Dung Beetles

James Ridsdill-Smith , Leigh W. Simmons , in Encyclopedia of Insects (Second Edition), 2009

Nesting

The beliefs for which the dung beetle is best known is the removal of dung from the pat and compaction in tunnels in the footing as provisioning for their offspring. A single egg is laid in each breed mass. 3 types of provisioning are observed. In teleocoprids (rollers), a sphere is fabricated from pieces of dung at the dung pat or from pellets of dung (Fig. 3A. The beetle rolls the ball away from the dung pat, commonly with the hind legs, and buries it in the soil, before laying a single egg. In paracoprids (tunnelers), the beetle digs a tunnel in the soil under a dung pat, cuts off pieces of dung using its front legs, head, and trunk, and carries them down the tunnel where they are packed into the finish to form a compacted brood mass, earlier laying a single egg. Soil is then placed over the brood mass and another brood mass is made. Branching tunnels may be fabricated containing many brood masses with eggs (Fig. 3B). The size and shape of the breed mass, and the depth of the brood mass in the soil, are characteristic for each species, but are besides influenced by soil moisture and soil hardness. Endocoprids (dwellers) construct brood balls in cavities inside the dung pat.

Figure 3. Dung protrude reproduction in cattle dung. (A) Dung pat with a beetle removing a ball of dung and burying it. (B) Dung pat with a protrude producing brood masses in tunnels beneath the pat. Brood mass containing: (C) egg, (D) larva, (Due east) pupa, and (F) immature adult.

 (Illustration by Tom Prentis from Waterhouse, 1974; reproduced with permission.)

Current phylogenetic analyses of the Scarabaeinae suggest that tunneling may accept been the bequeathed breeding system of all dung beetles, with multiple origins of ball rolling occurring throughout their evolutionary history. Intense contest for dung may have provided the evolutionary pressures generating switches in behavior. At that place appears to have been simply a unmarried evolutionary origin of dwelling from a tunneling antecedent, in the lineage leading to Oniticellus and Tragiscus.

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The Global Reject of Dung Beetles

Jean-Pierre Lumaret , ... Imelda Martínez-1000 , in Reference Module in Earth Systems and Ecology Sciences, 2020

Are dung beetles threatened?

Dung beetles represent a modest group of insects, with roughly 5000 described species, and their highest diversity is in tropical forests and savannahs ( Hanski and Cambefort, 1991). At that place are numerous threats to dung beetles (Nichols et al., 2008; Sánchez-Bayo and Wyckhuys, 2019). The strongest empirical evidence of dung protrude reject can be found in the Mediterranean surface area, where the range of most brawl rolling dung beetles in the Iberian Peninsula contracted in the second one-half of the 20th century (Lobo, 2001). In Italy, there was an overall, significant reduction of 31.4% in the number of records of roller species during the 20th century (Carpaneto et al., 2007). A similar reject in the abundance of dung beetles has been reported in French republic and Nifty United kingdom, and also in Finland where the reject of many dung beetle species has been documented since the 1960s; several species take virtually disappeared since that time, and are probably extinct (Biström et al., 1991). In tropical regions the situation is as well alarming, with an accelerated decline of numerous forest species due to the destruction of their habitat and the gross simplification of ecosystems, allowing a few generalist species living in open environments to supplant them.

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Copris nevinsoni Dung Beetle

Abdalbasit Adam Mariod , ... Ismail Hussein , in Unconventional Oilseeds and Oil Sources, 2017

Description

Dung beetles vest to family Scarabaeidae, order Coleoptera. Their sizes were varied co-ordinate to the species from approximately i.5 to four.v cm in length. Adult dung beetles vary significantly in size from small beetles no more than 1/eighth of an inch in length ( Aphodius pseudolividus) to large beetles measuring one¼ in. in length (Dichotomius carolinus). Most dung beetles are brown to black in color. Occasionally, a brilliant metallic green beetle appears and can be an hands identified as Phanaeus vindex. Many of the male dung beetles have distinct horns, for case, Onthophagus taurus horns resemble balderdash horns, while Onthophagus gazella has a brusque spike-like horns. Horn size is generally a product of larval nutrition. Major males accept large horns while minor males take short horns (Bertone et al., n.d.) (Fig. 43.1).

