Leptinotarsa decemlineata

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Geographic Range

Currently, Leptinotarsa decemlineata, the Colorado potato beetle, is distributed widely throughout North America east of the Rockies as well as some of Europe and Asia. Its distribution covers about 8 million km² in the Nearctic Region and about 6 million km² in the Palearctic and Oriental regions. Originally, Leptinotarsa decemlineata was found in the southwestern United States into Mexico. As potatoes were extensively planted for agriculture, the species spread into agricultural areas throughout North America, Europe, and Asia. It is predicted that Leptinotarsa decemlineata could occupy other regions including Korea, Japan, parts of Africa, and most of the temperate Southern Hemisphere. (Alyokhin, et al., 2008; Alyokhin, 2008; Jolivet, 1991; Vlasova, 1978; Worner, 1988)

Habitat

The Colorado potato beetle is found mostly in farm fields that specialize in growing agricultural crops in the family Solanaceae, such as potatoes, tomatoes, tobacco, eggplants and peppers. It can also be found on non-agricultural solanaceous plants in open grassland areas. (Alyokhin, et al., 2008; Casagrande, 1987; Kramer, et al., 2009)

Physical Description

Leptinotarsa decemlineata has the physical features typical of chrysomelid beetles such as 5-5-5 tarsi, an oval shape, and antennae shorter than the body. Adults can reach anywhere from 8 to 10 mm and have five bold, brown stripes along each elytron. The thorax has an intricate pattern of black spots on top of a deep orange complexion. Larvae typically have a row of black spots down the side of the abdomen, which is convex and very stout (large and plump) compared to the rest of the body. Eggs resemble footballs with an orange/yellow color. They are about 1.7 to 1.8 mm long and 0.8 mm wide. The dorsal and ventral surfaces are distinctly non-parallel and deep red in color. (Alyokhin, 2008; University of Kentucky, 2010)

  • Sexual Dimorphism
  • sexes alike
  • Range length
    8 to 10 mm
    0.31 to 0.39 in

Development

The last generation of adults of Leptinotarsa decemlineata each year move to the edge of fields, bury slightly in the soil and overwinter. A small percentage of adults remain in diapause and emerge the following season. Fields are colonized by overwintered adults that walk to the field from their overwintering sites or emerge from the soil within the field. Once they have colonized the field, the overwintered beetles first feed and then oviposit within 5 to 6 days depending on temperature.

Development from egg to adult is greatly affected by temperature, varying from 14 to 56 days. The larvae of L. decemlineata go through four instars, typically lasting around 21 days, while continuously feeding on the host plant. Once larvae are mature, they drop from the host plant where they burrow 2 to 3 cm into the soil. After about 2 days they pupate, and then emerge as adults after an average of 5.8 additional days. The optimal temperature for development is within the range of 25 to 32˚C. After development has been completed, adults start mating and laying eggs to complete another generation of beetles. By midsummer, generations are asynchronous and all life stages can be found. (Alyokhin, et al., 2008; Alyokhin, 2008; Fasulo, 2009; Ferro, et al., 1991; Hazzard, et al., 1991; University of Kentucky, 2010; Voss and Ferro, 1990)

Reproduction

Both sexes of Leptinotarsa decemlineata mate with multiple individuals over the course of their adult life. Last generation adults typically mate before overwintering, with females storing sperm that can be used in the spring. However, they also mate after emerging from overwintering, usually before entering the fields. Sperm from these spring matings show some precedence, fertilizing the majority of eggs. Males have slightly modified tarsal setae that although them to cling to the elytra of the female. This is a trade-off with the ability to cling to host plants, and females can adhere to hosts more effectively. (Alyokhin, et al., 2008; Roderick, et al., 2003; Voigt, et al., 2008)

After overwintering in crop fields, gardens, and field margins, the Colorado potato beetle becomes active in the spring, often in May. Adults feed for a very short time, then reproduce. Adult females have high fecundity, producing 300 to 800 eggs, which are laid on the underside of plant leaves. Eggs are clustered into groups of 10 to 30. Egg laying may last several weeks. (Alyokhin, et al., 2008; Ferro, et al., 1991)

  • Breeding interval
    Adults breed continuously after emerging from overwintering.
  • Breeding season
    Leptinotarsa decemlineata breeds from spring through summer.
  • Range eggs per season
    300 to 800

Leptinotarsa decimilineata has little to no parental investment in the offspring, other than provisioning of eggs by females.

