View source for Lytta vesicatoria

{{Short description|Species of beetle that produces a toxic blistering agent}}
{{About|the insect species|other uses of the term ''Spanish fly''|Spanish fly (disambiguation)}}
{{Good article}}
{{Use dmy dates|date=June 2021}}
{{Speciesbox
| image = Lytta-vesicatoria.jpg
| genus = Lytta
| species = vesicatoria
| authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])
}}

'''''Lytta vesicatoria''''', also known as the '''Spanish fly''', is an [[Aposematism|aposematic]] emerald-green [[beetle]] in the [[blister beetle]] family (Meloidae). It is distributed across [[Eurasia]].

The species and others in its family were used in traditional [[apothecary]] preparations as "Cantharides". The insect is the source of the [[terpenoid]] [[cantharidin]], a toxic blistering agent once used as an exfoliating agent, anti-rheumatic drug and an [[aphrodisiac]]. The substance has also found culinary use in some blends of the North African spice mix ''[[ras el hanout]]''. Its various supposed benefits have been responsible for accidental poisonings.

== Etymology and taxonomy ==

The [[Generic name (biology)|generic name]] is from the Greek λύττα (''lytta''), meaning martial rage, raging madness, [[Bacchantes|Bacchic frenzy]], or [[rabies]].<ref>{{cite web |last1=Liddell |first1=Henry George |last2=Scott |first2=Robert |title=λύττα, λυττάω, λυττητικός, etc., v. λυσς-. |url=https://logeion.uchicago.edu/%CE%BB%CF%8D%CF%84%CF%84%CE%B1 |publisher=Liddell & Scott |access-date=14 February 2017 |date=1940}}</ref><ref>{{cite web |last1=Liddell |first1=Henry George |last2=Scott |first2=Robert |title=λύσσα |url=https://logeion.uchicago.edu/morpho/%CE%BB%CF%8D%CF%84%CF%84%CE%B1 |publisher=Liddell & Scott|access-date=14 February 2017 |date=1940}}</ref> The [[Specific name (botany)|specific name]] is derived from Latin ''vesica'', blister.<ref>{{cite web |title=Latin definition for: vesica, vesicae |url=https://latin-dictionary.net/definition/38654/vesica-vesicae |website=Latin Dictionary & Grammar Resources |access-date=19 January 2022}}</ref>

''Lytta vesicatoria'' was formerly named ''Cantharis vesicatoria'',<ref name=FarlexDict12/> although the genus ''[[Cantharis]]'' is in an unrelated family, [[Cantharidae]], the soldier beetles.<ref>{{cite journal |last=Selander |first=Richardg B. |year=1991 |title=On the Nomenclature and Classification of Meloidae (Coleoptera) |url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1415&context=insectamundi |journal=Insecta Mundi |volume=5 |issue=2 |pages=65–94 }}</ref> It was classified there erroneously until the Danish zoologist [[Johan Christian Fabricius]] corrected its name in his ''Systema entomologiae'' in 1775. He reclassified the Spanish fly as the [[type species]] of the new genus ''[[Lytta]]'', in the family [[Meloidae]].<ref>{{cite journal |last=Selander |first=R. B. |year=1991 |title=On the nomenclature and classification of Meloidae (Coleoptera)] |journal=[[Insecta Mundi]] |volume=5 |issue=2 |pages=65–94 |url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1415&context=insectamundi}}</ref>

== Description and ecology ==

[[File:Spanish fly in Siberia (cropped).jpg|thumb|upright=1.7|An aggregation of Spanish fly adults in Siberia. The behaviour and their toxicity suggest that their conspicuous coloration is [[Aposematism|aposematic]].<ref name="Young 1984"/>]]

The adult Spanish fly is a slender, soft-bodied metallic and [[iridescent]] golden-green insect, one of the [[blister beetle]]s. It is approximately {{convert|5|mm|in|abbr=on|1}} wide by {{convert|20|mm|in|abbr=on|1}} long.<ref name=GrzimeksAnLifeEncycl>{{cite book |editor=Schlager, Neil |year=2004 |title=Grzimek's Animal Life Encyclopedia |chapter=Coleoptera (beetles and weevils) |edition=2nd |volume=3, Insects|location=Farmington Hills, Michigan |publisher=Thomson-Gale/American Zoo and Aquarium Association |isbn=978-0787657796 |page=331 |chapter-url=http://issuu.com/radu9990/docs/grzimek-s-encyclopedia-2nd-ed.---vol.-3---insects/348 |access-date=20 December 2015 }}</ref>

