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Harmonia axyridis

By Dylan Graves

Geographic Range

Harmonia axyridis, the Asian lady beetle, is native to 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 the majority 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 widely across South America, excluding Amazonian areas. 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)

Habitat

Harmonia axyridis can be found on many crop species in agricultural areas 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 areas, and meadows. (Borror and White, 1970; Brown, et al., 2011; Colunga-Garcia, et al., 1997; Hagan, 1962; Hodek, 1996; Koch, 2003)

Physical Description

Harmonia axyridis is oval shaped and convex. It is a polymorphic species, with three main color morphs: 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 morphs exist as well. Darker morphs are more common in its native range, with red or orange morphs more common in Europe and North America. The pronotum is generally white with four black spots that tend to form an "M" shape. This species has very distinct, square "shoulders." The last ventral segment on its abdomen differs between male and female individuals.

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 coloration.

The larvae of H. axyridis have long bodies and are black with orange coloration (including two long orange lines on the dorsal-lateral areas of the abdomen, which start to develop after the first instar and are complete by the fourth and final instar). They have double-branched spines (scoli) running down the length of the body. The larval stages often grow longer than the final adult individual, starting at an approximate length of 1.9 to 2.1 mm in the first instar and growing to 7.5 to 10.7 mm in the fourth instar. Larvae vaguely resemble tiny black and orange alligators.

Pupae are exposed and generally shaped like an elongated dome, often attached to leaves. They tend to be orange, with the fourth instar exoskeleton still attached at the point of substrate 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

Development

Harmonia axyridis is holometabolous, progressing from egg through four larval instars, to pupa and then adult. The mean duration of each immature stage is as follows: egg - 2.8 days, first instar - 2.5 days, second instar - 1.5 days, third instar - 1.8 days, fourth instar - 4.4 days, pupa - 4.5 days. The diet of the beetle is also known to have an effect on larval development and adult weight, as is temperature. Harmonia axyridis is generally considered bivoltine in much of the world, although up to four or five generations per year have been observed. Adults diapause 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)

Reproduction

Harmonia axyridis secretes pheromones to attract mates and at close distances may use sight. Reproduction is sexual, with internal fertilization of the ova. Studies show that many females are selective of male mates, often deciding on their mates based on the color of male elytra. This seems to be a selective preference, as certain morphs tend to evade predation more effectively. Even so, the species is generally polgynandrous, with individuals ultimately mating many times with many different individuals. After mating, males do not exhibit any apparent defense mechanisms to ensure the eggs are fertilized. (Hodek and Ceryngier, 2000; Koch, 2003; Stathas, et al., 2001)

Females of Harmonia axyridis will produce many eggs per season, averaging an approximate rate of 25 eggs per day. Individuals tend to lay egg clusters, with numbers ranging between 20 and 30 eggs per cluster. This species will breed continuously during its lifetime. Breeding is contingent on temperature, with lower temperatures increasing pre-mating and pre-oviposition periods. Females may lay unfertilized eggs along with the fertilized eggs, which are used by larvae as food sources in conditions where aphids and scale insects are scarce. (El-Sebaey and El-Gantiry, 1999; Hodek, 1996; Koch, 2003; LaMana and Miller, 1998; Perry and Roitberg, 2005)

  • Breeding interval
    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

There is virtually no parental investment in this species aside from females depositing nutrients in eggs. Additionally, females may lay unfertilized eggs along with the fertilized eggs, ensuring a food source for the larvae upon hatching. Otherwise, larval individuals are independent, surviving on their own after hatching. (Koch, 2003; Osawa, 1993; Stathas, et al., 2001)

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

Lifespan/Longevity

In its native range, Harmonia axyridis generally has two generations per year, but in some places five generations have been observed. Adults tend to live 30 to 90 days, contingent on temperature, although some have lived over three years. Whichever generation receives cues to diapause will generally live through winter, with reproduction then occurring in spring. (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

Behavior

Harmonia axyridis is a solitary species, although they often congregate over winter to hibernate. While no studies have been done on this species specifically, studies on other Coccinellidae species show that the main cue for diapause occurs once daylight hours drop below approximately 10 to 13 hours per day. It is a mobile predator, moving through its environment seeking its prey, which reside on plants. This species is mainly active during the day, and often flies from plant to plant to find prey species. (Banks, 1957; Biddinger, et al., 2009; Colunga-Garcia, et al., 1997; Hagan, 1962; Osawa, 2000)

Home Range

Harmonia axyridis does not generally exhibit territorial behavior, moving freely throughout its habitat. (Banks, 1957; Colunga-Garcia, et al., 1997; Hagan, 1962; Osawa, 2000)

