Editing Winter moth

{{Short description|Species of moth}}
{{self-reference|In North America, "winter moth" usually denotes the invasive species ''Operophtera brumata'', but may also refer to a native species, ''[[Erannis tiliaria]]'' (linden looper) or ''[[Operophtera bruceata]]'' (bruce spanworm).}}
{{Speciesbox
| image = Operophtera brumata01.jpg
| taxon = Operophtera brumata
| authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])
| synonyms = ''Cheimatobia brumata'' L.<ref name=cheimatobia>{{cite web |url=https://www.cabi.org/isc/datasheet/37695 |publisher=CABI Invasive species compendium |title=Operophtera brumata (winter moth)|date=2021 |doi=10.1079/cabicompendium.37695 |access-date=2019-07-23}}</ref><br />''Chimatobia brumata''<ref name=chimatobia>{{cite web |url=http://ubio.org/NZ/detail.php?uid=39237&d=1 |publisher=Nomenclator Zoologicus |title=Chimatobia|access-date=2019-07-23}}</ref><br />''Phalaena brumata'' L.<ref name=cheimatobia />
}}

The '''winter moth''' ('''''Operophtera brumata''''') is a [[moth]] of the family [[Geometridae]]. It is an abundant species in [[Europe]] and the [[Near East]] and a famous study organism for evaluating insect population dynamics.<ref>{{cite book |last1=Futuyma |first1=D.J. |title=Annual Review of Ecology, Evolution, and Systematics |date=2013}}</ref> It is one of very few [[lepidoptera]]ns of temperate regions in which adults are active in late autumn and early winter.  The females of this species are virtually wingless and cannot fly, but the males are fully winged and fly strongly. After the initial frosts of late fall, the females emerge from their pupae, walk to and up trees and emit [[pheromone]]s in the evening to attract males. After fertilization, they ascend to lay, on average, around 100 eggs each. Typically, the larger the female moth is, the more eggs she lays.

The winter moth is considered an [[invasive species]] in North America. [[Nova Scotia]], Canada, experienced the first confirmed infestations in the 1930s. It was later accidentally introduced to Oregon in the 1950s and the Vancouver area of British Columbia around 1970. Defoliation by the moth was first noted in eastern states of the United States in the late 1990s, and is now well established in [[Massachusetts]], [[Rhode Island]], [[New Hampshire]] and [[Maine]]. The winter moth is confirmed present in [[British Columbia]], [[Washington (state)|Washington]] and [[Oregon]].<ref>{{Cite news|last = Cairn |first = North |title = Experts: Destructive winter moths are spreading  | newspaper = Portland Press Herald  | location = Portland, Maine  | date = 8 December 2012  | url = http://www.pressherald.com/news/experts-destructive-winter-moths-are-spreading_2012-12-09.html  | access-date = 9 December 2012}}</ref> In Massachusetts, the moths have attracted the attention of several media outlets due to the severity of the infestation.<ref>{{Cite news   | last = Noonan  | first = Erica  | title = Mystery Pests  | newspaper = Boston Globe  | location = Boston, Massachusetts | date = 3 December 2009  | url = http://www.boston.com/news/local/articles/2009/12/03/invasion_of_winter_moths_has_scientists_residents_looking_for_answers/  | access-date = 9 December 2012}}</ref> Efforts at [[biological control]] are underway.<ref>{{cite web |last1=Elkinton |first1=Joseph |last2=Boettner |first2=George |last3=Liebhold |first3=Andrew |last4=Gwiazdowski |first4=Rodger |title=Biology, Spread, and Biological Control of Winter Moth in the Eastern United States |url=https://www.fs.fed.us/foresthealth/technology/pdfs/FHTET-2014-07_Biology_Control_Winter-Moth.pdf |publisher=USDA Forest Service |access-date=29 December 2018}}</ref><ref name=UMass>[https://ag.umass.edu/landscape/fact-sheets/winter-moth-in-massachusetts-history-biological-control Winter Moth in Massachusetts: History and Biological Control] University of Massachusetts Extension, The Center for Agriculture, Food and the Environment (2017). Accessed Dec 6, 2017.</ref>