  • Parental Investment
  • pre-hatching/birth
    • provisioning
      • female

Lifespan/Longevity

Most individuals live for at least a few weeks as adults during the summer. The last generation of the year overwinters as adults, and thus has a longer adult lifespan, although they are mostly inactive. A small percentage of adults will remain in diapause for a second winter or even a third winter (recorded in some populations). (Alyokhin, et al., 2008; Kramer, et al., 2009)

  • Range lifespan
    Status: wild
    24 (high) months
  • Typical lifespan
    Status: wild
    2 to 12 months

Behavior

Flight behavior of Leptinotarsa decemlineata seems to be greatly affected by mating status and sex. Any adults that emerge from diapause, or in crowded conditions will tend to move and fly distances up to several kilometers, increasing their chances of finding new hosts. Mated females tend to move less, optimizing host location, while mated males tend to move more actively, particularly within a field, to locate new mates.

While in a field, Leptinotarsa decemlineata moves freely throughout its habitat. This occurs by both extensive walking and short-distance flights. Adults and larvae generally become inactive at night, with feeding activity concentrated during daylight hours. Larvae exhibit predator avoidance behaviors, including walking away, rearing up, regurgitating, wiggling, and defecating. Larvae often become aggregated as they feed, and adults can also be found in large groups on host plants. (Alyokhin, et al., 2008; Hazzard, et al., 1991; Ramirez, et al., 2010)

Home Range

This species does not exhibit territorial behavior and has no defined home range.

Communication and Perception

The primary modalities of perception and communication are olfaction and vision. Chemical signals emitted by host plants can be vital for sexual communication and host selection. Leptinotarsa decemlineata uses plant volatiles for host location at close range, but seems to rely mostly on vision for host and mate location. It is known that adults can sense wavelengths from yellow through the ultraviolet, and probably also sense polarized light. After locating a suitable host, males then emit their aggregation pheromone, (S)-3,7-dimethyl-2-oxo-oct-6-ene-1,3-diol [(S)-CPB I], that attracts both males and females to areas of host plants. Colonizing adults use this pheromone as a signal for potential mates. This is one of the few species of insects where it is known that larvae also sense, and are attracted to adult aggregation pheromone. For mating, female Leptinotarsa decemlineata produce a pheromone to attract males. (Boiteau, et al., 2003; Dickens, et al., 2002; Dickens, 1992; Dickens, 2006; Hammock, et al., 2007; Oliver, et al., 2002; Otalora-Luna and Dickens, 2010)

Food Habits

Leptinotarsa decemlineata feeds primarily on Solanum plants, skeletonizing the plant and leaving only the roots and stems. The most suitable host for L. decemlineata is now the cultivated potato, Solanum tuberosum, thus the beetle's common name of the Colorado potato beetle. Other suitable hosts include Solanum rostratum and Solanum augustifolium, the insect's original hosts. A European species now widely distributed in North America, Solanum dulcamara is also commonly used in the wild. Solanum melongena (eggplant), Lycopersicon esculentum (tomato), peppers, tobacco, and other wild hosts such as S. carolinense, S. sarrachoides, S. elaeagnifolium, and Hyoscyamus niger are utilized occasionally. (Alyokhin, et al., 2008; Fernandez and Hilker, 2007; Hamilton and Lashomb, 1996; Hare, 1990; Hitchner, et al., 2008; Hough-Goldstein, et al., 1993; Mitchell and Low, 1994)