Communication and Perception

Like many insects, Harmonia axyridis communicates via vision and chemical/sensory signals through the release of various pheromones. Many pheromones are used for mating signals, as is the case with many insects. They also use these pheromones to detect other individuals for congregation and hibernation during the winter months. While researchers have found the effects that these pheromones have on behaviors, no research has been done regarding their identification. One identified pheromone, harmonine, is used for defense against predators. Harmonine is produced when attacked and is secreted by reflex bleeding from tarsal joints. Prey are generally found using sight or olfactory detection. (Colunga-Garcia, et al., 1997; Evans, 2003)

Food Habits

Harmonia axyridis is a predatory insectivore with chewing mandibles, primarily feeding on aphids and scale insects. They may also eat Thysanura species and mites. Butterfly and moth eggs may be eaten, as well. When other food sources are scarce, Harmonia axyridis has been known to eat other various Coccinellidae species. Occasionally, it may feed on grapes and similar fruits. Studies have seen that in some cases, this species will eat other sources, such as moths and pollen, but these are isolated incidences. Both adults and larvae of Harmonia axyridis will cannibalize eggs and smaller larvae. (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

Predation

Many species of bird prey on Harmonia axyridis. The pentatomid, Podisus maculiventris is known to prey on H. axyridis, as well as many species of ants, including the red imported fire ant, Solanopsis invicta. Interspecies predation also occurs, though only when the other coccinellids are larger than the H. axyridis prey. Cannibalism on eggs and larvae is very prevalent among Harmonia axyridis populations, with some studies showing up to 50% cannibalism of eggs.

To defend itself, Harmonia axyridis produces isopropyl methoxy pyrazine (IPMP), which it secretes from its tarsi when agitated. This is highly concentrated in the hemolymph and is used as a chemical defense mechanism. Further, this species has antimicrobial agents in its hemolymph that provide defense against Gram-positive and Gram-negative bacteria as well as yeast. This species displays aposematism, as its red and black elytra can deter predators, acting as an apparent warning sign. (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)

Ecosystem Roles

Harmonia axyridis is a host species for a variety of parasites and parasitoids, specifically from the orders Diptera (Strongygaster triangulifer) and Hymenoptera (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 it as a host species.

Outside its native range, this species can create severe ecological pressures on indigenous Coccinellidae species as they out-compete and utilize resources efficiently. Some of the species most affected are Coccinella transversoguttata, Adalia bipunctata, and Coccinella novemnotata, although others have certainly experienced pressures, as well. They also prey on herbivorous pest species, which helps maintain plant species. (Banks, 1957; Gordon, 1985; Kindlmann, et al., 2011; Lombaert, et al., 2010; Sloggett, 2008)

Commensal/Parasitic Species

Economic Importance for Humans: Positive

Harmonia axyridis serves as a biological control species for many agricultural crops, as their main dietary sources are major agricultural pests. Their worldwide introduction is due to their intense predatory nature. (Alhmedi, et al., 2010; Campbell and Cone, 1999; Davidson and Evans, 2010; Dixon, 2005; Gardiner and Landis, 2007; Sloggett, 2008)

  • Positive Impacts
  • controls pest population

Economic Importance for Humans: Negative

Harmonia axyridis can cause significant economic damages on fruit crops, specifically grapes. This species will crawl inside them to feed and, when the grapes are harvested, will reflex bleed, causing an unpleasant taste. This taste can even be detected in wine if H. axyridis is accidentally incorporated during the wine-making process. It is also known to nip humans when handled. (Botezatu, et al., 2013; Galvan, et al., 2009)

  • Negative Impacts
  • injures humans
    • bites or stings
  • crop pest

Conservation Status

Harmonia axyridis has no special conservation status.

Contributors

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

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

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.

World Map

Neotropical

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

World Map

Palearctic

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

World Map

agricultural

living in landscapes dominated by human agriculture.

aposematic

having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.

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.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

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

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

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.

insectivore

An animal that eats mainly insects or spiders.

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.

World Map

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

poisonous

an animal which has a substance capable of killing, injuring, or impairing other animals through its chemical action (for example, the skin of poison dart frogs).

polygynandrous

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

polymorphic

"many forms." A species is polymorphic if its individuals can be divided into two or more easily recognized groups, based on structure, color, or other similar characteristics. The term only applies when the distinct groups can be found in the same area; graded or clinal variation throughout the range of a species (e.g. a north-to-south decrease in size) is not polymorphism. Polymorphic characteristics may be inherited because the differences have a genetic basis, or they may be the result of environmental influences. We do not consider sexual differences (i.e. sexual dimorphism), seasonal changes (e.g. change in fur color), or age-related changes to be polymorphic. Polymorphism in a local population can be an adaptation to prevent density-dependent predation, where predators preferentially prey on the most common morph.

scrub forest

scrub forests develop in areas that experience dry seasons.

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

solitary

lives alone

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

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