Parasitic encyrtid wasps,
three times at 2-week intervals and at a
rate of three per ft2 or up to 1,000 para-
control methods
Sanitation
especially Metaphycus helvolus, have sites per acre. Scatter the parasites on
Remove heavily infested plants or spot-
been particularly effective. M. helvolus plants evenly throughout the infested
treat them chemically to keep crawlers
works well against hemispherical scale area. These parasites can search and find
from spreading to adjacent plants.
and has provided control of black scale the scales quite easily. The best time to
Wash off honeydew and dislodge
in several parts of the world, but it does release the wasps for control of hemi-
crawlers with water sprays or mists.
not reduce brown soft scale populations spherical scale is just before the scale
Inspect new plant material for pests
on some plants. However, it has reaches the third instar—which is
before placing it into general growing
provided satisfactory control of brown evident by the appearance of a ridged
areas. Starting with clean plant material
soft scale on poinsettia and in combina- “H” on the scale surface. Wasp emer-
will help prevent scale problems.
tion with other wasps on Ficus, certain gence holes should be visible after 2
cacti, and cymbidiums, as well as other weeks. Control should be achieved in Chemical control
orchids. This wasp is most effective in 2–3 months. Periodic additional releases Spot treat if there are two or more
semi-tropical conditions. may be necessary to maintain control. adult scales or large numbers of
For soft brown scale control, Microterys Predation by lady beetles, such as crawlers per leaf. Widespread applica-
flavus or Coccophagus lycimnia may be a Rhyzobius lophanthae or Chilocorus tion of traditional insecticides will
better choice than M. helvolus. However, nigritis, which feed on all developmental interfere severely with natural enemy
no research has been conducted on the stages of scales, complements parasite activity. Insecticidal soap or horticul-
use of either species in greenhouses. In activity, which tends to be more stage- tural oil will kill soft scale crawlers as
the field, competition from other para- or size-specific. Releasing a predator well as exposed natural enemies.
sites limits the effectiveness of M. flavus. along with a parasite may offer the Insect growth regulators are regis-
To improve control with natural strongest potential for control. R. lophan- tered for use on ornamentals for
enemies, avoid the use of residual insec- thae is most effective at cooler tempera- control of soft scales and other
ticides for a month. Reduce scale popu- tures (59°–77°F) and relative humidity of insects in a preventative program.
lations with insecticidal soap or horticul- 20–90%. Release adult beetles at a rate These have minimal effects on
tural oil, and wash scale honeydew from of three to six beetles per 10 ft2 for light natural enemies. Azadirachtin, a
plants before releasing wasps. Some infestations or five to seven beetles per commercial formulation of neem
scales must be present, however, to 10 ft2 for heavy infestations. seed extract, is an insect growth
regulator that can be used. It is sold
under several brand names.
Soft scales
Enemies egg ————— nymph ————— adult

Coccophagus

Metaphycus helvolus

Metaphycus luteolus

Microterys flavus

Chilocorus

Green lacewing

Rhyzobius

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S S C A L E S & M E A LY B U G S 53

Armored scales Description and life cycle


Armored scales are soft-bodied insects
Monitoring
Visual inspection of plants is the best

A
rmored scales get their name from
that live beneath a hard cover made of way to detect armored scales. They are
the hard, waxy coating that covers
wax and protein. They are smaller than generally smaller than soft scales, not
their bodies. They are not as eco-
soft scales, varying in size from 1⁄25–5⁄32 raised and bumpy, and are more easily
nomically damaging in greenhouses as
inch (1–4 mm). Armored scales vary in overlooked. You can find them on lower
are other scale insects because armored
shape from circular to elongated (ellipti- leaf surfaces and stems, often forming
scales infest limited types of plants,
cal or like an oyster shell), in texture thick crusts. New plant shipments
mainly trees and shrubs, and they don’t
from smooth to rough, and in color. The should be checked carefully.
produce honeydew. They can be impor-
insect body beneath the covering is
tant pests in conservatories.
usually yellow or orange. The males’ cov- Natural enemies
erings tend to be smaller and more Of the many parasites and predators
Damage elongated than those of the females. that are known to attack armored scales,
The plant damage caused by armored
Most females lay eggs beneath their only a few have been utilized in green-
scale feeding is similar to that caused by
scale, but certain species produce live house biological control efforts.
soft scales. They inject toxins while
young under the scale. The crawlers
feeding on leaf tissue. The toxins kill cells Parasites
move to other plant parts and settle for
around the feeding site, causing a Many parasites attack armored scales,
the remainder of their lives. A waxy
yellow or brown halo. Heavy feeding including various Aphytis species that
covering that incorporates the shed skin
results in premature leaf drop. However, have been used against several armored
of the crawler and subsequent stages
since armored scales don’t produce scales in the greenhouse. The two
builds up around the immature insect.
honeydew, sooty mold is not a problem. species mentioned are in the families
Unlike soft scales, the scale covering on
Aphelinidae and Encyrtidae, respectively.
armored scales is easily removed from
the scale’s body. The adult female, which Aphytis melinus. This ectoparasitic wasp
is legless, remains under its covering. Up from India and Pakistan attacks certain
to six overlapping generations may be species of armored scales, including
produced in a year, depending on the especially California red scale, but also
species and environmental conditions. ivy scale, San Jose scale and oleander
scale. The tiny yellow wasp lays its eggs
on the soft body under the waxy scale
of third-instar nymphs. After developing
as a larva for 2–3 weeks, the adult
parasite emerges through a round exit
hole cut in the scale. The adults can live
Robert F. Luck, University of California-Riverside

3–4 weeks, with each female killing


more than 30 scales. This wasp does
best at temperatures of 76°–85°F and
relative humidity of 40–50%. This wasp
is produced commercially for control of
California red scale in citrus groves, but
has not been evaluated for use in green-
houses. Pesticide-resistant strains are
available. Another species, Aphytis ling-
nanensis, sold commercially in Israel and
Australia for California red scale control,
has not been evaluated in greenhouses.
Adult Aphytis melinus parasitizing an armored scale.
54 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Comperiella bifasciata. This is another Chilocorus kuwanae—Korean twice- Chilocorus nigritis. This species, dis-
commercially available wasp that stabbed lady beetle. This is an efficient cussed in the soft scales section, is also a
attacks a variety of armored scales. predator of at least 23 species of good predator of certain armored
Females prefer to oviposit in second- armored scale in China, Korea, and Japan. scales. Two related species, C. baileyii and
instar female scales. The adult wasps It has been used in other countries for C. circumdatus, are also available com-
emerge from mature scales. It has not release in fruit orchards and is now mercially in other countries for armored
been investigated for greenhouse use. established in the eastern United States scale control, but not in North America.
for control of euonymus scale, Unaspis Rhyzobius (=Lindorus) lophanthae.
Predators
euonymi. The adults are 1⁄8 inch (3 mm) This polyphagous lady beetle occurs
Lady beetles are the most important
long and shiny black with one red spot across the southern United States from
predators of armored scales. Their life
on each wing cover. They feed on all Maryland south to Florida and west to
cycle and basic biology are discussed in
stages of the scale by chewing holes in California. It was introduced into
the section on predators of soft scales.
the scale covering or by pushing under California from Australia between 1889
Chilocorus bipustulatus. This beetle is the scale to feed on the eggs or body and 1892 to control black scale, but it
an important predator of armored scales beneath. As with other Chilocorus
became established as a predator of a
in the Mediterranean and Middle East. It species, the eggs are deposited singly or
wide variety of armored scales. The 1⁄16-
also attacks some soft scales. Its life in small numbers under empty scale cov- inch (2-mm) adults are reddish brown
cycle is similar to that of C. kuwanae, but erings or in cracks and other protected
with a faint, green metallic tint and yel-
tolerates higher temperatures than C. places. Shiny black larvae hatch from the lowish-brown head, legs, and underside.
kuwanae. C. bipustulatus is not commer- eggs in about 8 days. The larvae feed for
Adults and larvae consume all scale
cially available in the United States. 2–4 weeks, depending on the availability stages by chewing large, jagged holes in
of food. Each larva consumes several the scale wax armor. R. lophanthae may
hundred scales during its development. be sensitive to high temperatures
Pupation often occurs in small groups on during some of its life stages. It does
the plant where the larvae developed. best at temperatures of 59°–77°F, but
There are three generations per year can tolerate humidity from 20–90%. This
with extensive overlap of all stages commercially available beetle will also
during the growing season. These feed on soft scales.
beetles overwinter as adults that
Other predators. Limited control of
become active when outside
some armored scales may be provided
temperatures rise above 50°F. It
by green lacewing (Chrysopa and
is intolerant of high tempera-
Chrysoperla spp.) larvae.
John Davidson, University of Maryland

tures and cannot complete its


development above 90°F. It is
not commercially available.

An adult Korean twicestabbed lady beetle,


Chilocorus kuwanae.
Cliff Sadof

The larva of the Korean twicestabbed lady beetle,


Chilocorus kuwanae.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S S C A L E S & M E A LY B U G S 55

Alternative
Possibilities for effective
biological control
Mealybugs
M
ealybugs are common pests in
Selected species of armored scales
control methods should be susceptible to biological
control using Aphytis melinus and/or
greenhouses and interior
plantscapes. Unlike the soft and
Sanitation armored scales, most retain their legs
lady beetles. If your greenhouse crop is
Start with clean plant material. This throughout life and are free-moving. The
infested with an armored scale, try intro-
will help prevent scale problems, and citrus mealybug, which is a native of the
ducing these natural enemies to find
removal or chemical spot treatment tropics and subtropics, is the most
whether they will control it. No research
of heavily infested plants will reduce common and most damaging. It feeds
has been conducted on wasp release
the spread of crawlers to adjacent on numerous flowering plant species in
rates for armored scales in greenhouses
plants. over 25 families, especially soft-
(three applications at a rate of one wasp
stemmed and succulent plants such as
Chemical control per ft2 is suggested for outdoor use), but
coleus, fuchsia, croton, and cactus. The
Spot treatments should be applied it has been suggested that procedures
host range of the longtailed mealybug is
when less than half of the crawlers similar to those for the soft scale
more restricted, but still very diverse.
present are parasitized and the infes- parasite Metaphycus helvolus be used.
Dracena is the preferred host. Other
tation is spreading. Use insecticidal The wasps will disperse readily from the
foliar-feeding species found in green-
soap or horticultural oil if no preda- release site. You can release wasps by
houses include the cactus mealybug
tors or parasites are present; these opening the container and allowing
(Hypogeococcus festerianus), Mexican
treatments will kill exposed natural them to fly out as you walk through the
mealybug (Phenacoccus gossypii), and
enemies. greenhouse.
obscure mealybug (Pseudococcus
Insect growth regulators are regis- Suggested release rates for lady beetles, affinis). The ground mealybug (Rhizoecus
tered for use on ornamentals for such as R. lophanthae, are four to six falcifer) is the most common soil
control of armored scales and other beetles per 10 ft2 for light infestations, mealybug, occurring on the roots of
insects as a preventative treatment. and slightly more for heavy infestations. many house plants, especially African
These can be used when parasites or Make two releases about 3 weeks apart. violets.
predators are present. They do not
affect natural enemies but may
damage some plants. Azadirachtin
(several brands) can also be used. This
insect growth regulator is a commer-
cial formulation of neem seed extract.

Armored scales
Enemies egg ————— nymph ————— adult

Aphytis melinus

Comperiella bifasciata

Lady beetles

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness
56 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Damage Monitoring Anagyrus pseudococci. This


Most mealybugs feed on the above- Visual inspection is the best method for Mediterranean wasp is very similar to
ground parts of plants, especially stem detecting infestations of mealybugs on Leptomastix dactylopii (next page), but it
tips, leaf junctures, and new growth, but leaves and stems. Both the insects them- attacks half-grown to full-grown citrus
some are root feeders. Feeding causes selves and the eggs in their masses of mealybug nymphs. It is a solitary internal
stunting, foliar yellowing, defoliation, waxy threads may look like white cotton parasite.The 1⁄16-inch (1.5-mm) females
wilting and general plant decline. Citrus on the plant. Inspect these cottony are yellow with black markings and dis-
mealybugs inject a toxin while feeding masses carefully so as not to confuse tinctive white antennae.The males are
that causes plant malformation. mealybugs with the beneficial smaller (1⁄25 inch, 0.8–0.9 mm) and black
Secondary damage results from sooty mealybug destroyer larvae that with yellow markings. Each female lays
mold growth on honeydew excreted by resemble mealybugs. On some plants about 45 eggs.This wasp’s life cycle is a
the mealybug, which impairs photosyn- mealybugs concentrate on the growing day or two shorter than that of L. dacty-
thesis and reduces crop marketability. tips, and on other plants they are more lopii. It reproduces best at 86°F.This wasp
dispersed. The longtailed mealybug fre- is available commercially; a related species,
Description and life cycle quently conceals itself in leaf whorls. A. fusciventris is available in Europe.
Mealybugs are pink, soft-bodied insects Ants, wasps, or bees may be present as Leptomastidea abnormis. This wasp
covered with a white, waxy, cottony they are often attracted to the attacks only young, second-instar citrus
material. The wax filaments at the end of honeydew produced by the mealybugs mealybugs and develops as a solitary
the body of the longtailed mealybug are and the sooty mold on leaves. endoparasitoid. The adult wasp is 1⁄16
slender and as long as or longer than Underground infestations are more diffi- inch (2 mm) long, pale yellow-brown
the body, while those of the citrus cult to detect. Yellowed or wilting foliage and has distinct bands on the wings. The
mealybug are very short. The citrus may indicate the presence of mealybugs female produces more eggs than
mealybug females are wingless, on the roots. Small white cottony Anagyrus pseudococci and Leptomastix
1⁄ 16 inch (1–3 mm) long, and ovoid. They masses around the drainage holes of dactylopii. The species has a longer life
lay up to 600 small (1⁄100 inch or 0.3 mm pots also indicate the presence of cycle, requiring from 19 days at 86°F to
long), yellow eggs within a protective mealybugs. 46 days at 65°F. L. abnormis reproduces
mass of white, cottony threads. Egg pro- best at 75°F and has a longer life span at
duction is temperature dependent, with Natural enemies 86°F than L. dactylopii or A. pseudococci.
fewer eggs laid under warmer tempera- A number of natural enemies, including This wasp is better adapted to the wide
tures. The longtailed mealybug does not several parasitic wasps and predators, range of temperatures that occur in
lay eggs but produces live young, similar are known to attack mealybugs. temperate greenhouses and parasitizes
to aphids. After depositing the egg mass a higher percentage of pests than do
Parasites
or live young over a period of 5–10 days, A. pseudococci and L. dactylopii. It is com-
A wide range of host-specific wasps in
the female mealybug dies. The imma- mercially available, primarily from
the family Encyrtidae attack mealybug
tures search for feeding sites on which European suppliers.
species. All develop in a similar manner,
to settle. Male nymphs settle and spin
as solitary internal parasites within the
an elongated, white waxy cocoon.
body of the host mealybug. The female
Females have three instars and are
wasp stings an appropriately sized
mobile throughout their lives. Adult
mealybug and lays a single egg in it. The
males are tiny, winged insects. Under
wasp larva develops within the
normal greenhouse conditions, the pro-
mealybug body before pupating in the
duction of a generation takes 1–3
mummified host. Most research has
months.
centered on the control of the citrus
mealybug.
Cliff Sadof

The wasp Leptomastidea abnormis.


S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S S C A L E S & M E A LY B U G S 57
Leptomastix dactylopii. This Brazilian Pauridia (=Hungariella) peregrina. This Predators
wasp is the most widely used parasite wasp is a solitary endoparasitoid of the Cryptolaemus montrouzieri—
for control of citrus mealybug. It is a longtailed mealybug. It mainly attacks mealybug destroyer. This predatory
1⁄8-inch (3-mm) long, yellow-brown the first and second instars. It will attack Australian lady beetle is probably the
wasp that attacks large nymphs and a few other mealybug species but not most successful natural enemy of mealy-
adult female citrus mealybugs. It has a the citrus mealybug. It is very tolerant of bugs. Both adults and larvae feed on all
characteristic short, hopping flight and high temperatures. At higher humidity stages of mealybugs, and on aphids and
taps its long antennae on the surface as levels, the wasp is better able to tolerate immature scales when mealybugs are
it walks. Females produce 18–20 eggs a higher temperatures and its life span not available. The adult beetle is about
day, preferring to oviposit in third-instar lengthens. H. peregrina has been used 5⁄ 32 inch (4 mm) long and dark brown or
and young adult citrus mealybugs. successfully on various crops outdoors in black except for its orange head, wing
Parasitized mealybugs become legless, subtropical areas, and in greenhouses on tips, and abdomen. Females lay up to
brown swollen mummies covered with ferns and, in conjunction with the 500 eggs at a rate of 10 eggs per day,
fine threads. The emerging adult cuts an mealybug destroyer, on dracena. Because singly in mealybug egg masses. Eggs
opening at one end of the mummy. The it prefers small instars, it complements hatch in 6–7 days at 75°F. The larvae can
life cycle of this solitary internal parasite the action of other wasps that attack grow up to 1⁄2 inch (13 mm) long. They
takes from 18 days at 81°F to 45 days at later mealybug stages. It may be avail- resemble a mealybug with long waxy fil-
64°F. Adult females can live up to 35 able from some commercial suppliers. aments covering the body. The vora-
days. Maximum reproduction occurs at cious larvae consume an average of
86°F, but the females are fairly short- 1400 mealybug eggs, nymphs, and
lived at this temperature. This wasp is adults during their 12–20 day develop-
available commercially. A related ment period and provide better
species, L. epona, is available in Europe. mealybug control than adults. Young
beetle larvae and adults prefer
mealybug eggs and small nymphs, but
older larvae will consume mealybugs of
any size. Other insects, such as soft
scales, may be eaten if mealybugs are
scarce, or the larvae may eat each other.
Generation time ranges from 25 days at
temperatures around 86°F to 72 days at
around 65°F. They are inactive below
50°F, and the lower temperatures and
Ray Cloyd

The Brazilian wasp Leptomastix dactylopii is the most


widely used parasite for control of citrus mealybug.
Cliff Sadof

The mealybug destroyer, Cryptolaemus montrouzieri, is


probably the most successful natural enemy of mealybugs.
58 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

shorter days of winter seem to reduce Possibilities for effective Adult mealybug destroyers should be
their activity, although the mealybugs biological control released as soon as the shipment
will continue to increase under these Biological control of citrus mealybug is a arrives, although they can be stored in a
conditions. Beetles are available from promising alternative to chemical pest refrigerator for a short time. Release
several commercial insectaries. control on commercial greenhouse beetles in early morning or late evening
Other lady beetles. Many coccinellid crops. Mealybug species other than onto plants at various infestation sites.
beetles are important predators of citrus mealybug are more difficult to Vents and windows should be screened
mealybugs in other cropping systems, control biologically because parasitoids to prevent the beetles from escaping,
but few have been investigated for bio- are not as widely available. The natural and during distribution workers should
logical control of mealybugs in green- enemies of citrus mealybug tend to be avoid wearing white clothing, which will
houses. Diomus flavifrons is native to specific, and several effective ones are attract the beetles. Optimal release rates
southern Australia where it commonly available from commercial suppliers. will vary depending on the crop, but
feeds on the longtailed mealybug. The two to five beetles per 10 ft2 is a good
The mealybug destroyer has been suc-
beetle was introduced into Texas for general guideline. Researchers have
cessfully used against citrus mealybug
control of citrus mealybug on citrus. The achieved control with one beetle per
on various ornamentals, such as
small black adults consume about 800 plant on gardenia and one beetle per
gardenia and chrysanthemum, when
mealybug eggs over a 70-day period. two plants on chrysanthemum.
temperatures remain above 70°F. Plants
Scymnus (=Nephus) reunioni and S. Mealybug destroyers should control the
grown in large numbers or with dense
bipunctatus are two other coccinellids mealybug population in about 10
bushy growth are better suited to
available commercially for mealybug weeks. Periodic releases may be neces-
mealybug control with the mealybug
control in other countries. sary if other natural enemies are not
destroyer. Because the beetle needs to
being used.
Green lacewing larvae. Chrysopa and oviposit among mealybug egg masses,
Chrysoperla larvae will feed on young it may be less effective against long- Citrus mealybug can be effectively con-
mealybugs but will also feed on other tailed mealybug, which produces live trolled within 3 months using the para-
natural enemies that are present. young. It is ineffective against mealybug sitic wasp Leptomastix dactylopii. For
infestations on roots. To enhance best results, reduce heavy mealybug
control, avoid using residual pesticides populations first either by releasing
in the month prior to beetle release and mealybug destroyers or by hosing off
control any ants in the area by using a dense accumulations and then spraying
boric acid bait. Otherwise, the ants will with insecticidal soap (if no other
protect the mealybugs from predators natural enemies are present) or kino-
and parasites. To favor the mealybug prene. Also, adjust the greenhouse tem-
destroyer, adjust temperatures to peratures to 75°–81°F. No other residual
72°–77°F and relative humidity to pesticides should be used for a month
70–80% so that maximum control may prior to wasp release. Adult wasps
be achieved. The pest population does should be released at several sites
not need to be reduced to a lower level throughout the infested area at a rate of
before introducing the predator. five wasps per 10 ft2. Place the wasps as
close to the mealybug infestations as
possible. Periodic releases may be nec-
essary to maintain control.
University of Florida

