NC STATE UNIVERSITY
The
Ground Beetles
of Eastern North Carolina
Agriculture
Ground beetles (a.k.a.. carabid beetles or carabids) are a common, but often overlooked, beneficial insect in
agricultural areas. Because they typically live in or on the soil and are usually active only at night, most people don’t
notice them. However, they can be abundant and affect pest management. Some ground beetles are predators of
other insects in the soil or upon plants, while others feed on weed seeds. Although scientists have studied ground
beetles for years, because of their secretive habits there is still much to be learned about them. One of the first steps
to appreciating the value of these beetles is to know which species are present and how to recognize them.
This publication provides a quick guide to the various
ground beetles that inhabit agricultural fields of the piedmont
and eastern North Carolina. It offers Extension personnel,
consultants, growers, scientists, and the general public an
introduction to the identification and life histories of our
most abundant ground beetles. We focus on adult beetles
only, because they are much easier to collect and recognize
than are larvae. Reference material is provided at the end of
the article for those who are interested in learning more.
Ground Beetle Ecology and Habits
Ground beetles are predators that feed on insects, other
invertebrates (such as snails), and seeds. They are specialized for many modes of life in and on the ground, as well as
on plants, under bark, or as “miners” spending most of their
time digging underground. Since agricultural settings are
highly simplified environments, only a few of the many species of ground beetles are found there, though sometimes in
great numbers. Agricultural settings tend to support ground
beetles that are likely to feed on insect pests such as caterpillars and aphids, as well as weed seeds. Farmers have long
been encouraged to look upon carabid beetles as naturally
occurring pest control. Although it is difficult to say exactly how much pest control is provided by these beetles on
individual farms, it is safe to say that they have a meaningful
impact like other general predators such as ladybeetles and
spiders. Current research is attempting to answer this question and learn what practices best enhance ground beetle
activity in agriculture.
Conserving and Enhancing Ground Beetles
There are a number of strategies growers can employ that
show promise for conserving beneficial ground beetles
in crop fields. Ground beetles are generally vulnerable to
deep or inversion tillage (e.g. moldboard plow) and benefit
from reduced tillage regimes (Kromp 1999). Although adult
ground beetles may be fast enough to escape from large
disturbances like heavy plowing, their larvae and eggs are
not mobile and are very susceptible to these disturbances.
Increasing amounts of ground cover in crop fields benefit
ground beetles, and cover crops can enhance beetle numbers
(Shearin et al., 2008). However, cover crops are eventually
turned under and destroyed and ground beetles need refuges
where they can thrive and reproduce year round. Good
refuges for ground beetles can be created in the borders of
the field by planting a diverse mix of annuals and perennials
(Mennaled et al 2001). A similar strategy is to create these
“beetle banks” within the crop field itself. Other strategies,
such as intercropping and using organic fertilizers rather
than mineral fertilizers, seem to promote ground beetles as
well (Kromp 1999).
A specific strategy for weed seed eating ground beetles is
to delay fall cover crop planting and fall tillage. This practice
keeps weed seeds on the soil surface where they are more
likely to be eaten by ground beetles (Gallandt et al., 2005).
Classification of Ground Beetles
Unfortunately for the average reader, the names that are used
to describe the various types of ground beetles are based
on taxonomic jargon. Because most of these beetles don’t
have common names, we will also follow this convention.
We have organized the various species of beetles by tribe.
In addition to providing taxonomic organization, the tribe
classification, to some degree, separates beetles based on
their feeding habits. We have presented the basic hierarchical
naming system used in insect classification (taxonomy and
systematics):
THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
Order (e.g. Coleoptera, the beetles)
Family (e.g. Carabidae, the ground beetles)
Tribe (e.g. Cicindelini, the tiger beetles)
Genus (e.g. Megacephala, the big-headed tiger beetles)
Species (e.g. Megacephala carolina, the Pan-Ameri-
can big-headed tiger beetles)
The full species name of an insect includes the genus,
species, the name of the person who described the species
(the author), and the date the species was described, such as
Megacephala carolina (Linnaeus, 1767). For the well-known
authors, such as Linnaeus or Fabricius, the authors’ names
are shortened to the first letter of the name. In the scientific
literature (other than systematics), the date is usually omitted.
