Diabrotica | |
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Scientific classification![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Coleoptera |
Suborder: | Polyphaga |
Infraorder: | Cucujiformia |
Family: | Chrysomelidae |
Subfamily: | Galerucinae |
Tribe: | Luperini |
Subtribe: | Diabroticina |
Genus: | Diabrotica Chevrolat inDejean, 1836 [1] |
Diabrotica is a large, widespreadgenus ofbeetles in thefamilyChrysomelidae. Members of this genus include several destructiveagricultural pestspecies, sometimes referred to ascorn rootworms.
There are an estimated 400 species grouped into the genusDiabrotica.[2] WithinDiabrotica there are 3 series: fucata, virgifera, andsignifera groups. Thefucata series contains the majority ofDiabrotica diversity with 354 species.[2]Fucata are characterized as multivoltine, producing two broods of offspring per year.[2] Thevirgifera series comprises 24 species, and thesignifera series contains 11. Additionally,virgifera andsignifera are characterized as univoltine, and will only brood one set of offspring annually.[2]
There is very limited information on most species ofDiabrotica.[2] Majority of the research conducted on theDiabroticagenus focuses on investigating species of consequential economic importance, such asD. balteata,D. barberi,D. undecimpunctata howardi,D. virgifera,D. speciosa. MultipleDiabrotica species are considered major agricultural pests, therefore the control and management of populations is of significant importance to farm management strategies.[2]
TheDiabrotica genus arose in theCretaceous period and began to diversify and speciate ~60 million years ago, which culminated ~30-40 million years ago.[2] Contrary to previous suggestions, Diabrotica speciation was not linked to the onset of corn and other crop cultivation. Gene sequencing has led to a phylogenetic reconstruction of theDiabrotica genus that indicates monophagy is the ancestral trait. The genus began to diversify when certain lineages expanded their diets to include a multitude of different types of plants, leading to a polyphagous trait.[2] However, subsequent reversals back to monophagy or oligophagy have occurred in some evolutionary branches.
The diversification and evolution ofDiabrotica species is closely linked to their relationship with wild plant species in the familyCucurbitaceae (cucurbits), which characteristically produce cucurbitacin secondary compounds.[2] Cucurbitacins are bitter and toxic. While cucurbitacins deter most herbivores,Diabrotica beetles are attracted to cucurbitacins and compulsively feed on cucurbit species, especially the tissues that contain high concentrations of cucurbitacins, such as roots, seeds, and cotyledons.Diabrotica beetles favour cucurbitacin-containing plants to the extent that they will leave another nutritious plant host for a cucurbit plant.[2] The mouthparts ofDiabrotica beetles display receptors that bind cucurbitacins to stimulate this compulsive feeding behaviour. By feeding on cucurbit plants and sequestering cucurbitacin in their haemolymph,Diabrotica beetles are afforded some protective advantages, such as chemical defenses against natural predators. This is an example of pharmacophagy, in which insects consume plant metabolites for reasons besides nutrition.[2] The aforementioned demonstrates a situation of chemically mediated coevolution betweenDiabrotica and cucurbit plants, and evenDiabrotica species that have evolved to no longer rely on cucurbitacin-containing plants still demonstrate this compulsive feeding behavior in the presence of these plants.
Diabrotica species are separated into 3 groups:virgifera,fucata, andsignifera.[3] These are ‘groups of convenience’ as they are based on host range, diet, life history, and other ecological traits, rather than being supported by molecular and genetic data.
Diabrotica is a neotropical genus that evolutionarily originated in Central America and is native to North and South America.[2] Central America is the most rich inDiabrotica species (i.e. has the highest number ofDiabrotica species), but Mexico and Brazil are also high inDiabrotica diversity. Although the tropical areas are significantly more diverse inDiabrotica species, the USDiabrotica fauna has a greater proportion of pest species.[4] 4 out of the 7Diabrotica species native to the US are pests. However, only 6 out of the 338Diabrotica species found in the tropical regions are pests.
Climate is the main factor that puts constraints on the distribution of different groups ofDiabrotica species.[4] For example, the entirefucata group is incapable of overwintering. Consequently, members of thefucata group inhabit host plants primarily located in the tropics and subtropics.[2] However, variations in weather conditions from year to year lead to annual fluctuations in the geographic ranges offucata group species.[4]D. speciosa sensu lato, regarded as the best known pest species within thefucata group in South America, has a broad distribution covering the majority of the continent.Signifera group species are exclusively found within South America.[2] A multitude of species in thevirgifera group are capable of overwintering.[4] Specifically the USvirgifera group species overwinter as cold-resistant eggs.