The larva of the mealybug destroyer


resembles a mealybug.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S S C A L E S & M E A LY B U G S 59
The best biological control of citrus Researchers in the Netherlands used
mealybug has been achieved using a
combination of natural enemies, espe-
two wasps (L. dactylopii and L. abnormis)
and two beetles, mealybug destroyer,
Alternative
cially supplementing the mealybug
destroyer with parasitic wasps. However,
and Scymnus (=Nephus) reunioni, against
citrus mealybug on 50,000 Stephanotis
control methods
experimental results have been quite plants grown for cuttings in a commer- Sanitation
variable, depending on the plant cial greenhouse. They released over Starting with clean plant material will
species, the greenhouse temperature, 1300 adult beetle predators in areas help prevent mealybug infestations.
and the greenhouse size. When used with heavy mealybug infestation, and Removing heavily infested plants or
together in French greenhouses, the released 13,330 L. dactylopii and 15,520 spot treating them with chemicals will
mealybug destroyer and L. dactylopii L. abnormis approximately every other reduce the spread of mealybugs to
completely controlled citrus mealybug week from May through October. Citrus adjacent plants. Remove soil accumu-
on clivia (Kaffir lily) and crotons, and mealybug was controlled during the lations beneath plant pots where root
provided reasonable control on gerani- summer months, but not during the mealybugs are established to elimi-
ums, African violets, cattleya orchids, and winter, when temperatures were nate this potential source of future
Pilea. The wasp Leptomastidea abnormis, 55°–63°F and the parasites were infestations.
the mealybug destroyer, and three other inactive. Chemical control
natural enemies were used to reduce The predatory beetle Scymnus Spot treat female mealybugs when
high citrus mealybug infestations to less (=Nephus) reunioni, has been used with egg masses are present and natural
than 1% in gardenia within 3 months in some success in greenhouses in France, enemies are not, especially if growing
a California commercial greenhouse. but no published research on this tips are infested or the plant is sensi-
Releases of 19,300 L. dactylopii and 600 predator has been conducted in the tive to damage. Insecticidal soap or
mealybug destroyers over a 4-month United States. In combination with L. horticultural oil should be used only
period reduced citrus mealybug infesta- dactylopii, it achieved excellent control if no predators or adult parasites are
tion on commercially grown Stephanotis of citrus mealybug on bromeliads. present because the treatments will
from 66% to zero in a small California Similar control on a wide range of orna- kill exposed natural enemies.
greenhouse maintained at 70°F and mentals was noted in the United Use insect growth regulators (IGRs)
above. The mealybug destroyers were Kingdom. when parasitized mealybugs or pred-
able to maintain a small population in ators are present. IGRs are registered
the greenhouse after the disappearance for use on ornamentals to control
of citrus mealybug by using hemispheri- mealybugs, scales, and other insects.
cal scale as an alternate food source. They do not affect natural enemies
Even after an accidental re-introduction but may damage some plants.
of citrus mealybug into the greenhouse Azadirachtin (several brands), an
several months later, the mealybug pop- insect growth regulator, may be used.
ulation remained well below economi- It is a commercial formulation of
cally damaging levels. neem seed extract.

Mealybugs
Enemies egg ————— nymph ————— adult

Anagyrus pseudococci

Letpomastidea abnormis

Leptomastix dactylopii

Pauridia peregrina

Mealybug destroyer

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness
60 Thrips
T
hrips are important pests of cucum- Leaf feeding reduces plant vigor and
bers, peppers, and a broad range of diminishes the yields of vegetable and
Available natural enemies and
their potential for control ornamental greenhouse crops. ornamental crops. Feeding on the
There are some These very small insects commonly hide growing tips and blossoms of plants
predators, in flowers, buds, and leaf axils, and often causes deformities in the new leaves,
pathogens, and go unnoticed until damage appears. flowers and fruit, reducing the plant’s
parasites avail- Western flower thrips (the word “thrips” marketability. Tomato spotted wilt virus
able for control is both singular and plural) is of special and impatiens necrotic spot virus, trans-
of thrips. importance because it transmits the mitted by western flower thrips, cause
However, even when used in combina- tomato spotted wilt virus and impatiens lesions on leaf tissue and can reduce
tion, they will provide only moderate necrotic spot virus to many greenhouse crop yields. Viral symptoms vary depend-
control. Other remedies may be neces- floricultural and horticultural crops. It is ing on the plant species; on cyclamen
sary when pest populations are high. replacing many of the other thrips as the lesions appear as ringspots. Affected
the predominant thrips pest in green- ornamental plants are not marketable.
Order Thysanoptera: Thrips Thrips will also bite people, causing a
houses throughout the world. Thrips
Family Thripidae: Common thrips palmi, native to Southeast Asia, is a mild stinging sensation.
Western flower thrips, severe pest of numerous vegetable and
Frankliniella occidentalis ornamental crops. It is now present in Description and life cycle
Florida and may spread to other parts of Adult thrips are slender insects 1⁄16 inch
Onion thrips, Thrips tabaci
the United States. (1–2 mm) long with narrow fringed
Eastern flower thrips, wings. They may be yellow, tan, brown,
Frankliniella tritici Damage or black depending on the species or
Greenhouse thrips, Both larval and adult thrips have rasping strain. Western flower thrips are yellow
Heliothrips haemorrhoidalis mouthparts that they use to puncture to brown with an orange thorax. Onion
the plant surface. They feed on the sap thrips are a uniform tan or brown.
Thrips palmi
that is exuded from the resulting Greenhouse thrips are dark brown or
wound. Plants are also injured when black. T. palmi are clear yellow with black
female thrips lay their eggs in the plant hairs. Males are typically smaller and
tissue. The damage appears as white or lighter in color than females.
silver speckles on the leaf or flower Thrips can reproduce year-round in
petal, arranged in streaks rather than the greenhouses, producing 12–15 genera-
stipples caused by mite or aphid tions per year. The female thrips inserts
feeding. These spots eventually dry up minute, opaque eggs through holes cut
and turn tan or brown on some plants into leaf, flower, or fruit tissue with her
but remain silver on others. Thrips can sawlike ovipositor. They lay 60–100 eggs
enter flowers while still in the bud stage during their lives. The nymphs hatch in
and after color shows. Blossoms turn 2–7 days, feed during the first two
brown, or buds fail to open completely. instars, then pass through two nonfeed-
Petals become distorted, develop brown ing pseudopupal stages (the “prepupa”
edges, and stick together. As they feed, and the “pupa”) in protected parts of the
thrips excrete brown droplets on petals plant such as leaves or flowers, in leaf
and leaves, and the droplets turn to litter, or in the soil. Although thrips have
black spots when they dry. a simple metamorphosis, and thus don’t
actually pupate, these pseudopupal
stages are similar to the pupal stage of
insects with a complete metamorphosis.
Therefore, thrips nymphs are often
referred to as larvae. They do not feed or
move unless disturbed during the 2–5
day “pupal” stage.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S T H R I P S 61
Thrips often remain together in large flowers, light intensity, placement of Natural enemies
numbers on a single leaf or flower. traps relative to crops, and crop type. A few parasites and pathogens attack
Onion thrips often feed along promi- These factors can make population thrips, and so do many general preda-
nent veins. Western flower thrips are counts inaccurate. tors, including mites, minute pirate bugs,
usually found within developing Although sticky traps are very useful for lacewings, syrphid larvae, and other
terminal foliage or at the base of flower detecting the presence of thrips, plant predacious thrips. Predatory mites are
petals where they feed on pollen. part sampling can track thrips popula- the most commonly used natural
Greenhouse thrips are found primarily tions more accurately. For immature enemies to control thrips in green-
on developed leaves. T. palmi feed along thrips, counting the number found on houses.
leaf midribs. leaf samples is the best indicator of pop-
Parasites
ulation trends. Sample leaves from the
Monitoring A few trichogrammatid and mymarid
middle of the plant to save time without
Effective population monitoring is wasps parasitize thrips eggs. Most of the
reducing accuracy. Monitoring imma-
essential for timing the application of parasitic wasps that have been found to
tures is still more costly and time con-
biological controls and for gauging their attack thrips larvae are tropical or sub-
suming than adult monitoring.
effect. Early visual detection of thrips is tropical species in the family
Blossom sampling is the best method Eulophidae. Few are currently commer-
difficult because they are small and
for monitoring populations of adult cially available.
often hide within young, terminal
western flower thrips, which concentrate
foliage and developing flowers. Let Ceranisus (=Thripoctenus) spp. These
in flowers. To sample a flower, place it
previous experience guide you in eulophid wasps develop inside thrips
into a 25 ml (0.85 oz) plastic vial filled
deciding when to begin monitoring. larvae. After female wasps lay single
with 70% alcohol and seal it immedi-
Using sticky traps and sampling plant eggs in young thrips larvae, the thrips
ately. Pour the vial contents through a
parts, including flowers, can give you continue to appear and behave
funnel lined with filter paper. Then count
some idea of the thrips population in normally while the wasps develop inside
the flower thrips by examining the filter
your greenhouse. them. The thrips die when the parasites
paper under a dissecting micro-
Yellow or blue sticky traps are most scope. The flower may need to
useful for detecting the first infestation be dissected to find all the
of thrips and determining where they thrips. Although not as
are invading the greenhouse. The traps, accurate, a quick visual assess-
which are commercially available, are ment of the number of thrips
also useful for monitoring population per flower can be done by
trends within a greenhouse. Begin moni- merely blowing into open
toring early in the cropping cycle, espe- flowers. The carbon dioxide
cially before flower buds form. It is easier agitates thrips and causes them
to see dark-colored thrips on yellow to move around so they can be
traps, but blue traps are more attractive counted. A combination of
to some species of thrips and therefore sticky traps, blowing into
will detect lower populations. The blue flowers, and flower monitoring
traps will highlight light-colored species. allows quick detection of
Traps should be placed so that the potential problem areas.
bottom of the trap hangs 1–2 inches
above the crop canopy. In mature com-
mercial cucumber crops, place traps 8 ft
above the floor to catch the most
Susan E. Rice Mahr

western flower thrips. Be aware that


there is no correlation between trap
catch and actual plant damage or the
number of thrips in flowers. This is
because thrips catches may be influ-
Hang blue sticky traps 1–2 inches above the
enced by the presence and color of
crop canopy to detect thrips.
62 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

pupate inside their hosts. The adult Thripobius semiluteus. This Predatory mites
wasps also kill young thrips by feeding parthenogenic wasp was introduced in Euseius spp. Several phytoseiids in the
on body fluids from holes made with 1987 from Brazil and Australia as a genus Euseius are commercially avail-
the ovipositor. Several other species possible control agent for greenhouse able in Europe, but have not been evalu-
have potential as biological control thrips in avocado orchards in southern ated for use on greenhouse crops. E. del-
agents of thrips. C. brui occurs in Japan, California. Females lay 15–20 eggs singly hiensis (=rubini) will feed on thrips,
Indonesia, Europe, and the Caribbean. It in thrips larvae and the developing spider mites, broad mites, and whitefly
was introduced and established in wasp kills the thrips before it reaches eggs. E. hibisci is predatory on many
Hawaii in 1933–34 for control of onion the pupal stage. The adults also host small insects, including thrips. It has
thrips, the most important vector of feed on immature thrips, in addition to been used successfully in inoculative
yellow spot virus of pineapple, but has feeding on nectar and honeydew. Under field releases in California to control
not been evaluated as a biological optimal conditions (65°–75°F and citrus thrips. It also feeds on various
control agent for thrips in greenhouses. 50–60% relative humidity) development mites and sweetpotato whitefly eggs.
C. menes is a solitary endoparasitoid that from egg to adult takes about 3 weeks. E. scutalis is widely distributed in the
attacks the first instar of western flower It enters diapause when temperatures warm climates of North Africa and the
thrips. It occurs worldwide, and has been fall below 40°F. This species is commer- Middle East, where it occurs mainly on
found parasitizing western flower thrips cially available. trees and shrubs. It prefers to feed on
on alfalfa and roses in California. It is pollen and spider mite eggs but will
Predators
being investigated for western flower attack thrips, spider mites, and whitefly
Many generalist predators will feed on
thrips control in European greenhouses. eggs as well.
thrips in addition to other insects.
Thrips feeding on exposed plant parts Hypoaspis (=Geolaelaps) miles. This soil-
Numerous species of predatory mites
are likely to be attacked by this para- dwelling laelapid mite is a native of North
and a few bugs are more specific preda-
sitoid, but it may not be able to find America. It feeds on many soil-inhabiting
tors of thrips. Phytoseiid mites are the
thrips that enter flower and/or terminal arthropods, including thrips pupating in
most important predators.
buds. Researchers at the University of the soil. Adults can kill 15–30 prey per day,
Florida are studying another species of targeting more small prey than large prey.
Ceranisus from Thailand for possible Females deposit one to three eggs per
importation as a biological control day.The mites complete a life cycle in
agent of 13–22 days, during which each nymph
T. palmi in sub-
Jim McMurtry, University of California-Riverside

consumes 16–33 prey. It does not


tropical areas diapause in the winter.This mite is avail-
of the United able commercially.
States. None
are commer-
cially available.

Two thrips larvae; the larva on the left has been parasitized by
Thripobius semiluteus, the other is unparasitized.
Max Badgley

Euseius hibisci feeding on citrus thrips.


S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S T H R I P S 63
Neoseiulus barkeri. This phytoseiid Other predators
mite—formerly called Amblyseius Dicyphus tamaninii. This mirid bug is
mckenziei—is still listed by some com- an important predator of whiteflies, but
mercial suppliers, even though this it has also been shown to be effective
species is now considered to be a strain against western flower thrips in
of N. cucumeris. cucumber greenhouses. (See “Whiteflies”
Neoseiulus (=Amblyseius) cucumeris. for more information.)
This predatory phytoseiid mite, previ- Macrolophus caliginosus. This mirid
ously known as Typhlodromus thripsi, bug is primarily a predator of whiteflies,
feeds on young thrips. The pear-shaped but will consume thrips and other pests.
adults are a pale brown and are notice- (See “Whiteflies” for more information.)

Koppert B.V.
ably smaller and flatter than the spider Orius spp. These minute pirate bugs are
mite predator Phytoseiulus persimilis. common in gardens and many agricul-
Females lay an average of two small tural crops, and are often found in
white eggs per day over a period of 20 flowers. The black, 1⁄16–3⁄16 inch The predatory mite Neosiulus cucumeris.
days, often attaching them to plant (2–5 mm) adults are ovoid and
hairs. Adults live up to 30 days, consum- somewhat flat, with distinctively pat-
ing three to six immature thrips per day. terned black and white wings. The eggs
Although they prefer first-instar thrips, are laid in leaf tissues with one end
they will feed on spider mites and their barely sticking out. The nymphs are
eggs, broad mite adults, and pollen of pinkish-yellow to light brown. First-
various plants if thrips are not available. instar nymphs are about the same size
When no food is available they will as thrips larvae and are found in the
resort to cannibalism. This mite will same places. Both minute pirate bug
coexist with Phytoseiulus, although both nymphs and adults are very active and
prey on each other to some extent. feed on all active stages of thrips. They
Some strains of the mite enter diapause consume 5–20 thrips per day, living up
during the winter months in response to to 25 days as an adult. Because they are
shorter photoperiod (less than 12.5

Koppert B.V.
found in flowers, they are effective pred-
hours daylight) and temperatures below ators of western flower thrips. They also
69°F. Both diapausing and nondiapaus- feed on spider mites, aphids, whiteflies,
ing strains are available commercially. and caterpillar eggs, as well as pollen
Neoseiulus cucumeris prefers humidity Adult minute pirate bug, Orius sp.,
and plant juices. Several species of Orius feeding on thrips.
levels above 65%; eggs will not hatch at are available commercially.
lower humidity levels.
Neoseiulus (=Amblyseius) degenerans.
This species is very similar to N. cuc-
umeris in appearance and biology, but it
does not enter diapause. The dark
brown adults are found in flowers and
on the underside of leaves. Larvae have
a brown X shape on the back. This
species can tolerate lower humidities
than N. cucumeris can. It is commercially
available.
Les Shipp

Orius nymph feeding on western flower


thrips larva.
64 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Pathogens sporulate. Infected thrips often move to Nematodes


Several fungal pathogens and some an elevated part of the leaf before the Heterorhabditis bacteriophora (=helio-
nematodes infect thrips naturally. Onion fungus breaks through to facilitate thidis). This nematode has been shown
thrips are susceptible to Beauveria spore release. Sporulation occurs on to reduce adult emergence of western
bassiana, Metarhizium anisopliae, cloudy days when the humidity is flower thrips by 40%. In addition, infec-
Paecilomyces fumosoroseus, Verticillium greater than 80%, but ceases in bright tion may reduce the number of progeny
lecanii, and several species of sunshine. It is not commercially avail- produced by adults that survive, and
Entomophthora. However, natural fungal able. reduce the ability of thrips to transmit
infections do not provide effective Metarhizium anisopliae. This fungus virus diseases. H. bacteriophora has good
control despite high rates of mortality attacks many insects, including thrips searching capabilities and attacks thrips
because epizootics occur when thrips (see “Aphids” for more information). All pupae deeper in the soil profile than do
populations have already grown large mobile stages of thrips are susceptible other nematodes, such as Steinernema
and damaging. These pathogens are to this fungus, although the immature feltiae. Rates necessary to control thrips
useful only if the active stages of the stages are more resistant to infection. are extremely high and may not be eco-
fungi can be introduced early and under This fungus is not registered for use in nomically feasible. It is commercially
environmental conditions suitable for the United States on any greenhouse available.
infection. Beauveria is the only fungus crops. Thripinema nicklewoodii. This nematode
currently commercially available for infects both larval and adult thrips.The
Paecilomyces fumosoroseus. This
control of western flower thrips. infective stage of the nematode bores
fungus has been investigated in Florida
Fungi for control of western flower thrips on into the host thrips’ body and multiplies
Beauveria bassiana. This fungus attacks greenhouse ornamentals (see “Aphids” in its abdominal cavity.The mature
many insects, including thrips (see for more information). This fungus nematode burrows into the thrips’ diges-
“Aphids” for more information). At least requires humidity over 90% for infec- tive tract, then passes out with its frass.
two strains of the fungus are commer- tion, which limits its utility. It is commer- Nematode infection in adult female
cially available. cially available only in Europe. thrips prevents egg production. Up to
20% of western flower thrips in California
Entomophthora parvispora. This fungus Verticillium lecanii. This fungus controls
are parasitized by indigenous popula-
causes significant thrips mortality aphids and whiteflies and has strong
tions of this nematode. However, T. nickle-
outdoors in Europe, but in greenhouses it potential for controlling both onion
woodii has not been investigated for use
usually changes quickly to the resting thrips and western flower thrips.
in greenhouses and is not commercially
spore stage that does not infect or Although it is commercially available in
available.
spread. Shortly after the thrips is infected Europe, this product is not expected to
by a conidial spore, the body becomes be available in North America for use
filled with fungal growth that kills the against any greenhouse pests in the
host insect in 3–6 days. Active spore near future.
preparations could be useful for green-
house applications, but the fungus would
not spread beyond the initial application.
It is not commercially available.
Entomophthora thripidum. Fungal epi-
zootics in onion thrips in European
greenhouses are usually caused by this
fungus. Where it occurs, it can almost
eliminate a thrips population within a
Mike Brownbridge

