Editing Beetle (section)

==Ecology==
<!--[[File:Pegesimallus sp robberfly.jpg|thumb|Beetles such as this [[Chrysomelidae|chrysomelid]] (lower left) are preyed upon by insects such as this [[robber fly]].]]-->
[[File:Long-horn Beetle - Ecyrus dasycerus, Leesylvania State Park, Woodbridge, Virginia.jpg|thumb|upright|A [[camouflage]]d [[longhorn beetle]], ''[[Ecyrus dasycerus]]'']]

===Anti-predator adaptations===
Beetles, both adults and larvae, are the prey of many animal [[predator]]s including [[mammals]] from [[bat]]s to [[rodent]]s, [[birds]], [[lizard]]s, [[amphibians]], [[fishes]], [[dragonflies]], [[robberflies]], [[Reduviidae|reduviid bugs]], [[ant]]s, other beetles, and [[spider]]s.<ref name="Evans27">[[#refEvans|Evans & Bellamy (2000)]], pp. 27–28</ref><ref>{{cite book |author=Cott, H. B. |author-link=Hugh B. Cott |date=1940 |title=Adaptive Coloration in Animals |title-link=Adaptive Coloration in Animals |publisher=Methuen |page=[https://archive.org/details/adaptivecolorati00cott/page/414 414]}}</ref> Beetles use a variety of [[anti-predator adaptation]]s to defend themselves. These include [[camouflage]] and [[mimicry]] against predators that hunt by sight, toxicity, and defensive behaviour.

====Camouflage====
{{Further|Camouflage}}

Camouflage is common and widespread among beetle families, especially those that feed on wood or vegetation, such as [[leaf beetle]]s (Chrysomelidae, which are often green) and [[weevil]]s. In some species, sculpturing or various colored scales or hairs cause beetles such as the [[avocado]] weevil ''[[Heilipus apiatus]]'' to resemble bird dung or other inedible objects.<ref name="Evans27"/> Many beetles that live in sandy environments blend in with the coloration of that substrate.<ref name="Evans126">[[#refEvans|Evans & Bellamy (2000)]], p. 126</ref>

====Mimicry and aposematism====
[[File:Clyte bélier - MHNT - Vue dorsale.jpg|thumb|left|''[[Clytus arietis]]'' ([[Cerambycidae]]), a [[Batesian mimic]] of wasps]]
{{further|Mimicry|Aposematism}}

Some [[longhorn beetle]]s (Cerambycidae) are effective [[Batesian mimics]] of [[wasp]]s. Beetles may combine coloration with behavioural mimicry, acting like the wasps they already closely resemble. Many other beetles, including [[ladybirds]], [[blister beetle]]s, and [[Lycidae|lycid beetles]] secrete distasteful or toxic substances to make them unpalatable or poisonous, and are often [[aposematic]], where bright or contrasting [[animal coloration|coloration]] warn off predators; many beetles and other insects mimic these chemically protected species.<ref name="Evans"/>

[[File:Blister beetle (26390828032).jpg|thumb|Blister beetles such as ''[[Hycleus]]'' have brilliant [[aposematic]] coloration, warning of their toxicity.]]
[[File:Bloody -nosed Beetle^ Timarcha tenebricosa - Flickr - gailhampshire.jpg|thumb|The bloody-nosed beetle, ''[[Timarcha tenebricosa]]'', [[antipredator adaptation|defending itself]] by releasing a droplet of noxious red liquid (base of leg, on right)|left|172x172px]]

Chemical defense is important in some species, usually being advertised by bright aposematic colors. Some [[Tenebrionidae]] use their posture for releasing noxious chemicals to warn off predators. Chemical defenses may serve purposes other than just protection from vertebrates, such as protection from a wide range of microbes. Some species sequester chemicals from the plants they feed on, incorporating them into their own defenses.<ref name="Evans126"/>