Figure 43.1. Copris nevinsoni Dung beetle.

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Awarding to Sustainability of Ecosystem Services

Timothy D. Schowalter , in Insect Ecology (Quaternary Edition), 2016

Dung beetles and dung management in Australia

Detritivores tend to exist minor and concealed in soil and litter and, therefore, often overlooked. All the same, they play a critical office in the decomposition of plant litter, carrion, and dung and, thereby, contribute to the sustainability of numerous ecosystem services. Their importance to the sustainability of pasture and livestock production, and to biological command of nuisance flies, is demonstrated by Australian efforts to manage dung aggregating and biting fly populations following introduction of domestic livestock.

Dung beetles are instrumental in burial vertebrate dung within a few days and contributing to its decomposition. Individual species are relatively specific with regard to substrate conditions and host range of colonized dung (A. Davis, 1996). When cattle were introduced to Australia, their dung accumulated in pastures, without specialized dung beetles to consume it. Although Australia is home to 437 species of native dung beetles (Doube and Marshall, 2014), these species prefer the relatively dry, pelleted carrion of marsupials and were effective in shredding and burying cattle dung for merely a few weeks in spring and autumn (Tyndale-Biscoe, 1994). Ferrar (1975) reported that experimentally marked cattle dung survived at to the lowest degree 3 months and sometimes more than than a twelvemonth. Dung accumulation smothered pasture vegetation and increased reproductive habitat for two hematophagous flies, the buffalo fly, Haematobia irritans exigua, and the bush fly, Musca vetustissima, that became serious pests of cattle and humans ( Ferrar, 1975; Tyndale-Biscoe and Vogt, 1996 Ferrar, 1975 Tyndale-Biscoe and Vogt, 1996 ).

Kickoff in 1967, a number of African dung beetles were evaluated and introduced into Australia to accelerate dung disintegration and nutrient turnover and to manage wing populations (R. Hughes et al., 1978). Initial introductions resulted in substantially increased dung disintegration and burying, from < 7% calendar week−i at sites with simply 1 exotic species to 30% at sites with five exotic species, but fluctuated from 0% to 70% depending on beetle affluence (Tyndale-Biscoe, 1994). Suppression of fly reproduction occurred primarily through dung disturbance (R. Hughes et al., 1978), but the start exotic species were most agile during the warm monsoon season and relatively ineffective earlier when bush flies first appear ( A. Davis, 1996; Tyndale-Biscoe and Vogt, 1996 Davis, 1996 Tyndale-Biscoe and Vogt, 1996 ). Subsequent inquiry identified additional dung protrude species that could be active earlier (Ridsdill-Smith and Kirk, 1985) and demonstrated the importance of phoretic mites, for case, native Macrocheles glaber and exotic G. peregrinus, that prey on fly larvae in dung pads (J. Roth et al., 1988). Although nosotros do not know all the consequences of these introductions, mean abundances of native dung beetles have remained similar to their preintroduction abundances (Tyndale-Biscoe and Vogt, 1996). This experience demonstrates that dung beetles are capable of dramatically reducing accumulation of cattle dung and abundances of bitter flies thereby contributing to more sustainable commitment of important ecosystem services.

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The physiology of insect families: A door to the written report of social development

Stephen T. Trumbo , in Advances in Insect Physiology, 2019

five Natural and bogus selection of care

Dung beetle, O. taurus, mothers in Western Australia produce heavier brood balls than those in Eastern Due north America, and that difference has of import implications for the fettle of the young that develop (Beckers et al., 2015; Macagno et al., 2016). Casasa and Moczek (2018a) have related these populational variations to pre-existing variation in the European ancestral population and subsequent canalization in the descendant populations. In addition to maternal investment, information technology is evident that the threshold size at which a larva will subsequently develop exaggerated horns, itself, is nether selection (Moczek and Nijhout, 2003; Moczek et al., 2002).

The brusque generation times of insects, which can exist reared in small spaces, make them ideal candidates for experimental evolution (artificial selection). For example, Harano et al. (2010) examined intralocus sexual conflict in broad-horned flour beetles (Gnatocerus cornutus) past selecting laboratory populations for large or small male mandibles. When populations were selected for males with large mandibles, females had lower fettle than controls; when populations were selected for males with pocket-sized mandibles, females had higher fitness. The negative correlation between an exaggerated male trait and female fitness is a articulate marking of intralocus sexual conflict.