The female lays her fertilised eggs on the ground, near the nest of a ground-nesting solitary bee. The [[larva]]e 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, [[triungulin]]s (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 [[predator]]s and [[parasite]]s. The active larvae [[ecdysis|moult]] into very different, more typically [[Scarabaeoidea|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.<ref name="GrzimeksAnLifeEncycl"/><ref name="ITMA">{{cite web |title=Illustrated lecture notes on Tropical Medicine - Ectoparasites - Beetles |url=https://www.itg.be/files/docs/onderwijs/lntm2017.pdf |publisher=Institute of Tropical Medicine Antwerp |access-date=14 February 2017 |archive-date=20 January 2022 |archive-url=https://web.archive.org/web/20220120231450/https://www.itg.be/Files/docs/onderwijs/lntm2017.pdf |url-status=dead }}</ref>

The defensive chemical [[cantharidin]], for which the beetle is known, is synthesised 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.<ref>{{cite book |last1=Boggs |first1=Carol L. |editor1-last=Leather |editor1-first=S. R. |editor2-last=Hardie |editor2-first=J. |title=Male Nuptial Gifts: Phenotypic Consequences and Evolutionary Implications |date=1995 |publisher=[[CRC Press]] |pages=215–242 |url=https://www.researchgate.net/publication/233994025 }}</ref> Zoologists note that the conspicuous coloration, the presence of a powerful toxin, and the adults' aggregating behaviour in full view of any predators strongly suggest [[aposematism]] among the blistering meloid beetles.<ref name="Young 1984">{{cite journal |last=Young |first=Daniel K. |title=Cantharidin and insects: an historical review |journal=The Great Lakes Entomologist |volume=17 |issue=4 |year=1984 |pages=187–194 |url=https://scholar.valpo.edu/cgi/viewcontent.cgi?article=1518&context=tgle}}</ref>

== Range and habitat ==

The Spanish fly is found across Eurasia, though it is a mainly a southern European species,<ref name=Cutler>{{cite book |last=Cutler |first=Horace G. |title=Phytochemical Resources for Medicine and Agriculture |chapter=An Historical Perspective of Ancient Poisons |editor1-last=Nigg |editor1-first=Herbert N. |editor2-last=Seigler |editor2-first=David S. |year=1992 |page=3 |doi=10.1007/978-1-4899-2584-8_1 |isbn=978-1-4899-2586-2 }}</ref><ref name=GrzimeksAnLifeEncycl/><ref name=ITIS>{{cite book |editor-last=Guala |editor-first=Gerald |date=2015 |title=Lytta vesicatoria (Linnaeus, 1758), Taxonomic Serial No.: 114404 |chapter=Geographic Information: Geographic Division |location=Reston, Virginia |publisher=[[United States Geological Survey]], [[Integrated Taxonomic Information System]] |chapter-url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=114404# |access-date=20 December 2015 }}</ref> with some records from southern [[Great Britain]]<ref name=CGB>{{cite web |url=http://www.coleoptera.org.uk/species/lytta-vesicatoria |title=''Lytta vesicatoria'' (Linnaeus, 1758) |access-date=12 January 2018 |publisher=[[UK Centre for Ecology & Hydrology]] |work=UK Beetle Recording |archive-date=4 November 2017 |archive-url=https://web.archive.org/web/20171104082706/http://www.coleoptera.org.uk/species/lytta-vesicatoria |url-status=dead }}</ref> and Poland.<ref name=KSIB>{{cite web |url=http://baza.biomap.pl/en/taxon/species-lytta_vesicatoria_vesicatoria/mapb/tlm/list_species |title=''Lytta (Lytta) vesicatoria vesicatoria'' Linnaeus, 1758 |access-date=12 January 2017 |publisher=Polish Biodiversity Information Network (Krajowa Sieć Informacji o Bioróżnorodności)}}</ref>

Adult beetles primarily feed on leaves of [[ash tree|ash]], [[lilac]], [[amur privet]], [[honeysuckle]] and [[white willow]]. It is occasionally found on [[plum]], [[rose]], and [[elm]].<ref name=GrzimeksAnLifeEncycl/><ref>{{cite book |title=Medicines, their uses and mode of administration; including a complete conspectus of the three British Pharmacopoeias, an account of all the new remedies, and an Appendix of Formulae |last1=Neligan |first1=J. M. |last2=Macnamara |first2=R. |year=1867|page=297|url=https://play.google.com/store/books/details/John_Moore_NELIGAN_Medicines_their_uses_and_mode_o?id=_d5ZAAAAcAAJ |publisher=Fanin & Company }}</ref>