Other species have special glands to produce deterrent chemicals. The defensive glands of carabid ground beetles produce a variety of [[hydrocarbon]]s, [[aldehyde]]s, [[phenol]]s, [[quinone]]s, [[ester]]s, and [[acid]]s released from an opening at the end of the abdomen. African carabid beetles (for example, ''[[Anthia]]'') employ the same chemicals as ants: [[formic acid]].<ref name="Evans">[[#refEvans|Evans & Bellamy (2000)]]</ref> [[Bombardier beetle]]s have well-developed pygidial glands that empty from the sides of the intersegment membranes between the seventh and eighth abdominal segments. The gland is made of two containing chambers, one for [[hydroquinone]]s and [[hydrogen peroxide]], the other holding hydrogen peroxide and [[catalase]] enzymes. These chemicals mix and result in an explosive ejection, reaching a temperature of around {{convert|100|C|F}}, with the breakdown of hydroquinone to hydrogen, oxygen, and quinone. The oxygen propels the noxious chemical spray as a jet that can be aimed accurately at predators.<ref name=insenc/>

====Other defenses====
Large ground-dwelling beetles such as [[Carabidae]], the [[rhinoceros beetle]] and the longhorn beetles defend themselves using strong [[Mandible (insect)|mandibles]], or heavily sclerotised (armored) spines or horns to deter or fight off predators.<ref name="Evans126"/> Many species of weevil that feed out in the open on leaves of plants react to attack by employing a drop-off reflex. Some combine it with [[thanatosis]], in which they close up their appendages and "play dead".<ref name="insenc199">[[#refMcHugh|McHugh (2009)]], p. 199</ref> The click beetles ([[Elateridae]]) can suddenly catapult themselves out of danger by releasing the energy stored by a click mechanism, which consists of a stout spine on the prosternum and a matching groove in the mesosternum.<ref name="Evans27"/> Some species startle an attacker by producing sounds through a process known as [[stridulation]].<ref name="stridulation"/>

===Parasitism===
A few species of beetles are [[Parasitism#Basic concepts|ectoparasitic]] on mammals. One such species, ''[[Platypsyllus castoris]]'', parasitises beavers ([[Beaver|''Castor'']] spp.). This beetle lives as a parasite both as a larva and as an adult, feeding on epidermal tissue and possibly on skin secretions and wound exudates. They are strikingly flattened dorsoventrally, no doubt as an adaptation for slipping between the beavers' hairs. They are wingless and eyeless, as are many other ectoparasites.<ref>{{cite journal |author=Peck, Stewart B. |year=2006 |title=Distribution and biology of the ectoparasitic beaver beetle ''Platypsyllus castoris'' Ritsema in North America (Coleoptera: Leiodidae: Platypsyllinae) |journal=Insecta Mundi |volume=20 |url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1107&context=insectamundi |issue=1–2 |pages=85–94}}</ref> Others are kleptoparasites of other invertebrates, such as the [[small hive beetle]] (''Aethina tumida'') that infests [[honey bee]] nests,<ref>{{cite journal |doi=10.1051/apido:2004010 |author1=Neumann, P. |author2=Elzen, P. J. |name-list-style=amp |year=2004 |title=The biology of the small hive beetle (''Aethina tumida'', Coleoptera: Nitidulidae): Gaps in our knowledge of an invasive species |journal=Apidologie |volume=35 |issue=3 |pages=229–247 |doi-access=free }}</ref> while many species are parasitic [[inquiline]]s or [[myrmecophily in Staphylinidae|commensal in the nests of ants]].<ref>{{cite web |url=http://www.cals.ncsu.edu/course/ent425/compendium/coleop~1.html |title=Coleoptera |author=Meyer, John R. |date=March 8, 2005 |publisher=[[North Carolina State University]] |access-date=March 13, 2011 |archive-url=https://wayback.archive-it.org/all/20000524004320/http://www.cals.ncsu.edu/course/ent425/compendium/coleop~1.html |archive-date=May 24, 2000  }}</ref> A few groups of beetles are primary [[parasitoid]]s of other insects, feeding off of, and eventually killing their hosts.<ref>{{Cite book|title=Encyclopedia of Entomology|last1=Weber|first1=Donald C.|last2=Saska|first2=Pavel|last3=Chaboo|first3=Caroline S.|publisher=Springer Netherlands|year=2008|isbn=978-1-4020-6242-1|pages=719–721|chapter=Carabid Beetles (Coleoptera: Carabidae) as Parasitoids|doi=10.1007/978-1-4020-6359-6_492}}</ref>

===Pollination===
[[File:Protaetia cuprea - 2019 Zachi Evenor.jpg|thumb|An Israeli Copper Flower-Chafer (''[[Protaetia cuprea]] ignicollis'') on a crown daisy (''[[Glebionis coronaria]]'')]]