few weeks. The fungus completes its life


cycle in the insect host in 4 days. Only
the abdominal organs are infected, so
even after the fungus breaks through
abdominal coverings, the thrips remains
alive as long as the fungus continues to The fungus Verticillium lecanii infecting a thrips larva.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S T H R I P S 65
Possibilities for effective together may provide control over the Predatory mites are shipped in a carrier
biological control range of conditions characteristic of or packing material. If you receive the
Most of the research on the biological greenhouses. mites in loose packing material, sprinkle
control of thrips has focused on veg- The greenhouse crops themselves influ- the material (often cereal bran) onto
etable crops, especially pepper and ence the effectiveness of predatory plants. If the material is in slow-release
cucumber. Biological control can be mites. On peppers, N. cucumeris tends to packets (sachets), hang them on plants.
used on other greenhouse crops for establish easily, even without thrips. Because some strains of N. cucumeris
thrips control, but it may not suffice for Therefore, it can be introduced to diapause during the winter months,
ornamentals that cannot tolerate peppers in low numbers before thrips when the temperature drops below 70°F
cosmetic damage or the potential for are detected for earlier control. However, and daylight lasts less than 12.5 hours,
viral transmission. In vegetable crops, N. cucumeris is not as efficient a predator the mite is not usually released in green-
success often depends on the individual on cucumber, perhaps because the tri- houses in the upper Midwest until
grower’s tolerance for foliar damage chomes on cucumber leaves interfere March or April. However, warm night
during the initial control phase— with the mite’s ability to search. temperatures in the greenhouse, even
damage that will not affect fruit produc- Repeated releases are often necessary during winter, will prevent diapause in a
tion. Other limitations to the biological for successful establishment on this large proportion of the mites. N. degen-
control of thrips include reinfestation crop. Pollen is critical for optimal repro- erans can be released during fall and
from surrounding outdoor plants during duction of N. cucumeris, so it is not as winter.
warm weather, the propensity of successful on nonflowering plants and Success with these predators depends
western flower thrips to hide in flowers those that do not produce sufficient on releasing them as soon as thrips are
or buds, and the difficulty of protecting pollen. Thrips reproduction also can vary detected on sticky traps or plants. For
crops from tomato spotted wilt virus on different cultivars, and even a slight cucumbers, releases of 50 predatory
and impatiens necrotic spot virus if decrease in thrips reproduction may mites per plant plus an additional 100
present in a thrips population. allow the mites to prevail. per infested leaf are recommended. For
Nevertheless, if selected and released The greenhouse conditions into which peppers, recommended release rates are
appropriately for your thrips problem, predatory mites are released also influ- 10 predatory mites per plant plus an
predatory mites, minute pirate bugs and ence the mites’ effectiveness. Thrips that additional 25 per infested leaf. The initial
soil mites may control thrips. They may migrate from outdoors in summer can pest density does not seem to be impor-
work best when used in combination, as overwhelm the ability of the mites to tant as long as introductory rates are
detailed below. get the thrips under control. high. Repeated releases of predatory
The most commonly used natural Environmental conditions such as mites are recommended for good pro-
enemies for thrips control are the two relative humidity and temperature also tection of peppers or cucumbers. Make
phytoseiid predatory mites Neoseiulus affect predator efficiency and egg weekly releases until there is one preda-
cucumeris and N. degenerans. N. cuc- survival. Optimal conditions for preda- tory mite for every two thrips. The mites
umeris is effective against both the tory mites include moderate tempera- only attack thrips in their first and
western flower thrips and onion thrips, tures (around 86°F) and high humidity second instars, so it may take several
especially on cucumber. Most strains of (80–90%). Thrips thrive at 70–90% months for biological control to work.
N. cucumeris do not reproduce during humidity, but their mortality increases Making regular releases of predatory
short winter days with night tempera- above that humidity level. The mites will mites is also the only way to control
tures below 65°F. By contrast, N. degener- also be more efficacious if plant leaves thrips on ornamental crops, such as
ans does not enter diapause and will are touching, so that the mites are able chrysanthemum. You should release the
continue to develop under cool condi- to easily move from plant to plant. mites weekly, biweekly or monthly,
tions. The eggs of this species are also depending on the crop and the thrips
better able to tolerate low humidity, pressure. Before thrips are present, intro-
which can severely reduce hatching of duce the mites at 10–50 per ft2, depend-
N. cucumeris eggs. Both species are effi- ing on the crop. Good results have also
cient at low thrips densities and feed on been obtained on chrysanthemum
spider mites, broad mites, and pollen in using N. cucumeris at rates of 100 mites
the absence of thrips. The two species per individual cutting. Additional
66 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

releases of three mites per leaf are suffi- Minute pirate bugs are shipped as Minute pirate bugs enter a reproductive
cient where adult thrips pressure is adults in a carrier or packing material, diapause in late fall, so more regular
intense. Under less pressure, you can such as bran, rice hulls, or vermiculite, releases become necessary after late
gain control with fewer mites per leaf, along with a food source. Shake the October to keep populations high
but regular releases are still necessary. packing material onto the plants, and enough for thrips control. The use of
Remember, your own experience is your the bugs will readily disperse and locate supplemental blue light has been
best guide to the timing and rates of prey. Distribute the insects evenly shown to reduce the number of O.
natural enemy releases. throughout the greenhouse, but place insidiosus entering diapause, which may
Minute pirate bugs (Orius spp.) may be extra bugs in thrips hot spots, such as allow these bugs to suppress western
more effective than mites for control of open flowers. Release the bugs in the flower thrips in short-day flowering
thrips in some situations. Mites feed on early morning or late afternoon, or on a crops such as chrysanthemums.
thrips for only a small portion of their cloudy day, to reduce the chances they The predatory bug Dicyphus tamaninii
lives.The nymphal and adult minute will fly out of the greenhouse. Keep the was effective at controlling western
pirate bugs prey on both adults and vents closed or screened to prevent the flower thrips in a Mediterranean
immature stages of the thrips.They also bugs from leaving the greenhouse. cucumber greenhouse. A ratio of three
feed in flowers, where female western One minute pirate bug per plant was late-instar nymphs to 10 pests kept
flower thrips, which are directly responsi- sufficient to control western flower thrips under the economic injury level. A
ble for population growth, concentrate. thrips within 5 weeks in small-scale lower ratio was successful only when
Minute pirate bugs will also feed on experimental trials on greenhouse thrips populations were low. Releasing
other pests, such as spider mites, as well cucumbers. Release rates of one Orius late-instar nymphs would not be as
as pollen and plant juices, so they can insidiosus per 20 ft2 provided control of practical in a commercial greenhouse as
exist for long periods at low thrips densi- western flower thrips on peppers in releasing adults which could disperse to
ties. Orius are most effective when they some trials in European greenhouses, find the thrips, but might also leave the
reproduce in the crop, but the adults do but higher rates may be necessary on greenhouse.
not lay eggs equally well on all plants. O. other crops. On peppers an initial intro- The soil-dwelling predatory mite
insidiosus reproduces best on chrysan- duction of three to five Orius per 10 ft2 is Hypoaspis, which is not a phytoseiid, will
themum, gerbera, cucumber, and suggested in areas of pest activity. This attack thrips pupae in the soil when
peppers, but oviposition is so poor on should be followed by an introduction placed around the roots of each plant.
roses and carnations that the bugs of two Orius per 10 ft2 throughout the These mites work best when the soil or
provide no control when thrips are greenhouse 2–4 weeks later. On medium is moist, with an open struc-
present in the flowers. O. laevigatus and cucumber and eggplant, use 5–10 Orius ture, and soil temperature is at least
O. albidipennis are able to control western per 10 ft2 in specific areas of pest 59°F. It cannot be relied on as a sole
flower thrips on peppers and strawber- activity, followed 2–4 weeks later by a source of control for thrips in a commer-
ries, but not on cucumbers, due to the general introduction of one to two per cial greenhouse, but it could enhance
absence of pollen and the numerous 10 ft2. Additional research on a larger biological control by predators that feed
hairs on the cucumber leaves which scale is necessary to make recommen- on immature thrips and adults on
inhibit movement. Other Orius species dations for release rates on other crops leaves. In small-scale experiments this
may have different plant preferences. in commercial greenhouses. Minute mite reduced adult thrips by about 70%.
pirate bugs can also be used in conjunc-
These mites are also shipped in a carrier,
tion with predatory mites, although the
such as vermiculite, and are distributed
bugs will eat the mites if insufficient
by shaking the contents onto the soil or
food (thrips) is available.
potting medium. H. miles does not need
to be applied to every pot and/or flat of
plants because it will disperse some
distance on its own. It must be applied
early in the growing season to allow
establishment before thrips begin
pupating in the soil.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S T H R I P S 67
Using phytoseiid mites, minute pirate The parasite Thripobius semiluteus can Pathogens can also be used to suppress
bugs, and Hypoaspis together should be used for control of greenhouse thrips thrips populations. Adult western flower
offer the best control of thrips. The only. Because it only accepts this species thrips appears to be most susceptible to
predatory mites consume immature of thrips it is important to know what fungi possibly because they generally
thrips but do not kill adult thrips, which species you have before contemplating occur in flowers where humidity is
can continue to damage the plants for releasing this insect. The wasps are higher and conditions more favorable
several months until they die. Minute shipped in the pupal stage and should for infection. The higher humidity in the
pirate bugs will prey on adult thrips, as be held at 65°–75°F until adults emerge. flowers increases the potential for the
well as immatures on the foliage. The highly mobile adults will disperse fungus to sporulate and infect addi-
Hypoaspis will feed on thrips in the soil, throughout the greenhouse to find tional thrips. The commercially available
where the other predators do not occur. greenhouse thrips. Release rates vary Beauveria bassiana suppresses western
Releases of Hypoaspis and phytoseiids considerably depending on the crop, flower thrips on rose, gerbera, carnation,
together provided excellent control on thrips population levels, and other and other flowers as well as currently
chrysanthemum in Canadian green- natural enemies being used at the time registered insecticides do. However,
houses. Release the mites before thrips of release. Minimum rates begin at one none of these materials provide satisfac-
are detected, and release Orius in hot wasp per 20 ft2. Two releases should be tory control of western flower thrips. The
spots when thrips are found. If addi- made 2–3 weeks apart. Monitor for wasp fungus and predatory mites or Orius
tional releases are necessary, alternate reproduction and establishment by need to be used together to achieve
weekly between phytoseiids and Orius. observing immature thrips. Parasitized acceptable control.
The phytoseiid mites perform best thrips have a milky appearance. Once
under humid conditions and often die the parasite pupates inside the thrips,
out when humidity declines. Orius toler- the host body turns black and hard, but
ates drier conditions, so the two are still remains attached to the plant
complementary. surface.

Thrips
Enemies egg ————— nymph ————— adult

Ceranisus

Thripobius

Hypoaspis

Phytoseiid mites

Dicyphus

Macrolophus

Minute pirate bugs

Pathogens

Heterorhabditis

Thripinema

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness Enemy attacks hosts in soil only
68 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Several brands and formulations of


B. bassiana are available for use in green-
houses. The fungus takes 3–7 days to kill
Alternative control methods
an insect, so it will take some time to Sanitation Environmental control
suppress the thrips population when This is the most important cultural Misting to increase relative humidity
using these products. Thorough spray practice for preventing thrips near 90% will reduce thrips popula-
coverage is essential because fungal problems. Remove weeds that can tions on cucumber. Soil that is kept
spores must contact the insect for infec- act as a refuge for thrips or harbor wet and frequently waterlogged will
tion to occur. This fungus is susceptible viruses from inside and outside the drown thrips pupae or allow fungal
to some fungicides, so chemical fungi- greenhouse. Remove and immedi- epizootics to kill them. However, wet
cide applications should not be made ately destroy crop residues after soil may also increase plant disease,
within 48 hours of B. bassiana applica- harvest. In addition, remove any soil fungus gnat, and shore fly problems.
tions. Multiple applications are usually debris which could contain thrips
necessary to achieve control. Traps
pupae. Avoid wearing yellow and
Yellow or blue sticky traps can be
The whitefly strain of V. lecanii also blue clothing in the greenhouse
used for mass trapping of thrips. For
affects thrips. When humidity is high because they attract thrips and facili-
western flower thrips, which pupate
and temperatures are 64°–77°F, this tate the spread of the pest.
mainly in the soil, place the traps near
fungus can cause considerable mortality
Insect screens the soil to catch large numbers of
of thrips. In experiments on cucumbers,
Thrips migration into greenhouses newly emerged thrips before they
V. lecanii killed more than 80% of the
can be a continual problem. Monitor move into the crop. Blue traps may
thrips within 4–6 days. In comparative
the greenhouse with sticky traps to be more effective for mass trapping
assays, Metarhizium anisopliae caused
determine where thrips are entering, than yellow traps.
more adult mortality of western flower
and install screening to exclude
thrips than V. lecanii. The efficacy of M. Resistant cultivars
them. Screens on greenhouse vents
anisopliae increases as temperatures Certain chrysanthemum cultivars
and doors will help block thrips’ entry
increase, with an optimal range of appear to be more tolerant of
from outside. Screening just the side
75°–82°F. Paecilomyces fumosoroseus has western flower thrips than others.
of the greenhouse that faces into the
also been shown to be effective against Dutch researchers have found differ-
prevailing winds can also be effective
thrips in some tests. ences in susceptibility of cucumber
at reducing thrips, even for passive air
These pathogens are applied as spores cultivars. If possible, select varieties
intake systems. The maximum hole
suspended in water for spraying. High that do not show as much injury
size to exclude thrips is 192 µ. Thrips
humidity is necessary for infection. from thrips feeding.
can easily get through commercial
When humidity is low, the performance whitefly-proof screens or unwoven Chemical control
of the fungus is unpredictable. Humidity polyester filters. Spinosad is an insecticide derived
can be increased by dampening the from natural metabolites produced
plants with water sprays. Late afternoon Soil sterilization (pasteurization)
under fermentation conditions by the
applications reduce spore injury by Steam sterilizing the soil or the used
actinomycete Saccharopolyspora
ultraviolet light and desiccation, since planting medium between crops will
spinosa. It has a high level of contact
the greenhouse is more humid at night. kill immature thrips that are pupating
and oral activity and rapid speed of
However, these fungi may be practical in the soil.
action. It is very effective against
for use only in humid areas with thrips but has low to moderate
moderate temperatures, such as rooting impact on beneficials. It is registered
benches and shade-cloth covered areas for control of many pests on land-
used to induce inflorescence in chrysan- scape ornamentals and can be used
themums. Repeated applications will be on ornamentals in shade houses and
necessary if humidity is not high lath-houses.
enough to allow continuous infection.
Since these fungi do not impair natural
enemies, their use can be integrated
with predatory mites or bugs.
Weevils 69

T
he black vine weevil is a cosmopoli- greenhouse the length of the life cycle
tan pest that caused economic depends on temperature rather than
Available natural enemies and
their potential for control damage to greenhouse plants as time of year.
Insect-parasitic early as 1834 in Germany. It is an
nematodes can be very imported species but is established Monitoring
effective against weevil throughout the northern United States. Pitfall traps, trap boards, and burlap
larvae. It has a host range of over 140 plant traps have been developed for monitor-
species, including many common green- ing both black vine weevil and straw-
Order Coleoptera: berry root weevil adults. Inspect the
house ornamentals such as ferns,
Beetles traps once or twice per week. Record the
geranium, gloxinia, primrose, rose, and
Family Curculionidae: Weevils especially cyclamen, as well as straw- number of weevils in each trap to calcu-
Black vine weevil, Otiorhynchus berry, yew, and numerous other nursery late the average number of weevils per
sulcatus plants. The host range of strawberry root plant or determine where localized
weevil, also an introduced species, is infestations occur.
Strawberry root weevil,
Otiorhynchus ovatus more restricted.
Natural enemies
Damage No parasites or predators attack root
Larvae of both species feed on roots and weevils specifically, although some gen-
other underground portions of plants. eralist predators, such as carabid beetles
They feed on the smaller roots first, then and ants, may occasionally kill larvae.
tunnel through the larger roots and Nematodes are the only natural enemies
move up into the crown. This reduces developed for commercial use.
plant vigor and in heavy infestations
Pathogens
may kill the plant. The feeding damage
Several entomogenous nematodes
may also make plants susceptible to
occur naturally in the soil and parasitize
diseases. Adult weevils feeding on the
a variety of soil-inhabiting insects,
foliage may cause cosmetic damage, but
including weevil larvae. Nematodes are
otherwise they do little damage.
long, slender roundworms. They are
about 1⁄64 inch (0.5 mm) long, transpar-
Description and life cycle ent, and practically invisible to the
The adult weevils are 3⁄8 inch (8–9 mm)
naked eye. They seek out insects in
long. The shiny black beetles feed on
moist soil; moisture is essential to their
foliage or flowers at night and do not fly.
movement and persistence. They enter
Each female may lay over 600 eggs, scat-
an insect through natural openings and
tering them randomly over the soil
release a bacterium that kills the host
surface. The larvae hatch in approxi-
within 48 hours. The bacteria then serve
mately 2 weeks and feed on roots as
as food for the nematodes, which
they go through five to six instars. The
complete their development inside the
larvae are whitish, wrinkled, legless
dead insect. The next generation of
grubs up to 3⁄8 inch (1 cm) long. The
nematodes leaves the insect in search of
mature larvae hollow out earthen cells
new hosts. Unlike plant-parasitic nema-
in the soil and enter the prepupal stage,
todes, these nematodes do not damage
which lasts from 3 weeks to several
plants. Neither do they harm people,
months, depending on temperature.
animals, beneficial above-ground insects,
After 2–3 weeks in the pupal stage, the
or earthworms. Entomogenous nema-
adult remains in the pupal cell for an
todes are propagated and packaged to
additional week as the body hardens
use as biological insecticides.
and darkens before emerging. In the
70 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Heterorhabditis bacteriophora (=helio- Metarhizium anisopliae. This insect-


thidis). This nematode attacks black vineparasitic fungus has considerable poten-
and strawberry root weevils, as well as tial as a microbial control agent for
numerous other insects. Although weevils on a number of potted plant
larvae, pupae, and adults can become species, but it is not commercially avail-
infected, the larvae are the most suscep-able. Various fungal strains have been
tible. The bacteria carried by the nema- investigated that differ in efficacy
todes will kill larvae within a few days.against weevil larvae, the persistence in
The nematodes reproduce in the dead soil or potting media, and optimal tem-
host insect, and only the infective larval
perature ranges. Also, the host plant
stage lives free in the soil. Several affects the efficacy of the fungus.
thousand infective juvenile nematodes Control on some plants, such as
emerge from host cadavers within 2–3 cyclamen, is lower than on other plants,
weeks of the host’s death. This such as azalea, begonia, impatiens, and
nematode is commercially available. kalanchoe. The best control is achieved
Steinernema (=Neoaplectana) car- when the fungus is applied as a soil
pocapsae. This nematode has consider- drench before the eggs hatch. The larvae
able potential as a biological control quickly move down into the soil after
agent because of its rapid action and hatching and are easily infected when
environmental safety. It attacks weevils moving through the concentrated
in a manner similar to H. bacteriophora. spores in the upper layers of the soil. The
You can achieve good control with this fungus can also be incorporated into
nematode if they are well-distributed potting media before planting, and may
throughout the potting medium. Several provide better control of weevils on
commercial brands are available. certain plants than a drench will. This
fungus is not registered for use in the
United States on any greenhouse crops.
Ralph E. Berry, Oregon State University

Nematodes Heterorhabditis bacteriophora look


like spaghetti inside a black vine weevil larva. The
top larva is not infected.
Ray Cloyd

Steinernema carpocapsae nematodes.


S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S W E E V I L S 71
Possibilities for effective To apply the nematodes, simply mix the Nematodes should be applied to moist,
biological control package contents with water, as but not saturated, soil. This may require
Nematodes are an economical and directed on the package. Occasionally, watering the area to be treated before
effective method for controlling black improper storage or shipment has application. The soil and water tempera-
vine weevil and strawberry weevil on resulted in non-viable nematodes. After ture must be above 65°F for the nema-
potted plants in commercial green- adding the nematodes to water, todes to be effective. Application in the
houses. They are very cost effective, examine with a magnifying glass to be early evening or morning is recom-
readily available, and easy to apply with certain that the nematodes are moving. mended to avoid exposing the nema-
existing chemical application equip- You can apply nematodes as a soil todes to extreme heat and sunlight. It is
ment. Nematodes are as effective as or drench to potted plants. Nematodes can important to keep the treated pots
better than many chemical controls. withstand pressures of up to 300 psi, so moist after the application.
you can apply them with the same It may take 10–30 days for root weevil
To be effective, nematodes must be
equipment used for chemical pesticides. populations to decline, but control can
applied shortly after weevil eggs hatch.
Application rates of 40,000 nematodes occur 2–5 days after treatment if the
Applications made before weevil eggs
per 6-inch pot and 80,000 nematodes larvae are close to the soil surface. More
hatch provide no control. Unfortunately,
per 8-inch pot caused 100% weevil mor- than one application of nematodes may
timing the application is difficult
tality in experimental trials. Application be necessary if soil conditions are not
because adults do not begin laying eggs
to the soil surface is suitable for plants suitable for nematode persistence—that
until about a month after they emerge.
with shallow root systems, such as is, moist and moderately warm.
Egg laying occurs over a 4–6 week
cyclamen. At a rate of 20,000 nematodes
period. After that, the eggs take only
per liter (1 quart) of soil, a surface appli-
about 2 weeks to hatch, opening a
window of opportunity for biological
cation killed 90% of the beetle larvae in
pots. For plants with roots throughout
Alternative
control. Try applying nematodes 6–8
weeks after your peak weevil catch.
the pot, soil injection would be much
more effective, but it is twice as time
control methods
Most eggs should have hatched by then.
consuming as surface application. Sanitation
Sterilize soil to kill all stages of the
weevil. Eliminate old strawberry beds
in the vicinity of greenhouses that
serve as breeding places for the adult
weevils.

