Coccinellidae

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"Lady Bug" and "Ladybug" redirect here. For other uses, see Lady Bug (disambiguation).
"Ladybird" redirects here. For other uses, see Ladybird (disambiguation).

Ladybird, ladybug, lady beetle
Temporal range: Eocene—Present
Coccinella septempunctata
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Polyphaga
Infraorder: Cucujiformia
Superfamily: Coccinelloidea
Family: Coccinellidae
Latreille, 1807[1]
Subfamilies[1]

(traditional, but see below):

Synonyms
  • Cerasommatidiidae
  • Epilachnidae

Coccinellidae (

predators
that they taste bad.

Most coccinellid species are

hibernate and diapause during the winter; tropical species are dormant during the dry season
. Coccinellids migrate between dormancy and breeding sites.

Species that prey on agricultural pests are considered

beneficial insects. Several species have been introduced outside their range as biological control agents, with varying degrees of success. Some species are pests themselves and attack agricultural crops, or can infest people's homes, particularly in winter. Invasive species like Harmonia axyridis can pose an ecological threat to native coccinellid species. Other threats to coccinellids include climate change and habitat destruction. These insects have played roles in folklore, religion and poetry, and are particularly popular in nursery rhymes
.

Etymology

The name Coccinellidae, created by

true bugs.[9][10][11][12] Names in some other countries may be similar; for example, in Germany they are known as Marienkäfer meaning 'Marybeetle' or 'ladybeetle'.[9]

Description

Coccinellids range in size from 0.8 to 18 mm (0.03–0.7 in).

hindwings when the insects are not in flight. Their legs are relatively short,[16] with a tarsal formula of 4-4-4 or 3-3-3.[18] The tarsus (end of leg) has two claws at the tip.[16]

As adults, these beetles differ from their closest relatives with the following morphological characteristics:[18]

  • Five pairs of spiracles (holes) on the abdomen
  • A tentorium (internal supports inside the head) with separated branches at the front and no bridge
  • No line dividing the frons and clypeus (frontoclypeal suture)
  • Maxillary palps with non-needle-shaped tips,
  • Divided galea and lacinia (lobes at the end of the mouthparts)
  • Smaller molar (flattened) area of the mandible
  • Coxal cavities (holes where the leg articulates with the thorax) that open from the back in the front of the thorax and from the front in the middle of the thorax
  • Epimeron (corner plates) on the metathorax with parallel edges
  • Lines on the second abdominal sternum
  • Tube-shaped, siphon-like genitalia in the male

Coccinellids are often distinctively coloured and patterned. The elytron may be light with dark spots or dark with light spots. Light areas are typically yellow, red, orange or brown, and the spots vary in size and shape and numbers. Some species have striped or

checkered patterns. The pigment carotene creates the lighter colours, and melanins create darker colours. Other parts of the body also vary in colouration.[7][19] These colour patterns typically serve as warning colouration, but some can act as camouflage, attract mates or even regulate heat.[7][20] Several individual species may display polymorphism and even change colour between seasons.[7]

Coccinellid

setae, the abdominal segments, in particular, each having six divided into pairs, and one to three segmented antennae.[18][7] Their colouration varies from grey, blue-grey, grey-brown or brown and spotted with white, yellow, red or orange. They tend to brighten as they get closer to adulthood.[22]

Evolution

Fossil history

Over 6,000 living species of Coccinellidae have been described.

Serangium[25] and Rhyzobius as well as extinct genera belonging to the tribes Microweiseini (Baltosidis)[26] and Sticholotidini (Electrolotis).[23]

Phylogeny

Further information: List of Coccinellidae genera

The Coccinellidae are within the superfamily Coccinelloidea, which in turn is part of the infraorder Cucujiformia, a group containing most of the plant-eating beetles. The ladybirds form the majority of the species in the Coccinelloidea; many of the rest are fungus-feeding beetles or scavengers.[27]

Cucujiformia
Coccinelloidea

Bothrideridae and allies Bothrideres bipunctatus

Latridiidae Latridius porcatus

Akalyptoischiidae Akalyptoischion

Alexiidae Sphaerosoma pilosum

Corylophidae and allies Arthrolips obscura

Endomychidae Brachytrycherus bipunctatus.jpg

Coccinellidae Coccinella septempunctata

Coccinellidae have historically been divided into up seven subfamilies (Chilocorinae, Coccidulinae, Coccinellinae, Epilachninae, Microweiseinae, Scymninae and Sticholotidinae) and 35 tribes based on morphology. However, genetics studies have called into question the monophyly (single ancestry) of most of these subfamilies. The monophyly of Coccinellinae has the most support.[28][29]

A 2021 genetic study sampling many species, identified three subfamilies, Microweiseinae (with three tribes), Coccinellinae (26 tribes) and a newly identified group, the

angiosperm plants then encouraged the radiation of insects of the clade Sternorrhyncha such as aphids, on which ladybirds could feed.[30]