Handling Live Beetles
Only the largest of the ground beetles can manage a recognizable bite. However, many are capable of producing
noxious chemical defenses in the form of foul odors, as in
the Chlaeniini, or sometimes even acidic stinging spray as in
the large Carabini. These chemical defenses are not used for
attack but to dissuade their own predators (usually birds, or
sometimes raccoons and other small mammals) in much the
same manner as a skunk does.
After handling live beetles, it is always prudent to wash your
hands before touching your eyes or any other sensitive skin.
Otherwise, you may discover that the “Fiery Searchers”
(Calosoma sp.) are aptly named!
Collections
This publication is based on collections made near Goldsboro,
NC, in 2009 and 2010. The sites where we collected differ substantially in soil types, distance from water, and surrounding
vegetation (see Forehand et al. 2006). Our collections were not
geographically extensive throughout eastern North Carolina.
However, Kirk (1972) and Thiele (1977) indicate that even
relatively local, non-intensive collections will quickly reveal at
least the dominant species in an area. Our collecting regime
was intensive, conducted in three crops (corn, soybeans, hay)
in nine fields, and was year-round, so we feel confident that
we collected the vast majority (if not all) of the species that
typically occur in these agricultural habitats.
Table 1. List of carabid beetles found from pitfall sampling in nine agricultural fields near Goldsboro, N.C.
Feeding habits from Larochelle and Larivière (2003). Species in bold are common in our collections.
Tribe
General Feeding Habit (if known)
Species
Cicindelini
Predatory; stalk arthropod prey on the
ground
Megacephala carolina (Linnaeus, 1767)
Megacephala virginica (Linnaeus, 1767)
Cicindela punctulata Olivier, 1790
Harpalini
Herbivorous/Omnivorous: primarily
feed on weed seeds but will also feed
on other arthropods if needed.
Harpalus pensylvanicus (De Geer, 1774)
Harpalus caliginosus (Fabricius, 1775)
Acupalpus sp.
Amphasia sericea (T.W. Harris, 1828)
Anisodactylus merula (Germar, 1824)
Anisodactylus fervus LeConte, 1863
Anisodactylus sanctaecrucis (Fabricius, 1798)
Anisodactylus laetus Dejean, 1829
Anisodactylus sp.
Selenophorus palliatus (Fabricius, 1798)
Stenolophus infuscatus (Dejean, 1829)
Stenolophus lineola (Fabricius, 1775)
Stenolophus ochropezus (Say, 1823)
Stenolophus spretus Dejean, 1831
Cratacanthus dubius (Palisot de Beauvois, 1811)
(Table continued next page)
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THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
Table 1. continued
Tribe
General Feeding Habit (if known)
Species
Chlaeniini
Mainly feed on arthropods, and other
invertebrates, but also fungi
Chlaenius tomentosus (Say, 1823)
Chlaenius aestivus Say, 1823
Chlaenius tricolor DeJean, 1826
Chlaenius laticolis Say, 1823
Pterostichini
Mainly arthropod and other
invertebrate prey, especially caterpillars,
and beetle eggs
Poecilus chalcites (Say, 1823)
Poecilus lucoblandus (Say, 1823)
Cyclotrachelus sigillatus (Say, 1823)
Pterostichus sculptus LeConte, 1852
Carabini
Predatory; Mainly on caterpillars often
found in agricultural settings
Calosoma sayi DeJean, 1826
Platynini
Caterpillars, aphids, mealworms
Agonum pallipes (Fabricius, 1787)
Carabus vinctus (Weber, 1801)
Agonum punctiforme (Fabricius, 1823)
Agonum octopunctatum (Fabricius, 1798)
A. rigidulum (Casey, 1920)
Calathus opaculus LeConte, 1854
Scaratini
Caterpillars, wire worms, other insects,
and occasional plant matter
Scarites subterraneus Fabricius, 1775
Galeritini
Caterpillars and grass seeds
Galerita bicolor Drury, 1773
Liciniini
Caterpillars
Dicaelus elongatus Bonelli, 1813
Cyclosomini
Small insects
Tetragonoderus intersectus (Germar, 1824)
Zabrini
Plant seeds, caterpillars
Amara cupreolata Putzeys, 1866
Amara apricaria (Paykull, 1790)
Amara musculis (Say, 1823)
Bembidion affine Say, 1823
Bembidiini
Bembidion rapidum (LeConte, 1848)
Notiophilus novemstriatus LeConte, 1848
Notiophilini
Clivinini
Burrowing habit, tiny insects
The Beetle Species
Eleven tribes of carabid beetles were represented in pitfall
trap collections taken from July through December 2009 in
three corn fields, three soybean fields, and three hay fields
near Goldsboro. Thirty-one species representing twenty
genera and eleven tribes of carabids were identified from
these collections (Table 1). Of these, nine genera and eleven
species from six tribes made up approximately 99 percent
Clivina bipustulata (Fabricius, 1801)
of the species assemblage. Megacephala carolina alone accounted for 55.4 percent of the beetles collected, followed
by Harpalus pensylvanicus (26.5 percent); Anisodactylus spp.
