Coccinellidae

The name Coccinellidae, created by Pierre André Latreille in 1807, is derived from the Latin word meaning . The common English name ladybird originated in Britain where the insects became known as "Our Lady's birds". In the United States, the name was popularly adapted to ladybug. Entomologists prefer the names ladybird beetles or lady beetles to avoid confusion with true bugs. Names in some other countries may be similar; for example, in Germany they are known as meaning or . == Description ==

Coccinellids range in size from . Adult females tend to be slightly larger than males. They are generally oval with domed backs and flattened undersides. The tarsus (end of leg) has two claws at the tip. As adults, these beetles differ from their closest relatives with the following morphological characteristics: • 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. These colour patterns typically serve as warning colouration, but some can act as camouflage, attract mates or even regulate heat. Several individual species may display polymorphism and even change colour between seasons. Coccinellid larvae are elongated with square heads. They are covered in hairs or setae, the abdominal segments, in particular, each having six divided into pairs, and one to three segmented antennae. 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. File:BIEDRONA mirrored, cropped.png|Coccinella septempunctata, black spots on red|alt=A red ladybird File:Psyllobora.vigintiduopunctata.6920.jpg|Psyllobora vigintiduopunctata, black spots on yellow|alt=A yellow ladybird File:Coccinellidae - Vibidia duodecimguttata-1 (cropped).JPG|Vibidia duodecimguttata, whitish spots on brown|alt=A brown ladybird File:Brumoides suturalis (cropped).jpg|Brumoides suturalis, longitudinally striped|alt=A striped ladybird File:Rhyzobius chrysomeloides.jpg|Rhyzobius chrysomeloides, brown, unspotted|alt=An unspotted ladybird == Evolution ==

Fossil history Over 6,000 living species of Coccinellidae have been described. Although molecular clock estimates have placed their origin in the Cretaceous, the oldest fossils of the group are known from the Oise amber of France, dating to the Early Eocene (Ypresian) around 53 million years ago, which belong to the extant genera Rhyzobius and Nephus. The greatest number of fossils comes from the younger Eocene Baltic amber, including members of the extant genera Serangium and Rhyzobius as well as extinct genera belonging to the tribes Microweiseini (Baltosidis) and Sticholotidini (Electrolotis). }} Coccinellidae have historically been divided into 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. A 2021 genetic study sampling many species, identified three subfamilies, Microweiseinae (with three tribes), Coccinellinae (26 tribes) and a newly identified group, the Monocoryninae (one tribe). All three subfamilies were strongly supported, but the study noted that although the tribes are mostly monophyletic, their relationships are only weakly supported. The study suggests that the crown group appeared some 143 Mya in the Early Cretaceous, and that the group diversified rapidly during the Late Cretaceous, perhaps because the growth in diversity of angiosperm plants then encouraged the radiation of insects of the clade Sternorrhyncha such as aphids, on which ladybirds could feed. }} 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. == Biology and ecology==

