BioKIDS home

Kids' Inquiry of Diverse Species

Harmonia axyridis

What do they look like?

Harmonia axyridis is oval shaped. It has three main color forms: red or orange with zero to many black spots (form succinea), or black with either four orange spots (form spectabilis) or two (form conspicua). Several less common forms exist as well. Darker forms are more common in its native range, with red or orange morphs more common in Europe and North America. The area behind the head is usually white with four black spots that form an "M" shape. The last segment on the body is different between males and females.

The eggs of H. axyridis are approximately 1.2 mm in length and yellow in color. Just before hatching, eggs darken to a gray or black color.

The larvae of H. axyridis have long bodies and are black with orange coloration (including two long orange lines on the middle of their backs, which start to develop after the first larval stage, and are complete by the last larval stage). They have spines along the length of the body. Larvae are longer in length than adults, growing as long as 7.5 to 10.7 mm in the later larval stages. Larvae look like tiny black and orange alligators.

Pupae are exposed and generally shaped like a long dome, often attached to leaves. They are usually orange, with the exoskeleton from the fourth larval stage still attached at the point of attachment. (Arnett, 1993; Banks, 1957; Gordon, 1985; Hagan, 1962; Hodek, 1996; Sasaji, 1971)

  • Sexual Dimorphism
  • sexes alike
  • Range length
    5 to 8 mm
    0.20 to 0.31 in

Where do they live?

Harmonia axyridis, the Asian lady beetle, is originally from the Oriental region, found in China, ranging to the far south (Yunnan and Guangxi Provinces), Japan, Korea, Mongolia, and parts of the Palearctic region, from northern Kazakhstan, and eastern Russia west to the Altai Mountains and north to Siberia. It has been introduced to at least four other continents. Its range covers most of North America; it is found in Mexico, across the United States (excluding Alaska and Wyoming) and much of southern Canada. This species has been found across South America, excluding areas near the Amazon river. Its habitat ranges across southern and western Europe into Bosnia and Herzegovina and Romania, and recently was found in Tunisia, Egypt, and South Africa. (Brown, et al., 2011; Osawa, 2000)

What kind of habitat do they need?

Harmonia axyridis can be found on many crop species on farm lands where it has been introduced. It is often found on deciduous trees, flowering plants, and other plant species found in open areas. This species tends to live in open fields, agricultural fields, and meadows. (Borror and White, 1970; Brown, et al., 2011; Hagan, 1962; Hodek, 1996; Koch, 2003)

How do they grow?

Harmonia axyridis is holometabolous, meaning it goes through complete metamorphosis. It develops from an egg through four larval stages (instars), to pupa and then adult. The diet of the beetle is known to have an effect on larval development and adult weight, as is temperature. Harmonia axyridis usually has two generations per season in much of the world, although up to four or five generations per year have been observed. Adults go into hiding over winter and start laying eggs in early spring, whenever average temperatures begin to reach 12°C. (El-Sebaey and El-Gantiry, 1999; Kindlmann, et al., 2000; Koch, 2003; LaMana and Miller, 1998; Osawa, 2000)

How do they reproduce?

Harmonia axyridis produces pheromones to attract mates, and at close distances may use sight to identify mates. Reproduction is sexual, with internal fertilization. Many females are picky when choosing male mates. Some decide on their mates based on the color of the male elytra (which cover the wings). This may have to do with the fact that certain colors provide better camouflage from predators. By choosing males of certain colors to mate with, offspring will have better camouflage and will be more likely to survive. H. axyridis is usually polygynandrous, meaning that these beetles mate many times with many different beetles. After mating, males do not do anything to defend the female from other males. (Hodek and Ceryngier, 2000; Koch, 2003; Stathas, et al., 2001)

Females of Harmonia axyridis will produce many eggs per season, laying about of 25 eggs per day. Females tend to lay egg clusters, with numbers ranging between 20 and 30 eggs per group. This species will breed many times during its lifetime. Females may lay unfertilized eggs that will not hatch along with the fertilized eggs. When larvae hatch, they eat the unfertilized eggs, which is useful in case other food is not available. (El-Sebaey and El-Gantiry, 1999; Hodek, 1996; Koch, 2003; LaMana and Miller, 1998; Perry and Roitberg, 2005)