== Interaction with humans ==

[[File:Collecting cantharides.jpg|thumb|Collecting ''Cantharides'', 19th century]]

=== Preparation of cantharidin ===
{{main|Cantharidin}}

[[File:Cantharidin-2D.svg|thumb|[[Cantharidin]]]]

[[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]].<ref>{{cite journal |last=Robiquet |first=M. |year=1810 |title=Expériences sur les cantharides | journal=Annales de Chimie |volume=76 | pages=302–322 }}</ref><!--no current medical claim is being made here-->

Each beetle contains some 0.2–0.7&nbsp;mg of cantharidin, males having 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 as a blistering agent 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 [[iridescent]] reflections, 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''.<ref name=FarlexDict12>{{cite book |author=Anon |year=2012 |orig-year=2009 |title=Farlex Partner Medical Dictionary |chapter=Cantharide |location=Huntingdon Valley, Pennsylvania |publisher=Farlex |chapter-url=http://medical-dictionary.thefreedictionary.com/Cantharide |access-date=14 December 2015 }}</ref><ref name=Froberg10/><ref name=AggrawalAPC>{{cite book |editor-last=Aggrawal |editor-first=Anil |year=2007 |title=APC Textbook of Forensic Medicine and Toxicology |chapter=VII. Spanish Fly (Cantharides) |location=New Delhi, India |publisher=Avichal |isbn=978-8177394191 |chapter-url=https://books.google.com/books?isbn=8177394193 |access-date=14 December 2015 |page=652f }}</ref><ref name=SaundersVet>{{cite book |editor1=Blood, Douglas Charles |editor2=Studdert, Virginia P. |editor3=Gay, Clive C. |year=2007 |title=Saunders Comprehensive Veterinary Dictionary |edition=3rd |chapter=Cantharides |location=Philadelphia, PA, USA |publisher=Elsevier |isbn=978-0702027888 |chapter-url=https://books.google.com/books?isbn=070202788X |access-date=14 December 2015 |url-access=registration |url=https://archive.org/details/saunderscomprehe00doug }}</ref><ref name=MosbyDict>{{cite book |editor=Jonas, Wayne B. |year=2005 |title=Mosby's Dictionary of Complementary and Alternative Medicine |edition=3rd |chapter=Cantharides |location=Philadelphia, PA, USA |publisher=Elsevier Saunders |isbn=978-0323025164 |chapter-url=https://books.google.com/books?isbn=0323025161 |access-date=14 December 2015 }}</ref>

=== Toxicity and poisonings===

Cantharidin is dangerously toxic, inhibiting the enzyme [[PP2|phosphatase 2A]]. It causes irritation, blistering, bleeding and discomfort. These effects can escalate to erosion and [[hemorrhage|bleeding]] of [[mucosa]] in each system, sometimes followed by severe gastro-intestinal bleeding and [[acute tubular necrosis]] and [[Glomerulus (kidney)|glomerular]] destruction, resulting in gastro-intestinal and [[kidney dysfunction|renal dysfunction]], [[organ failure]], and death.<ref name=Froberg10/><ref name=EvansHooserCompTox10>{{cite book |last1=Evans |first1=T. J. |last2=Hooser |first2=S. B. |editor1=Hooser, Stephen |editor2=McQueen, Charlene |year=2010 |chapter=Comparative Gastrointestinal Toxicity (Ch. 16) |title=Comprehensive Toxicology |edition=2nd |isbn=978-0080468846 |pages=195–206 |location=London, England  |publisher=Elsevier Academic Press |chapter-url=https://books.google.com/books?isbn=0080468845 }}</ref><ref name=Gupta12>{{cite book |last1=Gwaltney-Brant |first1=Sharon M. |last2=Dunayer |first2=Eric |last3=Youssef |first3=Hany  |editor=Gupta, Ramesh C. |year=2012 |chapter=Terrestrial Zootoxins [Coleoptera: Meloidae (Blister Beetles) |title=Veterinary Toxicology: Basic and Clinical Principles |edition=2nd |isbn=978-0123859266 |pages=975–978 |location=London, England |publisher=Elsevier Academic Press |chapter-url=https://books.google.com/books?isbn=0123859263 |access-date=14 December 2015 }}</ref><ref name=karras>{{cite journal |last1=Karras |first1=David J. |title=Poisoning From "Spanish Fly" (Cantharidin) |journal=[[The American Journal of Emergency Medicine]] |year=1996 |doi=10.1016/S0735-6757(96)90158-8 |quote=While most commonly available preparations of Spanish fly contain cantharidin in negligible amounts, if at all, the chemical is available illicitly in concentrations capable of causing severe toxicity. |volume=14 |issue=5 |pages=478–483 |pmid=8765116 |last2=Farrell |first2=S. E. |last3=Harrigan |first3=R. A. |last4=Henretig |first4=F. M. |last5=Gealt |first5=L. |display-authors=3}}<!--paper reviews knowledge as well as reporting cases--></ref><ref name=WilsonCompTox10>{{cite book |last=Wilson |first=C. R. |editor=Hooser, Stephen |editor2=McQueen, Charlene |year=2010 |chapter=Methods for Analysis of Gastrointestinal Toxicants (Ch. 9) |title=Comprehensive Toxicology |edition=2nd |isbn=978-0080468846 |pages=145–152, esp. 150 |location=London, England |publisher=Elsevier Academic Press |chapter-url=https://books.google.com/books?isbn=0080468845 |access-date=14 December 2015 }}</ref>