Thewestern corn rootworm,D.virgifera, is the most damaging pest of corn crops in the US and was accidentally brought to Serbia.[2] As a result, its range expanded to threaten corn crops in the Eastern and Central areas of Europe, especially in Germany and Hungary. Belgium, Netherlands, and the UK were successful in extirpating in western corn rootworm. Through various strategies, such as crop rotation,Diabrotica beetles have been kept at non-damaging levels in Germany and France.[2]
Since adultDiabrotica are highly motile and have the ability to migrate, the genus occupies diverse habitats.[3]Diabrotica movement patterns are driven by the appeal and availability of food sources. Typically, the preferred food source of adultDiabrotica includes pollen and reproductive structures of plants. Species of thefucata group are polyphagous, which means that they feed on various plant species.[2] Species of thevirgifera andsignifera groups are oligophagous, which means that they consume only a few plant species.
Diabrotica species in thevirgifera group that feed on corn primarily inhabit huge maize monocultures.[2] BothD. virgifera andD. longicornis barberi are corn-feeding species within thevirgifera group; however, their feeding and egg-laying behaviours exhibit some key differences.[5] Both species feed on silks, pollen, and young kernels of corn. AdultD. virgifera also feed on corn leaves, butD. l. barberi do not. In fact,D. virgifera will remain in a corn field past the seasonal availability of pollen and silk to feed on the corn foliage, especially when their egg-laying period is prolonged. However, once silk and pollen are no longer available in a corn crop,D. l. barberi will move on and inhabit other fields of plants.[5] As a result, high densities ofD. l. barberi larvae are commonly present in crops where corn was not cultivated during the prior year. This is seldom observed forD. virgifera larvae, rather they usually remain in the corn field. Research has revealed thatD. virgifera populations tend to rise when corn is continuously planted in the same field each year, butD. l barberi populations tend to benefit more from crop rotation.[5] All together, it suggests thatD. virgifera are much more dependent on corn as a habitat and food source thanD. l. barberi.D. undecimpunctata howardi is a member of the fucata group that uses a broad range of plant species and does not rely on corn as long as other nutritive food sources are present.[3]
Adult femaleDiabrotica that are feeding on the leaves and pollen of the host plant release their eggs (~300-400 eggs) in the soil adjacent to the plant's roots.[2] Eggs can be deposited as deep as 15 cm into the soil. After the eggs hatch, larval development occurs. InDiabrotica larval development, there are 3instar larva, which are stages in Arthropod life cycles that occur between moults. Following the 3rd instar, is a maturepupa, which is the stage exhibiting complete metamorphosis to the adult form. As larvae develop, they feed on and thus damage the roots of the host plant.[2]
For multivoltine species (2 or more broods of offspring each year), the whole lifecycle (egg to adult) lasts around 30 days.[2] In addition, these species usually spend the winter as dormant adults suspended in their development (diapausing adults). In contrast, eggs overwinter in univoltine species. Thevirgifera group species are univoltine andfucata group species are multivoltine.[3]
The corn rootworms, which includeD. virgifera virgifera,D. virgifera zaea, andD. longicornis barberi, are the 3 most economically relevant taxa due to their damage on agricultural crops, especially corn.[5] These taxa are univoltine, thus they spend the winter as eggs deposited in soil and hatch in the late spring. Adults can be seen feeding on corn silk, leaves, and pollen from the midsummer through to the frost.