  • How often does reproduction occur?
    Individuals will breed often and continuously over the duration of adulthood.
  • Breeding season
    Harmonia axyridis will start to breed as soon as temperatures increase beyond approximately 12°C (50°F).
  • Range eggs per season
    1642 to 3819

Females provide nutrients in the eggs for the offspring to use to grow and develop. Also, females may lay unfertilized eggs that will never hatch along with the fertilized eggs. Larvae can eat the unfertilized eggs after they hatch. Otherwise, H. axyridis does not provide any parental care. (Koch, 2003; Osawa, 1993; Stathas, et al., 2001)

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

How long do they live?

In its native range, Harmonia axyridis has two generations per year, but it has five generations in some places. Adults usually live 30 to 90 days, although some have lived over 3 years. (El-Sebaey and El-Gantiry, 1999; Koch, 2003; LaMana and Miller, 1998)

  • Range lifespan
    Status: wild
    3 (high) years
  • Typical lifespan
    Status: wild
    30 to 90 days

How do they behave?

Harmonia axyridis lives alone, but it will gather in large groups to hibernate over winter. Studies show that for many lady beetle species, they will go into hibernation once daylight hours decrease to between 10 and 13 hours a day, since days get shorter as winter approaches. H. axyridis is an active predator, often flying from plant to plant to find prey species. It moves around mostly during the day. (Banks, 1957; Biddinger, et al., 2009; Colunga-Garcia, et al., 1997; Hagan, 1962; Osawa, 2000)

Home Range

Harmonia axyridis does not generally do anything to defend its home from other lady beetles. Instead, it moves freely through its habitat. (Banks, 1957; Colunga-Garcia, et al., 1997; Hagan, 1962; Osawa, 2000; Banks, 1957; Colunga-Garcia, et al., 1997; Hagan, 1962; Osawa, 2000)

How do they communicate with each other?

Like many insects, Harmonia axyridis communicates with chemicals and pheromones. Many pheromones are used to find and communicate with mates. They also use these pheromones to find other lady beetles to spend the winter together. One pheromone that Harmonia axyridis uses is called harmonine. It is used for protection against predators. When H. axyridis is in danger, it produces harmonine and releases it from the joints in its legs, an act called reflex bleeding. H. axyridis finds prey by using sight or smell. (Colunga-Garcia, et al., 1997; Evans, 2003)

What do they eat?

Harmonia axyridis is an insectivore. It mainly eats aphids and scale insects. It sometimes also eats other insects in the order Thysanura, as well as the eggs of butterflies and moths. It will also eat mites. When these prey are not around, H. axyridis will eat other lady beetle species. It will also feed on grapes and other fruits, and very rarely eats pollen. Cannibalism also occurs in this species. Both adults and larvae of Harmonia axyridis will eat eggs and smaller larvae of its own species. (Adriano, et al., 2009; Alhmedi, et al., 2010; Berkvens, et al., 2010; Burgio, et al., 2002; Campbell and Cone, 1999; Davidson and Evans, 2010; Dixon, 2005; Evans, 2003; Naoya, 2011; Osawa, 2000; Pervez, 2006; Sloggett, 2008)

  • Animal Foods
  • insects
  • terrestrial non-insect arthropods
  • Plant Foods
  • fruit
  • pollen

What eats them and how do they avoid being eaten?

Many species of bird prey on Harmonia axyridis. Many other insect species also feed on H. axyridis. The pentatomid bug, Podisus maculiventris is known to prey on H. axyridis, as well as many species of ants, including the red imported fire ant, Solanopsis invicta. Other lady beetle species will prey on H. axyridis, although only when the other lady beetles are larger. Cannibalism on eggs and larvae is also very common in this species.