Beetle-pollinated flowers are usually large, greenish or off-white in color, and heavily scented. Scents may be spicy, fruity, or similar to decaying organic material. Beetles were most likely the first insects to pollinate flowers.<ref>{{cite journal |last1=Livingston |first1=Stephanie |title=This amber-encased beetle may have been one of the first insects to pollinate flowers |journal=Science |date=November 2019 |doi=10.1126/science.aba1758 |s2cid=213876270 |url=https://www.science.org/content/article/amber-encased-beetle-may-have-been-one-first-insects-pollinate-flowers |access-date=10 January 2021}}</ref> Most beetle-pollinated flowers are flattened or dish-shaped, with pollen easily accessible, although they may include [[Pollination trap|traps]] to keep the beetle longer. The plants' ovaries are usually well protected from the biting mouthparts of their pollinators. The beetle families that habitually pollinate flowers are the [[Buprestidae]], [[Cantharidae]], [[Cerambycidae]], [[Cleridae]], [[Dermestidae]], [[Lycidae]], [[Melyridae]], [[Mordellidae]], [[Nitidulidae]] and [[Scarabaeidae]].<ref name="Gullan">{{cite book |last=Gullan |first=P. J. |author2=Cranston, P. S. |title=The Insects: An Outline of Entomology |publisher=Wiley, John & Sons |date=2014 |edition=5 |isbn=978-1-4443-3036-6 |page=314}}</ref> Beetles may be particularly important in some parts of the world such as semiarid areas of southern Africa and [[southern California]]<ref>{{cite journal |author1=Jones, G. D. |author2=Jones, S. D. |name-list-style=amp |year=2001 |title=The uses of pollen and its implication for entomology |journal=Neotropical Entomology |volume=30 |issue=3 |pages=314–349 |doi=10.1590/S1519-566X2001000300001|doi-access=free }}</ref> and the montane grasslands of [[KwaZulu-Natal]] in South Africa.<ref name="Ollerton_a">{{cite journal |author1=Ollerton, J. |author2=Johnson, S. D. |author3=Cranmer, L. |author4=Kellie, S. |name-list-style=amp |year=2003 |title=The pollination ecology of an assemblage of grassland asclepiads in South Africa |journal=[[Annals of Botany]] |volume=92 |pages=807–834 |doi=10.1093/aob/mcg206 |issue=6 |pmid=14612378 |pmc=4243623}}</ref>

===Mutualism===
[[File:Ambrosia beetle life cycle.svg|upright=1.2|thumb|
1: Adult [[ambrosia beetle]] burrows into wood and lays eggs, carrying fungal spores in its [[mycangia]].<br />
2: Larva feeds on fungus, which digests wood, removing toxins, to mutual benefit.<br />
3: Larva pupates.|left]]

[[Mutualism (biology)|Mutualism]] is well known in a few beetles, such as the [[ambrosia beetle]], which partners with fungi to digest the wood of dead trees. The beetles excavate tunnels in dead trees in which they cultivate fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases spores of its fungal [[symbiont]]. The fungus penetrates the plant's xylem tissue, digests it, and concentrates the nutrients on and near the surface of the beetle gallery, so the weevils and the fungus both benefit. The beetles cannot eat the wood due to toxins, and uses its relationship with fungi to help overcome the defenses of its host tree in order to provide nutrition for their larvae.<ref name="Malloch">{{cite book |last1=Malloch|first1= D. |last2= Blackwell |first2= M. |year=1993|chapter=Dispersal biology of ophiostomatoid fungi|pages= 195–206 |title=Ceratocystis and Ophiostoma: Taxonomy, Ecology and Pathology |editor1= Wingfield, M. J. |editor2= K. A. Seifert |editor3= J. F. Webber |publisher= APS |place=St. Paul |isbn=978-0-89054-156-2}}</ref> Chemically mediated by a bacterially produced polyunsaturated peroxide,<ref name=Scott2008>{{cite journal |last=Scott |first=J. J. |author2=Oh, D. C. |author3=Yuceer, M. C. |author4=Klepzig, K. D. |author5=Clardy, J. |author6= Currie, C. R. |title=Bacterial protection of beetle-fungus mutualism |journal=[[Science (journal)|Science]] |date=2008 |volume=322 |issue=5898 |page=63 |pmid=18832638 |pmc=2761720 |doi = 10.1126/science.1160423|bibcode=2008Sci...322...63S }}</ref> this mutualistic relationship between the beetle and the fungus is [[coevolve]]d.<ref name="Malloch"/><ref>{{cite book|author=Francke-Grossmann, H. |year=1967 |chapter=Ectosymbiosis in wood inhabiting insects|editor=M. Henry|title=Symbiosis|volume=2|publisher=[[Academic Press]]|location=New York|pages=141–205}}</ref>
[[File:Bug in Tharparkar District.jpg|alt=Beetle found in Tharparkar District|thumb|Tenebrionid beetle in the [[Tharparkar|Thar Desert]]]]