Diabrotica larvae, specificallyDiabrotica viridula, are prominent pests of agricultural cornfields,[2] as the larvae's preferred food source primarily consists of corn roots.[2] Adult females intentionally deposit the eggs of their brood in the soil of cultivated farm fields.[5] This oviposition upon successfully hatching, allows larvae the ability to easily locate and tunnel towards the roots of the host plant.[5] There exists a proportionality between the density of laid eggs around the targeted host plant, and the damage inflicted upon the roots of the plant.[6] The closer eggs are laid to the host results in greater propensity for damage to incur on the host as a result of larval feeding. Other Diabrotica species larvae within thefucata series are predominantly polyphagous in nature, feeding off a variety of diverse vegetation.[4]
AdultDiabrotica species are herbivores, with a diet predominantly consisting of foliage and pollen.[2] Due to the vast richness of diversity in the genus a broad variety of plant species are used as host plants and food sources. Host sources of economic importance in prairie habitats include, but are not limited to, corn, squash, beans, and soybean varieties.[2] ManyDiabrotica species exhibit a preference for plants containing cucurbitacin's as they can be metabolized by the beetle into an effective chemical defense mechanism.[2]
Chemical defense is a key mechanism implemented by manyDiabrotica species to act as an important mode of protection against predation.[2] Cucurbitacin is a bitter tasting compound found in certain plants that is often toxic to insects.[2]Diabrotica species will relentlessly ingest cucurbitacin compounds synthesized by their preferential food sources.[2] After acquiring the cucurbitacin compounds, they metabolize the consumed cucurbitacin into the haemolymph.[2] TheDiabrotica beetle will have then successfully incorporated the bitter tasting properties of cucurbitacin compounds into itself. Therefore, the adoption of the deterring bitter, toxic characteristics of the cucurbitacin's results in theDiabrotica species significantly reducing the likelihood of predation.[2]
Female members ofDiabrotica species will release sex pheromones and begin expressing a characteristic calling posture signaling their receptivity to commence mating.[6] The calling posture consists of exposing the membranes of the females abdominal segment's to facilitate the secretion of the aforementioned sex pheromones.[6] Reproducing males will then approach the female and engage in tactile stimulation of the female's abdomen.[6] This likely facilitates relaxation and increases the females receptiveness prior to and throughout the duration of copulation occurring.[6] The mating procedure may last for 1–6 hours, with copulation lasting approximately 10–60 minutes.[6] Mating behavior appears to be influenced by circadian rhythms, with the majority of mating events occurring around the dusk hours.[7]
Lek formation is a behavioral pattern observed inDiabrotica species that preferentially consume cucurbitacin rich foods.[2] During copulation, the male will transfer detoxified cucurbitacin compounds to their female mate.[2] This transplantation of metabolized cucurbitacin's decreases the metabolic burden on brooding females to sequester energy to both brood offspring and synthesize chemical defense mechanisms against predation.[2]
Due to the pestilent species of univoltineDiabrotica lifecycles being intrinsically tied to specific host plants, agronomic practices such as crop rotation, shifting sowing times, and alternative tilling techniques are viable methods to decrease the likelihood of infestation in cultivated fields.[8] Although these techniques are effective for the management of univoltineDiabrotica species, multivoltine species remain largely unaffected by the implementation of these measures.[8] To mitigate the damage of multivoltineDiabrotica pests, early plantation of the crops to allow ample time for maturation is moderately effective at avoiding the larval stage ofDiabrotica when the crop is in its most vulnerable stage of development.[8] This provides the crop adequate opportunity to grow and increase resilience against larval feeding attacks.[8] However, this method is difficult to execute effectively due to variable hatch periods of overwintered eggs, and the dependence on correct seasonal timing making it susceptible to failure.[8]
The most effective treatment against pestilentDiabrotica larvae stage in maize cultivation involves the application of organophosphate and phenyl pyrazole insecticides into the seed furrow during planting.[8] For potatoes, in-furrow application of neonicotinoids is also an effective pesticide for controlling larvae.[8] These insecticides disrupt the development of the beetle's larval stages, drastically decreasingDiabrotica population's biotic potential, ultimately contributing to effective infestation control.[8]
Genetically modified crops are another method to reduce economic loss due toDiabrotica pests.[8] High risk crops can be genetically modified to decrease susceptibility to root damage subjected by the feeding of larvae providing protection against pest-related damage.[8]
Artificial selection of chemical defenses when growing particular crops can be utilized to increase resistance toDiabrotica pests.[8] For example, leptins can act as insecticidal agents, and glycoalkaloids confer a natural resistance to specificDiabrotica species at both the adult and larval stages in potato species.[8] These resistance-increasing biological compounds can be selected for and subsequently increased through generations of crop plants.[8]
Nematodes may be used as strategy in the management ofDiabrotica larvae populations.[9] Multiple families of nematodes will attackDiabrotica pests in the soil of cultivated fields.Steinernema carpocapsae is most commonly used for larval control in the field.[9] This method of pest control is heavily influenced by the environmental conditions of the soil, as moisture levels and application of the nematodes may drastically affect their effectiveness.[9] Low-tillage and reduced weed control procedures must also be implemented as agronomic strategies when using the nematodes as pest control.[8]
The previous technique primarily targeted the larval stages ofDiabrotica development. In order to target the adult demographic during infestation, an effective method involves exploiting the pestilentDiabrotica's preference for cucurbitacin containing food sources.[8] The chemical composition of cucurbitacin's acts as a strong phagostimulant for adultDiabrotica beetles.[8] Therefore, the beetles are attracted to the compound and inclined to ingest the material containing it.[8] By coating cucurbitacin producing plants with an appropriate insecticide, a toxic trap can be easily created.[8] These attractants can then be dispersed through fields to act as lures, poisoning the beetles and reducing the overall infestation.[8]