L. vesicatoria is sometimes called
Cantharis vesicatoria,
[1] although the genus
Cantharis is in an unrelated family,
Cantharidae, the soldier beetles.
[2]
Description and etymology[edit]

Collecting cantharides, 19th century.
Lytta vesicatoria is a slender, soft-bodied metallic and
iridescent golden-green insect, one of the
blister beetles. It is approximately 5 mm (0.20 in) wide by 20 mm (0.79 in) long.
[3][4][5][6]
The generic and specific names derive from the Greek λύττα (
lytta) for martial rage, raging madness,
Bacchic frenzy, or
rabies,
[7][8] and Latin
vesica for blister.
[9]
Range and habitat[edit]
The Spanish fly is a mainly southern European species
[10][11] although its range of habitats is more completely described as being "[t]hroughout southern Europe and eastward to Central Asia and Siberia,"
[3] alternatively as being throughout Europe, and parts of northern and southern Asia (excluding China).
[12]
Life-cycle[edit]
The defensive chemical
cantharidin for which the beetle is known is produced only by males; females obtain it from males during mating, as the
spermatophore contains some. This may be a nuptial gift, increasing the value of mating to the female, and thus increasing the male's reproductive fitness.
[14]
The female lays her fertilised eggs on the ground, near the nest of a ground-nesting solitary bee. The
larvae are very active as soon as they hatch. They climb a flowering plant and await the arrival of a
solitary bee. They hook themselves on to the bee using the three claws on their legs that give the first
instar larvae their name,
triungulins (from Latin
tri, three, and
ungulus, claw). The bee carries the larvae back to its nest, where they feed on bee larvae and the bees' food supplies. The larvae are thus somewhere between
predatorsand
parasites. The active larvae
moult into very different, more typically scarabaeoid larvae for the remaining two or more instars, in a development type called
hypermetamorphosis. The adults emerge from the bees' nest and fly to the woody plants on which they feed.
[15][3]
Cantharidin[edit]
Cantharidin, the principal active component in preparations of Spanish fly, was first isolated and named in 1810 by the French chemist
Pierre Robiquet, who demonstrated that it was the principal agent responsible for the aggressively blistering properties of this insect's egg coating. It was asserted at that time that it was as toxic as the most violent poisons then known, such as
strychnine.
[16]
The active agent has been estimated present at about 0.2-0.7 mg per beetle, males producing significantly more than females. The beetle secretes the agent orally, and exudes it from its joints as a milky fluid. The potency of the insect species as a
vesicant has been known since antiquity and the activity has been used in various ways. This has led to its small-scale commercial preparation and sale, in a powdered form known as
cantharides (from the plural of Greek κανθαρίς,
Kantharis, beetle), obtained from dried and ground beetles. The crushed powder is of yellow-brown to brown-olive color with
iridescentreflections, is of disagreeable scent, and is bitter to taste.
Cantharidin, the active agent, is a
terpenoid, and is produced by some other insects, such as
Epicauta immaculata.
[4][1][17]
Cantharidin is dangerously toxic, inhibiting the enzyme
phosphatase 2A. It causes irritation, blistering, bleeding and discomfort. These effects can escalate to erosion and
bleeding of
mucosa in each system, sometimes followed by severe gastro-intestinal bleeding and acute tubular necrosis and glomerular destruction, resulting in gastro-intestinal and
renal dysfunction, by
organ failure, and death.
[17][18][19][20] [21][18][22]
Preparations from
L. vesicatoria and its active agent have been implicated in both inadvertent
[17] and intentional poisonings.
[17] Froberg notes a 1954 manslaughter case where
cantharidin was administered in a
coconut-flavoured candy as an intended aphrodisiac, resulting in illness and eventual death of two women (agent identified postmortem), and in facial blistering and criminal conviction of the perpetrator.
[17]
Culinary uses[edit]
In
Morocco and other parts of North Africa, spice blends known as
ras el hanout sometimes included as a minor ingredient "green metallic beetles", inferred to be cantharides from
L. vesicatoria, although sale of this in Moroccan spice markets was banned in the 1990s.
[23] Dawamesk, a spread or jam made in North Africa and containing
hashish,
almond paste, pistachio nuts, sugar, orange or
tamarind peel,
cloves, and other various spices, occasionally included cantharides.
[24]
Other uses[edit]
Noteworthy cases[edit]
The Venezuelan leader
Simón Bolívar may have been accidentally poisoned by application of Spanish fly.
[27]
Arthur Kendrick Ford was convicted and given a multiyear prison sentence in 1954 for the unintended deaths of two women surreptitiously given candies laced with
cantharidin, which were intended to act as an aphrodisiac.
[17]