===Tolerance of extreme environments===
[[File:Stenocara gracilipes.jpg|thumb|The fogstand beetle of the [[Namib Desert]], ''[[Stenocara gracilipes]]'', is able to survive by [[fog collection|collecting water from fog]] on its back.]]
{{Further|Insect thermoregulation|Insect winter ecology}}

About 90% of beetle species enter a period of adult [[diapause]], a quiet phase with reduced metabolism to tide unfavourable environmental conditions. Adult diapause is the most common form of diapause in Coleoptera. To endure the period without food (often lasting many months) adults prepare by accumulating reserves of lipids, glycogen, proteins and other substances needed for resistance to future hazardous changes of environmental conditions. This diapause is induced by signals heralding the arrival of the unfavourable season; usually the cue is [[photoperiodic]]. Short (decreasing) day length serves as a signal of approaching winter and induces winter diapause (hibernation).<ref>{{cite journal |last1=Hodek |first1=Ivo |title=Review Article: Adult diapause in Coleoptera |journal=Psyche: A Journal of Entomology |volume=2012 |pages=1–10 |date=2012 |doi=10.1155/2012/249081 |doi-access=free }}</ref> A study of hibernation in the Arctic beetle ''[[Pterostichus brevicornis]]'' showed that the body fat levels of adults were highest in autumn with the [[alimentary canal]] filled with food, but empty by the end of January. This loss of body fat was a gradual process, occurring in combination with dehydration.<ref>{{cite journal |title=Hibernation in the Arctic beetle, ''Pterostichus brevicornis'', in Alaska |author1=Kaufmann, T. |date=1971 |journal=Journal of the Kansas Entomological Society |volume=44 |issue=1 |pages=81–92}}</ref>

