Geranium Bronze
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The geranium bronze or ''brun des pélargoniums'' in French (''Cacyreus marshalli''), is a
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
in the family
Lycaenidae Lycaenidae is the second-largest family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfl ...
. The geranium bronze butterfly is native to
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
. The butterfly was first introduced to
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
in the late 20th century, where it has quickly spread to many southern and eastern European regions. Since its introduction to Europe, the geranium bronze butterfly has become a
pest Pest or The Pest may refer to: Science and medicine * Pest (organism), an animal or plant deemed to be detrimental to humans or human concerns ** Weed, a plant considered undesirable * Infectious disease, an illness resulting from an infection ** ...
to cultivated ''
Pelargonium ''Pelargonium'' () is a genus of flowering plants that includes about 280 species of perennials, succulents, and shrubs, commonly called geraniums, pelargoniums, or storksbills. '' Geranium'' is also the botanical name and common name of a separ ...
'' and ''
Geranium ''Geranium'' is a genus of 422 species of annual, biennial, and perennial plants that are commonly known as geraniums or cranesbills. They are found throughout the temperate regions of the world and the mountains of the tropics, but mostly in ...
'' plant species. Currently, efforts are being made to contain the spread of the geranium bronze butterflies as well as to determine the most effective
pesticide Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampri ...
for the species.


Description

The adult geranium bronze butterfly's
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan of ...
ranges from 15–23 mm in males and 18–23 mm in females. The wings are brown/bronze with a white border outlining the wings. The underside is a grey-brown with darker bands interlaced with white, creating an intricate pattern. The hindwings contain an eye spot in order to divert predators from attacking. The male and female are similar in appearance. File:Lycaenidae - Cacyreus marshalli-1.JPG, Imago, underside File:Lycaenidae - Cacyreus marshalli.JPG, Imago, underside File:Cacyreus marshalli 01 by-dpc.jpg, Imago,upperside


Geographic range

The geranium bronze was first recorded in regions of
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
, including
KwaZulu-Natal KwaZulu-Natal (, also referred to as KZN and known as "the garden province") is a province of South Africa that was created in 1994 when the Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu) and Natal Province were merged. It is locate ...
and the Cape Provinces, as well as in neighbouring countries such as
Eswatini Eswatini ( ; ss, eSwatini ), officially the Kingdom of Eswatini and formerly named Swaziland ( ; officially renamed in 2018), is a landlocked country in Southern Africa. It is bordered by Mozambique to its northeast and South Africa to its no ...
,
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label=Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kalahar ...
,
Zimbabwe Zimbabwe (), officially the Republic of Zimbabwe, is a landlocked country located in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Botswana to the south-west, Zambia to the north, and Mozam ...
,
Lesotho Lesotho ( ), officially the Kingdom of Lesotho, is a country landlocked country, landlocked as an Enclave and exclave, enclave in South Africa. It is situated in the Maloti Mountains and contains the Thabana Ntlenyana, highest mountains in Sou ...
, and
Mozambique Mozambique (), officially the Republic of Mozambique ( pt, Moçambique or , ; ny, Mozambiki; sw, Msumbiji; ts, Muzambhiki), is a country located in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi ...
. It was accidentally brought to Europe in 1978. After being recorded in
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
, a colony of geranium bronze was soon found in
Mallorca Mallorca, or Majorca, is the largest island in the Balearic Islands, which are part of Spain and located in the Mediterranean. The capital of the island, Palma, is also the capital of the autonomous community of the Balearic Islands. The Bal ...
in 1990, and has since spread to most of the regions in southern Europe. In 1996, the geranium bronze butterfly was first recorded in
Rome , established_title = Founded , established_date = 753 BC , founder = King Romulus (legendary) , image_map = Map of comune of Rome (metropolitan city of Capital Rome, region Lazio, Italy).svg , map_caption ...
,
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
, and rapidly spread along the
Adriatic The Adriatic Sea () is a body of water separating the Italian Peninsula from the Balkans, Balkan Peninsula. The Adriatic is the northernmost arm of the Mediterranean Sea, extending from the Strait of Otranto (where it connects to the Ionian Sea) ...
and Tyrrhenian coastal areas before extending inland. The geranium bronze butterfly now has established colonies in
Belgium Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to th ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
, the
Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
, Italy,
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
,
Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders with ...
,
Malta Malta ( , , ), officially the Republic of Malta ( mt, Repubblika ta' Malta ), is an island country in the Mediterranean Sea. It consists of an archipelago, between Italy and Libya, and is often considered a part of Southern Europe. It lies ...
,
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
,
Portugal Portugal, officially the Portuguese Republic ( pt, República Portuguesa, links=yes ), is a country whose mainland is located on the Iberian Peninsula of Southwestern Europe, and whose territory also includes the Atlantic archipelagos of ...
, southern
Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
,
Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, an ...
,
Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...
, and the French
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
.