Flight Coccinellids mostly fly during the day. 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. These beetles may migrate long distances to hibernation and breeding sites, and areas with more food. They appear to be drawn to recognisable landmarks. The more crowded an area is, the more individuals leave, but will remain if there are enough prey species to feed on. "Trivial flights" refer to flying while foraging or when finding a place to lay eggs. File:Coccinella transversalis 2.jpg|Coccinella transversalis, elytra in the open position|alt=Scale-like front wings, open File:Lady beetle taking flight.jpg|Full wings of a Harmonia axyridis taking flight|alt=Ladybird taking flight Life cycle and her eggs near a milkweed aphid colony. , eggs maturing, and emerging first instar larvae on narrow-leaf milkweed. 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. 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. After hatching, the larvae will begin eating, including the other eggs in their clutch. Larvae typically have four developmental stages with three moults between them. 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. In temperate areas, coccinellids may hibernate or enter diapause during the winter. Individuals during this period gather in clumps, large or small depending on the species. Overwintering insects can be found both in lowland areas, aggregating under dead vegetation, and at the tops of hills, hibernating under rocks and on grass tussocks. In areas with particularly hot summers, the insects experience summer dormancy or aestivation; in the tropics, coccinellids enter dormancy during the dry season. The majority of coccinellids are carnivorous and predatory, typically preying on Sternorrhyncha insects like aphids, scale insects, whiteflies, psyllids and adelgids. Some species feed on the larvae of moths and other beetles, as well as mites. Since much of their prey are agricultural pests, coccinellids are considered to be beneficial insects. Larvae and adults eat the same foods, unlike in other insect groups. Under pressure from coccinellid predation, aphid species have evolved to become more toxic, forcing coccinellids to develop immunities. Coccinellid predators of aphids need to defend themselves against ants that tend and defend aphids for their honeydew, and coccinellid eggs laid near aphids are disposed of. Cannibalism has been recorded in several species; which includes larvae eating eggs or other larvae, and adults feeding on individuals of any life stage. Some coccinellids are mostly non-predatory, such as some species in the genera Epilachna and Henosepilachna. The majority of predatory species may also supplement their diet with other sources of food both in their larval and adult stages. Non-animal matter consumed include leaves, pollen, nectar, sap, fungi, and honeydew. Members of the tribe Halyziini of the subfamily Coccinellinae are obligate fungus feeders. Coccinellids of any lifestage are preyed on by predators such as birds, spiders, ants and lacewings. File:Epilachna guttatopustulata01.jpg|Henosepilachna guttatopustulata, an herbivore, feeding on a potato leaf|alt=Ladybird eating a leaf File:Yellow Shouldered Ladybird (Apolinus lividigaster) with Aphis nerii.jpg|Yellow-shouldered ladybird, Apolinus lividigaster eating an aphid|alt=Ladybird eating an aphid File:Harmonia axyridis cannibalism.jpg|Harmonia axyridis larva cannibalism|alt=Ladybird larva eating another ladybird larva Defense '' reflex bleeding The bright warning colouration of many coccinellids discourage potential predators, warning of their toxicity. A 2015 study of five ladybird species found that their colouration honestly signalled their toxicity, implying the warning is genuine. Species with more contrast with the background environment tended to be more toxic. Coccinellid haemolymph (blood) contains toxic alkaloids, azamacrolides and polyamines, as well as foul-smelling pyrazines. Coccinellids can produce at least 50 types of alkaloids. When disturbed, ladybirds further defend themselves with reflex bleeding, exuding drops from their tibio-femoral (knee) joints, effectively presenting predators with a sample of their toxic and bitter body fluid. Predator-deterring poisons are particularly important for the immobile pupa. Access to food can affect the concentration of both pigments and toxins. The similarity of coccinellid patterning in red and orange with black markings has led to suggestions that they and some species of chrysomelids form Müllerian mimicry rings particularly to defend them from birds. 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. 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. == Distribution and status ==

'' Coccinellidae are found on every continent except Antarctica. Asian and African species are less studied than others. 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. == Relationship to humans ==

Biological control feeding on Icerya purchasi''|alt=Ladybird used as a biological control in horticulture 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. Several species have been introduced to areas outside their native range; the first being the vedalia beetle, Novius cardinalis. 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. There have been many further attempts to use ladybird species against pests, with varying degrees of success. As pests inside a farm building after the fall harvest in South Dakota|alt=Mass of ladybirds at rest Coccinellids can also act as pests. Harmonia axyridis is native to East Asia, but has been introduced to the Americas, Europe and Africa. After an abnormally long period of hot, dry weather in the 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. H. axyridis, C. septempunctata and Hippodamia convergens are the most common causes of ladybird taint in wine. As few as 1.3 to 1.5 coccinellids per of grapes can affect wine quality when they are present during the wine-making process. The Mexican bean beetle is an agricultural pest as it primarily feeds on plants, especially legumes, instead of insects. 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. 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. Coccinellids have been popularly featured in poems and nursery rhymes, the most famous being Ladybird! Ladybird!. This has come in several forms, including: File:Card cutout ladybirds for children's nature trail.jpg|Card cutout ladybirds for a children'snature trail|alt=Ladybird depictions for a nature trail File:Drawing, album (BM SL,5276.74) (ladybird flying over cyclamen).jpg|Ladybird flying over cyclamen, detail,by Maria Sibylla Merian, 1690s|alt=Historic ladybird illustration File:09. Ladybug, Ladybug - panoramio.jpg|Depiction of the Ladybird! Ladybird! rhyme in Hunter Valley Gardens, Australia|alt=Background painting and models used to illustrate a ladybird nursery rhyme == References ==