Coccinellidae

An earlier 2009 study concluded that consumption of

scale insects is the most basal diet of Coccinellidae. Aphid-eating evolved three separate times and leaf-eating evolved twice, one of which evolved from a clade that contains both aphid-eating and pollen-eating. The fungi-eating also evolved from aphid-eating.[28]

Biology and ecology

Flight

Coccinellids mostly fly during the day.[31] Springy, cylindrical veins in the hindwings stiffen when in flight and bend when folding. Folding of the wings is further aided by creases in the membrane.[32] These beetles may migrate long distances to hibernation and breeding sites, and areas with more food. They appear to be drawn to recognisable landmarks.[33] The more crowded an area is, the more individuals leave, but will remain if there are enough prey species to feed on.[34] "Trivial flights" refer to flying while foraging or when finding a place to lay eggs.[33] One study of species in Britain found that coccinellids can fly as far as 120 km (75 mi). They flew at speeds of 30 km/h (19 mph) and could reach altitudes close to 1,100 m (3,600 ft).[35]

Life cycle

In temperate climates, coccinellids typically breed from late spring to early summer. In warmer temperate regions, reproduction may occur in spring, fall and winter; tropical species reproduce during the wet season.[7][36] Mating is promiscuous. In some species, females appear to be selective in their partners, preferring males of a certain size and colour. Males produce sperm packets each with 14,000 sperm, and insert three of them into the female, even though she can only hold 18,000 sperm. This is likely a form of sperm competition.[7][37] Like other insects, coccinellids develop from egg, to larva, to pupa and finally adult. Eggs tend to be bright yellow, and the females lay them close together, standing upright and near where they can access food.[7] The number of eggs in a cluster can vary depending on the species; it is typically in the double digits but some species can lay over a thousand eggs in their lifetime.[38]

After hatching, the larvae will begin eating, including the other eggs in their clutch.[7] Certain species lay extra infertile trophic eggs with the fertile eggs, providing a backup food source for the larvae when they hatch. The ratio of infertile to fertile eggs increases with scarcity of food at the time of egg laying.[39] Larvae typically have four instar stages with three moults between them.[40] The larva eventually transitions into a pupa; which involves the development of a hunch, the fusion of the legs to the body, and the attachment of the posterior to the surface.[7][18][40]

Pupae may be uncovered, partially covered or fully covered by larval skin depending on the species. The pupa is mostly immobile, but the head can move in response to irritation. When the adult emerges, it has its hindwings, while the elytron starts out softer and lighter in colour, with no patterns.[41] The length of each development stage varies based on climate and between species. For Adalia bipunctata, eggs hatch after four to eight days, the larva stage lasts around three weeks and the pupa lasts seven to ten days.[7] Adult coccinellids develop much of their final colouration within hours, but may not fully darken for weeks or months.[42] The lifespan of an adult reaches up to a year.[7]

In temperate areas, coccinellids may

grass tussocks.[43] In areas with particularly hot summers, the insects experience summer dormancy or aestivation; in the tropics, coccinellids enter dormancy during the dry season.[7]

Trophic roles

Coccinellids act both as predators, prey and

metastudy by Hodek and Honěk found that aphid-eaters constituted around 68 percent of species that live in temperate areas but only 20 percent of species worldwide. Around 36 percent of total species mostly feed on scale insects.[47] Larvae and adults eat the same foods, unlike in other insect groups.[7]

Ladybird species vary in dietary specificity. An example of a specialist species is those of the genus Stethorus, which feed on spider mites. Aphid-eaters tend to be generalist; they have a high voracity and can multiply quickly in response to outbreaks, and switch to other prey when the ephemeral aphids become scarce. Predators of scale insects tend to be less voracious and are slower breeders and developers; matching their prey.[48] Under pressure from coccinellid predation, aphid species have evolved to become more toxic, forcing coccinellids to develop immunities.[49] Coccinellid predators of aphids need to defend themselves against ants that tend and defend aphids for their honeydew,[50] and coccinellid eggs laid near aphids are disposed of.[44] Some species including Coccinella magnifica and Diomus have adapted to grow within ant nests as larvae, and some like Diomus thoracicus are predators of the brood of the ant Wasmannia auropunctata.[51][52]

Cannibalism has been recorded in several species; which includes larvae eating eggs or other larvae, and adults feeding on individuals of any life stage.[53] Some coccinellids are mostly non-predatory, such as some species in the genera Epilachna and Henosepilachna.[54][55] The majority of predatory species may also supplement their diet with other sources of food both in their larval and adult stages.[56] Non-animal matter consumed include leaves, pollen, nectar, sap, fungi, and honeydew.[45][56][57] Members of the tribe Halyziini of the subfamily Coccinellinae are obligate fungus feeders.[58]

Coccinellids of any lifestage are preyed on by predators such as birds, spiders, ants and

hymenopterans and nematodes, and pathogens, including bacteria, fungi and protozoa.[44][60] Wolbachia bacteria infects eggs and kills male zygotes.[61] The promiscuity of Coccinellids has led to their being affected by sexually transmitted infections.[62]

