Large copper

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Large copper
Lycaena dispar02.jpg
Male
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Lycaenidae
Genus: Lycaena
Species:
L. dispar
Binomial name
Lycaena dispar
(Haworth, 1802)
Synonyms
  • Papilio disparHaworth, 1802

Large copper (Lycaena dispar) is a butterfly of the family Lycaenidae. Lycaena dispar is widely distributed across temperate Eurasia from western Europe to the Amur region, Korea and Manchuria. [2] While it is currently in severe decline in north-western Europe, it is expanding in central, northern and eastern regions. [3] [4]

Contents

Three European subspecies are commonly recognized: L. dispar dispar (single-brooded), the now-extinct English subspecies; L. d. batavus (usually single-brooded), the Dutch subspecies; and L. d. rutilus (double-brooded), which is widespread across the rest of the continent. [2] Several Asian subspecies extend the range eastward. The species has been declining in many European countries mainly as a result of habitat loss. [1]

Description

Lycaena dispar is a large lycaenid butterfly with a wingspan of 44–52 mm. [5] The species exhibits strong sexual dimorphism. Males have bright orange wings with narrow black margins and white fringes. The short androconia (sex brands) are located near the leading edge of the forewing. Females have broader dark brown margins on the forewings with a parallel row of dark spots.

Both sexes have distinctive silvery-blue hindwing undersides patterned with black spots and an orange submarginal band. This silvery underside distinguishes L. dispar from related species such as Lycaena virgaureae and L. hippothoe , which inhabit drier environments.

The subspecies can be differentiated by size, extent of black markings, and the length of the orange band on the hindwing underside. The extinct English subspecies (L. d. dispar) was notably variable in size and had larger spots on the underside than continental European forms.

Subspecies

Genetic diversity and phylogeography

European populations show distinct genetic structuring based on mitochondrial DNA analysis. A 2004 study identified ten distinct haplotypes across the continent, with populations from northern and central regions sharing close genetic relationships. This pattern indicates these populations likely descended from ancestors that recolonized northward from south-eastern European refugia following the last glacial period. Geographic analysis reveals that range expansion events have been the primary driver shaping current genetic distribution patterns. [6]

Distribution and populations

Current range

Lycaena dispar occurs between latitudes 40° and 60° across Eurasia. [2] The species is currently found in: Armenia, Austria, Azerbaijan, Belarus, Belgium, Bulgaria, China, Croatia, Czech Republic, Estonia, Finland, France, Georgia, Germany, Greece, Hungary, Italy, Kazakhstan, Republic of Korea, Latvia, Lithuania, Luxembourg, Moldova, Mongolia, Montenegro, Netherlands, Poland, Russian Federation, Serbia, Slovakia, Slovenia, Switzerland, Türkiye, Ukraine and Uzbekistan. [1] [7]

The species shows contrasting population trends across its range: severe decline in north-western Europe, stability or expansion in central and northern Europe, and mixed status in Asia. [3]

Regional variations

Western Europe

The species is regionally extinct in the United Kingdom (since 1864) and Ireland (failed reintroductions). [1] In the Netherlands, populations of L. d. batavus persist and are largely monophagous on R. hydrolapathum. [2] Western European populations have suffered from extensive drainage of wetland habitats. In Luxembourg, monitoring from 2007 to 2015 documented the species at 183 sites across 33 municipalities in the south-west and west of the country. Populations primarily utilize R. obtusifolius and R. crispus as host plants and correlate with grassland distribution and habitat mosaics. [8]

Northern Europe

Estonia represents the species' northern range expansion. Absent until the 20th century, L. dispar was first recorded near Tartu in 1947. [2] The species has since expanded north-westward and achieved widespread status, though it remains absent from Estonia's western islands. Estonian populations are univoltine (single-brooded) with flight period from late June to late July, and utilize primarily R. crispus and R. obtusifolius as host plants. [2]