All insects are [[poikilotherm]]ic,<ref name='Outline'>{{cite book |author1=Gullan, P. J. |author2=Cranston, P. S. |date=1994 |title=The Insects: An Outline of Entomology |publisher=Chapman and Hall |isbn=978-0-412-49360-7 |pages=103–104}}</ref> so the ability of a few beetles to live in extreme environments depends on their resilience to unusually high or low temperatures. The [[bark beetle]] ''[[Pityogenes chalcographus]]'' can survive {{gaps|−39|°C}} whilst overwintering beneath tree bark;<ref>{{cite journal |author1=Lombadero, Maria J. |author2=Ayres, Matthew P. |author3=Ayres, Bruce D. |author4=Reeve, John D. |title=Cold tolerance of four species of bark beetle (Coleoptera: Scolytidae) in North America |journal=Environmental Ecology |volume=29 |issue=3 |date=2000 |pages=421–432 |url=http://www.dartmouth.edu/~mpayres/pubs/Fina.Cold.pdf |archive-url=https://web.archive.org/web/20070417144045/http://www.dartmouth.edu/~mpayres/pubs/Fina.Cold.pdf |archive-date=2007-04-17 |url-status=live}}</ref> the Alaskan beetle ''[[Cucujus]] clavipes puniceus'' is able to withstand {{gaps|−58|°C}}; its larvae may survive {{gaps|−100|°C}}.<ref>{{cite journal |author1=Sformo, T. |author2=Walters, K. |author3=Jeannet, K. |author4=Wowk, B. |author5=Fahy, G. M. |author6=Barnes, B. M. |author7=Duman, J. G. |title=Deep supercooling, vitrification and limited survival to −100°C in the Alaskan beetle ''Cucujus clavipes puniceus'' (Coleoptera: Cucujidae) larvae |journal=[[Journal of Experimental Biology]] |date=2010 |volume=213 |issue=3 |pages=502–509 |doi=10.1242/jeb.035758 |pmid=20086136|doi-access=free }}</ref> At these low temperatures, the formation of ice crystals in internal fluids is the biggest threat to survival to beetles, but this is prevented through the production of antifreeze proteins that stop water molecules from grouping together. The low temperatures experienced by ''Cucujus clavipes'' can be survived through their deliberate dehydration in conjunction with the antifreeze proteins. This concentrates the antifreezes several fold.<ref>{{cite news |url=https://www.bbc.co.uk/nature/21923937 |title=The life of extremophiles: Surviving in hostile habitats |last1=Brooks |first1=Christopher |date=2013-03-26 |publisher=BBC Nature |access-date=2017-03-16}}</ref> The [[hemolymph]] of the mealworm beetle ''[[Tenebrio molitor]]'' contains several [[antifreeze protein]]s.<ref>{{cite journal |author1=Graham, L. A |author2=Liou, Y. C. |author3=Walker, V. K. |author4=Davies, P. L. |title=Hyperactive antifreeze protein from beetles |journal=[[Nature (journal)|Nature]] |volume=388 |issue=6644 |pages=727–728 |date=August 1997 |doi=10.1038/41908 |quote=The yellow mealworm beetle, ''Tenebrio molitor'', contains a family of small Cys-rich and Thr-rich thermal hysteresis proteins that depress the hemolymph freezing point below the melting point by as much as 5.58°C(ΔT=thermal hysteresis). Thermal hysteresis protein expression was evaluated throughout development and after exposure to altered environmental conditions. |pmid=9285581 |bibcode=1997Natur.388..727G|s2cid=205029622 |doi-access=free }}</ref> The Alaskan beetle ''[[Upis ceramboides]]'' can survive −60&nbsp;°C: its [[cryoprotectant]]s are [[xylomannan]], a molecule consisting of a [[sugar]] bound to a [[fatty acid]],<ref>{{cite journal |author1=Walters, K. R. Jr |author2=Serianni, A. S. |author3=Sformo, T. |author4=Barnes, B. M. |author5=Duman, J. G. |title=A nonprotein thermal hysteresis-producing xylomannan antifreeze in the freeze-tolerant Alaskan beetle Upis ceramboides |journal=[[PNAS]] |volume=106 |issue=48| year=2009 |pages= 20210–20215 |doi=10.1073/pnas.0909872106 |pmid=19934038 |pmc=2787118|bibcode=2009PNAS..10620210W |doi-access=free }}</ref> and the sugar-alcohol, [[threitol]].<ref>{{cite journal| title=Cryoprotectant biosynthesis and the selective accumulation of threitol in the freeze-tolerant Alaskan beetle, ''Upis ceramboides'' |author1=Walters, K. R. Jr.|author2=Pan, Q. |author3=Serianni, A. S. |author4=Duman, J. G. | journal=[[Journal of Biological Chemistry]] |year=2009 |volume= 284 |issue=25 | pages=16822–16831 | doi=10.1074/jbc.M109.013870 | pmid=19403530 | pmc=2719318|doi-access=free }}</ref>

Conversely, desert dwelling beetles are adapted to tolerate high temperatures. For example, the [[Tenebrionid]] beetle ''[[Onymacris rugatipennis]]'' can withstand {{gaps|50|°C}}.<ref>{{cite journal |author=Edney, E. B. |date=1971 |title=The body temperature of tenebrionid beetles in the Namib desert of southern Africa |journal=[[Journal of Experimental Biology]] |volume=55 |pages=253–272 |doi=10.1242/jeb.55.1.253 |url=http://jeb.biologists.org/content/jexbio/55/1/253.full.pdf |archive-url=https://web.archive.org/web/20170211080953/http://jeb.biologists.org/content/jexbio/55/1/253.full.pdf |archive-date=2017-02-11 |url-status=live}}</ref> Tiger beetles in hot, sandy areas are often whitish (for example, ''[[Habroscelimorpha dorsalis]]''), to reflect more heat than a darker color would. These beetles also exhibits behavioural adaptions to tolerate the heat: they are able to stand erect on their tarsi to hold their bodies away from the hot ground, seek shade, and turn to face the sun so that only the front parts of their heads are directly exposed.<ref>{{cite journal|doi=10.1093/aesa/83.5.911 |title=Seasonal activity and thermoregulatory behavior of ''Cicindela patruela'' (Coleoptera: Cicindelidae) |journal=Annals of the Entomological Society of America |volume=83 |issue=5 |pages=911–915 |year=1990 |last1=Knisley |first1=C. B. |last2=Schultz |first2=T. D. |last3=Hasewinkel |first3=T. H. }}</ref>