(6.9 percent—note that Anisodactylus species are difficult
even for experts to distinguish, and at least five species are
represented in this collection); Poecilus chalcites (3.5 percent); Calosoma sayi (3.1 percent); Agonum punctiforme
(1.7 percent); Cyclotrachelus sigillatus (1.3 percent); Cicindella punctulata (0.7 percent); and Chlaenius tomentosus
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THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
(0.7 percent). For practical purposes, beetles collected with a frequency of 0.5 percent or more were considered common and are both described and illustrated below.
Beetles with a collection frequency between 0.5 and 0.1 percent were considered
uncommon and are only illustrated in Figure 14. Those with a collection frequency
less than 0.1 percent were considered rare and are not illustrated.
Seasonality and Abundance
The life cycles of different types of carabid beetles vary, and so collections of adult
beetles will vary with the seasons. In our collections, Cicindelini, Carabini, Chlaeniini, and Pterostichini were most abundant in the summer months, but were
still found in smaller numbers in the fall. By late fall (December) they were rarely
encountered. Harpalini were most abundant in the fall, but were found in much
smaller numbers in the summer and late fall. Platynini never reached any appreciable numbers until late fall (December) when they became the dominant carabid
trapped, though in much smaller numbers than carabids dominant in summer. A
small number of Bembidiini species were found in a collection made in February
only. An impressive number (>3000) of Amara cupreolata (tribe Zabrini Fig. 12m.)
were found in one particular field in May, likely the result of a locally high density of
annual bluegrass that was allowed to mature in that particular field.
Figure 1. Threadlike antennae and
5-5-5 tarsal “formula” (circles) typical of
carabid beetles. Photo by Henri Goulet,
Agriculture & Agri-Food Canada.
Description of the Commonly Collected Ground Beetles
Ground beetles all share a few common features. They always have threadlike, long
antennae that do not come to a club or branch (Fig. 1). They have five tarsal (“foot”)
segments on each leg (Fig 1), and an expanded hind trochanter (”hip joint”), which
looks a bit like a jelly bean, lying next to the femur of the hind leg (Fig. 2). They
are all fast-running hunters, seed feeders, or both. The majority of carabid species
are nocturnal, preferring to hide under rocks, logs, or leaf-litter during daylight
hours. Like all beetles, the front wings of ground beetles are modified into distinctive protective shell-like coverings called elytra that protect the hind wings. Most
carabid larvae build burrows in which they live. The depth of these burrows appears
to be specific to each group. Predatory larvae attack passing insect prey, which they
drag into their burrows. Seed-feeding larvae may line the walls of their burrows with
stores of seeds.
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Figure 2. Expanded hind trochanters
(indicated by arrow) of the Carabidae.
THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
About the Images
In a perfect world, field-caught specimens would look just like the pictures in
this publication and make everyone’s job of identifying things a breeze. In the
real world, however, specimens are seldom identical to their pictures and often
require specific equipment for viewing. A good hand lens is indispensible, but the
most important aspect of viewing the coloration of beetles is lighting. While the
tiger beetles are often very easy to discern, some of the coloration on Chlaeniini
and other groups is much more subtle and requires strong directed light, such as
that provided by a dissecting-scope illuminator. Even with the appropriate lighting, shifting angles often produce a range of coloration. Also, coloration naturally
varies even in the same species, and sometimes dead specimens look a little less
vivid than live ones, which is why taxonomists try not to rely entirely on colors
for identifications. Lines beside the images in Figures 4 to 13 represent actual size,
when the page size in this article is 8.5 by 11 inches.
Figure 3. Tiger beetle mouthparts.
Tribe Cicindelini (Tiger beetles)
The adults of this group are easily recognized by their extremely long, almost
spidery legs and massive jaws (Fig. 3, 4, 5). The Cicindelini are the one tribe of
carabids in which the labrum (top lip) is wider than the antennal insertions (Fig.