To defend itself, Harmonia axyridis produces isopropyl methoxy pyrazine (IPMP), which it secretes from its leg joints when threatened. This is a chemical defense, making it poisonous to predators. This species also has antimicrobial agents in its body that protect against bacteria and yeast. The red and black coloring of H. axyridis is called aposematic coloration. Predators know that insects with bright colors like these are usually poisonous, and do not attack them as often. The colors act as a warning signal to predators. (Firlej, et al., 2005; Gross, et al., 2010; Katsoyannos and Aliniazee, 1998; Koch, 2003; Nalepa and Weir, 2007; Nalepa, et al., 1996; Riddick, et al., 2009; Roy, et al., 2008; Saito and Bjørnson, 2008; Sloggett, et al., 2011)

  • These animal colors help protect them
  • aposematic

What roles do they have in the ecosystem?

Many parasites and parasitoids use the body of Harmonia axyridis to grow and develop in. These species include flies (Strongygaster triangulifer) and wasps (Dinocampus coccinellae, Oomyzus scaposus, Homalotylus terminalis, Pachyneuron altiscuta). Mites of the genus Coccipolipus, protozoans (such as Microsporidia), nematodes, and fungal species (Hesperomyces virescens, Beauveria bassiana) may also use the body of H. axyridis as a host.

In areas where it is not originally from, Harmonia axyridis competes with native lady beetle species for resources. H. axyridis usually out-competes these other species by using resources more efficiently, causing decreases in the population sizes of these other lady beetles. Some of the lady beetle species most affected are Coccinella transversoguttata, Adalia bipunctata, and Coccinella novemnotata. H. axyridis also preys on insects that eat crops and other plants, which helps prevent plant damage. (Banks, 1957; Gordon, 1985; Kindlmann, et al., 2011; Lombaert, et al., 2010; Sloggett, 2008)

Commensal or parasitic species (or larger taxonomic groups) that use this species as a host

Do they cause problems?

Since Harmonia axyridis will sometimes eat grapes and other fruits, it can cause significant economic damage on these crops. These lady beetles will crawl inside the grapes to feed, and when threatened by farmers or machinery collecting the grapes, they bleed toxic chemicals from the joints in their legs. These chemicals that are normally meant for predators cause a bad taste in the fruit. These chemicals can also be tasted in wine if H. axyridis has fed in the grapes used during the wine-making process. H. axyridis is also known to bite humans if handled. (Botezatu, et al., 2013; Galvan, et al., 2009)

  • Ways that these animals might be a problem for humans
  • injures humans
    • bites or stings
  • crop pest

How do they interact with us?

Since Harmonia axyridis mainly eats aphids and other pest insects that can do serious damage to crops, it can control the populations of these pest insects. This prevents damage to crops. Since they are also intense predators, they are able to survive in many habitats, making them a very useful population control species. (Alhmedi, et al., 2010; Campbell and Cone, 1999; Davidson and Evans, 2010; Dixon, 2005; Gardiner and Landis, 2007; Sloggett, 2008)

  • Ways that people benefit from these animals:
  • controls pest population

Are they endangered?

Harmonia axyridis is not an endangered species.


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


Adriano, G., N. Vandenberg, J. McHugh, J. Forrester, S. Slipinski, K. Miller, L. Shapiro, M. Whiting. 2009. The evolution of food preferences in Coccinellidae. Biological Control, 51(2): 215-231.

Alhmedi, A., E. Haubruge, F. Francis. 2010. Intraguild interactions and aphid predators: biological efficiency of Harmonia axyridis and Episyrphus baltetus. Journal of Applied Entomology, 134(1): 34-44.

Alyokhin, A., K. Landry, C. Finlayson. 2008. Abundance of native and non-native lady beetles (Coleoptera: Coccinellidae) in different habitats in Maine. Annals of the Entomological Society of America, 101(6): 1078-1087.

Alyokhin, A., G. Sewell. 2004. Changes in a lady beetle community following the establishment of three alien species. Biological Invasion, 6: 463-471.

