Overview
Brief Summary
Introduction
Although officially called lady beetles, members of the family Coccinellidae are more commonly known as ladybugs (American) or ladybirds (Great Britain, Australia, Canada). This is the largest of the cucujoid families with nearly 6,000 described species and about 370 genera worldwide. They are among the most familiar and beloved of insects, and are widely recognized for their role in controlling voracious plant pests such as aphids and scale insects. Not surprisingly, in many cultures coccinellid beetles are considered harbingers of good luck and a bountiful harvest. Even the common name, which dates back to medieval times, is a testament of their sacred status as "beetles of Our Lady" (in reference to the virgin Mary).
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Comprehensive Description
Lady Beetles are small, round, with convex wing-covers. The head and pronotum are often shiny black with white markings, while the wing-covers are often shiny orange-red with black dots. Some species, however, are predominantly black, or gray with black dots, and may have fine hairs on their bodies. The adults usually feed on aphids, scale insects, and other small insects, while the larvae tend to feed on the same kinds of insects. Adults occasionally appear on flowers with exposed nectaries to feed on nectar. Lady Beetles are considered beneficial insects, although a few species feed on the foliage of crop plants.
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Hilty, J. Editor. 2010. Insect Visitors of Illinois Wildflowers. World Wide Web electronic publication. flowervisitors.info, version (09/2010).
See: Abbreviations for Insect Activities, Abbreviations for Scientific Observers, References for behavioral observations H
© John Hilty
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Color Pattern Illustrations
 striped |  checkered |  banded |
 ocellate |  metallic |  punctuation marks |
 ink blot |  radial lines |  spotted |
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Classification
 Carl Linnaeus |  L. Redtenbacher |  M. E. Mulsant |
Present day members of the family Coccinellidae first appeared in Linnaeus’ (1758) Systema Naturae under the genus Coccinella, which contained, at the time, 36 species. Redtenbacher (1844) provided the first internal classification system for the family by recognizing two biologically defined groups, the plant feeders and the aphid feeders. His phytophagous group corresponds to the subfamily Epilachninae (sensu Sasaji 1968) but the aphidophagous one spans the other currently recognized subfamilies. Mulsant’s (1846; 1850) interpretation of the family diversity divided Coccinellidae into the “Trichoisomides” (the hairy species) and the “Gymnosomides” (the glabrous ones), a system that was even more artificial than its predecessor. Mulsant, however, was the first author to recognize supra-generic categories for the family many of which correspond to the current tribes in Coccinellidae. Korschefsky’s (1931) classification recognized the subfamilies Epilachninae, Coccinellinae, and Lithophilinae. This system is consistent with that of Redtenbacher but further subdivides the entomophagous taxa (the so-called “aphidophaga”) on the basis of the tarsal structure. Despite these advances, the classification of Coccinellidae was still rudimentary and extremely artificial until the second half of the 20th century.
In the most influential series of works of the 20th century dealing with this topic, Sasaji (1968; 1971a, b) revised the classification of the family, proposing six subfamilies: Sticholotidinae, Coccidulinae, Scymninae, Chilocorinae, Coccinellinae and Epilachninae. This system was widely accepted and remains the primary reference for the family (e.g., Booth et al., 1990; Pakaluk et al., 1994; Lawrence and Newton, 1995; Kuznetsov, 1997). Additional subfamilies have since been recognized: Azyiinae and Exoplectrinae, elevated from tribal status (Gordon, 1994); Ortaliinae, for Ortaliini plus Noviini (Kovář, 1996); and Hyperaspidinae, for Hyperaspidini plus Brachiacanthini (Duverger 1989; 2001) (Giorgi et al. 2009). In the most recent major contribution to the systematics of Coccinellidae, Kovář (1996) proposed a classification of Coccinellidae worldwide, recognizing seven subfamilies and 38 tribes. Duverger (2003) attempted to apply a distinct subfamily name to each of the 18 major groupings in Kovář 's dendrogram (1996). Many of these names, however, were invalid (based on junior synonyms), misspelled, or applied inconsistently within the different sections of the same paper.