3). Most tiger beetles are active in either the spring and fall, or the summer. Summer active species dominate in the south, while spring/fall active species are more
common in the north (Pearson et al. 2006). Summer active adults die off at the
end of the year, but their larvae overwinter in burrows that are deep enough to
have moderate temperatures. Tiger beetles are thought to be valuable indicators
of ecosystem health due to their easy identification and collection, as well as their
specificity to an environ (Pearson and Cassola 1992).
Megacephala carolina (Pan-American big-headed tiger beetle, Carolina tiger
beetle) (Fig. 4) These beetles are predatory, nocturnal, often gregarious and
will scatter from the light of a flashlight. These were by far the most abundant
beetles in our collections.
Figure 4. Megacephala carolina.
Identification: They are easily recognized by their unique purple and green
metallic surface, and yellow comma-shaped markings on the elytra (circle). This
beetle is relatively large, usually near 20mm (0.79 inches).
Cicindela punctulata (punctured tiger beetle, sidewalk tiger beetle) (Fig. 5) isa
widely distributed upland beetle often found in agricultural settings. It is sometimes so common that it is believed to have developed some pesticide resistance
(Larrochelle and Lariviere 2003). It has a 1- to 2-year life cycle.
Identification: Dark shining olive or tar-colored dorsally, with slight white markings on the lateral edges of the elytra, and metallic blue ventrally. This beetle is
about 13mm (0.51 inches) long.
Figure 5. Cicindela punctulata.
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THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
Tribe Harpalini
Harpaline beetles are recognized by having one seta (hair) above each compound
eye, a mostly dark brown or black dorsal color, and all lack an interruption in the
lateral edge of the elytra (see tribe Pterostichini for description of this feature). Species are often difficult even for experts to discern.
Harpalus pensylvanicus (Fig. 6) is common in croplands and widely distributed
throughout the contiguous U.S. and Canada. This beetle has been observed feeding
on seeds of wheat, Timothy grass, and prairie grass. Gut content studies have found
ragweed pollen as well as ants and mites (Kirk 1974). Because of their preference
for seeds and because they are active in the fall months when summer annual weeds
have shed their seeds, Harpalus is an important weed seed predator. Harpalus species have been shown to consume 90% of the seeds of certain weed species (Zhang,
1993). Kirk (1971) lists this as the second most prevalent species in North Dakota
agricultural fields. There appears to be a preference for cultivated land, and aggregation in weedy grasses (foxtail grass) where the planter missed a row in corn fields
(Kirk 1974). Larvae may provision burrows with grass seeds pasted to the sides of
the tunnel-like tiles and pack such seeds around the opening (Kirk 1971).
Figure 6. Harpalus pensylvanicus.
Photo by Henri Goulet.
Identification: Light tan legs and a somewhat shining, tar colored body. It is 13 to
15.5mm (0.59 inches) in length.
Anisodactylus fervus, Anisodactylus merula and Anisodactylus sp. (Fig. 7) are a prevalent component of our collections in early to mid summer and are nearly replaced
by H. pensylvanicus in the fall. It is omnivorous, feeding on seeds, caterpillars, and
in laboratory on mealworms (Larochelle and Lariviere 2003). Laboratory experiments have shown that seed predation by these beetles can reduce weed pressure
from redroot pigweed (Amaranthus retroflexus) and lambsquarters (Chenopodium
album) (Kromp 1999). However, it is unclear what their effect is in agricultural fields
because they are prevalent before weeds shed their seeds in the fall. This nocturnal,
gregarious beetle, according to Larochelle and Lariviere (2003) favors human activities and is frequently found at roadsides, golf courses, and other disturbed areas.
Identification: Dark, black to rusty brown, and translucent at the posterior sides of
the pronotum. In many, the protibial apical spur (circle) is trifid (trident-like). These
beetles are about 8 to 13mm (0.51 inches) in length.
Figure 7. Anisodactylus merula
Tribe Pterostichini
These beetles are recognized by having two setae above each eye and an interruption
in the lateral to apical edge of the elytra (circle).
Poecilus chalcites (Fig. 8) was found in many of our pitfall samples. It is a common
predator on a number of crop pests including western corn rootworm, corn earworm, black cutworm, and armyworm. This species is recorded from 35 states east
of the Rocky Mountains and eastern Canada. It is often the most abundant ground
beetle during most of the growing season in Illinois farmland (Lundgren et al. 2005).