It is hoped that experimental evolution tin can uncover dynamics in family development and examination models of behavioural conflict and cooperation between interacting social partners. Unlike selection for the abiotic environment, option for adaptations to biotic environments (predator-prey, host-parasite, the family unit) are complicated past co-evolving agents. This selective interaction produces interesting evolutionary dynamics that tin either enhance or impede trait response to option (Jarrett et al., 2017; Royle et al., 2016; Smiseth and Royle, 2018). For example, if the event of the social environment on a particular trait is big and positive, the response of that trait to option is facilitated (Moore et al., 1997). Insect families can exist model systems for trait evolution in response to biotic environments because of the feasibility of experimental evolution and manipulable social interactions.

Experimental evolution to written report familial interactions has been used almost extensively in the burial protrude, N. vespilloides. Although post-hatching parental care is facultative in this species, larvae practise amend with care, which includes regurgitations from parents (Eggert et al., 1998). Using a laboratory population, Schrader et al. (2015) produced a No Care line in which immature never received post-hatching care. After four generations, larvae from the No Care and Full Care lines were both allowed to develop on a carcass without parents. Larvae from the No Care line survived significantly better (Schrader et al., 2015). Contempo work demonstrates that function of the No Intendance reward in a parent-less environment was that these larvae had evolved larger mandibles that more easily penetrated the skin of a carcass. The presence of a parent that pre-macerates food, therefore, permits selection for less investment in mandibles by young larvae, demonstrating that cooperative social interactions tin permit i partner to invest less when the other invests more than (Jarrett et al., 2018a). Interspecific comparisons besides point to the mandibles as a focus of selection. Nicrophorus species that can develop independently of care accept highly serrated mandibles at the earliest instar, which allow easier tearing and penetration of the carcass skin in the absence of parental feedings (Benowitz et al., 2018). Larvae from No Care selected lines also hatch more synchronously than Full Care larvae. Jarrett et al. (2018b) speculate that synchronous hatching allows siblings to benefit from each other'south attempts to penetrate the carcass, a mutualism that is vital in the absence of parental regurgitations (Magneville et al., 2018). Experimental development using this species also demonstrates that parental intendance alters the potential for trait evolution. When laboratory populations were selected for larger body size, the response to choice was clear and positive in populations that received Total Care but was negligible in populations under No Care. Choice for smaller body size, even so, was much more than rapid in the No Care line (Jarrett et al., 2017). Given the physiological detail that is emerging on the regulation of care in biparental burying beetles, this group is condign a model for the bear upon of parental care on the evolution of family dynamics.

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Origins of Evolutionary Novelty

Nelson R. Cabej , in Epigenetic Principles of Development (2d Edition), 2019

Evolution of Horns in Beetles

In that location are thousands of protrude species, in which a proportion of male individuals develop horns (Moczek and Nagy, 2005 ) every bit cuticle extensions. Some representatives of the dung beetles of the genus Onthophagus, consisting of more than 2000 known species, have been object of extensive studies on the development of horns in male beetles. Males utilise horns, which represent more than ten% of the body mass, as weapons in gainsay with other males to get admission to resources accumulated past females in tunnels in the soil below dung (Emlen, 2000). Horned males block the tunnel entrance, thus preventing other males from having access to the buried female person and the dung deposited at the lesser of the tunnel.

O. taurus, O. nigriventris, and other species of the Onthophagus genus, display two distinct morphs: individuals that develop horns later on attaining a threshold in body mass and hornless individuals. Horn Anlagen develop early on during the prepupal stage. Large male pupae develop a long pronotal horn (developing on the dorsal exoskeletal plate of the first thoracal segment), whereas pocket-sized male pupae only form a pronotal outgrowth. Such an outgrowth likewise develops in female pupae, only it disappears afterward. Whether male beetles develop horns or not depends on reaching a threshold body size; only males that attain that threshold get horned beetles (Fig. 10.12).