Food resources


Caterpillars

After hatching, the geranium bronze larvae feed on the flower buds, young leaves, and soft stems of their host plant, typically from the genera ''
Pelargonium ''Pelargonium'' () is a genus of flowering plants that includes about 280 species of perennials, succulents, and shrubs, commonly called geraniums, pelargoniums, or storksbills. '' Geranium'' is also the botanical name and common name of a separ ...
'' and ''
Geranium ''Geranium'' is a genus of 422 species of annual, biennial, and perennial plants that are commonly known as geraniums or cranesbills. They are found throughout the temperate regions of the world and the mountains of the tropics, but mostly in ...
''. In order to eat the stem of these plants, the larvae drill into the stems, causing damage to the plants.


Life history


Egg

Geranium bronze eggs are small with a slight green color that turns white then brown or light-yellow before the larva hatches. The eggs are 0.5 mm in diameter and 0.3 mm in height. The eggs are typically laid close to the flower buds, but are occasionally laid on the leaves. The laying of eggs usually occurs during summer months, while caterpillar activity has been recorded in the summer, fall, and winter. File:Cacyreus marshalli larva02.jpg, Geranium bronze larva File:Cacyreus marshalli egg.jpg, Cacyreus marshalli egg


Caterpillar

Once hatched, the larvae bore through the stem of the host plant, where they are typically found within the stem or flower buds. The plant stem usually turns black after being invaded by larvae. The larva feeds on the host plant, damaging the host until it enters the
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
stage. During the first
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
stage, the caterpillar's average length is 1 mm, which will increase to 2 mm during the following 8 days. Second instar caterpillars grow to an average of 3 mm; third instars to 6 mm; and fourth instars grow to 13 mm. The second, third, and fourth instar stages occur for a duration of 8, 8, and 9 days, respectively. The color of the caterpillars can vary. Most caterpillars are yellow or green, and may have pink markings.


Pupa

The pupa stage occurs at the bottom of the flower peduncle in many of the host species. The peduncles offer the best protection for the caterpillars, which is why the caterpillars will typically remain at the base of the flower until
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
occurs. It has been suggested that geranium bronze may overwinter as either a caterpillar or a pupa, although further data is needed to confirm this. The pupa color varies, but they are typically green, pale-yellow, or brown. The pupa are hairy and are typically 9 mm long.


Migration

Adults are on wing year-round in warmer areas, but usually from August to May in South Africa. At higher altitudes, adults are on wing in December and January. Geranium bronze have spread from their native regions of South Africa to numerous southern and eastern European regions. This was likely due to accidental transportation of plants infested with geranium bronze larvae and not due to any natural migration patterns. The geranium bronze butterflies fly for short periods of time and frequently rest.


Predators

The geranium bronze butterfly has not been reported as a pest in its native home and surrounding South African regions. It is believed that the butterfly has not become a pest in the South African regions because of an indigenous predator or
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
that has kept the butterfly population low. The predator has yet to be identified.


Parental care


Oviposition

Geranium bronze females prefer to lay their eggs on the flower buds of various ''Geranium'' species. They typically oviposit on top of and underneath leaves, but infrequently on stems. Geranium bronze butterflies are multivoltine, meaning they lay two or more broods per year.