==Description==
[[Image:Operophtera_brumata_L._copulation.jpg|thumb|225px|Mating. Males prefer larger females (more eggs).]]
The forewing ground colour of the winged males varies from grey-yellow to beige-brown or occasionally slightly reddish-tinted. The patterns are often band-shaped dark brownish, often indistinct. The fringe is yellowish. The hindwings are pale grey or yellow grey. The antennae are short and finely hairy. The female appears completely wingless, but in fact female winter moths have greatly reduced wings.<ref name=":1">{{Cite web|title=StackPath|url=https://www.northeastnursery.com/blogs/strategies-for-identifying-and-controlling-winter-moth|access-date=2021-12-07|website=www.northeastnursery.com}}</ref> The flightless female has a brownish-grey body with rudimentary wing stubs that are brown to grey and have dark bands. Body length for both sexes is approximately {{convert|1|cm||abbr=on|sigfig=1}}.<ref name="Umass2">[http://ag.umass.edu/landscape/fact-sheets/winter-moth-identification-management Winter Moth Identification & Management] UMass Extension The Center for Agriculture, Food and the Environment (2017) Accessed December 7, 2017.</ref> Larvae are {{convert|0.1|in|mm}} in length after hatching and will grow to {{convert|0.75|in|mm}} over a six-week period.<ref name="UnivMaine">[http://digitalcommons.library.umaine.edu/etd/2338/ The Relationship Between the Winter Moth (''Operophtera brumata'') and Its Host Plants] O'Donnell KM. University of Maine (2015)</ref> In North America, the winter moth can be confused with the related native species, the [[Operophtera bruceata|Bruce spanworm]] (''Operophtera bruceata''). In fact, the two species [[Hybrid (biology)|hybridize]].<ref name="Havill et al 2017">{{cite journal |last1=Havill |first1=N.P. |author-link1=Nathan Havill |last2=Elkinton |first2=J. |last3=Andersen |first3=J.C. |last4=Hagen |first4=S.B. |last5=Broadley |first5=H.J. |last6=Boettner |first6=G.J. |last7=Caccone |first7=A. |title=Asymmetric hybridization between non-native winter moth, ''Operophtera brumata'' (Lepidoptera: Geometridae), and native Bruce spanworm (''Operophtera bruceata''), in the Northeastern United States, assessed with novel microsatellites and SNPs. |journal=Bulletin of Entomological Research |date=2017 |volume=107 |issue=2 |pages=241–250 |title-link=Operophtera bruceata |doi=10.1017/S0007485316000857 |pmid=27876095 }}</ref>

==Distribution==
It is native to Northern and Central Europe; in the south, its range extends to Northern [[Italy]]; in the east to the [[Caucasus]] and [[Asia Minor]]; in the north to the [[Baltic region|Baltics]]. The genetic populations of the winter moth in Europe are a result of recolonization following the last glacial period.<ref name="Andersen et al. 2017">{{cite journal |last1=Andersen |first1=Jeremy |last2=Havill |first2=Nathan |last3=Caccone |first3=Adalgisa |last4=Elkinton |first4=Joseph |title=Postglacial recolonization shaped the genetic diversity of the winter moth (''Operophtera brumata'') in Europe |journal=Ecology and Evolution |date=2017 |volume=7 |issue=10 |pages=3312–3323 |doi=10.1002/ece3.2860 |pmid=28515868 |pmc=5433974 |bibcode=2017EcoEv...7.3312A }}</ref> As an invasive species, this moth is found in Nova Scotia, coastal New England (Massachusetts, Maine, Rhode Island, Connecticut, and New York) and also the Pacific northwest (including Vancouver Island).<ref>{{Cite web |title=Winter moth |url=https://bcinvasives.ca/invasives/winter-moth/ |access-date=2025-03-21 |website=Invasive Species Council of British Columbia |language=en-US}}</ref> In [[New England]], expansion inland and north appears to be curtailed by cold winter temperatures, so for example, coastal Maine but not inland. Locally milder winters, as part of global climate change, may be allowing expansion of afflicted territory.<ref name=UMass/><ref name=Netherlands/> A study conducted in Massachusetts documented that winter moth defoliation reduced the annual trunk diameter growth rate of oak trees by an average of 47% while not significantly impacting growth rates of the less defoliated maple trees.<ref name=Simmons>{{cite journal |vauthors=Simmons MJ, Lee TD, Ducey MJ, Elkinton JS, Boettner GH, Dodds KJ |title=Effects of Invasive Winter Moth Defoliation on Tree Radial Growth in Eastern Massachusetts, USA |journal=Insects |volume=5 |issue=2 |pages=301–18 |year=2014 |pmid=26462685 |pmc=4592589 |doi=10.3390/insects5020301 |doi-access=free }}</ref>