Arnett, R. 1993. American insects: a handbook of the insects of America north of Mexico. Boca Raton, FL, USA: CRC Press.

Banks, C. 1957. The behaviour of individual coccinellid larvae on plants. The British Journal of Animal Behaviour, 5(1): 12-24.

Berkvens, N., J. Bale, D. Berkvens, L. Tirry, P. De Clercq. 2010. Cold tolerance of the harlequin ladybird Harmonia axyridis in Europe. Journal of Insect Physiology, 56(4): 438-444.

Biddinger, D., D. Weber, L. Hull. 2009. Coccinellidae as predators of mites: Stethorini in biological control. Biological Control, 51: 268-283.

Borror, D., R. White. 1970. Peterson Field Guides: A Field Guide to the Insects of America North of Mexico. Boston, MA, USA.: Houghton Mifflin.

Botezatu, A., Y. Kotseridis, D. Inglis, G. Pickering. 2013. Occurrence and contribution of alkyl methoxypyrazines in wine tainted by Harmonia axyridis and Coccinella septempunctata. Journal of the Science of Food and Agriculture, 93/4: 803-810.

Brown, P., C. Thomas, E. Lombaert, D. Estoup, A. Jefferies, L. Lawson Handley. 2011. The global spread of Harmonia axyridis (Coleoptera: Coccinellidae): distribution, dispersal and routes of invasion. Biocontrol, 56(4): 623-641.

Burgio, G., F. Santi, S. Maini. 2002. On intra-guild predation and cannibalism in Harmonia axyridis (Pallas) and Adalia bipunctata L. (Coleoptera: Coccinellidae). Biological Control, 24(2): 110-116.

Campbell, C., W. Cone. 1999. Consumption of damson-hop aphids (Phorodon humuli) by larvae of Coccinella transversoguttata and Hippodamia convergens (Coleoptera: Coccinellidae). Biocontrol Science and Technology, 9(1): 75-78.

Castro, C., L. Almeida, S. Penteado. 2011. The impact of temperature on biological aspects and life table of Harmonia axyridis (Pallas). Florida Entomologist, 94(4): 923-932.

Colunga-Garcia, M., S. Gage, D. Landis. 1997. Response of an assemblage of Coccinellidae (Coleoptera) to a diverse agricultural landscape. Entomological Society of America, 26(4): 797-804.

Cottrell, T., K. Yeargan. 1998. Intraguild predation between an introduced lady beetle, Harmonia axyridis (Coleoptera: Coccinellidae), and a native lady beetle, Coleomegilla maculata (Coleoptera: Coccinellidae). Journal of the Kansas Entomological Society, 71(2): 159-163.

Davidson, L., E. Evans. 2010. Frass analysis of diets of aphidophagous lady beetles (Coleoptera: Coccinellidae) in Utah alfalfa fields. Environmental Entomology, 39(2): 576-582.

Day, W., K. Tatman. 2006. Changes in abundance of native and adventive Coccinellidae (Coleoptera) in alfalfa fields, in northern New Jersey (1993-2004) and Delaware (1999-2004), USA. Entomological News, 117(5): 491-502.

Dixon, A. 2005. Insect predatory-prey dynamics: ladybird beetles and biological control. Cambridge, UK: Cambridge University Press.

El-Sebaey, I., A. El-Gantiry. 1999. Biological aspects and description of different stages of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). Bulletin of the Faculty of Agriculture, Cairo University, 50: 87-97.

Elliott, N., R. Kieckhefer, W. Kauffman. 1996. Effects of an invading coccinellid on native coccinellids in an agricultural landscape. Oecologia, 105(4): 537-544.

Evans, E. 2003. Searching and reproductive behavior of female aphidophagous ladybirds (Coleoptera: Coccinellidae): a review. European Journal of Entomology, 100(1): 1-10.

Fauske, G., P. Tinerella, D. Rider. 2003. A list of the ladybeetles (Coleoptera: Coccinellidae) of North Dakota with new records from North Dakota and Minnesota. Journal of the Kansas Entomological Society, 76: 38-46.

