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Cockchafer

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Common cockchafer
Female
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Clade: Pancrustacea
Class: Insecta
Order: Coleoptera
Suborder: Polyphaga
Infraorder: Scarabaeiformia
Family: Scarabaeidae
Genus: Melolontha
Species:
M. melolontha
Binomial name
Melolontha melolontha
(Linnaeus, 1758)
Synonyms
  • Scarabaeus melolontha Linnaeus, 1758
  • Melolontha vulgaris bioculatus Papp, 1943
  • Melolontha vulgaris bipunctatus Papp, 1943
  • Melolontha vulgaris nigroapicalis Papp, 1943
  • Melolontha vulgaris nigromarginalis Papp, 1943
  • Melolontha vulgaris nigropygidialis Papp, 1943
  • Melolontha vulgaris unicolor Papp, 1943
  • Melolontha vulgaris unipunctatus Papp, 1943
  • Melolontha melolontha amblypyga Lomnicky, 1923
  • Melolontha redtenbacheri Dalla Torre, 1912
  • Melolontha melolontha colpopyga Petz, 1905
  • Melolontha nigritarsis Pfanneberg, 1905
  • Melolontha vulgaris marginata Kraatz, 1888
  • Melolontha vulgaris femoralis Kraatz, 1885
  • Melolontha vulgaris ruficeps Kraatz, 1885
  • Melolontha vulgaris funesta Westhoff, 1884
  • Melolontha vulgaris humeralis Westhoff, 1884
  • Melolontha vulgaris luctuosa Westhoff, 1884
  • Melolontha vulgaris melanopus Westhoff, 1884
  • Melolontha vulgaris obscuripes Westhoff, 1884
  • Melolontha vulgaris scapularis Westhoff, 1884
  • Melolontha vulgaris pulcherrima Dalla Torre, 1879
  • Melolontha vulgaris nigra Kellner, 1876
  • Melolontha albida Mulsant, 1842
  • Melolontha vulgaris discicollis Mulsant, 1842
  • Melolontha vulgaris lugubris Mulsant, 1842
  • Melolontha vulgaris ruficollis Mulsant, 1842
  • Scarabaeus majalis Moll, 1785
  • Melolontha vulgaris Fabricius, 1775

The common cockchafer (Melolontha melolontha), also colloquially known as the Maybug,[1][a] Maybeetle,[3] or doodlebug,[4] is a species of scarab beetle belonging to the genus Melolontha. It is native to Europe, and it is one of several closely-related and morphologically similar species of Melolontha called cockchafers, alongside Melolontha hippocastani (the forest cockchafer).

The cockchafer develops via metamorphosis, in which the beetle undergoes stages of eggs, larvae, pupae and adults.

The mating behaviour is controlled by pheromones. The males usually swarm during the mating season while the females stay put and feed on leaves.[5] The leaves release green leaf volatiles when they are fed on by females, which the male can sense and thus locate the female for mating opportunity.[6][7] The larvae use both the plant volatiles and CO2 to locate the plant root for food.[8]

This species is an important and nutritious food source for many species. The adults and larvae feed on plants, and are regarded as agricultural pests of crops such as grasses and fruit trees. Adults have harmful effects for the crop when they aggregate in large groups. The larvae can cause severe damage and kill the plant by gnawing the plant roots.[9]

Etymology

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The name "cockchafer"[10] derives from the late-17th-century usage of "cock"[11] (in the sense of expressing size or vigour) + "chafer"[12] which simply means an insect of this type, referring to its propensity for gnawing and damaging plants. The term "chafer" has its root in Old English ceafor or cefer, of Germanic origin and is related to the Dutch kever, all of which mean "gnawer" as it relates to the jaw. As such, the name "cockchafer" can be understood to mean "large plant-gnawing beetle" and is applicable to its history as a pest animal. The genus name "Melolontha" originates from ancient Greek which means "fig-sheep", because it tends to actively feed on wild figs.[13]

Description

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Adults

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Detail of a male cockchafer, showing the seven leaflike extensions on each antenna

