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Leptinotarsa decemlineata

What do they look like?

Adults of Leptinotarsa decemlineata are oval shaped, and can be 8 to 10 mm in length. They have five thick brown stripes running front to back on each elytra, which cover the wings. The thorax, which is between the head and body, is orange with a pattern of black spots. Antennae are shorter than its body. Larvae typically have a row of black spots down the side of the body, which is very large and plump, when compared to the head and thorax. Eggs look like orange or yellow footballs. They are about 1.7 to 1.8 mm long and 0.8 mm wide. (Alyokhin, 2008; University of Kentucky, 2010)

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

Where do they live?

Leptinotarsa decemlineata, the Colorado potato beetle, is currently found throughout North America east of the Rockies, as well as some of Europe and Asia. Originally, Leptinotarsa decemlineata was found only in the southwestern United States into Mexico. As potatoes were planted in more and more places for farming, the Colorado potato beetle followed the potatoes that it eats, spreading through farm land in North America, Europe, and Asia. In the future, L. decemlineata could also expand its habitat into Korea, Japan, parts of Africa, and much of the Southern Hemisphere. (Alyokhin, et al., 2008; Alyokhin, 2008; Jolivet, 1991; Vlasova, 1978; Worner, 1988)

What kind of habitat do they need?

The Colorado potato beetle lives in farm fields where crops in the family Solanaceae grow, such as potatoes, tomatoes, tobacco, eggplants and peppers. It can also be found on plants in the Solanaceae family that are not grown on farms, but grow in the wild in open grasslands. (Alyokhin, et al., 2008; Casagrande, 1987; Kramer, et al., 2009)

How do they grow?

Leptinotarsa decemlineata goes through complete metamorphosis, with egg, larva, pupa, and adult stages. Adults of Leptinotarsa decemlineata that are still alive when winter arrives move to the edge of fields and bury themselves in the soil. They stay there throughout the winter. Adults then emerge again in spring, and walk the short distance to the crop fields. They eat and lay eggs within 5 to 6 days. Development from egg to adult can take from 14 to 56 days, depending on the temperature. After the eggs hatch, the larvae go through four stages, called instars, where they constantly eat the host plant. After about 21 days, larvae drop off the plant to the ground, and burrow 2 to 3 cm into the ground. After about 2 days they pupate, and then come out of the ground about 5.8 days later as adults. Adults then mate and lay eggs to complete another generation of beetles. By the middle of summer, 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)

How do they reproduce?

Both males and females of Leptinotarsa decemlineata mate with many other beetles over their lives. Adults usually mate before going into hiding for the winter. Females can store the male sperm in their bodies during winter, and then use it to fertilize their eggs in the spring. However, females will also sometimes mate after emerging in the spring, even before walking to the crop fields. Males have hairs on their legs that they use to hold on to females while mating. However, these hairs make it harder to hold on to host plants, and females are better at holding on to host plants to feed. (Alyokhin, et al., 2008; Roderick, et al., 2003; Voigt, et al., 2008)

After spending the winter in crop fields and gardens, the Colorado potato beetle becomes active in the spring, often in May. Adults eat for a few days, and then lay eggs. Females can lay up to 300 to 800 eggs, which they lay on the under side of the plant leaves. Eggs are laid together in groups of 10 to 30. Eggs may be laid over several weeks. (Alyokhin, et al., 2008; Ferro, et al., 1991)

  • How often does reproduction occur?
    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 provides nutrients in the eggs for the offspring to grow and develop. Otherwise, adults provide no more parental care.

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

How long do they live?

Adults of Leptinotarsa decemlineata live for a few weeks during the summer after emerging from pupation. Adults that are still alive at the end of summer go into hiding for the winter, so these adults have a longer lifespan, though they are mostly inactive during the winter. A few adults even remain in hiding for a second or even third winter. (Alyokhin, et al., 2008; Kramer, et al., 2009)

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

How do they behave?

Adults of Leptinotarsa decemlineata can fly, but the amount they fly depends on a few things. Beetles that emerge in spring after hiding for the winter may fly to a new location. When there are many beetles gathered together, some beetles will also fly to new locations, as far away as several kilometers away. This increases the beetles chances of finding new host plants. Females that have mated do not fly or move as often, staying in areas where there are host plants. Males that have mated move around more than females, to try and find new mates.