Insect screens
Screen vents and doors to prevent
adult weevils from crawling into the
greenhouse.

Weevils
Enemies egg ——— larva ——— pupa adult

Nematodes

Metarhizium

Natural enemy attacks the host Effectiveness depends on species of enemy


Width of bar indicates degree of effectiveness
72 Whiteflies
W
hiteflies infest a wide range of On ornamentals, whiteflies are pests pri-
greenhouse crops. The green- marily because their presence reduces
Available natural enemies and
their potential for control house whitefly is a tropical the aesthetic value of a plant. Heavy
There are a few available species that has become established infestations of silverleaf whitefly can
parasites of whiteflies. worldwide in greenhouses. It is a partic- cause stem and bract whitening on red
Some predators and ularly destructive pest of cucumber, poinsettia cultivars.
pathogens can be used tomato, fuchsia, geranium, hibiscus, Whiteflies also act as efficient vectors of
along with the parasites gerbera, and poinsettia. It attacks several viral diseases. The greenhouse
to improve control. The hundreds of other vegetable and orna- whitefly can transmit the beet pseudo
potential for successful mental crops. The greenhouse whitefly is yellow virus to cucumbers, while the
biological control varies from the most common species found in sweetpotato whitefly can transmit
moderate to high. northern greenhouses. The sweetpotato several viruses such as those causing
whitefly, from tropical and warm tem- tomato yellow leaf curl and cucumber
Order Homoptera: Aphids, leafhop-
perate regions, is also worldwide in dis- vein yellowing.
pers, planthoppers, mealybugs,
tribution and attacks hundreds of orna-
scales, and whiteflies
mental plant species. The silverleaf Description and life cycle
Family Aleyrodidae: Whiteflies whitefly, formerly known as the B strain Adult whiteflies are about 1⁄16 inch (1–3
Greenhouse whitefly, Trialeurodes of the sweetpotato whitefly, attacks mm) long and covered with a white,
vaporariorum poinsettia, hibiscus, rose, and other waxy powder. Most species appear very
Sweetpotato whitefly, plants. It extracts much more sap than similar, but they can be distinguished
Bemisia tabaci does the sweetpotato whitefly and also from each other. Greenhouse whiteflies
causes squash silverleaf, phytotoxemia hold their wings rather flat over the
Silverleaf whitefly, in poinsettia, and irregular ripening dis- abdomen. Sweetpotato and silverleaf
Bemisia argentifolia orders in tomato. Until 1993 the silver- whiteflies hold their wings roof-like and
Bandedwing whitefly, leaf whitefly was not distinguished from close to the sides of the abdomen. The
Trialeurodes abutilonea the sweetpotato whitefly, so much of bandedwing whitefly has light gray bars
the literature concerning sweetpotato across the wings. The adults of all these
whitefly may actually refer to silverleaf species congregate on the undersides of
whitefly. The bandedwing whitefly is less leaves, usually at the top of the plant.
common than the greenhouse, sweet-
Female greenhouse whiteflies lay 6–20
potato, or silverleaf whiteflies but occurs
eggs daily, in a perfect circle or a portion
on many plant species. It commonly
of a circle, on the underside of the leaf.
enters the greenhouse in late summer
The cigar-shaped eggs are deposited as
and early fall.
the female moves around in a circle with
her mouthparts inserted in the plant as
Damage a pivot. Sweetpotato whitefly eggs are
Whiteflies damage plants by removing
laid randomly, in small groups or singly,
plant sap and excreting large quantities
which makes them more difficult to
of honeydew. The clear, sticky honeydew
detect. The eggs are white when first
is often seen on the lower leaves.
laid but later turn dark brown. They
Honeydew serves as a medium for
hatch in 7–10 days into mobile first-
growth of black sooty mold fungus
instar nymphs —”crawlers” that
which interferes with photosynthesis
resemble scale crawlers—which search
and transpiration, and detracts from the
for a suitable place to settle, seldom
beauty and marketability of the crops.
moving more than 3⁄4 inch from the
Large numbers of adults and nymphs
eggshell. The next two instars are sessile,
feeding on the leaves cause the leaves
translucent green or yellow, rather flat,
to wilt or turn yellow and drop, and may
scale-like, and inconspicuous. The fourth
seriously reduce the yield of vegetables.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S W H I T E F L I E S 73
and final instar, often referred to as the greenhouse crops at planting. Whiteflies Sticky traps are useful for detecting the
pupa (although this species undergoes are generally not found uniformly first invasion of whiteflies, and they
simple metamorphosis), becomes throughout a crop at first. They will be provide information on the relative
opaque yellow and mounded. concentrated in a few areas and many abundance of whitefly adults. But many
The nymphs and pupae of many species areas will be without insects. As the variables, such as trap size and shape,
have distinctive white, waxy spines, but population grows, the infested areas amount of sunlight, and air movement
the sweetpotato whitefly has no spines. expand and multiply. affect trap catches. Do not rely on traps
Greenhouse whitefly pupae are cake- You should monitor whiteflies in two alone to monitor whitefly populations. If
shaped with erect side walls, whereas ways. Use yellow sticky traps to detect adults are detected, eggs and nymphs
the sweetpotato and silverleaf whitefly the first adult whiteflies and to monitor will be present, and visual plant inspec-
pupae are mounded. In the greenhouse adult whitefly populations. In addition, tion should be performed.
the generations usually overlap com- visually examine plants to determine You can see whiteflies most often on rel-
pletely. Greenhouse whiteflies develop the number of immatures present. atively new or young tissue. Turn the
best at moderate temperatures around Yellow sticky traps placed throughout leaves over to examine the undersides
75°F, while sweetpotato and silverleaf the crop will detect adults at very low for whiteflies. Adult whiteflies can easily
whiteflies prefer temperatures over 81°F. levels—one adult for every 10 plants. be seen, but a 10X–15X magnifier or
Traps are commercially available or can hand lens is helpful for observing
Monitoring be made by the grower. Hang the traps immature stages. Whitefly populations
Biological control of whiteflies will be just above the crop canopy to detect will vary considerably on different plants
difficult unless the infestation is greenhouse whitefly. You can place the and even on different cultivars of the
detected early. The greenhouse should traps within the crop canopy, or even on same type of plant. Maintain scouting
be thoroughly inspected for problems the ground or on a greenhouse bench records of observed pest levels for all
before the crop is introduced. Check for detecting sweetpotato whitefly, plant species and cultivars. Review these
hanging baskets and weeds under the especially in tall crops. Also place traps records to determine which plants
benches. Begin monitoring susceptible near doors, air-intake vents, and among should be monitored more carefully.
newly arriving plants. The number On many crops there can be mixed
of traps needed depends on the infestations of different species of white-
crop, but on average space them flies. Determine which species are
45–60 ft apart (one per 1000 ft2). present before implementing control, as
Check the traps once or twice natural enemy efficacy can vary depend-
weekly. Counting the whiteflies on ing on the whitefly species present.
a 1-inch wide vertical band of
If you are using Encarsia formosa wasps
either side of the sticky traps
for biological control, monitor both the
gives a good approximation of
whitefly and the wasp populations.
total trap densities and saves
Some adult parasites will also be caught
time.
on yellow sticky traps. Learn to recog-
nize the tiny adult wasps and black par-
asitized nymphs. Record parasitism rates
weekly until the whitefly populations
are under control, and periodically
thereafter. Remove yellow sticky traps
before releasing wasps and then put
them back up 3–4 days after releases are
made.
Ray Cloyd

Place yellow sticky traps throughout the crop to


detect adult whiteflies.
74 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Natural enemies Parasites Encarsia formosa. This wasp originated


Natural enemies of whiteflies include Several tiny parasitic wasps that attack in the semitropical areas of the New
many parasitic wasps, predators, and whiteflies have been used successfully World and was one of the first recorded
several species of fungus. Exploration for for biological control. None of these parasites in greenhouses. It has been
and evaluation of new natural enemies, wasp families or species have common used since the 1920s to control the
including many not mentioned below, names. Those discussed here are in the greenhouse whitefly in warm green-
continues. families Aphelinidae and Platygastridae. houses. E. formosa will also parasitize
The taxonomy of these wasps is very sweetpotato whitefly in greenhouses,
confused, and the true identify of many but this whitefly is not a good host for
species or strains is questionable. The this wasp, so control is not as effective.
encyrtid Encarsia formosa is widely used The tiny females, about 1⁄40 inch (0.6
to control the greenhouse whitefly in mm) long, are black with a yellow
greenhouses. Most of the other wasps abdomen and opalescent wings. Males
listed below are not yet commercially are somewhat larger than females, com-
available. pletely black, and extremely rare. The
Amitus fuscipennis. This Central parthenogenic—or asexually reproduc-
American platygastrid wasp is less than tive—females deposit 50–100 eggs indi-
1⁄ 25 inch (1 mm) long. The adults are vidually inside the bodies of third-instar
black with reddish brown legs and nymphs or pupae. The wasp larvae
antennae, and brown wings. Their larvae develop through four instars in about 2
parasitize whitefly nymphs, and the weeks at optimal temperatures. As the
Cliff Sadof

adult wasps emerge from the whitefly larvae grow they kill the whiteflies.
pupae. The females are very short-lived, Parasitized pupae of greenhouse
but they can parasitize many hosts if whitefly turn black in about 10 days,
The black whitefly nymphs are parasitized while the pupae of parasitized sweet-
by the encyrtid wasp Encarsia formosa; the they are available during this time.
A. fuscipennis’ effectiveness against potato whiteflies turn amber brown.
white nymph is unparasitized.
greenhouse and sweetpotato whiteflies Both are easily distinguished from
on poinsettia is being studied. This wasp unparasitized pupae. Wasp larvae
is produced by a grower cooperative in pupate within the whitefly body. Adults
Colombia but is not commercially avail- emerge about 10 days later. The adult
able in the United States. wasps also kill whitefly nymphs by
feeding on them through holes made
with the ovipositor. This species is widely
available commercially.
Koppert B.V.

Ray Cloyd

Encarsia formosa is a tiny wasp that parasitizes green-


house whiteflies.
Encarsia formosa larvae have pupated and emerged as
adults from these whitefly nymphs.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S W H I T E F L I E S 75
Encarsia luteola (=E. deserti). This Other Encarsia species. Several other Eretmocerus eremicus (=californicus).
species resembles E. formosa but is species of Encarsia have been men- This aphelinid wasp, indigenous to
slightly smaller, lighter brown, and males tioned as possible biological control desert areas of California and Arizona, is
are produced regularly. In the field it is agents for whiteflies in greenhouses. The a parasite of greenhouse, bandedwing,
usually found on upper leaves. It will development of these species varies silverleaf, and sweetpotato whiteflies.
parasitize greenhouse whitefly in the only slightly from that of E. formosa. The 1⁄32-inch (0.75-mm) adult females
lab, but parasitized nymphs do not turn These potentially useful species include are lemon-yellow with green eyes. The
black. Researchers in southern California ■ E. inaron (=E. aleyrodis=E. parteno- males are yellow with light-brown
and Arizona are evaluating E. luteola for pea), a Mediterranean species that markings. Both sexes have very long
control of sweetpotato and silverleaf parasitizes the pupae of green- antennae. Females lay eggs beneath
whitefly. This species is commercially house and sweetpotato whitefly; whitefly nymphs of any stage but prefer
available. the second and third instars. They will
■ E. lutea parasitizes sweetpotato
Encarsia pergandiella (=E. bemisiae=E. not lay eggs near whitefly pupae. The
whitefly pupae;
tabacivora=E. versicolor). This aphe- first instar wasp larvae feed externally
■ E. meritoria, from California, will par- until the host pupates. Then the wasp
linid wasp, native to North America, par-
asitize bandedwing, greenhouse, larvae chew inside the host and
asitizes greenhouse, sweetpotato, and
and sweetpotato whiteflies but complete their development. The adults
bandedwing whiteflies. The 1⁄40-inch
prefers other species, such as the do not host feed on all species of
(0.5-mm) adults are yellow and brown.
iris whitefly (Aleyrodes spiraeoides). whitefly, but do host feed on sweet-
Over a 2-week period, females lay an
Females are golden yellow, and potato and silverleaf whiteflies on poin-
average of 50 eggs singly inside whitefly
males are brown with yellow heads settia. This wasp does not reproduce
nymphs, preferring second or third
and legs; well on sweetpotato whitefly on poin-
instars. Parasitized whitefly nymphs do
not turn black. The adults also feed on ■ E. transvena (=E. sublutea), a tropical settia, but it reproduces very well on the
body fluids of third-instar nymphs and species. The lemon-yellow adult same whitefly on other plants, such as
pupae. This wasp develops faster than E. females parasitize third-instar hibiscus. It does best under hot condi-
formosa under cool conditions. However, nymphs of greenhouse or sweet- tions as would be found in late summer
its use in greenhouses may be limited potato whiteflies and host feed on in desert valleys (80°–110°F), but its
because some strains hyperparasitize second-instar nymphs and pupae, as longevity is considerably reduced. At
their own larvae or those of other well as on whitefly honeydew; and 70°–75°F its development is too slow to
Encarsia species to produce males. This control whitefly outbreaks. This species
■ E. tricolor, a European parasite of the
wasp is not available commercially in is commercially available.
greenhouse whitefly that oviposits
the United States. in pupae, but whose efficiency in
greenhouses is lower than that of E.
formosa. It does not survive above
93°F.
None of these species is commercially
available yet.
Koppert B.V.

The female Eretmocerus eremicus lays an egg beneath a whitefly


nymph.
76 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Eretmocerus haldemani. This is prima- Predators Chrysoperla rufilabris. This species is


rily a parasite of sweetpotato and band- A handful of specialists and many similar to C. carnea in biology and devel-
edwing whiteflies, although sometimes general predators feed on various stages opment, but it is better adapted for
it will parasitize greenhouse whiteflies. of whiteflies. Only a few have been development under humid conditions
Females are lemon yellow and males are investigated for their ability to control than C. carnea. This generalist predator is
dark yellow with brown markings. whiteflies in greenhouses. commercially available.
Females lay eggs beneath whitefly Chrysoperla (=Chrysopa) carnea. The Delphastus pusillus. This small, black
nymphs. First-instar wasp larvae feed larvae of green lacewings feed primarily lady beetle feeds on bandedwing,
externally for 3–4 days, then enter the on aphids but will also feed on greenhouse, silverleaf, and sweetpotato
whitefly nymph and molt to second immature whiteflies and other insects in whiteflies. Females are all black, while
instars. The second instars continue the absence of aphids. They develop males have a brown head. The adult
development only after the host more slowly on whiteflies than on females and larvae feed primarily on
pupates. Its development is similar to aphids and rarely reproduce in green- whitefly eggs but will also eat whitefly
that of E. eremicus. It is also most effec- houses. Green lacewings are available nymphs and spider mites. However, they
tive under hot conditions. E. haldemani is from many commercial suppliers. (See do not reproduce well without whitefly
not commercially available. “Aphids” for a description and life cycle eggs. Females, which can live up to
Eretmocerus mundus. This information.) 2 months, consume more than 150
Mediterranean parasite of sweetpotato Chrysoperla comanche—Comanche whitefly eggs or nymphs per day. They
whitefly lays eggs under any nymphal lacewing. This is another green lay their eggs at the tops of plants near
whitefly stage, although it prefers lacewing species that feeds on aphids whitefly eggs. The beetle larvae eat
second and third instars. The eggs hatch and a variety of other insect pests. In the nearly 1000 eggs each during their
only after the host pupates. The wasp absence of aphids, the larvae feed vora- 2-week developmental period. They dis-
larvae then enter the whitefly bodies ciously on whitefly eggs and nymphs, criminate between parasitized whitefly
and develop. In the field they are usually and will occasionally consume adults. nymphs (those in which the parasite has
found on the lower leaves of the plants. This species is closely related to C. rufi- developed for at least a week) and non-
E. mundus is not commercially available. labris, but is more adapted to dry condi- parasitized ones, and avoid feeding on
tions. It is commercially available. the parasitized nymphs. They pupate on
the lower leaves, in leaf litter, or in other
protected locations. If they pupate in
saucers at the bottom of potted plants
or other places that hold water, the
beetles will drown when the plants are
watered. This beetle is commercially
Richard Lindquist, The Ohio State University

available.
Other lady beetles. Larvae and adults
of Coleomegilla maculata and the con-
vergent lady beetle (Hippodamia conver-
gens) have been reported as predators
of bandedwing whitefly, feeding on
eggs, nymphs, and pupae. C. maculata,
Coccinella septempunctata, Cycloneda
sanguinea, and several others have been
reported as predators of sweetpotato
whitefly. None of these is as efficient as
The tiny black lady beetle Delphastus pusillus adult the other described predators, and they
consumes more than 150 whitefly eggs and nymphs per do not reproduce in greenhouses.
day; larvae eat nearly 1000 eggs during development.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S W H I T E F L I E S 77
Dicyphus tamaninii. This Mediterranean contents, leaving only the skin behind. Predatory mites. Several phytoseiid
mirid bug eats a wide range of prey and This bug will feed on plant sap, but does mites are known predators of sweet-
is an important predator of whiteflies not cause damage on most crops. It can potato whitefly on cotton in Sudan.
and other insect pests on tomatoes in cause damage to gerbera flowers, and Amblyseius aleyrodis feeds on whitefly
southern Europe. The bugs prefer may reduce tomato fruit set on some eggs and nymphs, as well as spider mite
whitefly nymphs to other insects, cultivars when prey is limited. This eggs, pollen, and plant juices. Immature
tending to feed on prey on the under- predator is commercially available in mites require 15–20 whitefly eggs or
side of leaves. When prey is scarce they Europe where it is widely used to nymphs for their development and the
will feed on tomato fruit but not control whitefly, thrips, mites, and cater- adults consume up to three eggs or two
cucumber fruit. They require approxi- pillars. It is not currently permitted to be nymphs daily. Euseius delhiensis (=rubini),
mately 19 days to complete develop- imported into North America, although E. scutalis, and Amblyseius swirskii have
ment at 77°F. This bug is not commer- some Canadian suppliers advertise it. also been reported as predators of
cially available. Orius spp. Several species of minute sweetpotato whitefly eggs. None of
Geocoris punctipes. This 1⁄16–3⁄16 inch pirate bugs are generalist predators that these predatory mites has been investi-
(2–4 mm) bigeyed bug, endemic to the feed on whiteflies, thrips, spider mites, gated as whitefly predators in green-
southwestern United States, is usually aphids, and caterpillar eggs, as well as houses, but E. delhiensis may be com-
dark brown or black. During nymphal pollen. They are also cannibalistic under mercially available.
development and as adults, they crowded conditions. The black, 1⁄16–3⁄16
consume numerous aphids, small cater- inch (2–5 mm) long adults are ovoid and
pillars, nymphal and pupal stages of somewhat flattened, with distinctively
whiteflies, other small insects, and spider patterned black and white wings. Their
mites. Supplementary green plant eggs are laid in leaf tissues with one end
material and seeds in their diet sticking out. The tiny Orius nymphs are
improves development, reproduction, pinkish-yellow to light brown. Both
and survival. Sunflower seeds scattered nymphs and adults are very active and
on plants have enhanced Geocoris pop- feed on all stages of whiteflies. Most
ulations in experimental vegetable crop species enter diapause under short days
fields. This bug has not been investi- and lower temperatures. Several species
gated for use in greenhouses, but is of Orius are available commercially.
commercially available.
Macrolophus caliginosus. This preda-
tory mirid bug attacks all stages of
whitefly, but prefers eggs and nymphs. It
will also feed on aphids, and to a lesser
extent spider mites, moth eggs,
leafminer larvae, and thrips, but popula-
tions develop most rapidly on whiteflies.
The slender adult bugs are bright green,
about 1⁄4 inch (6 mm) long with long
legs and antennae. Females lay 100–250
eggs in leaves and stems, depending on
temperature and food. The eggs take
Ray Cloyd

about 2 weeks to hatch. The yellowish-


green nymphs are found mainly on the
underside of leaves where their prey is.
Adult Orius insidiosus.
Both nymphs and adults actively search
for their prey. Adults eat 40–50 whitefly
eggs per day. They insert their mouth-
parts into the whitefly and suck out the
78 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Pathogens Beauveria bassiana. This fungus infects Paecilomyces fumosoroseus. This


Several fungi kill whiteflies. Most require numerous species of insects, including fungus infects all stages of whiteflies, as
high humidity for infection and devel- whiteflies, thrips, and aphids (see well as thrips and aphids, but it is most
opment, so they will be most useful in “Aphids” for more information). All effective against whitefly nymphs (see
high-humidity greenhouses. Because stages of the whitefly may be infected. “Aphids” for more information).
they are considered pesticides, they Several brands and formulations of Subsequent reinfections are not
must be approved by the Environmental Beauveria are commercially available for common in greenhouses, so reapplica-
Protection Agency before they can be use in greenhouses. tion may be necessary. This fungus is
sold commercially. Metarhizium anisopliae. This soil-borne limited by the fact that it requires a
Aschersonia aleyrodis. This fungus, first fungus infects over 200 species of relative humidity of around 90% for
described on citrus whitefly nymphs in insects, including whiteflies and aphids infection. It has been investigated in
Florida, is highly specific to whitefly (see “Aphids” for more information). It is several states for use on greenhouse
nymphs. The older the whitefly nymph, not registered for use in the United ornamentals and is commercially avail-
the less likely it is to be infected by this States on any greenhouse crops. able in Europe, but is not available in
fungus. Infected whitefly nymphs typi- North America.
cally have a “fried egg” look, with a yel-
lowish discoloration in the center of the
body. Under humid conditions bright
orange, slimy spore masses form on the
outside of infected nymphs. The first
signs of infection, a change in color from
normal whitish to yellow, occurs in 24–48
hours. White external mycelial growth
begins in 4–6 days, and the orange color

University of Florida
develops under the appropriate condi-
tions in 7–9 days. Infection is limited by
relative humidity, just like for other fungi.
It tolerates a wide range of temperatures
(59°–86°F), but does best at 73°–77°F. It
does not appear to infect parasitized The fungus Paecilomyces fumosoroseus consumes its host
nymphs and is harmless to adult para- from the inside out. The powdery spores turn the infected
sites. It is not commercially available. insect white at first, then change to shades of pink.