Fig. 10.12

Fig. 10.12. Endocrine regulation of male and sexual dimorphism in the beetle O. taurus. Past the middle of the third larval instar, large and small males differ in circulating levels of juvenile hormone (JH): large males have lower concentrations than smaller males. JH levels are assessed during a brief sensitive period immediately before the cessation of feeding (vertical greyness bar), and relatively large males have JH concentrations beneath the disquisitional threshold (black horizontal line) at this time. (A) Cells in the developing horns of these individuals undergo a brief pulse of rapid proliferation during the prepupal catamenia, and these larvae mature into adult males with fully adult horns (inset). (B) Minor male person larvae have JH concentrations above the threshold during the sensitive period, and these animals experience a brief pulse of a second hormone, ecdysone (arrow in B). Ecdysone is known to initiate cascades of gene expression, and this tactic-specific pulse appears to bear on the fate of horn cells such that they subsequently undergo only minimal proliferation. Small males mature into adults with simply rudimentary horns (inset).

 From Emlen, D.J., Hunt, J., Simmons, L.W., 2005. Evolution of sexual dimorphism in the expression of protrude horns: phylogenetic evidence for modularity, evolutionary lability, and constraint. Am. Nat. 166 Suppl, S42–S66.

Expression of the gene Dll (distalless) in all horn Anlagen, like in all arthropod appendages, has been interpreted as indication that the proximo-distal development of horns and arthropod limb development may depend on the same ontogenic machinery (Moczek and Nagy, 2005). A slight increase in the levels of ecdysteroids and a drop in JH (juvenile hormone) titers occur in females and hornless males. This is consistent with the role of ecdysone every bit a downregulator of the Dll. In contrast, large pupae, which develop horns, prove an increase in the hemolymph level of JH at the end of the third larval phase. When methoprene, a JH analog, is topically applied, 80% of the hornless males also develop horns (Emlen and Nijhout, 1999). During a brusk period of about 30 hours loftier levels of JH induce proliferation of the epidermal cells and germination of horns in males of the dung beetles, Onthophagus taurus (Emlen, 2000).

From an evolutionary view, it is important to bespeak out that horn development in dung beetles is characterized by expression of transcription factors Dll (distalless) and al (aristaless) qualitatively similarly to the example of the evolution of insect appendages. Not only horn patterning genes just genes for the relevant hormones (ecdysteroids and JH) are unchanged in dung beetle species that evolved singled-out morphologies and no differences in genes exist between the horned males and hornless females and modest males. Thus the horn in the dung protrude O. taurus is epigenetically adamant past shifts in the trunk size threshold:

One major avenue of evolutionary modify in this group has involved shifts in the threshold body size regulating horn growth.

Emlen (2000)

Hence, an understanding of the mechanism of the evolution of horn size and morphology requires knowledge of

ane.

How these insects might change the body size threshold, and

2.

How can they alter expression of JH receptors in epidermal cells.

Nijhout believes that "suppression of horns in small males required the evolution of a size-sensing mechanism in the final larval instar" (Nijhout, 2003) and it is believed that the stretch stimulus, after the third larval stage is received by proprioceptive neurons in the soft regions of the cuticle and transmitted for processing in the insect brain (Gorbman and Davey, 1991) where the ready point for body size is believed to be in insects. The location of the set points in the insect brain is also suggested by the fact that all the morphological and physiological changes that follow the achieve of the growth threshold commencement with point cascades from the brain. Others trunk mass set points are identified in the brain of Peromyscus maniculatus (Adams et al., 2001) and other mammals (Baeckberg et al., 2003), the thermoregulation (Hammel et al., 1963; Boulant, 2000, etc.) and a number of components of body fluids (see Section Fundamental Control of Animal Physiology in Chapter one).

A comparative written report on the North American and Western Australian populations of dung beetle, O. taurus, has shown that these populations have undergone a very rapid development of the horn polyphenism. Individuals of this circum-Mediterranean species were introduced to the above regions by late 1960s of the 20th century. Within equally piddling as ~   xl years these races have diverged dramatically in the body size threshold for the development of horns in nature, while maintaining the original threshold nether laboratory weather for many generations. W Australian populations now switch to the horned morph at a much larger body size, and North American populations—at a much smaller body, than their Mediterranean ancestors (Moczek and Nijhout, 2003). Differences betwixt the Western Australian and North American races are of the magnitude of differences between species. These "race" differences are of a magnitude comparable to differences of the original form with some other dung beetle species, Onthophagus illyricus. This suggests that North American and West Australian populations take entered separate paths of evolving into two new species in the genus Onthophagus (Moczek and Nijhout, 2003) by only changing the torso size threshold via a size-sensing machinery, or "self-perception of torso size" (Ben-Nun et al., 2013), which, in all likelihood, is a neural machinery.