Pope (1989) and Vandenberg (2002) pointed out major caveats in the current classification of Coccinellidae, emphasizing the lack of morphological support for most of the recognized subfamilies. Ślipiński (2007) acknowledged the criticism and proposed a more conservative system, in which only two subfamilies are recognized, Microweiseinae (=Scotoscymninae Duverger =Sticholotidinae sensu Kovář 1996, in part), and Coccinellinae. None of these various classifications has received a universal following (see Table below for alternative classifications).
Many regional taxonomic monographs have been published in the last three decades, including: Fürsch (1967) [European fauna]; Gordon (1985) [North America]; Hoang (1982; 1983) [Vietnam]; Iablokoff-Khnzorian (1982) [Palearctic and Oriental regions]; Kuznetsov (1997) [Russian Far East]; Pang and Mao (1979) [China]; Pope (1989) [Australian Coccinellinae]; Sasaji (1971a) [Japan]; and Savoiskaya (1983) [Central Asia and parts of the former USSR].
Table 1: Comparison of high-level classification systems of Sasaji (1971a), Kovář (1996) and Ślipiński (2007)
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| Sasaji (1971a) | Kovář (1996) | Ślipiński (2007) (Australian taxa only) |
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- 1Includes Oryssomini
- 2Includes Exoplectrini, Scymnini, Stethorini, and taxa near Shirozuellini and Azyiini
- 3Includes Halyziini
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Characteristics
Lady beetles come in many different colors and patterns. Mouse over the following words to see more examples below: striped, checkered, banded, ocellate, metallic, punctuation marks, inkblot, radial lines.
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The charismatic red and black dappled members of the tribe Coccinellini are easily recognized by the layperson, but the family as a whole is somewhat difficult to characterize. Most species can be identified by the broadly rounded to elongate body form with convex dorsum and flattened venter, clubbed antennae, and the presence of a postcoxal line on the first abdominal ventrite (lacking in Paranaemia, Naemia, and Coleomegilla). The tarsal formula of most species is 4-4-4 with the third tarsomere minute and tucked within the broad triangular second (cryptotetramerous or pseudotrimerous), only a few have the tarsomeres more equal (truly tetramerous), and some have tarsi reduced to 3-3-3 (truly trimerous).
It can be difficult to recognize a lady beetle without studying some of its minute morphological features. The majority of species are very small, drab, and pubescent ("furry") like the image below.
Scymninae. © 2009 Explore some of the common body types found within the family Coccinellidae by mousing over the numbered legend below. What advantages do these different forms possess? What are their respective vulnerabilities? More elongate <----1--- -----2----- -----3----- -----4----- ---5----> More compact Coccinellids are quite similar to endomychid beetles, but lack a frontoclypeal suture and have the mandibular mola reduced or absent. They also lack the impressed lines on the base of the pronotum which are characteristic of many endomychids. The family Corylophidae also shares many affinities with Coccinellidae, but Corylophids posess a functional spiracle on abdominal segment 7 (lacking in Coccinellidae), have the galea and lacinia of the mouthparts fused into a single lobe (separate structures in Coccinellidae), and have a distinct form to the male genitalia.
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Ecology
Associations
Associations
Coccinellidae is predator of larva of Gastrophysa viridula
Animal / predator
larva of Coccinellidae is predator of Aphididae
Animal / parasitoid / endoparasitoid
larva of Medina separata is endoparasitoid of abdomen of imago of Coccinellidae
Animal / parasite / endoparasite
Parasitylenchus coccinellinae endoparasitises intestine of adult of Coccinellidae
In Great Britain and/or Ireland:
Animal / associate
pupal cocoon of Perilitus coccinellae is associated with paralysed adult of Coccinellidae
Animal / parasite / ectoparasite
several hundred nymph of Podapolipus ectoparasitises inside of elytra of adult of Coccinellidae
Animal / parasitoid / endoparasitoid
gregarious larva of Tetrastichus coccinellae is endoparasitoid of larva of Coccinellidae
© BioImages
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Evolution and Systematics
Evolution
Discussion of Phylogenetic Relationships
Coccinellidae and the Cerylonid Series
Molecular evidence supports Coccinellidae as belonging to the Cerylonid Series (Hunt et al., 2007; Robertson et al., 2008) but the closest relative of the family remains unclear. Early morphological studies supported hypotheses that the sister-group was a clade comprising Endomychidae plus Corylophidae (Crowson, 1981; Sasaji, 1971) or one comprising Endomychidae plus Alexiidae (Pakaluk and Ślipiński, 1991). These studies, however, were not based on formal phylogenetic analyses.