Lundgren et al. (2005) report that P. chalcites requires a diapause before becoming
sexually mature.
Figure 8. Poecilus chalcites.
Photo by Henri Goulet
Identification: This beetle has a distinctive smooth, bronze-green sheen. The green
color will not be as distinctive as in Fig. 8, except under special microscope lighting.
It is about 10 to 12mm (0.47 inches) in length.
Cyclotrachelus sigillatus (Fig. 9) found in the late fall is black and shining.
Identification: The pronotum is almost square and as wide as the elytra. The elytra
do not open, rendering this beetle incapable of flight. They are 13 to 18mm (0.64
inches) in length.
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Figure 9. Cyclotrachelus sigillatus.
THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
Tribe Carabini
These relatively large beetles are often called “hunters” or “searchers” and famously prefer caterpillars. They have relatively large jaws and are often brightly marked
or colored. They are capable of spraying noxious acidic defensive secretions.
Calosoma sayi (Fig. 10) is among the largest beetles in our collections. Often
called “fiery searcher” due to the acids it will spray on your hands if handled.
Identification: The characteristic elytral puncture-markings can be quite variable
in color from bronze-red to blue-green. This is the largest carabid in our trap collections, often over 25mm (1 inch) in length.
Figure 10. Calosoma sayi.
Tribe Zabrini
Only two genera are contained in the Zabrini, and only the genus Amara is found
in our area. The Amara are recognized by having characteristics of the Pterostichini: two setae above each compound eye and an interrupted lateral elytral
edge. They differ from the Pterostichini in being oval in shape and lacking any
punctures on the dorsal surface of the elytra. Amara also have many setae on the
second segment of the labial palpi where many Pterostichini have only two (however this character alone will not suffice in separating the tribes).
Amara cupreolata (Sun beetles) (Fig. 11) is common in our collections in May
and early June. It can often be seen running about vigorously on bright sunny
days. It is a known seed feeder preferring grasses.
Identification: Oval, shining, brownish-bronze colored body with dark legs. They
are 6.6 to 9mm (0.35 inches) in length.
Figure 11. Amara cupreolata.
Tribe Chlaeniini
These velvety beetles are often metallic and brightly colored. They have a single
seta (hair) above each eye like the Harpalini), but they also have interruption in
the lateral edge of elytra like the Pterostichini and Zabrini. The Chlaeniini are
famously foul-smelling.
Chlaenius tomentosus (Fig. 12) is found in rural areas and in cities. It seems to
prefer cultivated land, old fields, and vacant lots. This beetle emits a musky or
smoky odor when handled.
Identification: It is a shining bronze-green beetle, covered in fine hair, like velvet
on the elytra. It is 13 to 15mm (0.59 inches) in length.
Figure 12. Chaelius tomentosus.
Photo by Henri Goulet.
Tribe Platynini
These mostly black, shining beetles have two setae above the eye (like the Pterostichini and Zabrini) and never have the lateral edge of the elytra interrupted.
Agonum punctiforme (Fig. 13) was only apparent in our collections in the late fall
and in warm winter weather.
Identification: It is a shining black beetle with a well rounded pronotum with
two distinctive rounded dimples basally. The elytra are slightly less black than
the head and pronotum and can reflect bronze green under strong lights. There
are 3 dorsal punctures on the elytra in the third interval. It is 7 to 9mm (0.35
inches) in length.
Figure 13. Agonum punctiforme.
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THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
Figure 14. Occasionally encountered carabid beetles from eastern North Carolina agricultural fields. a. Cicindelini, Megacephala virginica.
b- g. Harpalini, Amphasia sericea; Harpalus caliginosis; Cratacanthus dubious; Selenophorus palliatus (Photo by Igor Sokolov, LSU); Stenolophus lineola; S. ochropezus. h. Pterostichini, Poecilus lucublandus. i. Platynini, Agonum octopunctatum. j. Chlaeniini, Chlaenius tricolor. k.
Galeritini, Galerita sp. l. Scaritini, Scarites subterraneum. m. Licinini, Dicaelus elongatus. n. Carabini, Carabus vinctus. Photos marked “HG” by
Henri Goulet Agriculture and Agri-Food Canada. Images are not to scale.
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THE GROUND BEETLES OF EASTERN NORTH CAROLINA AGRICULTURE
For Further Information.
