Cacyreus Marshalli
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The geranium bronze or ''brun des pélargoniums'' in French (''Cacyreus marshalli''), is a
butterfly Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
in the family
Lycaenidae Lycaenidae is the second-largest family (biology), 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 geranium bronze butterfly is native to
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. The butterfly was first introduced to
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
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 to cultivated ''
Pelargonium ''Pelargonium'' () is a genus of flowering plants that includes about 280 species of perennial plant, perennials, succulent plant, succulents, and shrubs, common name, commonly called geraniums, pelargoniums, or storksbills. ''Geranium'' is also ...
'' 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, with the gre ...
'' 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 used to control pests. They include herbicides, insecticides, nematicides, fungicides, and many others (see table). The most common of these are herbicides, which account for approximately 50% of all p ...
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 opposite wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingsp ...
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 Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
, including
KwaZulu-Natal KwaZulu-Natal (, also referred to as KZN) is a Provinces of South Africa, province of South Africa that was created in 1994 when the government merged the Zulu people, Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu language, Zulu) and ...
and the
Cape Provinces The Cape Provinces of South Africa is a biogeographical area used in the World Geographical Scheme for Recording Plant Distributions (WGSRPD). It is part of the WGSRPD region 27 Southern Africa. The area has the code "CPP". It includes the Sou ...
, as well as in neighbouring countries such as
Eswatini Eswatini, formally the Kingdom of Eswatini, also known by its former official names Swaziland and the Kingdom of Swaziland, is a landlocked country in Southern Africa. It is bordered by South Africa on all sides except the northeast, where i ...
,
Botswana Botswana, officially the Republic of Botswana, is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory part of the Kalahari Desert. It is bordered by South Africa to the sou ...
,
Zimbabwe file:Zimbabwe, relief map.jpg, upright=1.22, Zimbabwe, relief map Zimbabwe, officially the Republic of Zimbabwe, is a landlocked country in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Bots ...
,
Lesotho Lesotho, formally the Kingdom of Lesotho and formerly known as Basutoland, is a landlocked country in Southern Africa. Entirely surrounded by South Africa, it is the largest of only three sovereign enclave and exclave, enclaves in the world, t ...
, and
Mozambique Mozambique, officially the Republic of Mozambique, is a country located in Southeast Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west, and Eswatini and South Afr ...
. It was accidentally brought to Europe in 1978. After being recorded in
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
, a colony of geranium bronze was soon found in
Mallorca Mallorca, or Majorca, is the largest of the Balearic Islands, which are part of Spain, and the List of islands in the Mediterranean#By area, seventh largest island in the Mediterranean Sea. The capital of the island, Palma, Majorca, Palma, i ...
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 Rome (Italian language, Italian and , ) is the capital city and most populated (municipality) of Italy. It is also the administrative centre of the Lazio Regions of Italy, region and of the Metropolitan City of Rome. A special named with 2, ...
,
Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
, 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 Se ...
and Tyrrhenian coastal areas before extending inland. The geranium bronze butterfly now has established colonies in
Belgium Belgium, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...
,
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
, the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
, Italy,
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
,
Greece Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...
,
Malta Malta, officially the Republic of Malta, is an island country in Southern Europe located in the Mediterranean Sea, between Sicily and North Africa. It consists of an archipelago south of Italy, east of Tunisia, and north of Libya. The two ...
,
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
,
Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in continental Europe, Portugal borders Spain to its north and east, with which it share ...
, southern
Switzerland Switzerland, officially the Swiss Confederation, is a landlocked country located in west-central Europe. It is bordered by Italy to the south, France to the west, Germany to the north, and Austria and Liechtenstein to the east. Switzerland ...
,
Slovenia Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
,
Croatia Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...
, 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 east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
. The first record of the species in Palestine was made in the garden of the
Palestine Museum of Natural History The Palestine Museum of Natural History (PMNH; ) is a natural history museum in Bethlehem, West Bank, Palestine. Founded in 2013, it opened to the public in 2017. The museum collections focus on botany, zoology and entomology. The goals of the m ...
.


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 perennial plant, perennials, succulent plant, succulents, and shrubs, common name, commonly called geraniums, pelargoniums, or storksbills. ''Geranium'' is also ...
'' 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, with the gre ...
''. 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 (; : 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 life cycle, the stages th ...
stage. During the first
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
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 transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
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 Altitude is a distance measurement, usually in the vertical or "up" direction, between a reference datum and a point or object. The exact definition and reference datum varies according to the context (e.g., aviation, geometry, geographical s ...
, 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 Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
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 The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
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 Geraniaceae is a family of flowering plants placed in the order Geraniales. The family name is derived from the genus '' Geranium''. The family includes both the genus '' Geranium'' (the cranesbills, or true geraniums) and the garden plants call ...
. 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 name bloody crane's-bill or bloody geranium, is a species of hardy flowering herbaceous perennial plant in the cranesbill family Geraniaceae. It is the county flower of Northumberland. Etymology The genus name i ...
,
Geranium sylvaticum ''Geranium sylvaticum'', the wood cranesbill or woodland geranium, is a species of hardy flowering plant in the family Geraniaceae, native to Europe and northern Turkey. The Latin specific epithet ''sylvaticum'' means "of woodland", referring ...
, and
Geranium pratense ''Geranium pratense'', the meadow crane's-bill or meadow geranium, is a species of flowering plant in the family Geraniaceae, native to Europe and Asia. Forming a clump up to broad, it is a herbaceous perennial with hairy stems and lax saucer-sh ...
'' 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 The European and Mediterranean Plant Protection Organization (EPPO) is an Intergovernmental organization, intergovernmental organisation responsible for European cooperation in plant protection in the European and Mediterranean Sea, Mediterranean ...
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 valued 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 used to pest control, control pest (organism), pests. They include herbicides, insecticides, nematicides, fungicides, and many others (see table). The most common of these are herbicides, which account for a ...
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; ...
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, lambda-
cyhalothrin Cyhalothrin (ISO common name) is an organic compound that, in specific isomeric forms, is used as a pesticide. It is a pyrethroid, a class of synthetic insecticides that mimic the structure and properties of the naturally occurring insecticide pyr ...
, 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