Formal phylogenetic studies that provide insights about the closest relatives of Coccinellidae resulted, each, in different answers. In these studies, the following taxa were recovered as close relatives of Coccinellidae: Endomychidae (Tomaszewska, 2000), Anamorphinae (Endomychidae) plus Alexiidae (Hunt et al., 2007), Corylophidae (Tomaszewska, 2005), Leiestinae (Endomychidae) (Robertson et al. 2008, parsimony) and Endomychidae minus Anamorphinae or Corylophidae plus Anamorphinae (Robertson et al. 2008, Bayesian). Since each of these studies addressed different phylogenetic questions, the taxonomic sampling was fairly different, which might account for the noted disparities.
Coccinellidae Subfamilies
Sasaji (1968, 1971) provided the first attempt to address the higher-level phylogenetic relationships of Coccinellidae (see fig. below). Under Sasaji’s classical phylogenetic scheme, three main branches are recognized. At the base is the subfamily Sticholotidinae, considered the most primitive group of coccinellids. On the main branch, one lineage includes the subfamilies Scymninae and Chilocorinae, and other includes Coccidulinae, Coccinellinae and Epilachninae. In the latter group, Coccidulinae was considered the sister group to the coccinelline plus epilachnine branch.
Kovář’s (1996) hypothesis was very similar to Sasaji’s (1968, 1971), differing only in placing Coccidulinae at the base of the scymnine plus chilocorine branch. He also recognized a seventh subfamily, Ortaliinae, comprised of the former Coccidulinae tribe, Noviini, and the former Scymninae tribe, Ortaliini.
In spite of the relevance of these contributions, both Sasaji (1968, 1971) and Kovář’s (1996) studies have some limitations. In both studies the hypotheses were not generated by formal phylogenetic methods, thus none of the clades are supported by synapomorphies. Sasaji’s study did not include any neotropical coccinellids, being mostly restricted to the Japanese fauna. Kovář’s (1996) study was more comprehensive in terms of diversity.
The first formal cladistic analysis was conducted by Yu (1994). His study used adult and larval morphological characters. The partitions (adults vs. larvae) were treated separately, and produced drastically conflicting topologies. The adult-based topology had few points of agreement with previous hypotheses. Yu (1994) recovered all Sticholotidinae at the base, but not as a monophyletic group. As with Sasaji’s (1968) hypothesis, this study supports a clade containing the subfamilies Chilocorinae, Coccidulinae and Scymninae, but none of these subfamilies were recovered as monophyletic. Coccidulinae was recovered as paraphyletic with respect to Ortaliinae, Chilocorinae, and Scymninae. Chilocorinae and Ortaliinae were nested within Scymninae. The sister group relationship between Epilachninae and Coccinellinae was not supported in this study. Epilachninae diverged from the rest of the family at the next highest node, followed by Coccinellinae. A close relationship between Serangiini and Sukunahikonini was the only point of similarity between the adult and larval topologies. 
Giorgi et al. (2009) conducted a parsimony analysis using the nuclear ribosomal genes 18S rDNA and 28S rDNA to address the evolution of food preferences in Coccinellidae. As in the previous studies, the subfamily Sticholotidinae was supported as the sister group of the rest of the coccinellids, but only in part. Other Sticholotidinae included in the analysis, were supported as more advanced forms. The resulting topology of this study disagrees with Sasaji’s (1968) classic hypothesis in many points. Unfortunately the relationships between most of the major branches were not strongly supported.
Thus, an accurate definition of the main lineages of coccinellids as well as the relationships between them still remains to be established.