Firlej, A., G. Boivin, E. Lucas, D. Coderre. 2005. First report of Harmonia axyridis Pallas being attacked by Dinocampus coccinellae Schrank in Canada. Biological Invasion, 7: 553-556.

Folukemi, A., P. Yong-Lak. 2010. Visual and olfactory preference of Harmonia axyridis (Coleoptera: Coccinellidae) adults to various companion plants. Journal of Asia-Pacific Entomology, 13(4): 319-323.

Galvan, T., E. Burkness, R. Koch, W. Hutchinson. 2009. Multicolored Asian lady beetle (Coleoptera: Coccinellidae) activity and wine grape phenology: Implications for pest management. Environmental Entomology, 38(6): 1563-1574.

Gardiner, M., D. Landis. 2007. Impact of intraguild predation by adult Harmonia axyridis (Coleoptera: Coccinellidae) on Aphis glycines (Hemiptera: Aphididae) biological control in cage studies. Biological Control, 40: 386-395.

Gordon, R. 1985. The Coccinellidae of America north of Mexico. Journal of the New York Entomological Society, 93: 1-912.

Gross, J., A. Eben, I. Mueler. 2010. A well protected intruder: the effective antimicrobial defense of the invasive ladybird Harmonia axyridis. Journal of Chemical Ecology, 36(11): 1180-1188.

Hagan, K. 1962. Biology and ecology of predaceous Coccinellidae. Annual Review of Entomology, 7: 289-326.

Harwood, J., C. Ricci, R. Romani, J. Obrycki. 2006. Historic prevalence of a laboulbenialean fungus infecting introduced coccinellids in the United States. Antenna, 30: 74-79.

Hesler, L., M. Catangui, J. Losey, J. Helbig, A. Mesman. 2009. Recent records of Adalia bipunctata (L.), Coccinella transversoguttata richardsoni Brown, and Coccinella novemnotata Herbst (Coleoptera: Coccinellidae) from South Dakota and Nebraska. Coleopterists Bulletin, 63(4): 475-484.

Hesler, L., R. Kieckhefer. 2008. An annotated and updated species list of the Coccinellidae (Coleoptera) of South Dakota. Coleopterists Bulletin, 62(3): 443-454.

Hodek, I. 1996. Ecology of Coccinellidae. New York, NY: Dordecht, Kluwever Academic.

Hodek, I., P. Ceryngier. 2000. Sexual activity in Coccinellidae (Coleoptera): a review. European Journal of Entomology, 97: 449-456.

Husband, R. 1984. The taxonomic position of Coccipolipus (Acarina: Podapolipidae), a genus of mites which are parasites of ladybird beetles (Coccinellidae). Acarology, 1: 328-336.

Kajita, Y., E. Evans. 2009. Ovarian dynamics and oosorption in two species of predatory lady beetles (Coleoptera: Coccinellidae). Physiological Entomology, 34(2): 185-194.

Kajita, Y., E. Evans. 2010. Relationships of body size, fecundity, and invasion success among predatory lady beetles (Coleoptera: Coccinellidae) inhabiting alfalfa fields. Annals of the Entomological Society of America, 103(5): 750-756.

Kajita, Y., E. Evans, H. Yasuda. 2009. Reproductive responses of invasive and native predatory lady beetles (Coleoptera: Coccinellidae) to varying prey availability. Environmental Entomology, 38(4): 1283-1292.

Katsoyannos, P., M. Aliniazee. 1998. First record of Strongygaster triangulifera (Loew) (Diptera: Tachinidae) as a parasitoid of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in western North America. The Canadian Entomologist, 130: 905-906.

Kindlmann, P., O. Ameixa, A. Dixon. 2011. Ecological effects of invasive alien species on native communities, with particular emphasis on the interactions between aphids and ladybirds. Biocontrol, 56(4): 469-476.

Kindlmann, P., H. Yasuda, S. Sato, K. Shinya. 2000. Key life stages of two predatory ladybird species (Coleoptera: Coccinellidae). European Journal of Entomology, 97: 495-499.