While in a crop field, Leptinotarsa decemlineata moves freely throughout its environment. These beetles will both walk and fly for short distances. Adults and larvae do not move around much at night, with most of their feeding taking place during the day. Larvae and adults often gather in large numbers on host plants while they feed. When threatened, larvae do certain behaviors such as walking away, standing up, regurgitating food, wiggling, and defecating. (Alyokhin, et al., 2008; Hazzard, et al., 1991; Ramirez, et al., 2010)

Home Range

This species does not have a specific territory and does not do anything to defend it. They move freely in areas of host plants.

How do they communicate with each other?

To communicate with other beetles and to view its habitat, Leptinotarsa decemlineata uses sight and smell. To find host plants, it detects chemicals produced by the plants. Vision is important to identify plants and mates, and these beetles can sense ultraviolet light, as well as the visible colors that human see. They also can probably see polarized light. After a male beetle has found a host plant to eat, it releases a pheromone that attracts other males and females to the same plant. It is also used to attract potential mates. Larvae can also detect this pheromone, and use it to gather in groups to feed. Females produce a different pheromone to attract male mates. (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)

What do they eat?

Leptinotarsa decemlineata eats Solanum plants. It eats all the leaves, leaving only the roots and stems. L. decemlineata mostly eats the potato, Solanum tuberosum, which gives the beetle its common name, the Colorado potato beetle. This species originally ate Solanum rostratum and Solanum augustifolium. Other plants that it eats include Solanum melongena (eggplant), Lycopersicon esculentum (tomato), peppers, tobacco, and other wild hosts such as Solanum dulcamara, S. carolinense, S. sarrachoides, S. elaeagnifolium, and Hyoscyamus niger. (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

What eats them and how do they avoid being eaten?

There are many known predators that eat Leptinotarsa decemlineata. Many arachnids, including spiders and daddy long legs eat the eggs and larvae (Phalangium opilio, Xysticus kochi, Peucetia viridans, Misumena, 2 species in the family Thomisidae). Lacewings eat the eggs (Chrysoperla carnea and Chrysoperla rufilabris). Wasps of Polistes eat larvae, and ants of genus Formica eat adults and larvae.

True bugs, including stink bugs, eat the larvae and eggs (Perillus bioculatus, Podisus maculiventris, Oplomus dichrous, Oplomus severus, Stiretris anchorago, Perilloides confluens, Zicrona coerules, Pinthaeus sanguinipes, Nabis roseipennis, Nabis alternatus, Geocoris punctipes, and species of Deraecoris). Many beetles, including lady beetles, eat all life stages of the Colorado potato beetle (Lebia grandis and at least 8 other species of Lebia, Pterostichus chalcites, Calledia decora, Coleomegilla maculata, Hippodamia convergens, Coccinella septempunctata, Coccinella transversoguttata, Harmonia axyridis, and Aiolocaria miriabilis (in USSR), Collops quadrimaculatus).

In the United States, the lady beetle Coleomegilla maculata is a significant predator that eats the eggs and small larvae of L. decemlineata. C. maculata can kill up to 58.1% of eggs in a generation. Since L. decemlineata can destroy the plants that it feeds on, many species have been used to decrease the population size. Predaceous stink bugs such as Perillus bioculatus and Podisus maculiventris attack beetle larvae, and can decrease the population by 62%. Other beetles such as Lebia grandis feed on eggs and larvae of the Colorado potato beetle, while the larvae of Lebia grandis act as parasitoids on the pupae of Leptinotarsa decemlineata.

The Colorado potato beetle may produce a toxic substance, leptinotarsin, that protects the larvae and adults from predators. Unlike some other beetles, L. decemlineata does not get this toxic substance from the plants it eats. (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)

What roles do they have in the ecosystem?

Leptinotarsa decemlineata eats Solanum plants, especially the potato, Solanum tuberosum. It can also eat 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 crop pest, since it damages the plants by eating and removing the leaves.

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 causes paralysis and death within 2 to 7 days. Bacillus thuringiensis can be used to control population size of the Colorado potato beetle, since it kills larvae, and Beauveria bassiana, a fungus, infects larvae and adults.