  • Plant Foods
  • leaves

Predation

There are many known predators of Leptinotarsa decemlineata. Predators include arachnids (Phalangium opilio, a phalangid that eats eggs and small larvae; Xysticus kochi, a spider in USSR; Peucetia viridans, Misumena, 2 species in the family Thomisidae, spiders that eat eggs and larvae), Neuroptera (Chrysoperla carnea and Chrysoperla rufilabris, lacewings that eat eggs), Heteropterans (Perillus bioculatus, Podisus maculiventris, Oplomus dichrous, Oplomus severus, Stiretris anchorago, Perilloides confluens, Zicrona coerules, Pinthaeus sanguinipes, all stink bugs that eat larvae and eggs; Nabis roseipennis, Nabis alternatus, Geocoris punctipes, and species of Deraecoris that all prey on eggs), Coleoptera (Lebia grandis and at least 8 other species of Lebia that eat eggs, larvae, and pupae; Pterostichus chalcites and Calledia decora, which feed on larvae; Coleomegilla maculata, Hippodamia convergens, Coccinella septempunctata, Coccinella transversoguttata, Harmonia axyridis, and Aiolocaria miriabilis (in USSR), all Coccinellids that eat eggs and larvae; Collops quadrimaculatus, a Melyrid), Hymenoptera (wasps of Polistes that eat larvae; ants of genus Formica that eat adults and larvae).

In the United States, the lady beetle Coleomegilla maculata is a particularly significant predator that consumes eggs and small larvae of L. decemlineata. When present, C. maculata can kill up to 37.8% of eggs in the first generation and up to 58.1% of eggs for the second generation. Several species have been introduced into Leptinotarsa decemineata populations to suppress the substantial numbers. Predaceous stink bugs such as Perillus bioculatus and Podisus maculiventris attack beetle larvae, significantly decreasing the population by 62%, and reducing the destruction of foliage by 86%. Other beetles such as Lebia grandis feed on eggs and larvae of the Colorado potato beetle, while the larvae of the same species act as parasitoids on the pupae of Leptinotarsa decemlineata.

There is some evidence that the Colorado potato beetle produces a toxin, leptinotarsin, that protects the larvae and adults from predation. This is evidently not sequestered from the host plant, because no sequestered alkaloids have been found. The larvae and adults also have not been demonstrated to be aposematic. (Alyokhin, et al., 2008; Armer, 2004; Boiteau and McCarthy, 2010; Bruni, et al., 2000; Brust, 1994; Canas, et al., 2002; Coll, et al., 1994; Drummond and Casagrande, 1989; Drummond, et al., 1990; Gollands, et al., 1991; Greenstone, et al., 2010; Groden, et al., 1990; Hamilton and Lashomb, 1996; Hare, 1990; Hazzard, et al., 1991; Hillbeck and Kennedy, 1996; Hillbeck, et al., 1997; Hough-Goldstein and McPherson, 1996; Hough-Goldstein, et al., 1993; Hsiao and Fraenkel, 1969; Hu, et al., 1999; Ignoffo, et al., 1982; Klinger, et al., 2006; Long, et al., 1998; Lopez, et al., 1997; Lopez, et al., 1993; Matlock, 2005; Munyaneza and Obrycki, 1998; O'Neil, et al., 2005; Ramirez, et al., 2010; Saint-Cyr and Cloutier, 1996; Snyder and Clevenger, 2004; Weber, et al., 2006; Weber, 2012)

Ecosystem Roles

Leptinotarsa decemlineata feeds almost exclusively on Solanum plants, particularly the cultivated potato, Solanum tuberosum. Other suitable hosts include Solanum rostratum, Solanum augustifolium, Solanum dulcamara, Solanum melongena (eggplant), Lycopersicon esculentum (tomato), peppers, tobacco, Solanum carolinense, Solanum sarrachoides, Solanum elaeagnifolium, and Hyoscyamus niger. It is a major agricultural pest and has the potential to significantly defoliate its host plants.

Eggs, larvae, and adults of L. decemlineata can serve as hosts to a variety of parasites and parasitoids. Parasites of the Colorado potato beetle include a couple of mites, Chrysomelobia labidomerae, which feeds under the elytra, and Pyemotes tritici, the straw itch mite, which is an ectoparasite that causes paralysis and death within 2 to 7 days. At least two internal parasites are also known. Bacillus thuringiensis can be used as a control agent, killing larvae, and Beauveria bassiana, a fungus, infects larvae and adults. The parasitic wasp, Edovum puttleri, has been found to parasitize up to 71% to 91% of the eggs in a mass on eggplant hosts, killing 67% to 69% of the egg mass.