Mike Brownbridge
University of Florida

Spores of the fungus Beauveria bassiana on whitefly


produce the characteristic “white bloom” appearance.
Whitefly nymphs infected with the fungus Aschersonia aleyrodis.
In humid conditions bright orange spore masses form on the
outside of infected nymphs.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S W H I T E F L I E S 79
Verticillium lecanii. This insect-parasitic as insecticidal soap, horticultural oil, on the plants near whitefly infestations
fungus infects both aphids and white- natural pyrethrins (not synthetic as soon as possible after the shipment
flies. The fungal strain with large spores pyrethroids), or insect growth regulators arrives. Be sure to distribute the pupae
infects aphids; the one with smaller before releasing natural enemies. evenly throughout the greenhouse
spores infects whiteflies. V. lecanii grows Natural enemy species introduced alone because these very small wasps will
and multiplies in the greenhouse at may provide effective whitefly control in disperse only about 6 feet from the
temperatures of 59°–77°F. Fungal spore certain situations. However, a combina- release point. Place extra wasps in
germination and infection occurs only tion of parasite releases, predator whitefly hot spots. Save several cards
when the relative humidity is greater releases, applications of nonresidual from each shipment to check for the
than 90% for at least 10 hours per day. insecticides, and leaf removal are often percentage of wasps emerging from the
The spores germinate and bore through necessary to produce commercially pupal stage; quality can vary between
the whitefly, then grow inside it, killing acceptable crops. shipments and suppliers.
it. The fungus appears as cottony white The parasite Encarsia formosa is the Greenhouse growers typically introduce
fluff on infected whiteflies which most widely used insect for biological Encarsia wasps either by inundation or
detracts from the appearance of some control of whiteflies. It has been used seasonal inoculation. Inundation, in
ornamentals. The fungus also causes extensively for many years in commer- which massive numbers of the parasite
some mortality of whitefly parasites cial vegetable greenhouses throughout are released into the greenhouse at
inside the pest’s body, but it seems to be the world. It can control whiteflies on once, is the simplest but can be expen-
compatible with other parasites. A com- vegetable crops, including cucumber sive. The more common method uses
mercial formulation of the fungus and tomatoes, and several ornamental successive introductions of smaller
specific to whiteflies is available in crops, such as poinsettia. On vegetable numbers of parasites. Make the first
Europe. V. lecanii is not yet registered forcrops that can tolerate a few whiteflies, release as soon as, or before, the first
use in North America. such as tomato, a single inoculative adult whiteflies are detected, either by
release of Encarsia formosa may be suffi- visual inspection or on yellow sticky
Possibilities for effective cient for control. However, on most veg- traps. Continue to make regular releases
biological control etable crops or short-term crops with a weekly or every other week until
Although there are many different much lower tolerance for whiteflies, 60–80% of the whitefly pupae are para-
strategies for biological control of white- such as poinsettia cuttings, multiple sitized. It is important to know how to
flies in greenhouses, certain principles inundative releases will be necessary. recognize adult parasites and their para-
apply in most cases. Natural enemies are Many other promising natural enemies sitized whitefly pupae. Check the crop
most effective if introduced when can be used in conjunction with every 3–4 days to make sure adult
whitefly numbers are low. Biological Encarsia wasps. Encarsia are present. The first parasitized
control often fails if initiated after whitefly pupae should be visible 2–3
The best time to introduce Encarsia is
whitefly populations exceed one adult weeks after the first release. As a general
when the whitefly population is low.
per 10 upper leaves on vegetable crops guide, about 25% of the whitefly pupae
Wasp efficiency is seriously impaired
and fewer on ornamentals. Biological should be parasitized within 1 month of
when whitefly nymphs are too
control usually does not eradicate the first introduction, 50% after 2
numerous (30–60 per in2). The wasps
whiteflies but maintains the pest popu- months, and 80% after 3 months. To
spend more time cleaning the excessive
lation at a low density. On many crops, determine the percentage parasitized,
honeydew from themselves than
especially poinsettia, there may be a count the number of black (parasitized)
searching for and parasitizing whiteflies.
mixed infestation of greenhouse and sil- and white (nonparasitized) whitefly
Consult the section on monitoring
verleaf whiteflies. It is important, there- pupae on the underside of the leaves.
whiteflies for instructions on estimating
fore, to correctly identify the whiteflies Do not count pupae from which adult
whitefly populations.
in your crop so that the proper rates of whiteflies or parasites have emerged.
the most effective natural enemies can Encarsia wasps are shipped in the pupal
There is a third, do-it-yourself method
be released. Residual insecticides should stage, glued to small cards that can be
hung directly on the plants. Sometimes for introducing Encarsia to the crop—
not be used within a month of parasite
using banker plants. In an area that is
or predator release. If the whitefly popu- the adult Encarsia have already
emerged, so open the parasite shipment separated from the production area,
lation is high, reduce the number of
infest young tomato, cucumber, or
whiteflies with a nonresidual spray, such in the greenhouse. Distribute the cards
80 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

tobacco plants with whiteflies when the on specific situations. However, the trial, four Encarsia per ft2 controlled
plants have about six leaves. Three number of Encarsia that actually emerge greenhouse whitefly on gerbera so the
weeks later introduce enough Encarsia may be considerably different from the plants were only marginally infested at
adults to parasitize at least 85% of the number promised by the insectary. Use the end of the experiment.
whitefly nymphs. Check in 2–3 weeks to quality parasites from reliable insec-
Most of the information on the use of
be sure that the wasps have parasitized taries. Large differences from the
Encarsia was obtained against green-
large numbers of pupae. A week or two expected numbers can drastically affect
house whitefly, so modifications may be
before introducing the banker plants control. Check each parasite shipment
necessary for control of other whiteflies.
into the crop, treat the plants with a for the number of emerging wasps, and
Encarsia parasitizes sweetpotato and sil-
nonpersistent insecticide, such as adjust release rates as necessary. To
verleaf whitefly, for example, but not as
pyrethrum, to kill any adult whiteflies. check wasp emergence, count exit holes
effectively as it does greenhouse
Place the banker plants uniformly on several cards at initial release and 1
whitefly. On sweetpotato whitefly,
throughout the greenhouse no more week later. Or, place several cards within
Encarsia reproduces less, takes longer to
than 20 ft apart. This provides a continu- airtight containers with transparent lids,
mature, and produces lower-quality off-
ous source of parasites emerging over place the containers out of direct
spring. The fact that the wasp is
8–10 weeks that will disperse to sunlight, and check the number of
produced commercially almost exclu-
whitefly-infested crop plants. When only adults that emerge after 1 week.
sively on greenhouse whitefly may
a few whiteflies are present in the crop, In general, for very low initial infesta- explain these problems. The rearing
the emerging parasites survive on the tions of less than one whitefly per plant, history of Encarsia can affect its accept-
whiteflies and honeydew present on the release rates of 1.5 parasite per 10 ft2 are ance on subsequent hosts. Also, there
bankers. The banker plants must be recommended. For higher initial infesta- are different strains of Encarsia that vary
established early in the growing season tions (still less than one whitefly per in their effectiveness against different
so that a large wasp population, up to upper leaf ), releases of 3–9 parasites per whitefly species. Double or triple release
10,000 per plant, has developed by the 10 ft2 should be made at 10–14 day rates and regular introductions through-
time whiteflies infest the crop. This may intervals. For greenhouse tomatoes and out the cropping season may be neces-
not be practical in large commercial peppers, weekly releases of one wasp sary to compensate for these problems.
greenhouses. per every four plants is suggested. For
The host plant also influences the ability
Whatever your introduction method, be cucumber the rate is one wasp per every
of Encarsia to control whiteflies. Large,
aware that excessive leaf removal can two plants, and on poinsettias, two
dense hairs (trichomes) can interfere
eliminate many parasites when they are wasps per plant, released weekly.
with parasites searching for prey and
about to emerge. Check the leaves for Release rates are better established for
reduce its efficiency. Biological control
black parasitized pupae. If there are many vegetable crops than for ornamentals,
with Encarsia may be improved by
black pupae that lack exit holes, leave the which have to be virtually pest-free to
selecting plant varieties with fewer hairs
prunings in the greenhouse for a week or be marketable. Encarsia can provide
or none. In one experiment, wasps were
two to allow time for the Encarsia to control comparable to conventional
able to parasitize 20% more whiteflies
emerge. If there are more white than insecticides on many ornamentals,
on a hairless cucumber than on a
black whitefly nymphs, discard the although a final “clean up” treatment
normal, hairy cucumber. By contrast,
prunings. Otherwise, you should remove with an insecticide may be necessary
certain host plants are particularly good
leaves regularly and frequently so that prior to sale. Initial whitefly infestation
hosts for whiteflies, making it harder for
large numbers of leaves with parasites on poinsettia usually occurs during
the wasp to keep up with whitefly pop-
are not removed all at once. plant propagation. For greenhouse
ulations. On plants such as cucumber
whitefly, release one parasite per plant
Recommended release rates vary con- and eggplant, whiteflies produce more
per week on the stock plants to achieve
siderably depending on the whitefly eggs, develop faster, and live longer
long-term control of a light infestation.
density, the time of year, the crop and its than they do on plants such as tomato.
Weekly introductions of 50 Encarsia per
growth stage, and the type of whitefly. Large populations on favorable host
150 newly potted poinsettia cuttings
Most suppliers provide detailed instruc- plants produce excessive honeydew
provide good control of a low infesta-
tions for the release and use of Encarsia excretion, which interferes with normal
tion of greenhouse whitefly. This rate
and can make recommendations about parasite searching behavior. To compen-
should also be suitable for fuchsia,
the number of wasps to release based sate for the better pest development on
gerbera, and other ornamentals. In one
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S W H I T E F L I E S 81
highly susceptible plants, introduce Recommended release rates are similar numbers, so it is particularly useful for
more parasites. For example, release to those for Encarsia; release two wasps release in whitefly hot spots. It may
rates of 14 wasps per plant are sug- per 15 ft2 every 1–2 weeks as a preventa- eradicate whiteflies if the population is
gested for eggplant, but only five per tive measure. When whitefly infestations not very dispersed. A release rate of
plant are needed for tomato. are established, a minimum of three seven per 10 ft2 is suggested.
Encarsia wasps fly and lay eggs best in weekly introductions at rates of 3–9 The mirid Macrolophus caliginosus is
warm (61°–82°F), bright greenhouses. wasps per 10 ft2 (depending on whitefly another good predator of whiteflies, par-
For maximum whitefly control, keep density) is suggested. ticularly when used in combination with
relative humidity at 50–70% and Predators of whiteflies can also be effec- other natural enemies. It remains active
daytime air temperature around 75°F. tive in biological control programs. Most at relatively low temperatures, and
Low temperatures allow the whitefly to of them do not feed exclusively on therefore can successfully supplement
multiply faster than the parasite, result- whiteflies, but these predators will control by other species that are active
ing in poor control. Cool, cloudy condi- provide control if whiteflies are the pre- only at higher temperatures. The bugs
tions and the low light levels of winter dominant pest. The lady beetle must be released early in the season
also reduce Encarsia’s ability to control Delphastus pusillus is the most whitefly- because of a slow population build-up.
whiteflies. Dutch researchers are devel- specific predator available, although it They are supplied as adults and nymphs
oping a strain that is more effective at will feed on spider mites if no whiteflies mixed with vermiculite. The mixture can
lower temperatures. Under cool condi- are available. It prefers high whitefly be shaken directly from the container
tions, you may need to increase release
rates or complement control using other
Whiteflies
natural enemies or a fungal pathogen
such as Beauveria bassiana. If you release Enemies egg —— nymph —— pupa adult
more wasps, you may also need to Encarsia formosa
decrease the distance between release
sites to compensate for slower repro- Encarsia pergandiella
duction and remove adult whiteflies in Encarsia inaron
the tops of plants by vacuuming the
Encarsia transvena
upper foliage or by hanging numerous
yellow sticky traps or ribbons. Whiteflies Eretmocerus spp.
are strongly attracted to yellow sticky
Chrysoperla spp.
traps, while the parasites are less
attracted—as long as sufficient whitefly Delphastus pusillus
nymphs are present. If you find more Lady beetles
than 15 whiteflies caught on yellow
traps in vegetable crops, consider Dicyphus tamaninii
spraying with a nonresidual insecticide. Macrolophus caliginosus
Many wasps other than Encarsia are also Orius spp.
good parasites of whiteflies. Eretmocerus
eremicus is better than Encarsia against Predatory mites
silverleaf whitefly because of its greater Aschersonia aleyrodis
mobility and ability to attack more
nymphs. This species is also less suscepti- Beauveria bassiana
ble to pesticides than Encarsia. It is often Metarhizium anisopliae
shipped as pupae mixed in a carrier, such
Paecilomyces
as bran or vermiculite. Sprinkle the
mixture into leaf axils or around the base fumosoroseus
of plants. Be sure that ants, which can Verticillium lecanii
carry off the pupae, are not present.
Natural enemy attacks the host Natural enemy host feeds as an adult
Width of bar indicates degree of effectiveness
82 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