The fogstand beetle of the [[Namib Desert]], ''[[Stenocara gracilipes]]'', is able to [[fog collection|collect water from fog]], as its elytra have a textured surface combining [[hydrophilic]] (water-loving) bumps and waxy, [[hydrophobic]] troughs. The beetle faces the early morning breeze, holding up its abdomen; droplets condense on the elytra and run along ridges towards their mouthparts. Similar adaptations are found in several other Namib desert beetles such as ''[[Onymacris unguicularis]]''.<ref>{{Cite journal| doi = 10.1038/35102108| volume = 414| issue = 6859| pages = 33–34| last1 = Parker| first1 = Andrew R.| last2 = Lawrence| first2 = Chris R.| title = Water capture by a desert beetle| journal = Nature| date = 2001-11-01| pmid=11689930| bibcode = 2001Natur.414...33P| s2cid = 34785113}}</ref>

Some terrestrial beetles that exploit shoreline and floodplain habitats have physiological adaptations for surviving floods. In the event of flooding, adult beetles may be mobile enough to move away from flooding, but larvae and pupa often cannot. Adults of ''[[Cicindela togata]]'' are unable to survive immersion in water, but larvae are able to survive a prolonged period, up to 6 days, of [[Hypoxia (environmental)|anoxia]] during floods. Anoxia tolerance in the larvae may have been sustained by switching to anaerobic metabolic pathways or by reducing metabolic rate.<ref>{{cite journal |author1=Hoback, W. Wyatt |author2=Stanley, David W. |author3=Higley, Leon G. |author4=Barnhart, M. Christopher |title=Survival of immersion and anoxia by larval tiger beetles, ''Cicindela togata'' |journal=The American Midland Naturalist |volume=140 |issue=1 |pages=27–33 |doi=10.1674/0003-0031(1998)140[0027:SOIAAB]2.0.CO;2|year=1998 |s2cid=86163282 }}</ref> Anoxia tolerance in the adult carabid beetle ''[[Pelophilia borealis]]'' was tested in laboratory conditions and it was found that they could survive a continuous period of up to 127 days in an atmosphere of 99.9% nitrogen at 0&nbsp;°C.<ref>{{cite journal |title=Anaerobiosis in the overwintering beetle ''Pelophila borealis'' |author1=Conradi-Larsen, Else-Margrete |author2=Sømme, Lauritz |date=1973 |journal=Nature |volume=245 |issue=5425 |pages=388–390 |doi=10.1038/245388a0|bibcode=1973Natur.245..388C |s2cid=4288059 }}</ref>

===Migration===
{{Main|Insect migration}}

Many beetle species undertake annual mass movements which are termed as migrations. These include the pollen beetle ''[[Meligethes aeneus]]''<ref>{{cite web |url=http://www.fwi.co.uk/arable/half-of-the-uk-sees-start-of-pollen-beetle-migration.htm |title=Half of the UK sees start of pollen beetle migration |author1=Allison, R. |date=2015-03-17 |access-date=2017-03-16}}</ref> and many species of [[Coccinellidae|coccinellids]].<ref>{{cite journal |doi=10.1111/j.1469-185X.1962.tb01609.x |title=Migration of Terrestrial Arthropods in Relation to Habitat |journal=Biological Reviews |volume=37 |issue=2 |pages=171–211 |year=1962 |last1=Southwood |first1=T. R. E.|s2cid=84711127 }}</ref> These mass movements may also be opportunistic, in search of food, rather than seasonal. A 2008 study of an unusually large outbreak of Mountain Pine Beetle (''[[Dendroctonus ponderosae]]'') in [[British Columbia]] found that beetles were capable of flying 30–110&nbsp;km per day in densities of up to 18,600 beetles per hectare.<ref>{{cite book |title=Migration: The Biology of Life on the Move |author1=Dingle, H. |date=2014 |publisher=Oxford University Press}}</ref>