Parasitoids of Leptinotarsa decemlineata include species of Diptera (Myiopharus aberrans, Myiopharus australis, Myiopharus doryphorae, Myiopharus macella, all Tachinidae that are larval parasitoids, usually emerging from adults), Coleoptera (larvae of Lebia grandis act as parasitoids on pupae), and Hymenoptera (Edovum puttleri, a Eulophid egg parasitoid; and possibly Brachymeria truncatella, a chalcidoid wasp that may be a hyperparasitoid of the tachinids, and Anaphes fuscipennis, a Mymarid that parasitizes eggs). Leptinotarsa decemlineata is prey to many other insects, including species of Neuroptera, Heteroptera, Coleoptera, and Hymenoptera. Many arachnids also feed on L. decemlineata. (Alyokhin, et al., 2008; Drummond, et al., 1990; Gollands, et al., 1991; Groden, et al., 1990; Hamilton and Lashomb, 1996; Hare, 1990; Hough-Goldstein, et al., 1993; Hu, et al., 1999; Ignoffo, et al., 1982; Klinger, et al., 2006; Long, et al., 1998; Lopez, et al., 1997; Lopez, et al., 1993; O'Neil, et al., 2005; Weber, et al., 2006)

Species Used as Host
  • Potato, Solanum tuberosum
  • Solanum rostratum
  • Solanum augustifolium
  • Solanum dulcamara
  • Eggplant, Solanum melongena
  • Solanum carolinense
  • Solanum sarrachoides
  • Solanum elaeagnifolium
  • Hyoscyamus niger
  • Tomato, Lycopersicon esculentum
  • Peppers
  • Tobacco
Commensal/Parasitic Species

Economic Importance for Humans: Positive

There are no known positive effects of Leptinotarsa decemlineata on humans.

Economic Importance for Humans: Negative

Leptinotarsa decemlineata is considered one of the most serious pests and agricultural defoliators. The Colorado potato beetle causes significant damage to agricultural plants such as tomatoes, eggplants, and, of course, potatoes. Both larvae and adults feed on the foliage of host plants, skeletonizing the plant, leaving only roots and stems. Because of such devastation, insecticides have been implemented to decrease the destruction that Leptinotarsa decemlineata has on agricultural production, costing farmers millions of dollars each year. This species has been a huge pest problem throughout the country and is a problem annually for farmers. With such heavy insecticide use, populations of this species have developed resistance to nearly all classes of chemicals used as insecticides. It is estimated that this resistance costs growers between $44 and $69 per hectare each year, but no good alternatives are currently available. (Alyokhin, et al., 2008; University of Kentucky, 2010)

  • Negative Impacts
  • crop pest

Conservation Status

Leptinotarsa decemlineata is not listed as an endangered or threatened species on any local, state, national or international level.

Other Comments

Vast amounts of information is available for Leptinotarsa decemlineata because of its status as a major pest insect. The information provided here is a general introduction to the biology of Leptinotarsa decemlineata. Further information can be found using the references listed here and the many studies cited within those references.

Contributors

Brandon Bodnariuk (author), University of Michigan Biological Station, Brian Scholtens (author, editor), University of Michigan Biological Station, Angela Miner (editor), Animal Diversity Web Staff.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

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Neotropical

living in the southern part of the New World. In other words, Central and South America.

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Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

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agricultural

living in landscapes dominated by human agriculture.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

diapause

a period of time when growth or development is suspended in insects and other invertebrates, it can usually only be ended the appropriate environmental stimulus.

diurnal
  1. active during the day, 2. lasting for one day.
ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

folivore

an animal that mainly eats leaves.

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

hibernation

the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.

holarctic

a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.

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Found in northern North America and northern Europe or Asia.

internal fertilization

fertilization takes place within the female's body

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

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oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polarized light

light waves that are oriented in particular direction. For example, light reflected off of water has waves vibrating horizontally. Some animals, such as bees, can detect which way light is polarized and use that information. People cannot, unless they use special equipment.

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

suburban

living in residential areas on the outskirts of large cities or towns.

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

References

Alyokhin, A. 2008. "Colorado Potato Beetle Biology and Management" (On-line). PotatoBeetle.org. Accessed July 15, 2012 at http://www.potatobeetle.org/overview.html.