Central Europe

In central Europe, L. dispar has adapted to a broader range of habitats including drier areas, fallows and urban wetlands. [2] Populations in Germany and Austria are characteristically oligophagous on various Rumex species. [2] Vienna's urban populations demonstrate successful colonization of anthropogenic habitats: a 2012 study documented 2,457 eggs at 23 sites within the municipality. [3] In Croatia's Zagorje region, populations remain locally numerous but are declining due to invasive plants. [9]

Southern Europe

The species' range extends south to Mount Olympus in Greece, where it was observed in July 2011. [10] Southern populations may produce a partial third generation in favourable conditions. [2]

Eastern Asia

Field monitoring in South Korea demonstrated significant range expansion southward from the species' previously known north-western strongholds. The 2024 study documented populations at 15 sites spanning six provinces. Genetic diversity patterns, combined with field observations, demonstrate that South Korean populations maintain strong connectivity and are not experiencing the isolation previously assumed. These findings led researchers to recommend downgrading the species from its Near Threatened status in South Korea. [4]

Ecology

Habitat requirements

Lycaena dispar is primarily a wetland species, though habitat preferences vary geographically. [11] The species prefers undisturbed grasslands along riverbanks and stream banks, particularly areas with its larval food plants. [3] To avoid flooding mortality, L. dispar often utilizes plants growing away from water edges and among reed-fen vegetation. [11]

The butterfly shows preference for warmer microclimates which allow faster larval development. [3] Agricultural disturbance, particularly mowing shortly after egg-laying, causes catastrophic population losses by destroying eggs and depriving larvae of food plants. [3]

Host plants

The large copper utilizes various dock species ( Rumex ) as larval host plants. Host plant range varies geographically, with broader usage in eastern populations and narrower specialization in western ones. [3] Female butterflies select host plants based on multiple factors: plant height, size, phenological stage, absence of flowering/fruiting stems and low acidity levels. [3]

Primary host plants vary by region:

Additional confirmed host plants include R. aquaticus , R. conglomeratus , R. patientia , R. sanguineus and R. stenophyllus . [2] [3] The latter two species represent novel host plant records documented in urban environments. [3] Sorrel species containing oxalic acid (e.g. Rumex acetosa ) are rarely utilized. [2]

Larvae create a characteristic feeding pattern on host plants, eating only the underside of leaves to produce a 'window' effect where the upper leaf surface remains intact. [13]

Life cycle

Most European populations are bivoltine with two generations annually: May–June and late July–early September, with peak flight in July. [2] Northern populations (e.g. Estonia) are univoltine, while southern populations may produce a partial third generation. [2]

The species overwinters as half-grown third-instar larvae, which enter diapause triggered by temperatures below 15°C and photoperiod cues. [14] Larvae resume feeding in early May when temperatures exceed 25°C. Heavy mortality occurs during the overwintering period, particularly from prolonged flooding. [11]

Reproduction and development

Females produce approximately 32 eggs per batch and can lay around 714 eggs in their lifetime. [15] Eggs are laid on sunny host plants in optimal microclimates. Herbivore damage and fungal infections on host plants do not significantly affect oviposition choices. [14]

In late June, mature larvae migrate up to 25 cm from their host plant and approximately 10 cm above ground to pupate. During this migration, larvae change color from bright green to pale yellow-brown for camouflage. Pupation lasts 10–14 days. [15] [14]

Natural enemies

Mortality factors include invertebrate predators (particularly during pre-diapause), parasitoids and vertebrate predators including reed-nesting birds, amphibians and small mammals. [11]

The ichneumonid wasp Hyposoter placidus has been recorded as a specialized parasitoid of L. dispar. First documented in Poland in 2021, adult wasps were reared from caterpillars collected in north-eastern Poland feeding on Rumex confertus, R. hydrolapathum and R. obtusifolius. [16] This parasitoid may be a factor limiting abundance of L. dispar populations. [16]