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Molecular Biology and Genetics
Barcode
Locations of barcode samples
© Barcode of Life Data Systems
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Statistics of barcoding coverage
| Specimen Records: | 899 |
| Specimens with Sequences: | 722 |
| Specimens with Barcodes: | 655 |
| Public Records: | 140 |
| Species: | 123 |
| Species With Barcodes: | 112 |
© Barcode of Life Data Systems
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Barcode data
© Barcode of Life Data Systems
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Wikipedia
Coccinellidae
The Coccinellidae are a family of beetles, known variously as ladybirds (UK, Ireland, Australia, Sri Lanka, Pakistan, South Africa, New Zealand, India, Malta, some parts of Canada and the US), or ladybugs (North America). When they need to use a common name, entomologists widely prefer the names ladybird beetles or lady beetles[2] as these insects are not true bugs. Lesser-used names include God's cow, ladyclock, lady cow, and lady fly.[3]
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Overview
Coccinellids are small insects, ranging from 1 mm to 10 mm (0.04 to 0.4 inches), and are commonly yellow, orange, or scarlet with small black spots on their wing covers, with black legs, head and antennae. Such color patterns vary greatly however; for example, a minority of species, such as Vibidia duodecimguttata, a twelve-spotted species. has whitish spots on a brown background.
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In some species the spots are so large that they merge, leaving the remaining bright colours as spots on a black background. Many coccinellid species are mostly, or entirely, black, grey, or brown and may be difficult for non-entomologists to recognize as coccinellids at all. Conversely,non-entomologists might easily mistake many other small beetles for coccinellids. For example the tortoise beetles, like the ladybird beetles, look similar because they are shaped so that they can cling to a flat surface so closely that ants and many other enemies cannot grip them.
Coccinellids are found worldwide, with over 5,000 species described,[4] more than 450 native to North America alone.
The Coccinellidae are generally considered useful insects, because many species feed on aphids or scale insects, which are pests in gardens, agricultural fields, orchards, and similar places. However, some species do have unwelcome effects. Among these the most prominent are the subfamily Epilachninae, which are plant eaters. Usually Epilachninae are only mild agricultural pests, eating the leaves of grain, potatoes, beans and various other crops, but their numbers can increase explosively in years when their natural enemies are few, such as parasitoid wasps that attack their eggs. When that happens they can do major crop damage. They occur in practically all the major crop producing regions of temperate and tropical countries.
Harmonia axyridis (or the harlequin ladybird/bug) is an example of how an animal might be partly welcome and partly harmful. It was introduced into North America from Asia in 1916 to control aphids, but is now the most common species, out-competing many of the native species.[5] It has since spread to much of western Europe, reaching the UK in 2004.[5][6] It has become something of a domestic and agricultural pest in some regions and gives ecological cause for concern. It similarly has turned up in parts of Africa, where it has proved variously unwelcome, perhaps most prominently in vine-related enterprises.
A common myth, totally unfounded, is that the number of spots on the insect's back indicates its age.[7][8] In fact the number, shape, and placement of the spots all are determined by the species of the beetle, and are fixed by the time it emerges from its pupa. The same applies to the colour, except that it may take some days before the colour of the adult beetle to mature and stabilise. Generally the mature colour tends to be fuller and darker than the colour of the callow.
Etymology
Coccinelid is derived from the Latin word coccineus meaning scarlet.[9] The name ladybird originated in Britain where the insects became known as Our Lady's bird or the Lady beetle.[10][11] Mary (Our Lady) was often depicted wearing a red cloak in early paintings and the spots of the seven spot ladybird (the most common in Europe) were said to symbolise her seven joys and seven sorrows.[10] In the United States the name was adapted to ladybug. Common names in other European languages have the same association, for example, the German name Marienkäfer translates to Marybeetle.[12]
Physical appearance
Most coccinellids have oval, dome-shaped bodies with six short legs. Depending on the species, they can have spots, stripes, or no markings at all. Seven-spotted coccinellidae are red or orange with three spots on each side and one in the middle; they have a black head with white patches on either side.
Biology
See also : List of Coccinellidae genera
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Coccinellids are typically predators of Hemiptera such as aphids and scale insects, though larvae and eggs of their own species can also be important resources when alternative prey are scarce.
Members of the subfamily Epilachninae are herbivores, and can be very destructive agricultural pests (e.g., the Mexican bean beetle). While predatory species are often used as biological control agents, introduced species of coccinellidae (such as Harmonia axyridis or Coccinella septempunctata in North America) out-compete and displace native coccinellids and become pests in their own right.[13]
The main predators of coccinellids are usually birds, but they are also the prey of frogs, wasps, spiders, and dragonflies. The bright colours of many coccinellids discourage some potential predators from making a meal of them. This phenomenon is called aposematism and works because predators learn by experience to associate certain prey phenotypes with a bad taste. A further defense known as "reflex bleeding" exists in which an alkaloid toxin is exuded through the joints of the exoskeleton, triggered by mechanical stimulation (such as by predator attack) in both larval and adult beetles, deterring feeding.