Adults of M. melolontha reach sizes of 25 to 30 millimetres (1 to 1+14 inches) in length.[5] They have a dark head with two antennae which has multiple segments on each. Behind their heads, they have a dark pronotum covered with short hairs. This black coloration distinguishes them from their close relative M. hippocastani, whose pronotum is brown. The top of their bodies have reddish brown elytra and a black abdomen which is partly white on the sides. Male cockchafers have seven flattened leaflike extensions on their antennae, whereas the females have only six.[5]

Larvae

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M. melolontha larvae typically has three stages of development over the course of 3–4 years. In the first stage (L1), they are 10–20 mm long which then grows to 30–35 mm in the second year of development (L2), and then finally reaches a full size of 40–46 mm in their final year of development (L3) before emerging.[5] In some areas of Eastern Europe, the larvae develops for a fourth year (L4). It has a white body which curves into an arc with a black coloration at the abdomen and a long, hairy, and well developed legs.[5] It has a large orange head with strong, grabbing mandibles. On the head, it has 2 small antennae which it uses to smell and taste the surroundings while being underground.[8][citation needed]

Intestinal components and microbiome

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The gut enzymes and microbiota of M. melolontha larvae allow them to exploit a variety of ecological niches which include low energy foods, rotting organic matter and freshly growing roots in the soil.[14] There are two major compartments in the scarabaeid larvae intestinal tract. The first is a tubular midgut that secretes hydrolytic enzymes for macromolecule breakdown, and the second is a bulbous hindgut used for fermentation. In most individuals of M. melolontha, the bacterial diversity in the midgut was shown to be comparatively reduced than the same in diet, indicating lysis of the bacteria.[15]

In the midgut, glucose is broken down and absorbed by the epithelium.[citation needed] It has been shown that proteolytic breakdown of toxins is a common resistance mechanism for agricultural pests.[16] Proteolytic activity of enzymes in the midgut is hypothesised to increase resistance to entomopathogenic bacteria such as BT toxin in the beetle larvae. Trypsin-like enzymes from the midgut of M. melolontha have been found to break down certain bacterial toxins and inactivate them.[citation needed]

The hindgut in melolontha has a low concentration of glucose caused by high fermentation or low hydrolysis of recalcitrant residues such as cellulose.[15] Acetate is a major product of this fermentation, suggesting that much of the bacteria in the hindgut is homoacetogenic. High abundance of species in the bacterial genus Desulfovibrio in the hindgut wall suggests that sulphate reduction in the beetle is caused by this bacteria, but the source of this sulphate in the diet is unknown.[15]

Some research on the M. melolontha microbiome has been focused on increasing the entomopathogenic properties of nematodes used as pest control due to their symbiosis. Bacteria such as Xenorhabdus nematophila are transported by nematodes and released into the insect's midgut.[14] The bacteria will release lytic enzymes and other antimicrobial substances to decrease competition from the beetle's native microbiome. This creates an optimal environment for nematode development.[citation needed] Bacterial species in the midgut of melolontha such as Pseudomonas chlororaphis have been found to fight back, acting as antagonists to entomopathogenic bacteria. These bacteria have been identified differentially in different larval stages, with P. chlororaphis usually being found in the third and final larval stage.[citation needed]

Distribution and habitat

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Cockchafers are prevalent across Europe, including in Germany, France, and the United Kingdom. They are particularly prevalent in temperate regions with suitable soil conditions for larval development. However, they have also been reported in parts of Asia, including Turkey and the Caucasus region.[13][17] Geographical barriers, climatic conditions, and ecological factors may limit their dispersal to other continents for e.g., M. melolontha has never been reported to be in the mountainous regions of Tyrol because it cannot survive in the Alps' high altitude and low temperature during colder months. Also they do not fly more than 2 to 3 km from where they emerge and remain in close proximity of the field or grasslands where they occur.[18]

Life cycle

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Female M. melolontha Beetle.

Adults appear at the end of April or in May and live for about five to seven weeks. After about two weeks, the female begins laying eggs, which she buries about 10 to 20 cm deep in the earth. She may do this several times until she has laid between 60 and 80 eggs. Most typically, the female beetle lays its eggs in fields. The preferred food for adults is oak leaves, but they will also feed on conifer needles.