Experimental development of horns in dung beetles by application of JH at sensitive stages of the larval development suggests that the torso size threshold may be correlated with the increased secretion of JH (Moczek and Nijhout, 2003).

Emlen et al. (2005) envisage the bones mechanism of horn development in beetles also equally the evolutionary of import switching (horned/hornless) every bit a hierarchical mechanism consisting of a

one.

sensory apparatus,

ii.

secretion of a hormone (JH),

3.

temporally restricted expression of the receptor for that hormone

4.

the downstream expression of secondary hormones and transcription factors.

The fact that during the development at least offset two steps (one and two) and the 4th step (iv) reside/are determined in insect's encephalon suggests that the evolutionary switching from horned to hornless beetles and the reverse involved a corresponding change in the neural circuits that decide the body size threshold and regulate JH secretion. Indeed, in that location is no other manner an evolutionary change that involves no changes in genes to occur just via appropriate changes in development.

The neural decision of the "size-sensing mechanisms" could explain the exceptional evolutionary lability of horns in beetles: horn sexual dimorphism in beetles has been gained vii times, lost 13 times and regained once, whereas male horn dimorphism has been gained eight times and lost twelve times (Emlen et al., 2005).

Neo-Darwinian Explanation

Neo-Darwinian theory would predict that in order for horns, equally a new graphic symbol, to evolve in dung beetles, one or a number of "useful" mutations or new genes would have been necessary. In that location is neither evidence nor a hint that such changes take occurred. On the opposite, all the genes and their protein products involved in horn development are functionally well conserved. Hence, the basic neo-Darwinian prediction on the development of horns in dung beetles is patently refuted.

Epigenetic Caption

From the epigenetic view it would be predicted:

1.

No changes in genes relevant to development of horns are necessary for evolution of head horns in dung beetles.

2.

Evolution of horns in dung beetles is result of epigenetically determined changes in the patterns of expression of genes that are essentially involved (ecdysone, JH, Dll, etc.) in the development of horns.

three.

Evolution of horn sexual dimorphism and horn intrasexual polyphenism in dung beetles of the genus Onthophagus involved an epigenetic change the body size threshold.

four.

Epigenetic information necessary for inducing point cascades determining these specific changes in expression patterns of genes involved in horn development originates in the beetles' CNS.

The first prediction is validated by the experimental evidence that the products of genes (ecdysone, JH, Dll) involved in the evolution of horns are conserved and have not changed their function.

The second prediction also is validated by the experimental evidence presented in this department. That evidence shows a clear stardom in the patterns of expression of genes (ecdysteroids and juvenile hormone and Dll) involved in the procedure of proliferation of epidermal cells during formation of horns in large size beetles compared to pocket-sized size male beetles, despite the fact that all of them share a common genetic groundwork.

The third prediction is likewise substantiated by the evidence already presented in this section. The regulatory, that is, epigenetic changes are the only changes that have scientifically been demonstrated to systematically occur in the process of horn evolution in dung beetles. The most relevant difference in determining the horned/hornless culling is the body size, as perceived in the insect encephalon. This body size threshold of the aforementioned circum-Mediterranean species O. taurus, in response to different environmental atmospheric condition, in Northward America and Australia, was respectively lowered and elevated, leading thus to formation of two incipient species (Moczek and Nijhout, 2003), within an evolutionary instant of less than 40 years.

The fourth prediction is that the signal cascade for horn development originates in the CNS. The indicate cascades for production of ecdysteroids and juvenile hormone, which are the just known signal cascades that activate genes involved in the development of dung beetle horns, start in the brain (plainly after the processing in neural circuits of the stretch input coming from proprioceptors).

All the evidence accumulated and then far shows that not any changes in genes but

The brain ultimately controls the alternative developmental pathways that atomic number 82 to the polyphenism.