Alyokhin, A., M. Baker, D. Mota-Sanchez, G. Dively, E. Grafius. 2008. Colorado potato beetle resistance to insecticides. American Journal of Potato Research, 85: 395-413.

Armer, C. 2004. Colorado potato beetle toxins revisited: evidence the beetle does not sequester host plant glycoalkaloids. Journal of Chemical Ecology, 30: 883-888.

Boiteau, G., A. Alyokhin, D. Ferro. 2003. The Colorado potato beetle in movement. Canadian Entomologist, 135: 1-22.

Boiteau, G., P. McCarthy. 2010. Is there a role for stripes of adults and colour of larvae in determining the avoidance of the Colorado potato beetle by the American toad?. Canadian Journal of Zoology, 88: 468-478.

Bruni, R., J. Sant'Ana, J. Aldrich, F. Bin. 2000. Influence of host pheromone on egg parasitism by scelionid wasps: comparison of phoretic and nonphoretic parasitoids. Journal of Insect Behavior, 13: 165-173.

Brust, G. 1994. Natural enemies in straw-mulch reduce Colorado potato beetle populations and damage in potato. Biological Control, 4: 163-169.

Canas, L., R. O'Neil, T. Gibb. 2002. Population ecology of Leptinotarsa undecimlineata Stal (Coleoptera: Chrysomelidae): population dynamics, mortality factors, and potential natural enemies for biological control of the Colorado potato beetle. Biological Control, 24: 50-64.

Casagrande, R. 1987. The Colorado potato beetle: 125 years of mismanagement. Bulletin of the Entomological Society of America, 33: 142-150.

Coll, M., L. de Mendoza, G. Roderick. 1994. Population structure of a predatory beetle: the importance of gene flow for intertrophic level interactions. Heredity, 72: 228-236.

Dickens, J. 2006. Plant volatiles moderate response to aggregation pheromone in Colorado potato beetle. Journal of Applied Entomology, 130: 26-31.

Dickens, J. 1992. Sexual contact influences orientation to plant attraction in the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). Naturwissenschaften, 94: 847-852.

Dickens, J., J. Oliver, B. Holister, J. Davis, J. Klun. 2002. Breaking a paradigm: male-produced aggregation pheromone for the Colorado potato beetle. Journal of Experimental Biology, 205: 1925-1933.

Drummond, F., R. Casagrande. 1989. Effect of the straw itch mite on larvae and adults of the Colorado potato beetle. American Potato Journal, 66: 161-163.

Drummond, F., Y. Suhaya, E. Groden. 1990. Predation on the Colorado potato beetle (Coleoptera: Chrysomelidae) by Phalangium opilio (Opiliones: Phalangidae). Journal of Economic Entomology, 83: 772-778.

Fasulo, T. 2009. "University of Florida Institute of Food and Agricultural Studies" (On-line). Accessed July 15, 2012 at http://www.entemdept.ufl.edu/creatures/veg/leaf/potato_beetles.htm.

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Forgash, A. 1985. Insecticide resistence in the Colorado potato beetle. Research Bulletin, Massachusetts Agricultural Experiment Station, 704: 33-52.

Gollands, B., M. Tauber, C. Tauber. 1991. Seasonal cycles of Myiopharus aberrans and M. doryphorae (Diptera: Tachinidae) parasitizing Colorado potato beetles in upstate New York. Biological Control, 1: 153-163.

Greenstone, M., Z. Szendrei, M. Payton, D. Rowley, T. Coudron, D. Weber. 2010. Choosing natural enemies for conservation biological control: use of the prey detectability half-life to rank key predators of Colorado potato beetle. Entomologia Experimentalis et Applicata, 136: 97-107.

Groden, E., F. Drummond, R. Casagrande, D. Haynes. 1990. Coleomegilla maculata (Coleoptera: Coccinellidae): its predation upon the Colorado potato beetle (Coleoptera: Chrysomelidae) and its incidence in potatoes and surrounding crops. Journal of Economic Entomology, 83: 1306-1315.

Hamilton, G., J. Lashomb. 1996. Comparison of conventional and biological control intensive pest management programs on eggplant in New Jersey. Florida Entomologist, 79: 488-496.