The larvae, known as "chafer grubs" or "white grubs", hatch four to six weeks after being laid as eggs. They feed on plant roots, for instance potato roots. The grubs develop in the earth for three to four years, in colder climates even five years, and grow continually to a size of about 4–5 cm, before they pupate in early autumn and develop into an adult cockchafer in six weeks.[5]

The cockchafer overwinters in the earth at depths between 20 and 100 cm. They work their way to the surface only in spring.

Because of their long development time as larvae, cockchafers appear in a cycle of every three or four years; the years vary from region to region. There is a larger cycle of around 30 years superimposed, in which they occur (or rather, used to occur) in unusually high numbers (10,000s).

Behaviour and ecology

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Mating

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Male M. melolontha Beetle.

Males leave the soil when the temperature is favourable in April or May. Sexual dimorphism is observed as male beetles, at dusk, will begin to swarm and locate around groups of trees at forest edges, which ends in darkness.[5] On the other hand, females will stay in place and feed on leaves until they reach sexual maturity. Males primarily fly around the branches looking for females to mate with.[6][7] The mating behaviour on the trees lasts for several hours. The male beetles engage in the process for about 10-20 days.[5] These swarms typically have minimal damage to the trees, but they are occasionally harmful in cherry or plum orchards because of their consumption of blossoms. Once the females have matured and mated, they return to the fields to lay their eggs 15 to 25 cm (0.49 to 0.82 ft) in the soil in a batch of 10 to 38 eggs. The females stay 2 to 4 days on the soil to protect the eggs. Only a third of females will survive this trip, but any survivors will make a second, and occasionally third, swarming trip and return to the field to lay eggs again. Number of days for M. melolontha eggs to mature depends on temperature—5 to 8 days at 27°C and 23 to 32 days at 32°C.[5]

Green leaf volatiles (GLVs) are a series of saturated and monounsaturated six-carbon aldehydes, alcohols, and esters released by vascular plants in response to stresses.[19] GLVs have been found to act as a sexual kairomone, that helps M. melolontha adults in mate finding, which is a compound released by the adult beetle that only benefits the receiver.[6][7] This enhances the attractiveness of toluquinone, a sex pheromone in scarab beetles which is released by the female to attract the male beetles.[20] Only male melolontha are attracted to GLVs particularly (Z)-3-hexen-1-ol, (E)-2-hexen-1-ol and 1-hexanol, using its release to identify leaves that female beetles are feeding on. Female melolontha are not attracted to any of the GLV compounds. [6][7] M. melolontha males are more sensitive to lower GLV concentrations, possibly due to the anatomical differences between male and female antennae.[7] Due to this phenomenon, sexual dimorphism can be observed in flight behaviour. This makes the melolontha beetle the first reported insect of needing both the sex pheromone and plant volatiles for mating. During swarming behaviour, males will hover around the foliage while females remain on twigs and branches to feed.[5] Males then use GLVs to identify which leaves have females that they can mate with.[7] GLVs are being investigated as a possible pest control technique to attract males and prevent mating.[citation needed]

Pest

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Though adults can damage some fruit trees, M. melolontha larvae are the primary agricultural pests.[5] Larva hatch from their eggs 4–6 weeks after being laid and develop into adults over the course of 3–4 years. Immediately after hatching, larvae will gnaw on small roots. It will continue feeding on roots, particularly grasses, cereals, and other crops, during its three larval stages, only pausing to burrow deep into the soil for winter hibernation.[5]

In their first stage, M. melolontha larvae depend on soluble compounds such as sugar, amino acids and isoflavonoids which is released by plant roots in rhizosphere. Hence it gets attracted by the CO2 released by respiring plant roots and soil microorganisms.[5][8] Above a density of 1000 larvae per m2, the damage to soil by melolontha larvae is noticeable.[5] In their second stage, larva will cause the most damage to crops such as Strawberry.[8] In their third stage, larva will do less but still severe damage to crops. They most prominently use structures on their antennae called pore plates (or lamellae) to smell. This structure is a thin layer of cells that covers a number of sensory units consisting of dendrite bundles. These and other olfactory organs namely maxillary and labial palp on the head of the larva can identify CO2 and plant volatiles. They've also been found to push their heads into the walls of their burrows and probe with their antennae, likely to taste the soil with bristle-like sensilla.[8]