onto leaves. Release 5–10 bugs per 10 in conjunction with Encarsia. Unlike par- ventional chemicals do not. It is compat-
ft2 into the first whitefly hot spots on a asites, fungi are able to control high ible with Encarsia, Eretmocerus, and the
weekly basis. Continue releasing until whitefly populations. Fungal applica- predator Delphastus pusillus and is
populations of M. caliginosus reach 0.5–2 tions may be useful in reducing whitefly tolerant of some fungicides.
per 10 ft2 throughout the greenhouse populations before introducing Encarsia Aschersonia aleyrodis requires several
(at least 2 months). Do not use this mirid or in conjunction with the parasite. applications in order to control white-
on gerbera because the bug can cause Under the appropriate conditions, fungi flies. This fungus is effective mainly
damage to the flowers. may eliminate the need for insecticides against the younger nymphal stages of
The green lacewings Chrysoperla carnea, to control whiteflies. However, the whitefly and does not affect Encarsia
C. rufilabris, and C. comanche are most B. bassiana must be used within an IPM formosa, which oviposits in old nymphs
efficient when leaves of adjacent plants framework because all fungi may be or pupae, so the two are very compati-
touch because the larvae are better able killed by certain fungicides used to ble for use together in greenhouses.
to spread among the plants. Releases of control plant diseases.
Fungi are applied as spores suspended
8–25 C. rufilabris larvae per plant con- The fungi Aschersonia aleyrodis, for spraying in water that contains a
trolled sweetpotato whitefly on hibiscus Metarhizium anisopliae, Paecilomyces wetting agent and other adjuvants. The
in Texas greenhouses—well enough fumosoroseus, and Verticillium lecanii are first diseased whiteflies appear 3–7 days
that all of the plants were marketable. promising biological control agents but after the spore application, but whitefly
However, lacewing development was are not currently available for use in the numbers do not decline until a week or
not normal, indicating that the white- United States. Whiteflies can be sup- two after spraying. High humidity is nec-
flies alone were not an adequate food pressed for several months following a essary for fungal sporulation and infec-
source. Orius spp. also provide some single application of V. lecanii if the tem- tion. When humidity is low, performance
control of whiteflies and other pests. perature remains between 59° and 77°F is unpredictable. You can increase
Many of these predators can be released and the humidity is greater than 90% for humidity by dampening the plants with
in conjunction with E. formosa. at least 10 hours per day. This fungus water sprays (be cautious; this may
Supplementary releases of green can kill 80–97% of the nymphs, and sub- encourage plant disease). Late afternoon
lacewings or the lady beetle Delphastus sequent infection kills many adults applications reduce spore injury by
pusillus will control whiteflies on many emerging from surviving nymphs. V. ultraviolet light and desiccation, since
crops, especially vegetables. These pred- lecanii is pathogenic to Encarsia when the greenhouse is relatively more humid
ators may feed on the same instars para- applied directly, but whiteflies are even at night. Humid areas with moderate
sitized by the wasp, but the beetles more susceptible, so the fungus and temperatures, such as rooting benches
avoid feeding on whiteflies in the late wasp can coexist. and shade-cloth covered areas used to
stages of parasitism. If the beetles are Experimental sprays of Paecilomyces induce inflorescence in chrysanthe-
released 2 weeks after the parasite, the fumosoroseus significantly reduced mums, may be the only practical areas
wasps should be developed enough sweet potato whitefly populations on for application. Multiple sprays are nec-
that beetles avoid them. Otherwise, you ornamentals (hibiscus, mandevilla, poin- essary for A. aleyrodis, B. bassiana, and P.
may wish to release these predators settia, and crossandra) when applied fumosoroseus, which will not spread in
only in areas of high whitefly numbers. weekly. P. fumosoroseus kills sweetpotato the greenhouse, and for V. lecanii if
The fungus Beauveria bassiana has been whitefly more quickly and with higher humidity is not high enough to allow
effective in controlling whiteflies under mortality than other fungi. Under continuous infection. These fungi are
a wide range of commercial production normal greenhouse conditions infection killed by many fungicides.
conditions. The fungus has controlled is detectable 7–10 days after applica-
whitefly on hibiscus, mandevilla, poin- tion. However, many of the whiteflies
settia, and tropical foliage plants in the killed by the fungus do not show the
greenhouse. It worked as well as or typical coloration unless placed in
better than the best conventional insec- humid conditions to allow sporulation.
ticide programs, even where infestations P. fumosoroseus also provides better
had reached critical levels. Beneficial control than conventional insecticides
insects are not affected unless sprayed because it kills all whitefly stages, includ-
directly, so this pathogen could be used ing adults and eggs, whereas most con-
Alternative control methods 83
Sanitation Plant selection Chemical control
This is the most important cultural Whiteflies prefer certain plant species, Localized infestations and high popula-
practice for preventing whitefly listed in the introduction to this tions that need to be reduced before
problems. Remove weeds, hanging section, so if you have a choice, avoid predator or parasite introduction
baskets, or “pet plants” that serve as growing these species. Within the should be spot sprayed.
alternate hosts for whiteflies from both same plant species there are also dif- Insecticidal soap can be used to
inside and outside the greenhouse. ferences in whitefly preference for and reduce whitefly populations with no
Whiteflies can easily move from an survival on distinct cultivars. For residual effects on natural enemies.
infested crop to an uninfested one. Be example, whiteflies prefer geranium Soap sprays are most effective against
cautious about introducing new plant cultivars with larger, less hairy leaves. nymphs and adults, but pupae are
material into greenhouses. Remove Poinsettia cultivars with white bracts killed as well. Whitefly eggs are
and immediately destroy crop residues are often more heavily infested than tolerant to soap sprays. Direct applica-
after harvest. Wait several days before those with pink or red bracts, but such tion to natural enemies will kill some,
bringing in new crops after an infested cultivar preferences are not consistent but once the soap dries it is nontoxic.
crop has been completely removed and may actually be based on plant Treat shortly before the initial release
from the greenhouse. To starve any odor rather than bract or foliage color. of parasites or predators. Thorough
remaining adult whiteflies, maintain Although all poinsettia cultivars are coverage of infested surfaces and
the greenhouse at production temper- readily infested, whitefly nymph several applications are necessary to
atures and free of whitefly hosts numbers vary consistently among suppress whiteflies. Follow label direc-
during this time. Hang strips of heavy certain cultivars. Some of the cultivars tions to avoid plant damage.
plastic in doorways between ranges to most susceptible to whiteflies are also
Horticultural oils can kill whiteflies and
restrict whitefly movement. Rogue out the most susceptible to infestation by
other insects, but they may be toxic to
severely infested plants. mealybugs and spider mites. Cultivar
some plants. Various brands are regis-
differences may also affect Encarsia
Insect screens tered for use on vegetables and orna-
performance.
Screens on vents and doors will help mentals in greenhouses to control
prevent whiteflies from moving into Fertilization many pests, including whiteflies.
the greenhouse from plants outside, Nutrient inputs can affect pests as well Azadirachtin is the active ingredient in
but often the most effective screens as the plant. Avoid over- or under-fer- the commercial formulation of extract
reduce air flow. To counteract air-flow tilizing, as whiteflies may develop from seeds of the neem tree
problems, the surface area of intake better on stressed or lush plants than (Azadirachta indica). Sold under various
vents can be expanded by building on normally fertilized plants. brand names, this natural compound is
screen boxes around the vents’ an insect growth regulator and repel-
Traps
openings. Screens must also be lent that has very low mammalian
Yellow sticky traps collect large
cleaned regularly. The maximum hole toxicity. It has provided good control
numbers of whiteflies and on vegeta-
size to exclude whiteflies is 462 µ. of sweetpotato whitefly when applied
bles effectively keep whitefly popula-
Repellents tions low when used from the begin- at 7-day intervals on poinsettia. It is
SunSpray 6E Plus horticultural oil ning of the crop. Canadian experi- registered for use on greenhouse veg-
showed some repellency against ments suggest one trap per 8–10 etables and ornamentals.
greenhouse whitefly for at least 11 tomato plants. In Hungarian green- Kinoprene is a synthetic insect growth
days after treatment on chrysanthe- houses, Encarsia was more effective regulator registered for use on orna-
mums. Adults landing on treated when introduced after using traps mentals for control of whiteflies and
plants quickly left, so fewer eggs were than when used alone. other insects. It is especially effective
laid on treated foliage. In another Whitefly movement within a green- as a spot treatment to control local
experiment, rooted poinsettia cuttings house with multiple crops may also be populations. It has minimal effects on
surrounded by white plastic mulch reduced by erecting plastic barriers, natural enemies but may damage
harbored significantly fewer green- perhaps coated with vegetable oil. some plants.
house whiteflies than plants with Such barriers around mature crops
red or black plastic mulch. during harvest will trap whiteflies
when they are disturbed.
84 Miscellaneous pests
S
lugs, symphylans, springtails, and 10 days or less, and the slugs mature in
sowbugs do not occur regularly in 3 months to a year, depending on the
Available natural enemies and
their potential for control greenhouses. They often move into species. Eggs are resistant to desiccation
greenhouses from outside during the and will persist in the soil for a long
warmer months and become a problem time.
only when populations are high or
Few effective natural enemies are when they contaminate plants for sale. Monitoring
available for slugs, symphylans, spring- In addition to the minor pests listed, Silvery slime trails indicate the presence
tails, and sowbugs. Cultural or other numerous other insect and other arthro- of slugs. Relative population levels can
controls can alleviate problems with pod pest species may infest green- be determined by counting the number
these minor greenhouse pests. houses and cause damage. These four of slugs that are attracted each night to
pests are among the most likely to shallow pans of beer placed at ground
Slugs level or boards placed on moist soil.
become nuisances on your greenhouse
Phylum Mollusca, Class Gastropoda crop. Growers should determine their own
Order Stylomatophora: Snails and threshold levels for initiating control
measures.
slugs
Family Limacidae: Slugs
Slugs
Slugs are not insects but mollusks, a Natural enemies
Symphylans group that includes snails, clams, Although there are many parasites,
Class Symphyla: Symphylans scallops, oysters, squid, and octopus. predators, and pathogens of slugs and
Slugs are omnivorous but prefer to eat snails, few have been investigated for
Family Scutigerellidae: Symphylans
vegetation, and they thrive in moist commercial control of these pests.
Garden symphylan, Scutigerella places like the humid greenhouse envi-
immaculata ronment. Several species of both native Predators
and imported slugs may cause damage Euthycera spp. The larvae of several
Springtails
in greenhouses on most vegetable and species of flies in the family Sciomyzidae
Order Collembola: Springtails are known to be slug-killers, but none
ornamental crops, especially orchids.
Various families has been investigated for use in green-
Sowbugs Damage houses. The European Euthycera cribata
Slugs are nocturnal feeders that eat lies in wait, poised for attack. When it
Class Crustacea: Crustaceans
seedlings or chew holes in succulent encounters a slug it rapidly attaches
Order Isopoda: Pillbugs and sowbugs itself with its mouth hooks, climbs onto
leaves, stems, or roots. During the day
Family Asellidae: Sowbugs they hide in dark, damp places beneath the prey and enters into the tissues. In a
benches, pots, or litter on the ground. few minutes it disappears almost com-
Like caterpillars, slugs damage plants by pletely, with only its breathing tube
rasping away plant tissue when young remaining visible. First-instar larvae feed
and by eating irregular holes when older. until the slug is dead, then move on to
The slime trails they leave on plants another prey; later instars continue
reduce the salability of ornamentals. feeding on the slug after it dies. This
species has only one generation per
Description and life cycle year. Two other European species of
Slugs range in length from 1⁄2 to 4 Euthycera also attack slugs. No Euthycera
inches (1.3 to 10 cm) and are usually species are commercially available.
gray or brown. They leave behind a char- Rumina decollata. This predatory snail
acteristic mucous trail as they move. The attacks, kills and consumes slugs,
hermaphroditic adults (individuals common brown snails, and garden snails.
have both male and female organs) lay It is a burrowing species, normally occur-
clusters of 20–100 gelatinous eggs in ring in the upper inch of the soil. It has a
moist soil crevices protected by debris brown, elongated spiral shell tapering to
or covered by containers. Eggs hatch in a blunt end. These snails prefer to feed
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S M I S C E L L A N E O U S P E S T S 85
on slugs and organic matter but will eat Pathogens use against slugs. The predatory
living vegetation if these foods are not Nematodes. The nematodes nematode Phasmarhabditis sp. that
available. It may take a few years for the Steinernema carpocapsae, S. feltiae, and attacks snails is commercially available
snails to provide control outdoors. Their Heterorhabditis bacteriophora will infect, in Europe. Research is underway to find
use in greenhouses has not been investi- kill and develop in slugs under labora- slug-parasitic nematodes that could be
gated but decollate snails are commer- tory conditions. However, other species commercialized in the United States.
cially available. They are legally sold only of nematodes may be more effective for
to residents of certain southern
California counties but might be
obtained with a letter from your county Alternative control methods
agricultural commissioner or similar local
Sanitation Hand picking
official stating that possession of the
Keep areas beneath benches dry and Removing all visible slugs by hand will
decollate in your county or state is not
clean. Remove plastic, boards, and have a noticeable effect on the popu-
illegal and that interstate shipments of
debris that may serve as hiding places lation, although this method would
decollates are permissible.
for slugs. Eliminate excess moisture. If not be practical on a large scale. Since
Staphylinus (=Ocypus) olens. This empty pots or flats must be stored slugs feed at night, that is the best
European rove beetle, often called the under benches, stack them on a clean, time to collect them.
devil’s coach horse in England, is a dry, wooden pallet on their sides, and
promising predator of slugs and of the Traps
keep them dry. Gravel under the
brown garden snail in California. Both Place shallow pans of beer or rubbing
benches allows for good drainage and
the 11⁄4-inch (3.2-cm) black adults and alcohol at ground level in the evening
is not a suitable habitat for slugs.
the black bristly larvae, which grow to 1 to attract slugs. They fall into the
Inspect all new plant material carefully liquid and die. You can also place
inch (2–2.5 cm) at maturity consume at
before bringing it into the green- boards or bricks in damp spots on the
least their weight in slugs or snail bodies
house. Quarantine any possibly soil. The slugs that collect under these
every day. They pupate in earthen cells
infested shipments to prevent con- can be smashed or killed with hot
in the soil. Little research has been con-
tamination of the entire range. water. Commercial traps are also avail-
ducted on this insect as a biological
control agent of slugs and it is not com- Barriers able.
mercially available. Slugs receive a mild shock when they Poison baits
Tetanocera spp. The larvae of the come in contact with copper, so they A bait that contains an extract of
widely distributed species of the usually do not cross copper bands. quackgrass, identified previously as a
sciomyzid fly, T. elata, uses fresh slime Tack or staple copper strips or bands slug-specific molluscicide, was very
trails to track slugs and immobilize their to the frames of raised beds or green- effective in field tests against the slug
prey with a toxic injection. The larvae are house benches, or wrap the copper Arion subfuscus, an important pest
host-specific as first-instar larvae, but around containers to prevent slugs species. However, at present no com-
this selectivity later disappears, and third from moving into the beds. Slugs are mercial product contains this material.
instars attack many different genera of less likely to cross gravel than they are
Metaldehyde or methiocarb pelleted
slugs and even snails. Each larva can kill moist organic soil, organic mulch, or
bran baits are commonly used to
four to nine slugs during its develop- soil overgrown with vegetation. To
control slugs, but these baits have
ment. There are two to three genera- capitalize on this, spread gravel
several drawbacks. They mold rapidly
tions per year. Three other North around the perimeter of the green-
and must be replaced frequently. Also,
American species of Tetanocera have the house to discourage entry.
some slug populations have devel-
same feeding method. This fly may be Repellents oped resistance to metaldehyde.
commercially available. A repellent called Snailproof, Predatory snails are susceptible to
composed of sawdust and shavings of poison baits, so you must quit using
the incense cedar, repels snails and baits at least 2 months before releas-
slugs. They do not like to travel over ing predatory snails.
this material but will if it becomes
packed down.
86 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Symphylans Monitoring
Examine the soil where infestations are
Pathogens
Several pathogens infect garden sym-
Symphylans are very active arthropods phylans and may cause up to 90% mor-
suspected. Symphylans tend to run away
found in damp soils rich in organic tality under optimal conditions. Neither
from the light when disturbed and
matter. They are closely related to cen- of the best pathogens—Entomophthora
escape quickly into the soil, so observe
tipedes and millipedes, so are not coronata and Metarhizium anisopliae—
carefully. Start control measures when
insects. They are often confused with has been developed for control of sym-
you find 10 or more symphylans on a
springtails, but symphylans are larger, phylans or other pests in greenhouses.
single root system. Symphylans in
have more legs, move faster, and do not
greenhouses tend to be more abundant Nematodes, such as Steinernema car-
jump. They tend to be found in moist
and destructive during the fall and pocapsae, will invade and kill active sym-
soils. Symphylans are general feeders
winter. phylans within 24 hours. Their use in
and may attack many vegetable and
ornamental crops. Garden symphylans controlling symphylans in greenhouses
Natural enemies has not been specifically investigated.
injure germinating seeds and seedlings
Several natural enemies attack garden (See “Fungus Gnats and Shore Flies” or
in particular, and they can be a serious
symphylans, including predatory mites, “Weevils” for more information on
problem on African violets when the
beetle larvae, centipedes, and diseases, nematodes.)
pots are placed in moist sand or soil on
but none has been used effectively in
the bench.
greenhouses.

Damage
These animals normally feed on algae,
Predators
Lamyctes spp. These centipedes are
Alternative
fungi, and decaying organic matter in
the soil and usually are no more than a
small, reddish-brown and asexual. In one
experiment, five adult centipedes each in
control methods
nuisance. However, any stage of sym- 4-inch pots quickly cut a symphylan pop- Sanitation
phylan may feed on sprouting seeds or ulation of 40 adults per pot in half, and Prevent symphylans from entering
plant roots. They prefer to eat root hairs greatly reduced symphylan injury to the the greenhouse on the roots of
or chew holes in larger roots and plants.The centipedes do not feed on plants or in soil. Symphylans
crowns. This causes wilting and a blue plants, even in the absence of prey.These commonly occur outdoors in com-
discoloration, and also encourages infec- and other centipedes, including Lithobius posted materials, such as manure
tion by disease organisms. Infested forficatus and L. bilabiatus, have also been piles and leaf mold. Steam sterilize
plants are stunted and do not respond reported as predators of symphylans but this material to kill symphylans and
to fertilization. are not available commercially. their eggs before using it in the
Pergamasus quisquiliarum. This preda- greenhouse.
Description and life cycle ceous mite was observed feeding on If possible, grow plants on raised
Symphylans are slender, white or symphylans in the field in Corvallis, beds to prevent pots from touching
translucent myriapods, 1⁄4–3⁄8 inch Oregon. In laboratory experiments each the soil. If crops are grown in ground
(6–8 mm) long with 15 body segments, mite consumed an average of seven beds, put solid bottoms in the
12 pairs of legs, and long antennae. symphylans during their 12-day devel- ground beds to permit steam sterili-
Females deposit clusters of up to 20 opment. Each adult female laid an zation of the soil mass.
eggs in the soil. They hatch in about 10 average of 33 eggs and consumed an
days. At hatching the immatures have average of 14 symphylans, although Cultivation
only 10 segments and six pairs of legs. they ate more at higher pest popula- Cultivate in-ground beds thoroughly
Each time the symphylan molts it adds a tions. P. quisquilarum has not been inves- to disrupt the symphylans’ movement
pair of legs until it has 12 pairs. The tigated for use in greenhouses and is through the soil.
immatures reach maturity in 3–6 not commercially available.
months under greenhouse conditions,
and the adults may live for 4 or more
years. Populations increase fastest at
about 75°F.
S E C T I O N 2 — P R A C T I C A L A P P R O A C H E S M I S C E L L A N E O U S P E S T S 87

Springtails Natural enemies


Many natural enemies attack springtails
Alternative
Springtails are small, jumpy arthropods
in natural ecosystems. Most are oppor-
that are similar to insects. When dis-
turbed, they use a forked structure on
tunistic general predators, but a few are
specific to springtails or have been
control methods
the bottom of the abdomen to jump
observed attacking springtails in green- Sanitation
3⁄ 4–2 inches. They are common and
houses. Prevent springtails from entering into
abundant in the litter layer of natural
the greenhouse on the roots of
ecosystems. In greenhouses, a few Predators plants or in soil. Steam sterilize soil to
species cause problems occasionally on Hypoaspis (=Geolaelaps) spp. These
kill springtails and their eggs before
seedlings and young plants. They are tiny mites feed upon small, soil inhabit-
using it in the greenhouse.
unlikely to be a problem for plants ing insects, mites, and all stages of
grown in a soilless medium. springtails. They are aggressive preda-

Damage
tors, consuming 15–30 prey daily, killing
more small prey than large prey. H. miles
Sowbugs
Sowbugs are crustaceans, related more
Springtails feed on decaying matter, and a Geolaelaps sp. are offered com-
closely to crayfish and crabs than to
algae, and fungi and are normally only a mercially, although little is known about
insects. They are common throughout
nuisance. Some species, however, will their ability to control springtails in
North America, although they came
feed on living plants. They chew greenhouses.
from Europe originally. When disturbed
pinholes in seedlings and young plants Other predators. It is hard for active they tend to curl up into a ball. They are
or scrape their foliage. Springtails may predators to catch springtails because mainly scavengers feeding on decaying
also feed on roots, causing plants to wilt they jump when they are touched. vegetation, rotting wood, manure, and
and increasing susceptibility to plant However, three genera of ground beetles soil arthropods. Sowbugs also feed on
pathogens. from the Northern Hemisphere (Loricera, the roots and leaves of plants. They will
Leistus, and Notiophilus) feed specifically feed on almost all greenhouse plants,
Description and life cycle on springtails. Some other arthropods, but cause problems especially on
Springtails are small and wingless. They such as spiders, mites, and pseudoscorpi- orchids.
vary in color, but most are white. ons, are known springtail predators.
Females lay up to 120 smooth, spherical,
A grower in Finland found that Damage
cream-colored eggs in small groups in
numerous Pardosa amentata spiders Sowbugs are nocturnal feeders that
the soil. The immatures resemble the
were living on springtails which were hide during the day under pots and
adults but are smaller. They grow
very abundant in the peat used as a debris, in damp, sheltered places. They
through six to eight instars (depending
growing substrate for the lettuce crop in collect in pots or beds and feed on the
on the species) in about 11⁄2 months
the greenhouse. fibrous roots of plants. Their feeding
and attain sexual maturity before they
may retard plant growth. Sowbugs also
reach maximum size. Springtails tend to None of these predators is commercially
may chew through the stems of
mass together in enormous numbers. available or has been investigated for
seedlings at or below soil level, and they
use in greenhouses.
occasionally chew holes in foliage.
Monitoring Pathogens
Large populations of springtails may be
The nematodes Steinernema feltiae
visible on the soil surface. They can also
(=Neoaplectana bibionis) and
be detected by floating the soil or sub-
Heterorhabditis bacteriophora (=helioth-
merging potted plants in a bucket of
idis) infected one species of springtail in
water. The springtails will come out of
laboratory experiments. Because spring-
the soil and be visible on the surface of
tails live in the soil, there is good poten-
the water.
tial for nematodes to infect them. These
nematodes are commercially available.
(See “Fungus Gnats and Shore Flies” or
“Weevils” for more information on
nematodes.)
88 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Description and life cycle Monitoring Nematodes, such as Steinernema spp.