Parasitoids are similar to parasites, but kill their host, while parasites keep their host alive. Many insects act as parasitoids of Leptinotarsa decemlineata. These include species of flies (Myiopharus aberrans, Myiopharus australis, Myiopharus doryphorae, Myiopharus macella, all Tachinidae that are parasitoids on larvae, usually emerging from adults), beetles (larvae of Lebia grandis act as parasitoids on pupae), and wasps (Edovum puttleri, an egg parasitoid, Brachymeria truncatella, and Anaphes fuscipennis, which parasitizes eggs). Leptinotarsa decemlineata is prey to many other insects, including species of lacewings, true bugs, beetles, and wasps and ants. 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 (or larger taxonomic groups) used as hosts by this species
  • 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 or parasitic species (or larger taxonomic groups) that use this species as a host

Do they cause problems?

Leptinotarsa decemlineata is considered one of the most important crop pests. Since the Colorado potato beetle feeds almost entirely on crop plants and removes the leaves when eating, it can do serious damage to a large area of crops. These crops include tomatoes, eggplants, and, of course, potatoes. Both larvae and adults feed on the leaves of host plants, leaving only roots and stems. To stop the damage, farmers use insecticides to kill the beetles, which costs farmers millions of dollars each year. This species has been a huge pest problem throughout the country and is a problem each year for farmers. Because farmers have used so much insecticide, some groups of the Colorado potato beetle have become resistance to the insecticides, meaning that the insecticides no longer kill these populations. This creates a new problem, as these populations continue to reproduce, producing more beetles that can not be killed by the traditional insecticides, which means farmers have to develop new ways to fight the crop damage these beetles cause. (Alyokhin, et al., 2008; University of Kentucky, 2010)

  • Ways that these animals might be a problem for humans
  • crop pest

How do they interact with us?

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

Are they endangered?

Leptinotarsa decemlineata is not an endangered species.

Some more information...

Huge amounts of information is available for Leptinotarsa decemlineata because it is a major crop pest. The information here is an introduction to the biology of Leptinotarsa decemlineata. Further information can be found using the references listed here.

Contributors

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

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.

Fernandez, P., M. Hilker. 2007. Host plant location by Chrysomelidae. Basic and Applied Ecology, 8: 97-116.

Ferro, D., A. Tuttle, D. Weber. 1991. Ovipositional and flight behavior of overwintered Colorado potato beetle (Coleoptera: Chrysomelidae). Environmental Entomology, 20: 1309-1314.

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.

Hammock, J., B. Vinyard, J. Dickens. 2007. Response to host plant odors and aggregation pheromone by larvae of the Colorado potato beetle on a servosphere. Arthropod-Plant Interactions, 1: 27-35.

Hare, J. 1990. Ecology and management of the Colorado potato beetle. Annual Review of Entomology, 35: 81-100.

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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Oliver, J., J. Dickens, T. Glass. 2002. (S)-3,7-dimethyl-2-oxo-6-octene-1,3-diol: an aggregation pheromone of the Colorado potato beetle, Leptinotarsa decemlineata (Say). Tetrahedron Letters, 43: 2641-2643.

Otalora-Luna, F., J. Dickens. 2010. Spectral preference and temporal modulation of photic orientation by Colorado potato beetle on a servosphere. Entomologia Experimentalis et Applicata, 138: 93-103.

Ramirez, R., D. Crowder, G. Snyder, M. Strand, W. Snyder. 2010. Antipredator behavior of Colorado potato beetle larvae differes by instar and attacking predator. Biological Control, 53: 230-237.

Roderick, G., L. De Mendoza, G. Dively, P. Follett. 2003. Sperm precedence in Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae): temporal variation assessed by neutral markers. Annals of the Entomological Society of America, 96: 631-636.

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Snyder, W., G. Clevenger. 2004. Negative dietary effects of Colorado potato beetle eggs for the larvae of native and introduced ladybird beetles. Biological Control, 31: 353-361.

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Bodnariuk, B. and B. Scholtens 2013. "Leptinotarsa decemlineata" (On-line), Animal Diversity Web. Accessed July 23, 2014 at http://www.biokids.umich.edu/accounts/Leptinotarsa_decemlineata/

BioKIDS is sponsored in part by the Interagency Education Research Initiative. It is a partnership of the University of Michigan School of Education, University of Michigan Museum of Zoology, and the Detroit Public Schools. This material is based upon work supported by the National Science Foundation under Grant DRL-0628151.
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