Ecological impact

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Environmental factors such as prevailing soil temperature, humidity, and plant type have a considerable impact on the existence and behaviour of cockchafers in wooded environments. It indicates that cockchafer populations are strongly influenced by climatic conditions, with warmer temperatures and higher humidity level favouring their occurrence but adversely and severely affects the larvae as it causes desiccation. Additionally, specific vegetation types, including deciduous trees and shrubs, provide suitable habitats for cockchafers, facilitating their survival and reproduction within forest stands.[21][22]

History of pest control

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Larva (grub)

Middle ages

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In the Middle Ages, pest control was rare, and people had no effective means to protect their harvest. This gave rise to events that seem bizarre from a modern perspective. In 1320, for instance, cockchafers were brought to court in Avignon and sentenced to withdraw within three days onto a specially designated area, otherwise they would be outlawed. Unsurprisingly, since they failed to comply, they were collected and killed. Similar animal trials also occurred for many other animals in the Middle Ages.[23]

19th century

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Both the grubs and adults have a voracious appetite and thus have been and sometimes continue to be a major problem in agriculture and forestry. In the pre-industrialised era, the main mechanism to control their numbers was to collect and kill the adult beetles, thereby interrupting the cycle. They were once very abundant: in 1911, more than 20 million individuals were collected in 18 km2 of forest.[1] Collecting adults was an only moderately successful method.

It has been reported that cockchafers were consumed in parts of Europe during the 19th century. In 1844, German medical officer Johann Joseph Schneider reported that in his hometown of Fulda in Hesse cockchafers were regularly consumed raw, as well as sugared, candied and in soup. Schneider described the flavour of the soup as similar to crab soup. However, it is unclear how prevalent the consumption of cockchafers including in soup really was (all recipes for cockchafer soup in German are based on Schneider's 1844 version), and is has been suggested that its prevalence has been exaggerated since the 19th century by newspaper stories due to its shocking and sensational nature, leading to it being described as a "myth".[24]

In Sweden, the peasants looked upon the grub of the cockchafer as furnishing an unfailing prognostic whether the ensuing winter will be mild or severe; if the animal has a bluish hue (a circumstance which arises from its being replete with food), they affirm it will be mild, but if it is white, the weather will be severe: and they carry this so far as to foretell, that if the anterior be white and the posterior blue, the cold will be most severe at the beginning of the winter. Hence they call this grub Bemärkelse-mask—prognostic worm.[25]

Modern times

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Only with the modernisation of agriculture in the 20th century and the invention of chemical pesticides did it become possible to effectively combat the cockchafer. Combined with the transformation of many pastures into agricultural land, this has resulted in a decrease of the cockchafer to near-extinction in some areas in Europe in the 1970s.

Since the 1970s, agriculture has generally reduced its use of pesticides. Because of environmental and public health concerns (pesticides may enter the food chain and thus also the human body) many chemical pesticides have been phased out in the European Union and worldwide. In recent years, the cockchafer's numbers have been increasing again, causing damage to agricultural use of over 1,000 square kilometres (400 sq mi) of land all over Europe (0.001% of land).