Nijhout, H.F., McKenna, Grand.Z., 2018. The distinct roles of insulin signaling in polyphenic development. Curr. Opin. Insect Sci. 25, 58–64.

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Thermoregulation

Bernd Heinrich , in Encyclopedia of Insects (Second Edition), 2009

Against Competitors

Competition competition or fighting over food is rare in insects, but at least ii species of African dung beetles, Scarabaeus laevistriatus and Kheper nigroaeneus, engage in combat over dung balls that they make to feed on and/or to serve every bit sexual attractants. An elevated thoracic temperature plays a crucial role in these contests on the footing. The more than a beetle shivers to go along warm (with its flight muscles), the higher the temperature of the leg muscles adjacent to the flight muscles in the thorax and the faster its legs can move and construct the dung into assurance and roll it away. Endothermy thus aids in the scramble competition for food, and information technology reduces the duration of exposure to predators. Additionally, hot beetles take the edge in competition competitions over dung balls made by other beetles; in fights over dung assurance, hot beetles almost invariably defeat libation ones, oftentimes despite a big size disadvantage.

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Microbiome of wood tree insects

Juliana A. Ugwu , ... Fred O. Asiegbu , in Forest Microbiology, 2021

iii.four Decomposition: Organic waste product

Organic waste decomposition is a vital ecosystem process that is principally done by microorganisms and some insect species. Many dung beetle species take been reported to play an of import role in the decomposition of manure. Dung beetles live in many habitats, including planted forests. Many dung beetles feed on decomposable fruits, leaves, and mushrooms while besides eating the dung of omnivores and herbivores. Furthermore, the larvae of beetles, termites, ants, and flies make clean upwardly dead plant matter past breaking it down through feeding by gut microbes. Animal tissues provide nutrient to diverse groups of insect detritivores such as flies and beetles ( Merritt and De Jong, 2015). Many of these insects contribute to soil health by improving the essential nutrients, micronutrients or total protein content in the soil (Macfadyen et al., 2015).

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Origins of Evolutionary Novelty

Nelson R. Cabej , in Epigenetic Principles of Development, 2012

Neo-Darwinian Explanation

Neo-Darwinian theory would predict that in order for horns, as a new character, to evolve in dung beetles, one or an undefined number of "useful" mutations need to occur in DNA or genes, whose products play central roles in the development of horns, such as the patterning genes Distalless and aristaless or the genes for ecdysteroids and JH. There is neither evidence nor a hint that such changes accept occurred. On the opposite, all the above genes and their poly peptide products are functionally well conserved. Hence, the basic neo-Darwinian prediction on the development of horns in dung beetles is patently refuted.

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Biodiversity and Pest Control Services

Azucena Lucatero , Stacy Thou. Philpott , in Reference Module in Life Sciences, 2021

Noncrop Habitats

Hedgerows, live fences, and other linear habitats within agronomical systems provide habitat for birds, bats, dung beetles, and butterflies ( Harvey et al., 2005) and specifically for several groups of invertebrate natural enemies (Bianchi et al., 2006). As agricultural habitats are constantly disturbed, hedgerows and other crop margins provide stable resource bases for natural enemies (Bianchi et al., 2006). The characteristics of noncrop habitats that benefit natural enemies include providing alternate prey, nectar and pollen, nesting sites, and host plants necessary for reproduction and life-cycle completion (Landis et al., 2000; Bianchi et al., 2006), and are like to benefits provided by vegetation diversity within crop fields. Hedgerows and field margins increase predator movement beyond agricultural landscapes similarly to how a loftier-quality matrix may increase movement of organisms between forest fragments (Vandermeer and Carvajal, 2001; Tscharntke et al., 2005). In some cases, increases in predator diversity in hedgerows can increment pest command. Linear vegetation strips in vineyards in California facilitate motility of natural enemies in the grapes, and thereby increase pest command (Benton et al., 2003). Field margins and hedgerows can tiresome motility of fungal pathogens and can serve as barriers to the movement of pests, thereby improving pest control (Altieri, 1999). Still, if field margins provide culling habitat for beneficial insects and other predators such that they forage more in the margins than in the crop, or prefer the crop-margin habitats more than the natural habitats, then this may harm pest control (Benton et al., 2003; Bianchi et al., 2006).

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