Sowbugs are gray, oval, flat crustaceans Observe sowbugs at night or trap them and Heterorhabditis spp., will infect and
about 1⁄2 inch (14 mm) long. The body is to determine relative population levels. kill sowbugs in laboratory experiments.
covered with a series of overlapping See “Alternative Control Methods” below Because sowbugs live in the soil, nema-
armored plates, and seven pairs of legs for trapping instructions. todes may be able to control them.
attach to the underside. Females carry However, most nematodes prefer insects
broods of 25–75 in a pouch on the Natural enemies to sowbugs because the body disinte-
abdomen for 1–2 months. The young are The sowbugs’ heavy exoskeleton grates quickly, restricting nematode
released from the maternal brood protects them from natural enemies. development. Several species of nema-
pouch about 7 days after the eggs Only a few carabid beetles are known todes that may infect sowbugs are com-
hatch. The young, which are similar to predators of sowbugs, and a few species mercially available.
the adults in appearance, take about a of tachinid flies have been found to par-
year to reach maturity. They may live asitize these isopods in the United Possibilities for effective
several years and several generations States. But no parasites, predators, or biological control
may occur each year in greenhouses. pathogens have been used to control Most of these miscellaneous pest do not
sowbugs outdoors or in greenhouses. offer many possibilities for biological
Desiccation seems to be the most control. Although some natural enemies
important cause of death for outdoor have been identified, none has been
populations. researched for implementation in green-
houses and most are not commercially
available. Sanitation and other cultural
controls are probably the best methods
Alternative control methods for dealing with these pests.
Where allowed, predatory snails might
Sanitation Baits be used for the control of slugs. These
Use sterile soil and take precautions Sowbugs are attracted to sweet baits, molluscs have provided very effective
to prevent sowbugs from being intro- such as brown sugar bait and bran biological control of brown garden snail
duced into the greenhouse. For baits. Brown sugar bait is simply outdoors in southern California, but
example, check pots brought in from brown sugar mixed with an insecti- their use in greenhouses has not been
outside. Also, remove objects that can cide. An old recipe for making a bran explored. Because they will feed on
serve as hiding places for sowbugs bait requires moistening 12 pounds living plants in the absence of slugs or
during the day. of bran with a mixture of 2 pints organic matter, they may not be a good
molasses, 1 ounce banana oil (amyl choice for use in greenhouses.
Traps
acetate), and an insecticide, then
Scatter wood or bark chips on the soil The general mite predator Hypoaspis
placing all these ingredients in a
surface to provide hiding places for miles, discussed in “Fungus Gnats and
gallon of water.
the sowbugs. For traps you might Shore Flies,” feeds on many soil-dwelling
also use inverted, hollowed potato Chemical control pests and might provide some control
halves or damp pots left on the Pyrethrum dusts or diatomaceous of springtails or symphylans.
ground. The sowbugs will collect earth applied to beds, walkways, and Nematodes are often effective for the
under these traps, then you can crush other surfaces may be effective control of pests in soil and have poten-
them or kill them with hot water. against sowbugs. tial for controlling symphylans, spring-
tails, and sowbugs. Nematodes have not
been investigated specifically for control-
ling these pests in greenhouses, so direc-
tions for their use and suggested appli-
cation rates have not been developed.
SECTION

3
Overview of biological control
of greenhouse pests
Contemporary Increasingly, greenhouse growers
around the world are replacing pesti-
biological control cides with biologically based controls.
Why? Biological control offers a way to

B
iological control of insects and
control pests that are resistant to insec-
mites in greenhouse crops began
ticides, may reduce the cost of insect
in England during the late 1920s
control, often achieves better control of
when Encarsia formosa was used to
the pests, and usually increases yields by
control greenhouse whitefly on
eliminating the subtle deleterious
tomatoes. The use of Encarsia stopped,
effects of repeated applications of
however, after the development of syn-
chemicals. In addition, public demand
thetic organic pesticides in the 1940s.
for pesticide-free produce is growing
Using natural enemies for pest control in
and governmental regulation is
greenhouses became popular again in
reducing the options for agricultural
the 1960s when twospotted spider mite
pesticides. Pesticides are still the main
populations in European greenhouses
means of insect control in most U.S.
became resistant to many pesticides
greenhouses, but the loss of effective
and devastated cucumber crops.
chemicals through regulation, increas-
Growers introduced predatory mites to
ing pest resistance, and the escalating
control the spider mites, and because
cost of developing new pesticides is
pesticide use had to be limited, Encarsia
encouraging greater adoption of biolog-
was again utilized for whitefly control on
ical control methods.
cucumbers and tomatoes. Researchers
developed most of the early techniques Integrated pest management is now
necessary for biological control in used in many European and Canadian
greenhouses at the Glasshouse Crops commercial vegetable greenhouses.
Research Institute in Littlehampton, Biological control has been very suc-
England and the Research Station for cessful on vegetable crops, such as
Vegetables and Fruit Under Glass in tomatoes and cucumbers, because these
Naaldwijk, The Netherlands. Today, crops can tolerate some insect or mite
researchers in North America also play damage without affecting yield.
an important role in developing and Vegetable growers also use biological
implementing biological control strate- control because it is easier to maintain
gies for greenhouse production. harvest schedules without pesticide re-
entry restrictions. Bumble bees can be
used for more cost-effective pollination
when pesticides are not used, and
produce without pesticide residues may
be more marketable or sold for higher
prices.
90 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

However, certain obstacles must be developed for biological control of Some of the best candidates of floricul-
overcome for the continued adoption of certain pests on many ornamentals. tural crops for biological control are
biological control in greenhouses. The Multicropping in the same greenhouse rose, alstroemeria, chrysanthemum and
greenhouse industry is shifting from also presents a challenge: maintaining gerbera for fresh cut-flowers, plus poin-
seasonal production to nearly year- biological control on several different settia and numerous foliage plants.
round production in the same green- plant species simultaneously. Certain These plants are attacked by a limited
houses. This may influence the effective- plant production programs, or crop number of pest species or can tolerate
ness of biological control. There is no mixes within a greenhouse, may prohibit some pest injury, although aphids and
longer a host-free period in which pests or severely restrict biological control. thrips can be difficult to control in flori-
starve to death, so newly introduced Among the biggest obstacles to the cultural crops. Biological control is most
plant material may become infested implementation of biological control in likely to succeed on cut flowers, such as
almost immediately. Natural enemies greenhouses are the negative attitudes chrysanthemums or gerbera daisies,
may move from one crop to the next of some growers or the unrealistic because only the top portion of the
along with the pests, but there will likely expectations of others. Many growers plant or flower stalk is harvested. About
be a lag time and hence some damage are unwilling to invest adequate time to the top 32 inches of the chrysanthe-
to the new crop. Worse, some of the learn how to succeed with biological mum plant is marketed and the bottom
available natural enemies enter control. It takes a fair amount of time 5–16 inches of each stem is stripped of
diapause under winter greenhouse con- and effort to convert from reliance on all leaves. Damage to the lower portion
ditions—short days and cooler nights— pesticides to using natural enemies. of the plant when young (the first 4–6
and will not provide adequate control Every situation is unique, and specific weeks of plant growth) does not affect
during this time. To prevent natural protocols must be developed for each the marketable portion when harvested.
enemies from entering diapause, you greenhouse if biological control is to be With gerbera daisies grown for cut
can increase night temperatures or effective. Experimentation may be flowers, only flower stalks that totally
extend daylength by using artificial required to adapt recommended release lack foliage are harvested.
lighting. Increasing release rates, using rates and timing to each situation. Even for crops where biological control
nondiapausing natural enemies or inte- Sometimes growers’ first attempts to of foliage and flower pests is difficult,
grating selective pesticides can also implement biological control fail. But there exist excellent opportunities for
help to control pests during this period. these failures often lead to a greater biological control of soil pests such as
Biological control may not be practical understanding of how to utilize natural fungus gnat larvae.
in the production of certain ornamental enemies successfully—if the grower Permanent plant collections, such as
plants or in multicropping. On ornamen- sticks to it. conservatories for public display or hor-
tals the presence of any insects or mites, Some growers believe that pesticides ticultural greenhouses for research or
whether pests or natural enemies, are the only way to provide sufficient educational purposes, are also excellent
reduces the aesthetic value and salabil- control for high plant quality. Certainly places for implementing biological
ity of a plant. An IPM or biological there are excellent pesticides and appli- control. Marketability of conservatory
control program on ornamentals must cation methods that can sometimes plants is not a factor, so tolerance for
be modified to eliminate both the pests provide virtually pest-free plants. injury may be higher.
and the natural enemies prior to sale. In However, even ornamental crops
addition, since the entire ornamental produced under conventional chemical
plant is often sold, it can tolerate very control—particularly those seen in
little vegetative damage. Control mass-marketing outlets—are not pest-
measures must reduce pest populations free. Biological control can be used
before they damage the marketable during at least a portion of the produc-
portion of the crop. But there is usually a tion of many ornamental plants to
delay between natural enemy introduc- produce plants that are equal to those
tion and the reduction of pest popula- produced with chemical pesticides.
tions, during which plant damage can Pesticides may be necessary just for the
occur. Effective commercial natural final clean-up.
enemies may not be available and
specific recommendations may not be
S E C T I O N 3 — O V E R V I E W 91

Biological control in the hobby greenhouse


A
great diversity of plants are The collection of plant species grown Another potential problem with small
grown in hobby greenhouses, in a hobby greenhouse might be very greenhouses is that occasionally it may
and these plants are equally as specialized, such as orchids, cactus, or be necessary to control a pest species
susceptible to pest attack as crops in palms, or a bit more generalized, such for which there are no acceptably
commercial greenhouses. Often, the as tropicals from a variety of plant effective natural enemies. For some
pest species present in commercial families, or very eclectic, including pests, the only alternative may be a
and hobby greenhouses are the same. both ornamental and food plants. broad-spectrum house plant insecti-
Many of these are general feeders, Some hobby greenhouses are used cide available from the local garden
such as greenhouse whitefly, aphids, primarily to start bedding or vegetable store. If you are using biological
mealybugs, and twospotted spider plants to be set out into the garden in controls, use such insecticides as care-
mite. The natural enemies that are spring. The nature of the plant collec- fully as possible because they will also
commercially available can be used in tion may impact the success of biolog- harm the natural enemies. If possible,
the hobby greenhouse, often quite ical control. More specialized pests remove those plants that need to be
successfully. may occur in specialized plant collec- treated from the greenhouse and
tions, and these pests may not have don’t return them to the greenhouse
However, the hobby greenhouse may
been researched for their biological for a few days after treatment to allow
present some logistical problems
control potential, or appropriate residues to dissipate. If it is not
when biological control is attempted.
natural enemies may not be commer- possible to remove plants for treat-
Many natural enemies require protec-
cially available. Therefore, some pests ment, try to leave protected refuges
tion from heat, intense light, and low
may require other approaches that can for natural enemies by treating only
humidity. The small hobby greenhouse
vary from hand removal to the use of those plants that are infested. Even
may not have sophisticated environ-
conventional pesticides. As with pests using this approach, it may be neces-
mental controls found in commercial
in any crop or situation, the first step in sary to make new natural enemy
greenhouses, and therefore natural
determining control options is to cor- releases starting a few weeks after the
enemies may experience unfavorable
rectly identify the pest. Your local pesticide application.
or even lethal conditions. At the
county or state university Extension As with all types of biological control,
extreme are hobby greenhouses that
service should be able to help in this no easy recipes exist for success in the
are set up specifically for growing cacti
regard. Once the pest is identified, hobby greenhouse. However, there are
and succulent plants; these may have a
contact a commercial natural enemy often many useful natural enemies
lower humidity and higher light inten-
supplier to determine the availability available for release, and the level of
sity, and possibly higher temperatures
of effective natural enemies. Keep in success can be high. The best way to
than are tolerated by many natural
mind that for certain more obscure determine the potential for success in
enemies. Under such conditions,
pests, even though specialized natural your own greenhouse is to learn about
certain natural enemies may not be
enemies may not be available, general- the options, discuss possibilities with
reliably effective and other controls
ist natural enemies such as lacewings, commercial natural enemy suppliers,
may be necessary. Smaller green-
pirate bugs, or insect pathogens may and give it a try.
houses also may not provide the diver-
provide some control.
sity of habitats and nutritional
resources (such as nectar and pollen)
necessary to maintain natural enemies
over an extended time period; there-
fore more frequent releases may be
necessary to achieve effective control.
92 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Integration of The intensive labor associated with


pruning plants provides growers with
Chrysanthemum
Management of greenhouse cut and
biological control frequent opportunities to monitor for
pests before they become widespread.
potted chrysanthemums is most suc-
cessful when the specific standards for
of all pests: Examples When small infestations of other pests
such as aphids and spider mites are
each of these crops is considered. Cut
chrysanthemums are managed to
Tomatoes found, they can be controlled by either produce flowers, whereas potted
The production of fresh-market physically removing the pests, or by chrysanthemums are managed for both
tomatoes poses a considerable chal- spot-spraying pesticides with low the foliage and flowers. Consequently,
lenge to greenhouse producers inter- residual toxicity then releasing their managers of cut chrysanthemums are
ested in using biological control natural enemies. primarily concerned with insects such as
because of the industry’s high cosmetic Disease problems such as leaf mold and western flower thrips that damage
standards. Whiteflies are the worst pest Botrytis blights can occur under humid flower buds and open flowers. Insects
for many growers. They reduce fruit size greenhouse conditions. These can be that feed on flowers reduce the crop
and coat fruit with honeydew, which controlled by increasing air movement quality directly through feeding, or indi-
serves as a medium for the growth of and by applying a fungicide that does rectly, by opening wounds that allow
black sooty mold fungus. At present, the not kill Encarsia wasps. Smoke formula- disease organisms, such as Botrytis, to
greenhouse whitefly is the most tions of chlorothalonil have been used enter. In contrast, managers of potted
commonly encountered problem on with some success. chrysanthemums need to protect the
tomatoes in many northern and central foliage and flowers. Insects can damage
Greenhouse tomato producers who
U.S. greenhouses. Sweetpotato whitefly foliage directly by removing tissue, as
have used biological control enjoy more
is not found frequently. caterpillars do, by distorting their shape,
than just the benefits of high-quality
Dribble (inoculative) releases of the par- as thrips do, or by discoloring them, as
produce. They reduce their labor greatly
asitic wasp Encarsia formosa have been leafminers and thrips do. Indirect
by employing bumble bees rather than
used successfully by many tomato damage to foliage occurs when sucking
human workers to pollinate the
growers to control greenhouse whitefly. insects, such as aphids and whiteflies,
tomatoes. In the absence of pesticide
This regular release of small numbers of cover foliage with honeydew that
residues, colonies of Bombus impatiens
wasps throughout the growing season fosters growth of black sooty mold.
and B. occidentalis can survive in green-
allows natural enemy populations to houses and pollinate tomato flowers. Biological control of chrysanthemum
build before whiteflies get out of Instructions for their use vary by region pests will be most successful when
control. The success of this method is and are available from some suppliers of combined with proper cultural practices,
partly due to the fact that growers are Encarsia. a regular monitoring and record-
starting plants in winter when green-
houses are closed. This allows an
extended period of plant growth prior
to the entrance of pests from outdoors
during spring and summer. However,
this requires the use of transplants that
are completely free of whiteflies.
S E C T I O N 3 — O V E R V I E W 93
keeping program, and early releases of Poinsettia A good pest monitoring routine will
natural enemies. You can reduce the Many growers are interested in integrat- detect potential problems before they
flow of new pests into the greenhouse ing biological control into their poinset- develop into severe problems. Establish
by purchasing quality chrysanthemum tia IPM program. The primary insect and test a monitoring program for at
cuttings from reputable suppliers that pests of poinsettia are greenhouse least one poinsettia crop before begin-
have been checked for insects. Removal whiteflies, sweetpotato whiteflies, silver- ning your biological control program.
of weeds, plant debris, and old blooms is leaf whiteflies and fungus gnats. Once you start the biological control
critical, because these act as reservoirs Whiteflies are
for insects, which can migrate onto the pests primarily
main crop. Screens on greenhouse side because their
vents, ridge vents, and side walls prevent presence
migrating insects from entering the diminishes the
greenhouse in the summer. appearance of
Monitoring techniques should be imple- the plant.
mented along with biological control. Fungus gnats
Growers must inspect both flowers and are nuisance
foliage weekly to determine the pests as
numbers of pests and natural enemies. adults, but the
In addition, yellow sticky traps should be larvae can
examined on a weekly basis. When damage the
recording the number of pests and roots of the
natural enemies, be sure to note the plant directly
temperature, month, time of day, and are
numbers caught per sticky trap, and capable of
location of plants and traps. transmitting
certain plant pathogens. Both whiteflies program, keep track of what your para-
All the major insect pests of cut and
and fungus gnats can be managed with sites are doing. For example, record
potted chrysanthemums, including
biological control. numbers of adult parasites and dead
aphids, caterpillars, leafminers, western
and parasitized whitefly nymphs on the
flower thrips, fungus gnats, and white- Successful whitefly biological control
plants in the scouting reports.
flies, can be controlled by a variety of programs are built on a foundation of
natural enemies described in earlier sanitation, regular monitoring, and early Begin biological control with a clean
chapters.The application of natural releases of natural enemies.The natural greenhouse planted only with poinset-
enemies in cut and potted chrysanthe- enemy used most commonly to control tia. The greenhouse should be inspected
mum production should follow an inocu- whiteflies is the parasitic wasp Encarsia for sources of whiteflies well before the
lative release program.This involves formosa. Another whitefly parasite, poinsettias are introduced. Any other
periodic releases before pest populations Eretmocerus eremicus, which is available plants, including weeds or hanging
build up. Order natural enemies from a commercially, may be more useful than E. baskets, should be eliminated. Inspect
reliable supplier at least 3 weeks in formosa against silverleaf whitefly. Fungal the incoming cuttings for whitefly adults
advance of the date when pests have pathogens, such as Beauveria bassiana, and nymphs. If necessary, reduce
appeared in previous years. Release the can also be used for whitefly control. whitefly numbers with pesticides that
natural enemies when insect pests are Growers must rely primarily on releases do not leave a residue that is harmful to
first detected.To control leafminers on of these or other natural enemies for the natural enemies. Parasites and pred-
chrysanthemum, for example, make early control, and use insecticides only as a ators will not be able to control white-
and weekly releases of the parasite back-up. A typical spray program for flies adequately if the initial infestation
Diglyphus isaea.This improves the whiteflies and fungus gnats will not be is too severe.
chances of delivering sufficient live compatible with predators or parasites.
natural enemies to the crop at the appro- Plan an initial small-scale trial for the first
priate time. Late releases are ineffective. season to gain familiarity with the natural
enemies before implementing biological
control on a large scale.
94 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

Release one to five Encarsia formosa per Summary


plant per week beginning soon after
planting the crop. Hang the cards con-
Biological control can be a highly effec-
tive means of managing many of the
Additional reading
taining parasite pupae on foliage near pests that attack greenhouse crops,
Costello, R. A., D. P. Elliot, L. A. Gilkeson,
the base of the plant canopy but not on although biological management
and D. R. Gillespie. 1992.
the soil, distributing the cards evenly programs have not been developed yet
Integrated control of greenhouse
throughout the greenhouse. When hot for all crops. Biological control is not
pests. British Columbia Ministry
spots of whiteflies are detected during based on recipes containing set
of Agriculture, Fisheries and
routine monitoring, be prepared to responses to particular pest problems.
Food. 19 pp.
release the predatory beetle Delphastus Many variables contribute to the effec-
pusillus or spot treat with compatible tiveness of biological control, and every Hunter, C.D. 1997. Suppliers of benefi-
insecticides. greenhouse presents a unique combina- cial organisms in North America.
tion of these variables. Therefore, California EPA, Department of
Fungus gnats can be controlled easily
growers must become familiar with the Pesticide Regulation,
with natural enemies. If released at the
specific conditions of their operations. Environmental Monitoring and
start of the crop, the soil-dwelling,
What problems have occurred in the Pest Management. 31 pp.
predaceous mite Hypoaspis miles can
past? How much damage can be toler- (Revised periodically.)
become established in the growing
media and provide season-long larval ated before the market value of the crop Mahr, D. L. and N. M. Ridgway. 1993.
control. Drenches of Bacillus thuringien- begins to drop? Are natural enemies Biological control of insects and
sis var. israelensis (Gnatrol) can be commercially available for controlling mites: An introduction to benefi-
applied for additional larval control, and the pests? These are just a few of the cial natural enemies and their use
entomogenous nematodes are also questions to consider as you plan your in pest management (NCR 481).
available. biological control system. Consider all Madison: University of
the options and evaluate whether they Wisconsin-Extension. 91 pp.
Many growers report successful use of
are compatible with each other and Powell, C. C. and R. K. Lindquist. 1997.
biological control on their stock plants
with your chosen control practices. The Ball pest & disease manual:
and on the blooming crop until late
best approach may be to coordinate Disease, insect and mite control on
October, when smokes and aerosols are
several control methods rather than flower and foliage crops. Ball
used for final clean-up. These growers
relying on a single solution. Publishing, Geneva, IL. 332 pp.
often comment that plants in biological
control greenhouses look healthier and Weinzierl, R. and T. Henn. 1991.
more attractive than those in regularly Alternatives in insect manage-
sprayed greenhouses. ment: Biological and biorational
approaches (NCR 401). Urbana:
University of Illinois-Extension.
73 pp.
95