==Life cycle==
[[File:Operophtera brumata (caterpillar) focus stacking-20230508-RM-131624.jpg|thumb|right|225px|Caterpillar will reach length of about 0.75 inches]]
Winter moth larvae ([[caterpillar]]s) emerge in early spring from egg masses laid near leaf buds after a series of days in which the daytime high temperatures reach around 10&nbsp;°C (50&nbsp;°F).<ref name=Umass2/> Recently hatched larvae feed on expanding leaf buds, often after having burrowed inside the bud, and later on foliage, for approximately six weeks. In addition to feeding on the tree where they hatched, young larvae will also produce silk strands to 'balloon' (be wind-blown) to other trees.<ref name="UnivMaine" /> By mid-May the larvae, green in color and about an inch long, descend to the ground. Very little mortality due to disease has been noted in winter moth larvae in North America.<ref name="Broadley et al. 2017">{{cite journal|last1=Broadley|first1=H.J.|last2=Boucher|first2=M.|last3=Burand|first3=J.P.|last4=Elkinton|first4=J.S.|date=2017|title=The phylogenetic relationship and cross-infection of nucleopolyhedroviruses between the invasive winter moth (''Operophtera brumata'') and its native congener, Bruce spanworm (''O. bruceata'')|journal=Journal of Invertebrate Pathology|volume=142|pages=61–68|doi=10.1016/j.jip.2016.11.016|pmid=27916614|doi-access=free|bibcode=2017JInvP.143...61B }}</ref><ref name="Donahue et al. 2018">{{cite journal|last1=Donahue|first1=K.L.|last2=Broadley|first2=H.J.|last3=Elkinton|first3=J.S.|last4=Burand|first4=J.P.|last5=Huang|first5=W.F.|last6=Andersen|first6=J.C.|date=2018|title=Using the SSU, ITS, and Ribosomal DNA Operon Arrangement to Characterize Two Microsporidia Infecting Bruce spanworm, ''Operophtera bruceata'' (Lepidoptera: Geometridae)|journal=Journal of Eukaryotic Microbiology|volume=66|issue=3|pages=424–434|doi=10.1111/jeu.12685|pmid=30136341|doi-access=free}}</ref>