Due to legal provisions from the European Union for the sustainable use of pesticides, aerial treatment, which had been used to successfully control M. melolontha populations, is now banned.[26] Light traps have been successful in attracting M. melolontha adults, particularly males, when put at a height of 4 metres (10 ft) above the ground. If a peak swarming time can be identified, shaking isolated trees and collecting feeding adults can reduce population, though it is time consuming.[26] Azadirachtin is a chemical that inhibits maturation feeding and egg development, and not immediate mortality, but low persistence and difficulty spraying it high enough in trees prevents widespread use.[26] Another pest control method, soil tilling, can be used for reducing crop damage by melolontha but it's effectiveness depends on the biological cycle of the beetle e.g., it is effective for grubs and not adults.[26][27] Pre-cropping as a pest control method works by buckwheat mixed with mustard or other leguminous plants to reduce a significant number of grubs by both weight and population before the crop of interest is planted.[26] Sex pheromones have been used for mass trapping, mating disruption, and "Attract and Kill" methods. The unlikelihood of developing resistance due to the sex pheromones being produced by the beetles makes this a promising method of pest control.[5]

Entomopathogens

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Entomopathogenic organisms—organisms that produce disease in insects—are an active area of research for the control of M. melolontha grub populations.[28] Entomopathogenic fungi is currently being studied as a way to control M. melolontha grub populations. Beauveria brongniartii has been found to be overall ineffective on the Melolontha species because of a lack of high quality fungi preparation and its inability to blend with the soil,[27] and B. bassiana has been successful with other agricultural pests.[28] B. brongniartii has not been allowed in the EU to be used as a commercial plant protection method and hence not used in cockchafer control.[28] Entomopathogenic nematodes such as Steinernematidae and Heterohabditis, have been found to be particularly successful ways of reducing populations, particularly when larvae are in the first (L1) and second (L2) stage.[27] Entomopathogenic bacteria such as Paenibacillus popillae and several species of Serratia have been found to be varyingly effective on melolontha for e.g., P. popillae alone was ineffective on the beetle but combined with B. brogniartii fungi, the disease incidence in the organism had increased.[27] The focus on entomopathogenic bacteria has been on their symbiosis with entomopathogenic nematodes and their ability to act together as a larval control strategy.[27] Poor results with the application of these methods have stemmed an intensive research into the proteolytic enzymes and microbiome of M. melolontha to determine if these factors are acting as a defense against entomopathogenic organisms.[16]

Interaction with other species

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Diet

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Cockchafer feeds on deciduous plant and fruit tree leaves, including oaks, maple, sweet chestnut, beech, plum, and walnut trees. The feeding behaviour of larvae can cause severe damage to the plants. They feed on both the small roots of field plants such as grain, grass, tree, beet roots and the large part of crop rootlets. Larvae can gnaw the root for 30 cm each day, which quickly kills the plant.[9]

Predators

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The European mole is a natural predator of cockchafers. Moles are known to feed on cockchafer larvae. They can detect them using their keen sense of smell and specialised digging behaviour. This predation can help regulate cockchafer populations in mole-inhabited areas.[29]

M. melolontha adults are predated by ground beetles (Carabidae) and ants (Formicidae). Larvae are predated by click beetles (Elateridae) while underground. Starlings, crows, and gulls also predate M. melolontha larvae, often after a field has been plowed.[5]

Parasites

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Dexia rustica is a parasitic fly that uses M. melolontha larvae as their hosts. D. rustica eggs hatch underground and search for hibernating cockchafer larvae. Their presence will ultimately kill the beetle larvae in spring when the fly hibernates. About 30 to 35 cm (0.98 to 1.15 ft) below the ground, over 10 percent of the grub (or larvae) can be parasitised by one to six D. rustica.[5]

Relationship with humans

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Max and Moritz shaking cockchafers from a tree
A group of cockchafers in Ukraine

Children since antiquity have played with cockchafers. In ancient Greece, boys caught the insect, tied a linen thread to its feet and set it free, amusing themselves to watch it fly in spirals. English boys in Victorian times played a very similar game by sticking a pin through one of its wings.[30] Nikola Tesla recalls that as a child he made one of his first "inventions", an "engine" made by harnessing four cockchafers in this fashion.[31]

Cockchafers appear in the fairy tales "Thumbelina" by Hans Christian Andersen and "Princess Rosette" by Madame d'Aulnoy.

The cockchafer is featured in a German children's song similar to the English "Ladybird, Ladybird":

Maikäfer, flieg!
Der Vater ist im Krieg,
die Mutter ist in Pommerland,
Pommerland ist abgebrannt –
Maikäfer flieg!