Coccinellids in temperate regions enter diapause during the winter, so they often are among the first insects to appear in the spring. Some species (e.g., Hippodamia convergens) gather into groups and move to higher elevated land, such as a mountain, to enter diapause. Most coccinellids overwinter as adults, aggregating on the south sides of large objects such as trees or houses during the winter months, dispersing in response to increasing day length in the spring.[14]
It is thought that certain species of Coccinellids lay extra infertile eggs with the fertile eggs. These appear to provide 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.[15]
Predatory coccinellids are usually found on plants which harbour their prey. They lay their eggs near their prey, to increase the likelihood the larvae will find the prey easily. In Harmonia axyridis, eggs hatch in 3–4 days from clutches numbering from a few to several dozen. Depending on resource availability, the larvae pass through four instars over 10–14 days, after which pupation occurs. After a teneral period of several days, the adults become reproductively active and are able to reproduce again, although they may become reproductively quiescent if eclosing late in the season. Total life span is 1–2 years on average.
Infestations and impacts
In the United States, Coccinellids usually begin to appear indoors in the fall. They leave their summer feeding sites in fields, forests and yards looking for a place to spend the winter. Typically when temperatures warm to the mid-60s Fahrenheit in the late afternoon, following a period of cooler weather, they will swarm onto or into buildings illuminated by the sun. Swarms of Coccinellids fly to buildings in September through November depending on location and weather conditions. Also, homes or buildings near fields or woods are more prone to infestation.[16]
The presence of Coccinellids in grape harvests can cause ladybird taint in wines produced from the grapes.
After an abnormally long period of hot dry weather in the Summer of 1976 in the UK, there was a marked increase in the aphid population followed by a "plague" of Coccinellids, with many reports of people being bitten as the supply of aphids dwindled.[17][18] Recent studies suggest that Coccinellids can also cause allergic reactions, such as eye irritation or asthma.[19]
UK ladybird survey
The ladybird atlas Ladybirds (Coccinellidae) of Britain and Ireland published in 2011[20] showed a decline of more than 20% in native species due to environmental changes and competition from foreign invaders. The distribution maps, compiled over a twenty year period with help from thousands of volunteers, showed a decline in the numbers of the common 10-spot and 14-spot ladybirds and a number of other species including the 11-spot, 22-spot, Cream-spot, Water and Hieroglyphic ladybirds, Coccidula rufa, Rhyzobius litura and Nephus redtenbacheri. Conversely, increases were seen in the numbers of Harlequin, Orange, Pine and 24-spot ladybirds as well as Rhyzobius chrysomeloides. The Kidney spot ladybird was recorded in Scotland for the first time in recent years, probably due to climate change, and the once-extinct 13-spot was found to have recolonised Cornwall, Devon and the New forest. The most commonly recorded species was the 7-spot, closely followed by the Asian Harlequin — an invader that arrived from continental Europe in 2003 after being introduced to control pests. An 'explosion' in the number of Orange ladybirds, which feed on mildew, is thought to be have been due to the warmer damper conditions that now prevail in parts of England.[21]
In Popular Culture
Folk History
Coccinellids are and have been for very many years a favorite insect of children. The insects had many regional names (now mostly disused) in English, such as the lady-cows, may-bug, golden-knop, golden-bugs (Suffolk); and variations on Bishop-Barnaby (Norfolk dialect) – Barnabee, Burnabee, the Bishop-that-burneth, and bishy bishy barnabee.[22] The etymology is unclear but it may be from St. Barnabas feast in June, when the insect appears or a corruption of "Bishop-that-burneth", from the fiery elytra of the beetles.[23]
The ladybird was immortalized in the still-popular children's nursery rhyme Ladybird, Ladybird:
Ladybird, ladybird, fly away home
Your house is on fire and your children are gone
All except one, and that's Little Anne
For she has crept under the warming pan.
This poem has its counterpart in German as Marienwuermchen, collected in Des Knaben Wunderhorn, and set to music by Robert Schumann as Op. 79, No. 13.