Pupation occurs in the soil in late May.<ref name="Simmons" /> Adults emerge from the soil in mid-late November. The female winter moths are flightless but release a sex pheromone to attract males.<ref name=":0">{{Cite web|date=2015-03-06|title=Winter Moth Identification & Management|url=https://ag.umass.edu/landscape/fact-sheets/winter-moth-identification-management|access-date=2021-12-06|website=Center for Agriculture, Food, and the Environment|language=en}}</ref> After mating, the female lays between 150 and 350 tiny eggs in bark crevices, on branches, in lichen, and under bark scales.<ref name=Simmons/><ref name=":0" /> With such a long pupal period, the winter moth is vulnerable to numerous pupal predators and parasitoids.<ref>{{cite journal|last1=Roland|first1=J.|last2=Embree|first2=D.G.|year=1995|title=Biological control of the winter moth.|journal=Annual Review of Entomology|volume=40|pages=475–492|doi=10.1146/annurev.en.40.010195.002355}}</ref><ref>{{cite thesis|last1=Broadley|first1=Hannah|date=January 2018|title=Impact of native natural enemies on populations of the invasive winter moth, (''Operophtera brumata'' L) in the northeast United States|url=https://scholarworks.umass.edu/dissertations_2/1327/|journal=Doctoral Dissertations| doi=10.7275/12760419 |access-date=29 December 2018}}</ref>

Research conducted in the Netherlands indicated that as climate warming is causing spring temperatures to become warmer sooner, some of the winter moth eggs were hatching before tree leaf buds – first food for the caterpillars – had begun to open. Early hatchlings starved. Late hatchlings survived. Because hatch timing is genetically controlled, the moths are evolving to resynchronize with bud opening by delaying the response to the temperature trigger by 5 to 10 days.<ref name="Netherlands">[https://www.sciencedaily.com/releases/2007/06/070619174647.htm Winter Moth Proves Able To Adjust To Climate Change.] Netherlands Institute of Ecology (2007).</ref> The larvae, like the adults, can withstand below freezing temperatures at night. Larval dispersal is the dominant source of density-dependent larval mortality and likely regulates high density population dynamics of winter moth in New England.<ref name="Pepi et al 2016">{{cite journal |last1=Pepi |first1=Adam |last2=Broadley |first2=Hannah |last3=Elkinton |first3=Joseph |title=Density‑dependent effects of larval dispersal mediated by host plant quality on populations of an invasive insect |journal=Oecologia |date=2016 |volume=182 |issue=2 |pages=499–509 |doi=10.1007/s00442-016-3689-z|pmid=27378097 |bibcode=2016Oecol.182..499P |s2cid=24424176 }}</ref> Larvae prefer [[oak]] and [[apple]], but also feed on [[maple]], [[birch]], [[hornbeam]], [[chestnut]], [[hazel]], [[quince]], [[beech]], [[larch]], [[Populus|poplar]], [[cherry]], [[pear]], [[rose]], [[raspberry]], [[blueberry]], [[willow]], [[elm]], and other leafy trees and shrubs.<ref name="Umass2" /><ref name="UnivMaine" />

== Invasive impact ==
The winter moth is native to Europe, but invaded the northeast of the [[United States|US]] in the late 1990s. As of 2007, winter moths have spread to coastal Maine, [[Greater Boston|eastern Massachusetts]], [[Rhode Island]], [[southeastern Connecticut]], and [[East End (Long Island)|eastern Long Island]]. The moths feed on a wide range of [[deciduous]] leaves, and some [[conifer]]s. This has caused defoliation of the trees across the states in which the moths live. Between 2003 and 2015 winter moths caused defoliation of forests and shade trees from between 2,266 and 36,360 hectares per year in Massachusetts alone.<ref>Elkinton, J. S., Boettner, G. H., and Broadley, H. J.. 2021. Successful biological control of winter moth, ''Operophtera brumata'', in the northeastern United States. ''Ecological Applications'' 31( 5):e02326. 10.1002/eap.2326</ref>