Translation:

Cockchafer, fly!
Father is at war,
Mother is in Pomerania,
Pomerania is burned to the ground –
Cockchafer, fly!

The verse dates back to the Thirty Years' War in the first half of the 17th century, in which Pomerania was pillaged and suffered heavily. Since World War II, it is associated in Germany with the closing months of that war as well, when Soviet troops advanced into eastern Germany.

Example of decorative trim, sourcils de hanneton, which translates to 'cockchafer eyebrows'.

According to one source, the dumbledore in Thomas Hardy's 1899 poem An August Midnight[32] is a cockchafer.[33] However, in his novel The Mayor of Casterbridge, Hardy uses the dialect word dumbledore to mean a bumble bee.[34]

There have been four Royal Navy ships named HMS Cockchafer.

Decorative trims of women's dresses between the 1750s and 1780s, were known as sourcils de hanneton, translating to 'cockchafer eyebrows'. This referred to the beetles fan-like antennae that resembled the decorative trim made with knotted silk floss.[35]

See also

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Explanatory notes

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  1. Other names include: bracken clock, bummler, chovy, cob-worm, dorrs, dumbledarey, dumbledore, humbuz, June bug, kittywitch, billy witch, may-bittle, midsummer dor, mitchamador, oak-wib, rookworm, snartlegog, spang beetle, tom beedel and chwilen y bwm (Welsh).[1][2]