Considerations when ordering and using commercially available


natural enemies for biological control
Before ordering ■ Provide your supplier with a safe ■ Attempt to assess the condition of
■ Know the specific pests that you delivery address, where the shipment the natural enemies. This is easier to
need to control. Identifying the pest will be cared for as soon as it arrives do with larger insects shipped in
as simply “whitefly” or “aphid” is not and where it will not be exposed to mobile stages than with very tiny
sufficient because different species of temperature extremes. insects or mites, or with insects
these pests require different natural ■ Understand proper release practices shipped as inactive eggs or pupae.
enemies. so that you will be prepared to make Mobile natural enemies should not
the release as soon as the shipment be stressed. It’s not unusual for a few
■ Determine the general level of infes-
arrives. to die during shipping, but most
tation. If it is very high, it may be nec-
should be in good condition.
essary to use other methods to ■ Know proper storage conditions in
reduce the pest population to allow case release can not be made imme- ■ Make release as soon as possible
biological control a better chance to diately after arrival. after receipt.
be effective. ■ Store under proper conditions if it is
■ Be sure all greenhouse vents are ade-
■ Know the best natural enemies avail- quately screened to prevent escape necessary to delay release. In many
able for the specific pests. In some of the released natural enemies cases, predatory and parasitic insects
cases, combinations of natural (important for most insects, but not can be stored only a few days before
enemies may work better than indi- for predatory mites). they start to lose vigor and effective-
vidual species. ness.
■ Do not apply broad-spectrum insec-
■ Know the proper timing of release of ticides for at least 2-3 weeks prior to ■ Make releases based upon recom-
natural enemies, based on the life an intended release of natural mendations of your supplier. Releases
cycles of both the pest and the enemies. should generally be made during a
predator or parasite. Often in green- cooler part of the day.
■ If you have questions about any of
houses there may be overlapping ■ As you are making releases, check
these matters, be sure to discuss
and continuous generations of the once again for quality characteristics.
them with your supplier. If you are
pest, resulting in the presence of the
just starting a biological control
susceptible stage in its life cycle at all
program, contact several suppliers After release
times. ■ Check during the following few days
initially and pick one that seems
■ Know the proper release rate for competent and professional in to see if the natural enemies appear
each natural enemy. dealing with your specific pest to be active and searching for the
problems. pests.
■ Calculate the amount of natural
enemies needed, based on the ■ Attempt to evaluate the impact by
release rate, the level of infestation, When the shipment arrives monitoring changes in the pest pop-
and the area to be covered. ■ Minimize exposure to hot or cold ulation and by checking for evidence
temperatures. of predation or parasitism (such as
■ If multiple releases are necessary,
■ Inspect the shipping container and mummified aphids or parasite
know the recommended frequency
contents for damage. cocoons).
of release. A program of multiple
shipments can be arranged with a ■ Determine if you received the species ■ Keep records of your releases and
call to your supplier. and quantities you ordered. their results. Biological control
programs sometimes need modifica-
tion and adjustment and your
previous records will be essential in
this process.
96 Glossary
Abiotic control. Those natural environ- Conservation of natural enemies. One Host-feeding. A form of predation.
mental factors that help control the of the three general approaches to Many adult parasitic wasps feed from
numbers of a pest population that do biological control. It is the provision the same types of hosts as they
not involve living organisms or life of food, shelter, and other needs for deposit eggs in. Although such wasps
processes. Weather events that kill natural enemies, and the avoidance are usually described as parasites or
many insects, such as severely cold of practices that kill natural enemies parasitoids, these terms properly
winters and heavy rains, are good or interfere with their beneficial refer to the feeding method of the
examples of abiotic controls. activities, such as the use of broad- larval stage. The adult wasps may also
Action threshold. A level of pest popu- spectrum insecticides. be beneficial by preying on other
lation at which controls should be Cosmopolitan. Occurring throughout individuals of the same species used
applied to prevent economic most of the world. as hosts for their offspring.
damage to the crop. See also Diapause. A period of prolonged inac- Hyperparasite. A parasite whose host is
economic injury level (EIL); economic tivity in insects. another parasite.
threshold (ET). Importation of natural enemies. One
Economic injury level (EIL). The popu-
Augmentation of natural enemies. lation level at which pests cause of the three general approaches to
One of the three general approaches economic damage if left untreated. biological control. Undertaken prima-
to biological control. It is the periodic rily by the U.S. Department of
Economic threshold (ET). The pest
release of captured or commercially Agriculture, universities, and state
population level at which control
produced natural enemies to supple- departments of agriculture, importa-
measures should be initiated to
ment those that occur naturally; it tion involves seeking natural enemies
prevent the population from exceed-
increases the effectiveness of biologi- in the native home of the pest, and
ing the economic injury level (EIL).
cal control. Also called augmentative introducing and permanently estab-
biological control. See also inocula- Ectoparasite. A parasite that feeds and lishing these natural enemies in the
tion; inundation. develops on the outside of its host. pest’s present habitat. This approxi-
Banker plant. An alternate host plant Endoparasite. A parasite that feeds and mates permanent natural control.
used to raise hosts and natural develops inside its host. Inoculation. A preventive method of
enemies in large numbers in a green- Entomopathogenic. Capable of augmentative biological control in
house for later movement into the causing disease in insects. which relatively small numbers of
crop. natural enemies are released periodi-
Exoskeleton. The external skeleton of
Biological control. The management of cally for sustained management of
an insect, composed of hard cuticle
pest populations by the purposeful the pest population below damaging
(the “skin”).
manipulation of beneficial organisms levels. Also called inoculative release.
Facultative. Organisms that normally See also augmentation of natural
called natural enemies. See also aug-
are free-living but have the ability to enemies; inundation.
mentation of natural enemies; con-
adapt to a parasitic or semi-parasitic
servation of natural enemies; impor- Insectary. A facility for rearing insects.
mode of life.
tation of natural enemies; natural The term is often used in biological
enemies. Frass. Fecal material and food fragments control for companies that mass-
produced by an insect in feeding. produce beneficial predators and
Cocoon. A silken case formed by an
insect larva as protection for Gregarious parasite. A species in which parasitic insects for release in aug-
pupation. numerous immature individuals mentative biological control.
develop within a single host. Instar. The stage of an insect between
Complete metamorphosis. Type of
insect development in which the Hermaphroditic. Having both male and successive molts.
insect passes through a pupal stage female reproductive organs; the
before becoming an adult. nature of an individual possessing
both ovaries and testes.
97
Integrated pest management (IPM). Nymph. The immature form of an insect Pheromone. A chemical substance
The use of all available and appropri- that undergoes simple metamorpho- secreted by an organism that causes
ate methods to control pest popula- sis. Between hatching and the a specific reaction by other individu-
tions in an effective, economic, and winged adult stage. Compare with als of the same species. Often used in
environmentally sound manner. larva. traps to lure insects for pest monitor-
Inundation. A curative method of Oviposit. To lay or deposit eggs. ing purposes.
natural enemy augmentation that Ovipositor. The egg-laying structure of Phloem. Plant tissue which transports
uses large-scale releases of natural a female insect. food over long distances within
enemies for the immediate reduction vascular plants.
Parasite. An organism that derives its
of pest populations that are at or near Proleg. A fleshy, unsegmented abdomi-
food from the body of another
damaging levels. Also called inunda- nal leg of caterpillars.
organism (the host). A parasitic insect
tive release. See also augmentation of
spends its immature stages in or on Pupa (pupae). A nonfeeding, inactive
natural enemies; inoculation.
the body of a host, which dies just stage during which an insect
Larva (larvae). The immature form of before the parasite pupates. See par- changes from a larva to an adult.
an insect that undergoes complete asitoid. Puparium (puparia). A protective case
metamorphosis; the stage between
Parasitoid. An insect that parasitizes created by the hardening of the
the egg and pupa. Compare with
and kills other insects. Many biologi- larval skin in which the pupa
nymph.
cal control workers prefer this term develops. Produced by flies.
Microbial insecticide. A commercial over parasite, which more properly Secondary pest outbreak. A rapid
preparation of living microorganisms refers to those types of organisms, increase in the population of one
(such as bacteria, viruses, or fungi) such as fleas and lice, that do not kill pest that occurs after treatment for
that are pathogenic to specific their hosts. another type of pest. The increase
groups of insects. These preparations
Parthenogenesis. Egg maturation results from the elimination of the
can be mixed with water and applied
without fertilization. In some insects, natural enemies of the secondary
with conventional pesticide-applica-
offspring are commonly produced pest, as after insecticide application.
tion equipment. Microbial insecti-
without the need of egg fertilization. Compare with pest resurgence.
cides are regulated as pesticides by
This commonly occurs in the parasitic Simple metamorphosis. Type of insect
the Environmental Protection
wasps and their relatives, where development in which the insect
Agency, so users must follow specific
female offspring are derived from fer- does not pass through a pupal stage
labeling and use guidelines.
tilized eggs and male offspring are before becoming an adult. Immatures
Molt. The process of shedding the skin derived from unfertilized eggs. are similar in shape to adults, but
between developmental stages
Pathogen. An organism capable of they are smaller and lack wings. Also
(instars).
causing disease. called incomplete metamorphosis.
Natural enemy. A beneficial organism
Pest resurgence. A pest outbreak that Solitary parasite. A species in which
that kills or interferes with pests.
results from the elimination of the only one individual develops within a
These are the biological components
pest’s natural enemies, such as after single host. Compare with gregarious
of natural control; when manipulated
an insecticide application. Even parasite.
by people they are the essential com-
though the pesticide may initially Species. A group of interbreeding indi-
ponents of biological control. The
control the target pest, the pest can viduals or populations, similar in
natural enemies of insect pests
recolonize and reproduce rapidly structure and physiology, that are dif-
include predators (such as predatory
because its natural enemies have ferent from all other groups and
insects), parasitic insects, and
been eliminated. Compare with sec- produce fertile offspring.
microorganisms that cause insect
ondary pest outbreak.
diseases. Thorax. The body region between the
head and abdomen. Bears the wings
and legs.
98
Index beetle D
Numbers in boldface indicate pages
with photos. carabid, 69, 88 Dacnusa sibirica, 35, 38, 39, 40
coccinellid, 58 Delphastus pusillus, 76, 81, 82, 94
A European rove, 85
Deraeocoris brevis, 18, 23
Adalia bipunctata, 18 ground, 87
Diaeretiella rapae, 16, 22
Amblyseius. See also Euseius, Neoseiulus bigeyed bug, 77
aleyrodis, 77 Dicyphus tamaninii, 63, 66, 77
Braconidae, 15
mckenziei, 63 Diglyphus spp., 38
swirskii, 77 C begini, 24, 36, 37, 38
Amitus fuscipennis, 74 cabbage loopers, 26, 27 intermedius, 36, 38
caterpillars, 26–29, 92–93 isaea, 36, 38, 39, 93
Anagyrus pulchripes, 36, 38
fusciventris, 56 centipedes, 86
pseudococci, 56 Diomus flavifrons, 58
Ceranisus (= Thripoctenus) spp., 61
Aphelinidae, 15, 74 brui, 62 Diversinervus elegans, 51

Aphelinus menes, 62
E
abdominalis, 15, 22 Chilocorus Encarsia spp., 75
asychis, 15 bailyii, 51, 54 formosa
flavipes, 15, 23–24 bipustulatus, 54 whiteflies, control of, 73, 74, 79–82
semiflavus, 15 circumdatus, 51, 54 uses, examples of, 89, 92, 93, 94
aphid midge, 21 infernalis, 51 luteola (=deserti), 75
kuwanae, 54 pergandiella (=bemisiae=tabacivora
Aphidiidae, 15 nigritis, 51, 52, 54 =versicolor), 75
Aphidiinae, 15 Chrysocharis. See Oscinidius Encyrtidae, 50, 56
Aphidius Chrysopa, 54, 58
colemani, 15, 22 Encyrtus
See also Chrysoperla infelix, 51
ervi, 15, 22
matricariae, 16, 22, 23, 24 Chrysoperla spp., 54, 58 lecaniorum, 51
carnea (=Chrysopa carnea), 17, 27, 76, 82 Entomophthora
aphidlions, 17, 23 comanche, 17, 76, 82 coronata, 86
Aphidoletes aphidimyza, 16, 21, 24 rufilabris, 17, 27, 76, 82 parvispora, 64
aphids, 14–25, 92–93 Coccinella septempunctata, 18, 76 thripidum, 64
Aphytis Coccoidea, 49 Ephedrus cerasicola, 16, 23, 24
lingnanensis, 53 Coccophagus lycimnia, 50, 51, 52 Eretmocerus
melinus, 53, 55 eremicus (=californicus), 75, 81, 93
Coleomegilla maculata, 18, 76
Aschersonia aleyrodis, 78, 82 haldemani, 76
Comperiella bifasciata, 54 mundus, 76
B corn earworms, 26 Eulophidae, 61
Bacillus thuringiensis (Bt), 28
Cryptolaemus montrouzieri, 51, 57 European corn borers, 26
var. israelensis (Bti), 31, 32, 33, 94
Crytopeltis modestus, 37 Euseius (=Amblyseius) spp., 62
banker plants, 22, 24, 47, 80, 95
cutworms delhiensis (=rubini), 44, 62, 77
Beauveria bassiana hibisci, 44, 62
black, 26
aphids, control of, 20, 24 scutalis, 44, 62, 77
bran bait, recipe for, 29
integrated biological control, stipulatus, 42–43, 44
variegated, 26
example, 93
thrips, control of, 64, 67–68 Cycloneda sanguinea, 18, 76 Euthycera spp., 84
whiteflies, control of, 78, 81, 82 cribata, 84

beet armyworms, 26, 29


99
F Leistus, 87 mites, 4
Feltiella, 47, 48 Lemnia biplagiata, 18 acarid, 41–42
acarisuga (=Therodiplosis persicae), 45 broad, 42–43
Leptomastidea abnormis, 56, 59 bulb, 41–42
minuta, 45
Leptomastix cyclamen, 42–43
flies
dactylopii, 56, 57, 58–59 laelapid, 31, 41, 62
hover, 19, 22
epona, 57 phytoseiid. See mites, predatory
sciomyzid, 85
Lindorus. See Rhyzobius predatory
syrphid, 19
caution about insecticidal soap, 25
tachinid, 88 Lithobius mites, control of, 42–48
fungus gnats, 30–33, 93–94 bilabiatus, 86 thrips, control of, 61–63, 65, 67
forficatus, 86 whiteflies, control of, 77
G Loricera, 87 symphylans, control of, 86
Galendromus (=Metaseiulus, tarsonemid, 42–43
Lysiphlebus testaceipes, 16, 23, 24
=Typhlodromus) occidentalis, 44, 46 spider, 43–48, 89, 92
Ganaspidium utilis, 36 M moths. See caterpillars
Geocoris punctipes, 77 Macrolophus
Geolaelaps sp., 87
caliginosus, 19, 45, 63, 77, 81-82 N
costalis, 19 nematodes
See also Hypoaspis
nubilis, 45 fungus gnats, control of, 32–33
integrated biological control,
H–I mealybug destroyer, 51, 57–59
example, 94
Heterorhabditis spp., 32, 88 mealybug, 55–59 leafminers, control of, 37, 39
bacteriophora (=heliothidis), 64, 70, 85, 87
Mesoseiulus (=Phytoseiulus) longipes, 44 slugs, control of, 85
megadis, 32
Metaphycus sowbugs, control of, 88
Hippodamia convergens, 18, 51, 76 springtails, control of, 87
helvolus, 50, 51, 52, 55
Hungariella. See Pauridia luteolus, 50 symphylans, control of, 86
zebratus, 50 thrips, control of, 64
Hypoaspis spp., 87
weevils, control of, 69–71
aculeifer, 41–42 Metarhizium anisopliae
miles aphids, control of, 20, 24 Neoaplectana. See Steinernema
fungus gnats, control of, 31, 33 thrips, control of, 64, 68 Neoseiulus (=Amblyseius)
integrated biological control, weevils, control of, 70 barkeri, 63
example, 94 whiteflies, control of, 78, 82 californicus, 44, 46, 47
springtails, control of, 87, 88 cucumeris, 43, 63, 65–66
Metaseiulus. See Galendromus
thrips, control of, 62, 66–67 degenerans, 63, 65
vacua, 41 Micromus angulatus, 19 fallacis, 44
indicator plants, 44, 46, 47 Microterys flavus, 51, 52 Nephus. See Scymnus
midge, 16, 21–22, 45, 47–48 Notiophilus, 87
L minute pirate bug
lacewings, 22
brown, 19
aphids, control of, 19 O
caterpillars, control of, 27 Ocypus. See Staphylinus
Comanche, 17, 76
thrips, control of, 63, 65, 66, 67 Opius spp., 40
green
whiteflies, control of, 77 dimidiatus, 37, 38
aphids, control of, 17, 21, 22
caterpillars, control of, 29 mirid bug dissitus, 37
scales and mealybugs, control of, aphids, control of, 18, 23 pallipes, 37
51, 54, 58 mites, control of, 45 Orius spp.
whiteflies, control of, 76, 82 thrips, control of, 63 aphids, control of, 19
whiteflies, control of, 77, 81-82 thrips, control of, 63, 66, 67
lady beetles, 27, 55
convergent, 18, 22, 51, 76 whiteflies, control of, 77, 82
Korean twicestabbed, 54 albidipennis, 66
insidiosus, 66, 77
Lamyctes spp., 86
laevigatus, 66
leafminers, 34–40, 92–93
Oscinidius (=Chrysocharis) parksi, 37, 38
100 B I O L O G I C A L C O N T R O L O F I N S E C T S A N D O T H E R P E S T S O F G R E E N H O U S E C R O P S

P sticky traps, 12
Paecilomyces fumosoroseus aphids, monitoring, 15
aphids, control of, 20, 24 fungus gnats and shore flies,
thrips, control of, 64, 68 monitoring, 31
whiteflies, control of, 78, 82 integrated biological control, use in, 93
leafminers, monitoring, 34–35, 38, 40
Pardosa amentata, 87
thrips, monitoring, 61, 65, 68
Pauridia (=Hungariella) peregrina, 57 whiteflies, monitoring, 73, 79, 81, 83
Pergamasus quisquiliarum, 86 symphylans, 86
Phasmarhabditis sp., 85
Phytoseiulus. See also Mesoseiulus
T
Tetanocera spp., 85
macropilis, 45, 46
elata, 85
persimilis, 44, 45, 46–47, 48, 63
Tetradonema plicans, 32
Platygastridae, 74
Therodiplosis persicae, 45
predatory gall midge, 45, 47
Thripinema nicklewoodii, 64
R Thripobius semiluteus, 62, 67
Rhyzobius (=Lindorus) lophanthae,
Thripoctenus. See Ceranisus
51, 52, 54, 55
thrips, 60–68, 92–93
Rumina decollata, 84
sixspotted, 45, 47, 48
S tomato bug, 37
scale Trichogramma spp., 27, 29
armored, 53–55 minutum, 27
soft, 49–52 platneri, 27
Sciomyzidae, 84 pretiosum, 27
Scolothrips sexmaculatus, 45 Typhlodromus thripsi, 63
See also Galendromus
Scymnus (=Nephus) spp., 51
bipunctatus, 58
reunioni, 58, 59
V
Verticillium lecanii
shore flies, 30–33 aphids, control of, 20, 22, 23, 24
slugs, 84–85 scales, control of, 51
thrips, control of, 64, 68
snails, 84–85, 88
whiteflies, control of, 79, 82
sowbugs, 87–88
bran bait, recipe for, 88 W
springtails, 87 wasps
aphelinid, 74–75
Staphylinus (=Ocypus) olens, 85
braconid, 14–16, 35, 37
Steinernema (=Neoaplectana) spp., 88 encyrtid, 50–52, 74
carpocapsae eucoilid, 36
fungus gnat, control of, 32 eulophid, 24, 36, 37, 61
leafminers, control of, 37, 39 mymarid, 61
slugs, control of, 85 platygastrid, 74
symphylans, control of, 86 trichogrammatid, 61
weevils, control of, 70
weevils, 69–71
feltiae (=bibionis), 32, 64, 85, 87
riobravis, 32 whiteflies, 72–83, 89, 92–94
Stethorus spp., 46
punctillum, 46
punctum, 8
Copyright © 2001 by the Board of Regents of the University of Wisconsin System doing
business as the division of Cooperative Extension of the University of Wisconsin-
Extension. Send inquiries about copyright permission to: Director, Cooperative Extension
Publishing, 201 Hiram Smith Hall, 1545 Observatory Dr., Madison, WI 53706.
Authors: Susan E. Rice Mahr is entomology specialist and Daniel L. Mahr is professor of
entomology, College of Agricultural and Life Sciences, University of Wisconsin-Madison
and University of Wisconsin-Extension, Cooperative Extension. Raymond A. Cloyd is assis-
tant professor and Extension specialist of ornamental entomology and integrated pest
management, Department of Natural Resources and Environmental Sciences, University
of Illinois. Clifford Sadof is associate professor of entomology, Purdue University.
Produced by Cooperative Extension Publishing, University of Wisconsin-Extension,
Cooperative Extension.
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U.S. Department of Agriculture and Wisconsin counties, publishes this information to
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NCR581 Biological Control of Insects and Other Pests of Greenhouse Crops


I-09-01-3.5M-2500