==Biological control==
[[File:WinterMothLeafDamage.jpg|thumb|225px|Maple tree leaves showing damage by winter moth larvae]]
In Europe, where winter moths are native, two [[parasitism|parasitic]] species, a [[parasitoid wasp|wasp]] (''[[Agrypon flaveolatum]]'') and a fly (''[[Cyzenis albicans]]'') prey on winter moth caterpillars. The wasps insert eggs into the larvae. The flies lay eggs on leaves, which the larvae then ingest when eating the leaves.<ref name=Capinera>{{cite book|author=Capinera, John L.|title=Encyclopedia of Entomology|url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA4278 |year=2008 |publisher=Springer Science & Business Media|isbn=978-1-4020-6242-1 |page=4278}}</ref> As a [[biological pest control|biological control]], the wasp was introduced in Canada but is not being pursued in the United States because there is not sufficient evidence that the wasp would not lay eggs in larvae of other moth species.<ref name=UMass/> Introduction of ''C. albicans'', which is species-specific to preying on winter moths, has proven successful in strongly reducing, although not eliminating, winter moth infestation in Nova Scotia. Test introductions were begun in eastern Massachusetts in 2005. This proved successful, so more fly releases have taken place at sites from southeast [[Connecticut]] to coastal [[Maine]]. As moth densities can reach ten million larvae per acre, several years are needed before parasitism by the flies reaches 20–50%. Once this is achieved the winter moth density will decline and leaf defoliation will be reduced to levels not as harmful to the trees.<ref name=UMass/> The delay in establishment is likely the result of heavy mortality from predators and [[Hyperparasite|hyperparasitoids]] on the pupating ''C.&nbsp;albicans''.<ref name="Broadley et al. 2018">{{cite journal |last1=Broadley |first1=H.J. |last2=Kelly |first2=E.A. |last3=Elkinton |first3=J.S. |last4=Kula |first4=R.R. |last5=Boettner |first5=G.G. |title=Identification and impact of hyperparasitoids and predators affecting ''Cyzenis albicans'' (Tachinidae), a recently introduced biological control agent of winter moth (''Operophtera brumata'' L.) in the northeastern U.S.A. |journal=Biological Control |date=2018 |volume=121 |pages=99–108 |doi=10.1016/j.biocontrol.2018.01.011 |s2cid=90918769 |doi-access=free |bibcode=2018BiolC.121...99B }}</ref> Management of the winter moth in Massachusetts using biological control with ''C.&nbsp;albicans'' is showing positive results.<ref name="Elkinton et al. 2018 FHTET">{{cite web |last1=Elkinton |first1=Joseph |last2=Boettner |first2=George |last3=Broadley |first3=Hannah |last4=Reardon |first4=Richard |last5=Weeks |first5=Ronald |title=Biological Control of Winter Moth in Northeast North America |url=https://www.fs.fed.us/foresthealth/technology/pdfs/FHAAST-2018-03_Biology_Control_Winter-Moth.pdf |publisher=USDA Forest Service |access-date=29 December 2018}}</ref>

There are different ways to successfully control the winter moth population depending on the time of year. During the late fall and early winter, a dormant oil stray can be used to suffocate the eggs. When the eggs are newly hatched in late March and early April Bt (''[[Bacillus thuringiensis]]'') is a safe and effective way to control caterpillars. As the caterpillars get bigger, [[spinosad]] is also effective.<ref name=":1" />

==References==
{{Reflist}}

== Further reading ==
* Chinery, Michael, ''Collins Guide to the Insects of Britain and Western Europe'', 1986 (Reprinted 1991)
* Skinner, Bernard, ''Colour Identification Guide to Moths of the British Isles'', 1984

==External links==
{{Commons category|Operophtera brumata}}
* [https://www.ukmoths.org.uk/species/operophtera-brumata/ UKmoths] Winter Moth ''Operophtera brumata'' (Linnaeus, 1758)
* {{in lang|de}} [http://www.lepiforum.de/lepiwiki.pl?Operophtera_Brumata Lepiforum.de] ''many excellent photos''

{{Taxonbar|from=Q164002}}
{{Authority control}}

[[Category:Operophtera]]
[[Category:Agricultural pest insects]]
[[Category:Insect pests of temperate forests]]
[[Category:Moths described in 1758]]
[[Category:Moths of Europe]]
[[Category:Moths of Iceland]]
[[Category:Moths of North America]]
[[Category:Moths of Asia]]
[[Category:Taxa named by Carl Linnaeus]]