Citations

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  1. 1 2 3 "Common Cockchafer". Bug Life. Archived from the original on 2019-04-08. Retrieved 2016-07-04.
  2. Marren, Peter; Mabey, Richard (2010). Bugs Britannica. Chatto & Windus. ISBN 978-0-7011-8180-2.
  3. "Cockchafer | insect". Encyclopedia Britannica. Archived from the original on 2021-07-09. Retrieved 2021-07-01.
  4. "7 things you never knew about the cockchafer". Discover Wildlife. 8 April 2014. Archived from the original on 23 March 2016. Retrieved 4 July 2016.
  5. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Huiting, H. F.; Moraal, L. G.; Griepink, F. C.; Ester, A. (2006). Biology, control and luring of the cockchafer, Melolontha melolontha: literature report on biology, life cycle and pest incidence, current control possibilities and pheromones. Praktijkonderzoek Plant & Omgeving. BO-10-006-21. Archived from the original on 2023-12-01. Retrieved 2024-02-28.
  6. 1 2 3 4 Reinecke, Andreas; Ruther, Joachim; Tolasch, Till; Francke, Wittko; Hilker, Monika (2002-06-01). "Alcoholism in cockchafers: orientation of male Melolontha melolontha towards green leaf alcohols". Naturwissenschaften. 89 (6): 265–269. Bibcode:2002NW.....89..265R. doi:10.1007/s00114-002-0314-2. ISSN 0028-1042. PMID 12146792. S2CID 25772038.
  7. 1 2 3 4 5 6 Reinecke, Andreas; Ruther, Joachim; Hilker, Monika (April 2005). "Electrophysiological and behavioural responses of Melolontha melolontha to saturated and unsaturated aliphatic alcohols". Entomologia Experimentalis et Applicata. 115 (1): 33–40. Bibcode:2005EEApp.115...33R. doi:10.1111/j.1570-7458.2005.00274.x. ISSN 0013-8703. S2CID 84471627. Archived from the original on 2024-11-11. Retrieved 2024-02-28.
  8. 1 2 3 4 5 Eilers, Elisabeth J.; Talarico, Giovanni; Hansson, Bill S.; Hilker, Monika; Reinecke, Andreas (2012-07-25). "Sensing the Underground – Ultrastructure and Function of Sensory Organs in Root-Feeding Melolontha melolontha (Coleoptera: Scarabaeinae) Larvae". PLOS ONE. 7 (7) e41357. Bibcode:2012PLoSO...741357E. doi:10.1371/journal.pone.0041357. ISSN 1932-6203. PMC 3405142. PMID 22848471.
  9. 1 2 Fraval, A. (1998). "HYPP Zoology".
  10. cockchafer (n.) Archived 2022-04-14 at the Wayback Machine www.etymonline.com accessed 30 May 2021
  11. cock (n.1) Archived 2021-12-04 at the Wayback Machine www.etymonline.com accessed 30 May 2021
  12. chafer (n.) Archived 2022-04-13 at the Wayback Machine www.etymonline.com accessed 30 May 2021
  13. 1 2 Ashworth M, Darwin Tree of Life Barcoding collective, Wellcome Sanger Institute Tree of Life programme et al. The genome sequence of a cockchafer, Melolontha melolontha (Linnaeus, 1758) [version 1; peer review: 2 approved, 2 approved with reservations]. Wellcome Open Res 2023, 8:222 (https://doi.org/10.12688/wellcomeopenres.19434.1)
  14. 1 2 Skowronek, Marcin; Sajnaga, Ewa; Pleszczyńska, Małgorzata; Kazimierczak, Waldemar; Lis, Magdalena; Wiater, Adrian (2020-01-16). "Bacteria from the Midgut of Common Cockchafer (Melolontha melolontha L.) Larvae Exhibiting Antagonistic Activity Against Bacterial Symbionts of Entomopathogenic Nematodes: Isolation and Molecular Identification". International Journal of Molecular Sciences. 21 (2): 580. doi:10.3390/ijms21020580. ISSN 1422-0067. PMC 7013910. PMID 31963214.
  15. 1 2 3 Egert, Markus; Stingl, Ulrich; Dyhrberg Bruun, Lars; Pommerenke, Bianca; Brune, Andreas; Friedrich, Michael W. (August 2005). "Structure and Topology of Microbial Communities in the Major Gut Compartments of Melolontha melolontha Larvae (Coleoptera: Scarabaeidae)". Applied and Environmental Microbiology. 71 (8): 4556–4566. Bibcode:2005ApEnM..71.4556E. doi:10.1128/AEM.71.8.4556-4566.2005. ISSN 0099-2240. PMC 1183286. PMID 16085849.
  16. 1 2 Wagner, Wolfgang; Möhrlen, Frank; Schnetter, Wolfgang (July 2002). "Characterization of the proteolytic enzymes in the midgut of the European Cockchafer, Melolontha melolontha (Coleoptera: Scarabaeidae)". Insect Biochemistry and Molecular Biology. 32 (7): 803–814. Bibcode:2002IBMB...32..803W. doi:10.1016/S0965-1748(01)00167-9. PMID 12044497. Archived from the original on 2024-04-18. Retrieved 2024-02-28.
  17. UK, CAB International (2005-12-13). "Melolontha melolontha . [Distribution map]". Distribution Maps of Plant Pests. doi:10.1079/DMPP/20056600193. ISSN 1369-104X.
  18. Pedrazzini, Chiara; Strasser, Hermann; Zemp, Niklaus; Holderegger, Rolf; Widmer, Franco; Enkerli, Jürg (2023). "Spatial and temporal patterns in the population genomics of the European cockchafer Melolontha melolontha in the Alpine region". Evolutionary Applications. 16 (9): 1586–1597. Bibcode:2023EvApp..16.1586P. doi:10.1111/eva.13588. ISSN 1752-4571. PMC 10519412. PMID 37752964.
  19. Matsui, Kenji; Engelberth, Jurgen (2022-10-31). "Green Leaf Volatiles—The Forefront of Plant Responses Against Biotic Attack". Plant and Cell Physiology. 63 (10): 1378–1390. doi:10.1093/pcp/pcac117. ISSN 0032-0781. PMID 35934892.
  20. Reinecke, Andreas; Ruther, Joachim; Hilker, Monika (2002-03-10). "The scent of food and defence: green leaf volatiles and toluquinone as sex attractant mediate mate finding in the European cockchafer Melolontha melolontha". Ecology Letters. 5 (2): 257–263. Bibcode:2002EcolL...5..257R. doi:10.1046/j.1461-0248.2002.00318.x. ISSN 1461-0248. Archived from the original on 2022-02-27. Retrieved 2026-06-21.
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Cockchafer
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