Koch, R. 2003. The multicolored Asian lady beetle, Harmonia axyridis: a review of its biology, uses in biological control, and non-target impacts. Journal of Insect Science, 32(3): 1-16.

Koch, R., W. Hutchinson. 2003. Phenology and blacklight trapping of the multicolored Asian lady beetle (Coleoptera: Coccinellidae) in a Minnesota agricultural landscape. Journal of Entomological Science, 38: 477-480.

Koch, R., W. Hutchinson, R. Venette, G. Heimpel. 2003. Susceptibility of immature monarch butterfly, Danaus plexippus (Lepidoptera: Nymphalidae: Danainae), to predation by Harmonia axyridis (Coleoptera: Coccinellidae). Biological Control, 28(2): 265-270.

LaMana, M., J. Miller. 1998. Temperature-dependent development in an Oregon population of Harmonia axyridis (Coleoptera: Coccinellidae). Environmental Entomology, 27: 1001-1005.

Lombaert, E., T. Guillemaud, J. Comuet, T. Malausa, B. Facon, A. Estoup. 2010. Bridgehead effect in the worldwide invasion of the biocontrol harlequin ladybird. Public Library of Science, 5: e9743. Accessed July 12, 2012 at

Lundgren, J., M. Seagraves. 2011. Physiological benefits of nectar feeding by a predatory beetle. Biological Journal of the Linnean Society, 104(3): 661-669.

Majerus, M., I. Geoghegan, T. Majerus. 2000. Adaptive preferential selection of young coccinellid hosts by the parasitoid wasp Dinocampus coccinellae (Hymenoptera: Braconidae). European Journal of Entomology, 97: 161-164.

Majerus, T., M. Majerus, B. Knowles, J. Wheeler, D. Bertrand, V. Kuznetzov, H. Ueno, G. Hurst. 1998. Extreme variation in the prevalence of inherited male-killing microorganisms between three populations of Harmonia axyridis (Coleoptera: Coccinellidae). Heredity, 81: 683-691.

Michaud, J. 2000. Development and reproduction of ladybeetles (Coleoptera: Coccinellidae) on the citrus aphids Aphis spiraecola Patch and Toxoptera citricida (Kirkaldy) (Homoptera: Aphididae). Biological Control, 18(3): 287-297.

Nalepa, C., K. Kidd, K. Ahlstrom. 1996. Biology of Harmonia axyridis (Coleoptera: Coccinellidae) in winter aggregations. Annals of the Entomological Society of America, 89(5): 681-685.

Nalepa, C., A. Weir. 2007. Infection of Harmonia axyridis (Coleoptera: Coccinellidae) by Hesperomyces virescens (Ascomycetes: Laboulbeniales): role of mating status and aggregation behavior. Journal of Invertebrate Pathology, 94: 196-203.

Naoya, O. 2011. Ecology of Harmonia axyridis in natural habitats within its native range. Biocontrol, 56(4): 623-641.

Obrycki, J., T. Kring. 1998. Predaceous Coccinellidae in biological control. Annual Review of Entomology, 43: 295-301.

Obrycki, J., M. Tauber. 1981. Phenology of 3 coccinellid species (Coleoptera: Coccinellidae) Adalia bipunctata, Coccinella transversoguttata, Coccinella septempuncatata – thermal requirements for development. Annals of the Entomological Society of America, 74(1): 31-36.

Okuda, T., P. Ceryngier. 2000. Host discrimination in Dinocampus coccinellae (Hymenoptera: Braconidae), a solitary parasitoid of coccinellid beetles. Applied Entomology and Zoology, 35: 535-539.

Osawa, N. 1993. Population field studies of the aphidophagous ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae): life tables and key factor analysis. Researches on Population Ecology, 35: 335-348.

Osawa, N. 2000. Population field studies on the aphidophagous ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae): resource tracking and population characteristics. Population Ecology, 42(2): 115-127.