Other parasitoids include Phryxe vulgaris (Diptera: Tachinidae), which affects post-diapause larvae and results in death of late instar larvae. [11]

Female large copper Lycaena dispar female Obersulm 20070718 5.jpg
Female large copper
Egg of L. dispar Lycaena dispar egg 2.jpg
Egg of L. dispar

History in Britain and Ireland

Illustration from John Curtis's British Entomology Volume 5 Britishentomologyvolume5Plate12.jpg
Illustration from John Curtis's British Entomology Volume 5

Extinction in England

The British subspecies, L. d. dispar, was first recorded from the Huntingdonshire fens in 1749 and formally described in 1759. [17] The butterfly was once common in several English fenland localities including Whittlesea Mere, using food-plant Rumex hydrolapathum .

Drainage of the fens for agriculture led to rapid population decline:

The value of specimens increased dramatically post-extinction:

Reintroduction attempts

Multiple reintroduction attempts have been made:

Failures are attributed to captive-bred stock being poorly adapted to wild conditions. [11] The Dutch subspecies L. d. batavus, described in 1915 and nearly identical to the extinct English form, has been used in reintroduction attempts without lasting success. [17]

Conservation

Lycaena dispar is classified as Near Threatened by the IUCN, [1] though regional status varies considerably. The species is protected under the Bern Convention on the Conservation of European Wildlife and Natural Habitats and EU Habitats Directive (Annexes II and IV). [2]

Primary threats include:

Conservation recommendations include:

The contrasting population trends across Europe – decline in the west, expansion in the north and east – suggest the species may be responding to climate change as well as habitat availability. [2]