Many cultures consider Coccinellids lucky and have nursery rhymes or local names for the insects that reflect this. For instance, the Turkish name for the insect is uğur böceği, literally meaning good luck bug. In many countries, including Russia, Turkey, and Italy, the sight of a coccinellid is either a call to make a wish or a sign that a wish will soon be granted.
In Christian areas, Coccinellids are often associated with the Virgin Mary and the name that the insect bears in the various languages of Europe corresponds to this. Though historically many European languages referenced Freyja, the fertility goddess of Norse mythology, in the names, the Virgin Mary has now largely supplanted her, so that, for example, freyjuhœna (Old Norse) and Frouehenge have been changed into marihøne (Norwegian) and Marienkäfer (German), which corresponds with Our Lady's Bird.[24] Sometimes, the insect is referred to as belonging directly to God (Irish bóín Dé, Romanian vaca-Domnului, Polish boża krówka, all meaning "God's [little] cow").[25] In Dutch it is called lieveheersbeestje, meaning "little animal of our Good Lord".
In both Hebrew and Yiddish, it is called "Moshe Rabbenu's (i.e. Moses's) little cow" or "little horse", apparently an adaptation from Slavic languages. Occasionally, it is called "Little Messiah".[26]
Popular Media
"Ladybugs" or "ladybirds" feature in a range of children's literature as well as in art, TV, and film. They are often depicted in an anthropomorphic or otherwise fictionalised manner.
In Roald Dahl's children's book James and the Giant Peach, The Ladybird (among a cast of anthropomorphic minibeast characters grown to human size) becomes perhaps the closest friend of the protagonist James, treating him with motherly affection. In the American-produced 1996 animated film version, she was voiced by Jane Leeves and renamed The Ladybug, despite Leeves' northern English interpretation.
In the animated film A Bug's Life, Francis the Ladybug (voiced by Dennis Leary) is an aggressive Coccinellid and the clown in P.T. Flea's circus. The contrast between him being a male and a 'lady'bug is a recurring joke in the film.
Tree Fu Tom features ladybirds as "cows" being ranched by a southern-belle cowgirl.
Ledyba and Ledian of the Pokémon series are based on Coccinellids.
As a logo
Bold colors and simple shape have led to use as a logo for a wide range of organizations and companies including:
- Ladybird Books (owned by Pearson PLC)
- Ladybird range of children's clothing sold by Woolworths.co.uk and formerly by the (now defunct) Woolworth's chain store in the UK
- Polish supermarket chain Biedronka
- Atmel AVR Studio software logo
- Estonian mobile operator EMT[27]
- Software development firm Axosoft.
- Symbol of the Finnish Swedish People's Party.
- The ladybird street tile (pictured) is a symbol against senseless violence in The Netherlands and is often placed on the sites of deadly crimes.[28]
In addition, it has been chosen as
- State insect of: Delaware, Massachusetts, New Hampshire, New York, Ohio, and Tennessee, though only New York has selected a species native to the United States (Coccinella novemnotata); the other states have all adopted an invasive European species (Coccinella septempunctata).
- An "official national mascot"[29] for Alpha Sigma Alpha, a national sorority in the United States.
References
- ^ "Chrysomelidae". Integrated Taxonomic Information System. http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=114509.
- ^ White, R.E. 1983. A field guide to the beetles of North America. Peterson Field Guide Series #29.
- ^ Definition of lady cow, Webster's Revised Unabridged Dictionary (1913), provided by die.net. Retrieved 14 November 2008.
- ^ Judy Allen & Tudor Humphries (2000). Are You A Ladybug?, Kingfisher, p. 30
- ^ a b Anon (5 October 2004). "'Deadly ladybird' sighted in UK". BBC News (BBC). http://news.bbc.co.uk/1/hi/england/essex/3715120.stm. Retrieved 17 June 2010.
- ^ Anon. "The Harlequin Ladybird has landed!". The Harlequin ladybird survey. http://www.harlequin-survey.org/. Retrieved 17 June 2010.
- ^ "Everything Ladybug! The source for Ladybug Stuff!". Everything-ladybug.com. http://everything-ladybug.com/ladybug-spots.html. Retrieved 2010-06-22.