References Cited
Ciegler, J. C. 2000. Ground beetles and Wrinkled Bark Beetles
of South Carolina (Coleoptera: Geadephaga: Carabidae
and Rhysodidae). Biota of South Carolina. Vol. 1. Clemson University, Clemson, S. C.
Forehand, L.M., D.B. Orr, and H.M. Linker. 2006. Evaluation of
a Commercially Available Beneficial Insect Habitat for Management of Lepidoptera Pests in Organic Tomato Production. J. Econ. Entomol. 99: 641-647.
Pearson, D. L., C. B. Knisley, and C.J. Kazilek. 2006. A Field
Guide to the Tiger Beetles of the United States and Canada.
Oxford University Press.
Shearin, A. F., Reberg-Horton, S. C. , and Gallandt, E. R. 2008.
Cover crop effects on the activity-density of the weed seed
predator Harpalus rufipes (Coleoptera: Carabidae). Weed
Science 56(3):442-450.
Thiele, H. U. 1977. Carabid Beetles in Their Environments.
Zoophysiology and Ecology 10. Springer-Verlag. Berlin
Heidelberg New York.
Gallandt, E. R., Molloy, T., Lynch, R. P., and Drummond, F. A.
2005. Effect of cover-cropping systems on invertebrate seed
predation. Weed Sceince 53:69-76.
Web Resources
Kirk, V. M. 1971a. Ground beetles of cropland in South Dakota.
Ann. Entomol. Soc. Am. 64: 238-41.
Ground Beetles of Canada Image Library
http://www.cbif.gc.ca/spp_pages/carabids/phps/image_e.php
Kirk, V. M. 1974. Biology of a Ground Beetle, Harpalus pensylvanicus. Ann. Entomol. Soc. Am. 66(3): 513-518.
Image Gallery “Carabidae of the World”
http://www.carabidae.ru/foto/index.php
Kromp, B. 1999. Carabid beetles in sustainable agriculture:
a review of pest control efficacy, cultivation impacts and
enhancement. Agriculture, Ecosystems and Environment 74:
187-228.
Larochelle, A. and M.C. Larivière. 2003. A Natural History of
the Ground-Beetles (Coleoptera: Carabidae) of America
north of Mexico. Pensoft Sofia, Bulgaria.
Lundgren J. G., J. J. Duan, M. S. Paradise and R. N. Wiedenmann 2005. Rearing Protocol and Life History traits for Poecilus chalcites (Coleoptera: Carabidae) in the Laboratory.
J. Entomol. Sci. 40(2): 126-135.
Menalled, F. D., J. C. Lee, and D. A. Landis. 2001. Herbaceous
filter strips in agroecosystems: Implications for ground
beetle (Coleoptera : Carabidae) conservation and invertebrate weed seed predation. Great Lakes Entomologist 38(2):
472-483.
Pearson, D. L., F. Cassola. 1992. World-Wide Species Richness
Patterns of Tiger Beetles (Coleoptera: Cicindelidae): Indicator Taxon for Biodiversity and Conservation Studies. Conservation biology 6(3): 376-91.
Books of Interest
Acorn, J. H. 2001. The Tiger Beetles of Alberta. Alberta Insects
Series. The University of Alberta Press. Edmonton, Alberta.
120pp.
Ball, G. E., and Y. Bousquet 2000. Carabidae. In Arnett, R. H.
Jr. (Editor), and M. C. Thomas (Editor) American Beetles,
Volume I. Archostemata, Myxophaga, Adephaga, Polyphaga:
Staphyliniformia CRC Press. 464pp.
Ciegler, J. C. 2000. Ground beetles and Wrinkled Bark Beetles
of South Carolina (Coleoptera: Geadephaga: Carabidae and
Rhysodidae). Biota of South Carolina. Vol. 1. Clemson University, Clemson, S. C. 149pp.
Pearson, D. L., C. B. Knisley, and C.J. Kazilek. 2006. A Field
Guide to the Tiger Beetles of the United States and Canada.
Oxford University Press. 227pp.
White, R. E. 1983. A Field Guide to the Beetles of North
America. The Peterson Field Guide Series. Houghton Mifflin
Company. New York, NY. 368pp.
Prepared by
Geoff Balme, Department of Entomology, NC State University
David Orr, Department of Entomology, NC State University
Aaron Fox, Department of Crop Science, NC State University
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