Plant selection for egg laying

Geranium bronze typically lay eggs on plants from genera ''Pelargonium'' and ''Geranium'', both in the family Geraniaceae. Once hatched, these plants act as hosts to the larvae. Quacchia et al. (2008) found that geranium bronze butterflies in Italy displayed preference for egg laying on ''
Geranium sanguineum ''Geranium sanguineum'', common names bloody crane's-bill or bloody geranium, is a species of hardy flowering herbaceous perennial plant in the cranesbill family Geraniaceae. It is also the county flower of Northumberland. Geranium sanguineum ...
, Geranium sylvaticum, and Geranium pratense'' Data was collected from a 2-year observation in various regions of Piedmont and the Aosta Valley, Italy. Plant preference and offspring fitness (using wingspan as a marker for fitness) were analyzed, but no statistically significant correlation was found.


Interaction with humans


Pest of plants

Geranium bronze butterflies pose a large threat to ''Pelargonium'' species in Europe. European and Mediterranean Plant Protection Organization currently lists geranium bronze as an A2 quarantine pest for Europe, and an A1 quarantine pest for Turkey. If geranium bronze numbers in Italy and other regions of Europe continue to rise, it is possible that they will become a larger threat for not only the native flora, but also for local biodiversity. Geranium bronze may start to be in competition with indigenous
lycaenid Lycaenidae is the second-largest family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfl ...
species whose only food source is ''Geranium'' plants, such as ''E. eumedon'' and ''A. nicias.'' Unfortunately, as pests, the geranium bronze butterflies have caused economic troubles as well. In countries like Spain, the ''Geranium'' plant species are highly bought in Spanish homes as ornaments. According to Sarto i Monteys et al., these plants are highly important in both sales and in employment for both production and marketing (Sarto i Monteys V 1991). In Spain alone, there are four major ''Geranium'' plant companies, together collecting over $30 million a year in the market. Companies have already seen drops in sales as the ''Geraniums'' have been affected by these butterflies. Currently, research is being done to find effective ways to prevent the attacks by these butterflies, but unfortunately many possible insecticides are too chemically potent on the plants that control of these butterflies has been difficult.


Use of pesticides on geranium bronze

There is a large body of research on the use of
pesticides Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampric ...
on geranium bronze butterflies. Contact pesticides were found to have no effect on the butterfly, because the larva spends the majority of its life within the plant, along with other
endophytic An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; h ...
habits. Herrero et al. found that ''
B. thuringiensis ''Bacillus thuringiensis'' (or Bt) is a gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. ''B. thuringiensis'' also occurs naturally in the gut of caterpillars of various types of moths and butterflie ...
'' may be effective in controlling the geranium bronze . In addition to ''Bacillus thuringiensis'', the insecticides diflubenzuron, flufenoxuron, hexaflumuron, lamda-cyhalothrin, alphamethrin and benfuracarb are also effective.


References

* J. Kleinekuhle: ''Zur Indigenität und Ausbreitung des afrikanischen Bläulings Cacyreus marshalli in Südeuropa''. Atalanta 26: 209-214, 1995 * J. H. R. Thiele, W. A. Nässig: ''Der Pelargonienbläuling auch in Deutschland''. Nachrichten des Entomologischen Vereins Apollo, Frankfurt am Main, N. F. 20 (3/4): 290, 2000 *Tom Tolman, Richard Lewington: ''Die Tagfalter Europas und Nordwestafrikas'' Franckh-Kosmos Verlags-GmbH & Co, Stuttgart 1998, *G. Tiberglien, « '' Les Lycène des Géraniums, '' Cacyreus marshalli '': état 2002 d'une espèce invasive'' », dans « Bull. Soc. Sc. Nat. Ouest Fr. N. S. », n° 24, 4, 2002 (p. 205-214) *Sarto Monteys, V., 1992. Spread of the Southern African Lycaenid butterfly, Cacyreus marshalli Butler, 1898, (LEP: Lycaenidae) in the Balearic Archipelago (Spain) and considerations on its likely introduction to continental Europe. ''J. Res. Lepid'', 31(1–2), pp. 24–34. Available at: http://lepidopteraresearchfoundation.org/pdf/pdf31/31-024.pdf ccessed October 4, 2017 * * {{Taxonbar, from=Q1768348 Butterflies described in 1897 Cacyreus Butterflies of Europe Butterflies of Africa Taxa named by Arthur Gardiner Butler