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Hazzard, R., D. Ferro, R. Vandriesche, A. Tuttle. 1991. Mortality of eggs of Colorado potato beetle (Coleoptera, Chrysomelidae) from predation by Coleomegilla maculata (Coleoptera, Coccinelidae). Environmental Entomology, 20: 841-848.

Hillbeck, A., C. Eckel, G. Kennedy. 1997. Predation on Colorado potato beetle eggs by generalist predators in research and commercial potato plantings. Biological Control, 8: 191-196.

Hillbeck, A., G. Kennedy. 1996. Predators feeding on the Colorado potato beetle in insecticide-free plots and insecticide-treated commercial potato fields in eastern North Carolina. Biological Control, 6: 273-282.

Hitchner, E., T. Kuhar, J. Dickens, R. Youngman, P. Schultz, D. Pfeiffer. 2008. Host plant choice experiments of Colorado potato beetle (Coleoptera: Chrysomelidae) in Virginia. Journal of Economic Entomology, 101: 859-865.

Hough-Goldstein, J., G. Heimpel, H. Bechmann, C. Mason. 1993. Arthropod natural enemies of the Colorado potato beetle. Crop Protection, 12: 324-334.

Hough-Goldstein, J., D. McPherson. 1996. Comparison of Perillus bioculatus and Podisus maculiventris (Hemiptera: Pentatomidae) as potential control agents of the Colorado potato beetle (Coleoptera: Chrysomelidae). Journal of Economic Entomology, 89: 1116-1123.

Hsiao, T., G. Fraenkel. 1969. Properties of leptinotarsin a toxic hemolymph protein from the Colorado potato beetle. Toxicon, 7: 119-130.

Hu, J., D. Gelman, R. Bell. 1999. Effects of selected physical and chemical treatments of Colorado potato beetle eggs on host acceptance and development of the parasitic wasp, Edovum puttleri. Entomologia Experimentalis et Applicata, 90: 237-245.

Ignoffo, C., C. Garcia, M. Kroha. 1982. Susceptibility of the Colorado potato beetle Leptinotarsa decemlineata to Bacillus thuringiensis. Journal of Invertebrate Pathology, 39: 244-246.

Jolivet, P. 1991. The Colorado beetle menaces Asia (Leptinotarsa decemlineata Say) (Col. Chrysomelidae). L'Entomologiste, 47: 29-48.

Klinger, E., E. Groden, F. Drummond. 2006. Beauveria bassiana horizontal infection between cadavers and adults of the Colorado potato beetle, Leptinotarsa decemlineata (Say). Environmental Entomology, 35: 992-1000.

Kramer, M., D. Weber, Z. Szendrei. 2009. Habiat manipulation in potato affects Colorado potato beetle dispersal. Journal of Applied Entomology, 133: 711-719.

Long, D., F. Drummond, E. Groden. 1998. Susceptibility of Colorado potato beetle (Leptinotarsa decemlineata) eggs to Beauveria bassiana. Journal of Invertebrate Pathology, 71: 182-183.

Lopez, E., D. Ferro, R. Van Driesche. 1993. Direct measurement of host and parasitoid recruitment for assessment of total losses due to parasitism in the Colorado potato beetle Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) and Myiopharus doryphorae (Riley) (Diptera: Tachinidae). Biological Control, 3: 85-92.

Lopez, E., L. Roth, D. Ferro, D. Hosmer, A. Mafra-Neto. 1997. Behavioral ecology of Myiopharus doryphorae (Riley) and M. aberrans (Townsend), tachinid parasitoids of the Colorado potato beetle. Journal of Insect Behavior, 10: 49-78.

Matlock, R. 2005. Impact of prey size on prey capture success, development rate, and survivorship in Perillus bioculatus (Heteroptera: Pentatomidae), predator of the Colorado potato beetle. Environmental Entomology, 34: 1048-1056.

Mitchell, B., R. Low. 1994. The structure of feeding behavior in the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). Journal of Insect Behavior, 7: 707-29.

Munyaneza, J., J. Obrycki. 1998. Searching behavior of Coleomegilla maculata larvae feeding on Colorado potato beetle eggs. Biological Control, 13: 85-90.

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