Defense

Coccinella septempunctata reflex bleeding

The bright warning colouration of many coccinellids

reflex bleeding, exuding drops from their tibio-femoral (knee) joints, effectively presenting predators with a sample of their toxic and bitter body fluid.[59] Predator-deterring poisons are particularly important for the immobile pupa.[65] Access to food can affect the concentration of both pigments and toxins.[66]

The similarity of coccinellid patterning in red and orange with black markings has led to suggestions that they and some species of chrysomelids[67] form Müllerian mimicry rings[68] particularly to defend them from birds.[69] Despite their chemical defenses, coccinellids are preyed on by some clerid beetles in the genus Enoclerus, several species of which are brightly coloured in red and black, and which possibly sequester the toxins of the prey to defend themselves against other predators.[70]

As an anti-predator defense, spiders of the genus Eresus, known as ladybird spiders, have evolved to replicate the patterns of coccinellids. This is a form of Batesian mimicry, as the spiders lack the chemicals. This resemblance is limited to adult male spiders which are actively searching for females and exposed – unlike the females and young, which remain sheltered in burrows.[71]

Distribution and status

The widespread and invasive Harmonia axyridis

Coccinellidae are found on every continent except Antarctica.

endemic and possibly threatened.[73]

Threats to coccinellids include

pesticides. Invasive threats include other coccinellids, particularly C. septempunctata in North America and H. axyridis globally.[73] These invaders outcompete the native species as well as eat their eggs.[73][75]

As of 2022, the IUCN Red List does not list the conservation status for any coccinellid, though there is an IUCN SSC Ladybird Specialist Group. Conservationists have suggested several measures for protecting the insects, including citizen science and education programs, habitat preservation and restoration, prevention of the spread of invasive species and a global monitoring program.[73]

Relationship to humans

Biological control

Further information: Biological pest control
Ladybird used as a biological control in horticulture
Biological control: larval Novius cardinalis feeding on Icerya purchasi

Coccinellids have been valued in biological pest control, as they prey on agricultural pests such as aphids and scale insects. Their importance in controlling pests was noted as far back as 1814 in England.[7] Their efficiency can vary: sometimes they have a relatively small effect on aphid populations; at others they cause significant seasonal declines.[76]

Several species have been

cottony cushion scale. The project was markedly successful, costing $1,500 in 1889,[77] making it "a textbook example of the great potential of classical biological control as a tactic for suppressing invasive pests." The beetle was then used in 29 countries, again with success; reasons for this include its high prey specificity, fast development, multiple generations each year, efficient discovery of host patches, and larval development completed on a single host insect.[78]

There have been many further attempts to use ladybird species against pests, with varying degrees of success.[79][7] Scale insect-eating coccinellids have been more successfully used than aphid predators.[7] Out of 155 deliberate introductions meant to control aphids by the year 2000, only one was deemed to be "substantially successful". This is due to aphid-eating species being fast-breeding, generalist and voracious, and thus difficult to control.[80]

As pests

Mass of ladybirds at rest
Cluster of invasive Asian lady beetles inside a farm building after the fall harvest in South Dakota

Coccinellids can also act as pests. Harmonia axyridis is native to East Asia, but has been introduced to the Americas, Europe and Africa.

summer of 1976 in the UK, a marked increase in the aphid population was followed by a "plague" of the native Coccinella septempunctata; there were many reports of people being bitten as the supply of aphids dwindled.[83][84][85]

H. axyridis, C. septempunctata and

ladybird taint in wine. As few as 1.3 to 1.5 coccinellids per 1 kilogram (2.2 lb) of grapes can affect wine quality when they are present during the wine-making process.[86] The Mexican bean beetle is an agricultural pest as it primarily feeds on plants, especially legumes, instead of insects.[87]

In culture

Coccinellids have had important roles in culture and religion, being associated with luck, love, fertility and prophecy. "Ladybird" is an affectionate term for someone, such as a loved one. In European folklore, an insect acts as a matchmaker, crawling on a woman and then flying to their true love. Coccinellids have been said to predict the future, particularly weather conditions and how well the crops will grow.[88][89] Contrary to a popular saying, the age of a lady bug cannot be determined by counting its spots.[90]

In Christianity, coccinellids have been seen as the literal gatekeepers of Heaven. A Swedish name for the insects, Himmelska nycla, means "Keys of Heaven". Jews have referred to the insects as the "Cow of Moses our Teacher". The Cherokee have revered them as the "Great Beloved Woman"; this was used as a title for the highest-ranking woman in the government, who would be painted in the colours and patterns of the insect during ceremonies.[91]

Coccinellids have been popularly featured in poems and

Ladybird! Ladybird!. This has come in several forms, including:[92]

Ladybird, ladybird, fly away home,
Thy house is on fire, thy children all roam,
Except little Nan, who sits on her pan,
Weaving gold laces as fast as she can.


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Sources

External links
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