Perry, J., B. Roitberg. 2005. Ladybird mothers mitigate offspring starvation risk by laying trophic eggs. Behavioral Ecology and Sociobiology, 58(6): 578-586.

Pervez, O. 2006. Ecology and biological control application of multicoloured Asian ladybird, Harmonia axyridis: a review. Biocontrol Science and Technology, 16(2): 111-128.

Riddick, E., T. Cottrell, K. Kidd. 2009. Natural enemies of the Coccinellidae: parasites, pathogens, and parasitoids. Biological Control, 51(2): 306-312.

Roehrich, C., C. Ngwa, J. Wiesner, H. Schmidtberg, T. Degenkolb, C. Kollewe, R. Fischer, G. Pradel, A. Vilcinskas. 2012. Harmonine, a defence compound from the harlequin ladybird, inhibits mycobacterial growth and demonstrates multi-stage antimalarial activity. Biology Letters, 8(2): 308-311.

Roy, H., P. Brown, P. Rothery, R. Ware, M. Majerus. 2008. Interactions between the fungal pathogen Beauveria bassiana and three species of coccinellid: Harmonia axyridis, Coccinella septempunctata and Adalia bipunctata. Biocontrol, 53: 265-276.

Ruan, C., D. Wen-Mei, W. Xiu-Mei, Z. Jun-Jie, Z. Lian-Sheng. 2012. Effect of long-term cold storage on the fitness of pre-wintering Harmonia axyridis (Pallas). Biocontrol, 57(1): 95-102.

Saito, T., S. Bjørnson. 2008. Effects of a microsporidium from the convergent lady beetle, Hippodamia convergens Guérin–Méneville (Coleoptera: Coccinellidae), on three non-target coccinellids. Journal of Invertebrate Pathology, 99(3): 294-301.

Sasaji, H. 1971. Coccinellidae. Tokyo: Academic Press of Japan.

Shapiro-Ilan, D., T. Cottrell. 2005. Susceptibility of lady beetles (Coleoptera: Coccinellidae) to entomopathogenic nematodes. Journal of Invertebrate Pathology, 89: 150-156.

Sloggett, J. 2008. Weighty matters: Body size, diet and specialization in aphidophagous ladybird beetles (Coleoptera: Coccinellidae). European Journal of Entomology, 105: 381-389.

Sloggett, J., A. Magro, F. Verheggen. 2011. Chemical ecology of Harmonia axyridis. Biocontrol, 56(4): 643-661.

Sloggett, J., M. Majerus. 2000. Habitat preferences and diet in the predatory Coccinellidae (Coleoptera): an evolutionary perspective. Biological Journal of the Linnean Society, 70(1): 63-88.

Stathas, G., P. Eliopoulos, D. Kontodimas, J. Giannopapas. 2001. Parameters of reproductive activity in females of Harmonia axyridis (Coleoptera: Coccinellidae). European Journal of Entomology, 98: 547-549.

Stathas, G., D. Kontodimas, F. Karamaouna, S. Kampouris. 2011. Thermal requirements and effect of temperature and prey on the development of the predator Harmonia axyridis. Environmental Entomology, 40(6): 1541-1545.

Turnock, W., I. Wise, F. Matheson. 2003. Abundance of some native coccinellines (Coleoptera: Coccinellidae) before and after the appearance of Coccinella septempunctata. Canadian Entomologist, 135(3): 391-404.

Yasuda, H., E. Evan, Y. Kajita, K. Urakawa, T. Takizawa. 2004. Asymmetric larval interactions between introduced and indigenous ladybirds in North America. Oecologia, 141(4): 722-731.

University of Michigan Museum of ZoologyNational Science Foundation

BioKIDS home  |  Questions?  |  Animal Diversity Web  |  Cybertracker Tools

Graves, D. 2013. "Harmonia axyridis" (On-line), Animal Diversity Web. Accessed April 21, 2024 at

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.
Copyright © 2002-2024, The Regents of the University of Michigan. All rights reserved.

University of Michigan