References

  1. 1 2 3 4 5 Gimenez Dixon, M. (1996). "Lycaena dispar". IUCN Red List of Threatened Species . 1996: e.T12433A3347854. doi: 10.2305/IUCN.UK.1996.RLTS.T12433A3347854.en . Retrieved 19 November 2021.
  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Lindman, Ly; Remm, Jaanus; Saksing, Kristiina; Sõber, Virve; Õunap, Erki; Tammaru, Toomas; Leather, Simon R.; DeVries, Phil (2015). "Lycaena dispar on its northern distribution limit: an expansive generalist". Insect Conservation and Diversity. 8: 3–16. doi:10.1111/icad.12087. S2CID   84861272.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 Strausz, Martin; Fiedler, Konrad; Franzén, Markus; Wiemers, Martin (2012). "Habitat and host plant use of the Large Copper Butterfly Lycaena dispar in an urban environment". Journal of Insect Conservation. 16 (5): 709–721. doi:10.1007/s10841-012-9456-5.
  4. 1 2 Park, Jeong Sun; Lee, Sung Hwan; Pyo, Ji-Young; Jeong, Heon Cheon; Kim, Sung-Soo; Kim, Iksoo (2024). "Field monitoring and genetic diversity of the large copper butterfly Lycaena dispar (Lepidoptera: Lycaenidae)". Journal of Biological Research - Bollettino della Società Italiana di Biologia Sperimentale. 97 (1). doi:10.4081/jbr.2024.12411.
  5. ↑ Emmet, A.M., J. Heath et al. (Ed.) (1990) The Butterflies of Great Britain and Ireland. The Moths and Butterflies of Great Britain and Ireland Vol. 7 Part 1 (Hesperiidae to Nymphalidae). Harley Books, Colchester, UK. 370p.
  6. ↑ Lai, Bo-Chi G.; Pullin, Andrew S. (2004). "Phylogeography, genetic diversity and conservation of the large copper butterfly Lycaena dispar in Europe". Journal of Insect Conservation. 8 (1): 27–36. doi:10.1023/B:JICO.0000027478.35309.46.
  7. ↑ https://www.biomonitoring.sk/Registration/AtlasAnimal/detail/70305
  8. ↑ Proess, Raoul; Rennwald, Erwin; Schneider, Simone (2016). "Zur Verbreitung und Ökologie des Großen Feuerfalters (Lycaena dispar Haworth, 1803) im Südwesten und Westen Luxemburgs". Bulletin de la Société des naturalistes luxembourgeois. 118: 89–110.
  9. ↑ Koren, Toni; Lauš, Boris; Šašić, Martina; Mihoci, Iva; Štih, Ana; Bralić, Petra; Gomboc, Stanislav (2017-12-31). "Contribution to the Knowledge of the Butterfly Fauna (Lepidoptera: Papilionoidea) of Hrvatsko Zagorje, Croatia". Natura Croatica: Periodicum Musei Historiae Naturalis Croatici. 26 (2): 167–196. doi:10.20302/NC.2017.26.15. eISSN   1848-7386.
  10. ↑ Wagner, Wolfgang. "Lycaena dispar ecology and distribution". Pyrgus.de. Retrieved 26 August 2025.
  11. 1 2 3 4 5 6 7 Webb, Mark R.; Pullin, Andrew S. (1996). "Larval survival in populations of the large copper butterfly Lycaena dispar batavus". Ecography. 19 (3): 279–286. doi:10.1111/j.1600-0587.1996.tb01255.x. JSTOR   3682914.
  12. ↑ Proess, Raoul; Rennwald, Erwin; Schneider, Simone (2016). "Zur Verbreitung und Ökologie des Großen Feuerfalters (Lycaena dispar Haworth, 1803) im Südwesten und Westen Luxemburgs". Bulletin de la Société des naturalistes luxembourgeois. 118: 89–110.
  13. ↑ "Large Copper". Butterfly Conservation. Retrieved 26 August 2025.
  14. 1 2 3 4 Kim, Seong-Hyun; Kim, Nam Jung; Hong, Seong-Jin; Lee, Young-Bo; Park, Hae-Chul; Je, Yeon-Ho; Lee, Kwang Pum (2014). "Environmental induction of larval diapause and life-history consequences of post-diapause development in the Large Copper butterfly, Lycaena dispar (Lepidoptera: Lycaenidae)". Journal of Insect Conservation. 18 (4): 693–700. Bibcode:2014JICon..18..693K. doi:10.1007/s10841-014-9678-9. S2CID   17887375.
  15. 1 2 Bink, F. A. (1986). "Acid stress in Rumex hydrolapathum (Polygonaceae) and its influence on the phytophage Lycaena dispar (Lepidoptera; Lycaenidae)". Oecologia. 70 (3): 447–451. Bibcode:1986Oecol..70..447B. doi:10.1007/bf00379510. JSTOR   4218073. PMID   28311934. S2CID   694523.
  16. 1 2 Kostro-Ambroziak, Agata; Sielezniew, Marcin (2021). "First record of Hyposoter placidus (Desvignes, 1856) (Hymenoptera: Ichneumonidae) in Poland: the specialized parasitoid of Lycaena Fabricius, 1807 butterflies (Lepidoptera: Lycaenidae)". Polish Journal of Entomology. 90 (3): 153–159. doi:10.2478/pjen-2019-0007.
  17. 1 2 3 4 Duffey, Eric (1968). "Ecological studies on the large copper butterfly Lycaena dispar Haw. batavus Obth. at Woodwalton Fen National Nature Reserve, Huntingdonshire". Journal of Applied Ecology. 5 (1): 69–96. Bibcode:1968JApEc...5...69D. doi:10.2307/2401275. JSTOR   2401275.
  18. ↑ Hugh D. Loxdale; Belinda J. Davis; Robert A. Davis (February 2016). "Known knowns and unknowns in biology". Biological Journal of the Linnean Society . 117 (2): 386–398. doi: 10.1111/bij.12646 .
  19. 1 2 3 4 5 Eeles, Peter (2023). British & Irish Butterfly Rarities: Migrants, extinctions & introductions. Newbury: Pisces Publications. p. 235. ISBN   978-1-913994-10-5.
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