- ^ Anon. "Ladybird spotters". UK Ladybird survey. http://www.ladybird-survey.org/lots_about.aspx. Retrieved 17 June 2010.
- ^ Brown, L., ed. (2007). The Shorter Oxford English Dictionary. 1 (6 ed.). pp. 441.
- ^ a b Anon. "Why are ladybirds so-called?". UK Ladybird survey. http://www.ladybird-survey.org/lots_about.aspx. Retrieved 15 October 2010.
- ^ Samaha, John M.. "Marian Roots of the Name". Our Lady's Bug. Dayton, Ohio: International Marian Research Institute. http://campus.udayton.edu/mary/questions/yq/ladybug.html. Retrieved 15 October 2010.
- ^ Anon. "common name: ladybirds, ladybird beetles, lady beetles, ladybugs (of Florida)". Featured creatures. University of Florida. http://entnemdept.ufl.edu/creatures/beneficial/lady_beetles.htm. Retrieved 8 April 2010.
- ^ http://www.treknature.com/gallery/photo268028.htm
- ^ A. Honek, Z. Martinkova & S. Pekar (2007). "Aggregation characteristics of three species of Coccinellidae (Coleoptera) at hibernation sites". European Journal of Entomology 104 (1): 51–56. http://www.eje.cz/pdfarticles/1197/eje_104_1_051_Honek.pdf.
- ^ J. Perry & B. Roitberg (2005). "Ladybird mothers mitigate offspring starvation risk by laying trophic eggs". Behavioral Ecology and Sociobiology 58 (6): 578–586. doi:10.1007/s00265-005-0947-1.
- ^ University of Kentucky-College of Agriculture Cooperative Extension Service
- ^ Anon (5 July 2001). "Phew, what a scorcher!". The Northern Echo. Newsquest media Group. http://archive.thenorthernecho.co.uk/2001/7/5/166513.html. Retrieved 8 April 2010.[dead link]
- ^ Wainright, Martin (17 May 2006). "The great drought". THe Guardian (London: Guardian News and Media Limited). http://www.guardian.co.uk/environment/2006/may/17/water.ethicalliving. Retrieved 8 April 2010.
- ^ DMinSite, a division of Lagniappe Marketing. "Do Ladybugs BITE? And How Can You Keep Them Out of the House?". Gardensalive.com. http://www.gardensalive.com/article.asp?ai=649. Retrieved 2010-06-22.
- ^ Roy, Helen; Peter Brown, Robert Frost, Remy Poland (15 June 2011). Ladybirds (Coccinellidae) of Britain and Ireland. The Field Studies Council. p. 204. ISBN 978-1-906698-20-1.
- ^ Sample, Ian (15 June 2011). "Spot check finds Britain's native ladybirds struggling to compete with alien invaders". The Guardian (Guardian Newspapers ltd.): p. 3. http://www.guardian.co.uk/environment/2011/jun/15/british-ladybird-species-struggling. Retrieved 17 June 2011.
- ^ Timmins, Nicholas (14 October 1994). "The Tories in Bournemouth: Teachers promised support as Shephard calls truce". The Independent (London). http://www.independent.co.uk/news/uk/the-tories-in-bournemouth-teachers-promised-support-as-shephard-calls-truce-1442798.html.
- ^ Lewie C. Roache (1960) Ladybug, Ladybug: What's in a Name? The Coleopterists Bulletin 14(1):21-25.
- ^ "Bishop Barnaby". Notes and Queries 9. 1849-12-29. http://www.gutenberg.org/files/13521/13521-h/13521-h.htm#page132.
- ^ [1][dead link]
- ^ Born to Kvetch, Michael Wex, St. Martin's Press, New York, 2005, ISBN 0-312-30741-1.
- ^ "Parima kvaliteediga kõned". emt.ee. http://www.emt.ee. Retrieved 2010-06-22.
- ^ Anon (April 2004). "Speech by Mrs Maud de Boer-Buquicchio on the occasion of the placement of a ladybird tile at the Council of Europe". Council of Europe. http://www.coe.int/t/e/com/files/pa-sessions/april-2004/disc-coccinelle.asp. Retrieved 21 September 2011.
- ^ ASA.osdcms.com
http://kids.nationalgeographic.com/kids/animals/creaturefeature/ladybug/
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