Preparations of Spanish fly and its active agent have been implicated in both inadvertent<ref name=Froberg10>{{cite book |last=Froberg |first=Blake A. |editor1=Holstege, Christopher P. |editor2=Neer, Thomas |editor3=Saathoff, Gregory B. |editor4=Furbee, R. Brent |year=2010 |chapter=Animals |title=Criminal Poisoning: Clinical and Forensic Perspectives |location=Burlington, Massachusetts |publisher=Jones & Bartlett |isbn=978-1449617578 |pages=39–48, esp. 41, 43, 45ff |chapter-url=https://books.google.com/books?isbn=1449617573 |access-date=16 December 2015 }} Note: ''the active agent appears variously as [[cantharidin]],{{rp|41}} and "cantharadin"{{rp|43,45ff}} or "canthariadin"{{rp|238}}'' (sic).</ref> and intentional poisonings.<ref name=Froberg10/> Arthur Kendrick Ford was imprisoned in 1954 for the unintended deaths of two women surreptitiously given candies laced with [[cantharidin]], which he had intended to act as an [[aphrodisiac]].<ref name=Froberg10/> It has been suggested that [[George Washington]] was treated with Spanish fly for [[epiglottitis]], the condition which caused his death.<ref>{{cite book |last=Henriques |first=Peter R. |year=2000 |title=The Death of George Washington: He Died as He Lived |publisher=Mount Vernon Ladies' Association |location=Mount Vernon, Virginia |pages=27–36 |isbn=978-0-931917-35-6 }}</ref>

Currently the cantharidin in US, in the form of collodion, is used in the treatment of warts and [[molluscum]].{{cn|date=May 2024}}

=== Culinary uses ===

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 ''L. vesicatoria'', although its sale in Moroccan spice markets was banned in the 1990s.<ref name="Davidson">{{cite book |last=Davidson |first=Alan |editor-last=Jaine |editor-first=Tom |others=Vannithone, Soun (illustrator) |title=The Oxford Companion to Food |location=Oxford, England |publisher=[[Oxford University Press]] |year=1999 |isbn=978-0-19-211579-9 |url=https://archive.org/details/oxfordcompaniont0000davi |url-access=registration |access-date=13 December 2015 |page=671ff }}</ref> [[dawamesc|''Dawamesk'']], a spread or jam made in North Africa and containing [[hashish]], [[almond paste]], pistachio nuts, sugar, orange or [[tamarind]] peel, [[clove]]s, and other various spices, occasionally included cantharides.<ref>{{cite news |last=Green |first=Jonathon |title=Spoonfuls of paradise |url=https://www.theguardian.com/books/2002/oct/12/featuresreviews.guardianreview34 |access-date=14 February 2017 |newspaper=[[The Guardian]] |date=12 October 2002}}</ref>

=== Other uses ===

In [[History of China#Ancient China|ancient China]], the beetles were mixed with human excrement, [[arsenic]], and [[Aconitum|wolfsbane]] to make the world's first recorded [[stink bomb]].<ref name="Theroux_1">{{cite book |last=Theroux |first=Paul |author-link=Paul Theroux |title=[[Riding the Iron Rooster]] |publisher=Ivy Books |year=1989 |isbn=978-0-8041-0454-8 |page=[https://archive.org/details/ridingironrooste00ther_0/page/54 54]}}</ref>

In [[ancient Greece]] and [[Medicine in ancient Rome|Rome]], Spanish fly was used to attempt to treat skin diseases, while in medieval Persia, [[Medicine in the medieval Islamic world|Islamic medicine]] applied Spanish fly, named ''ḏarārīḥ'' (ذراریـح), to attempt to prevent [[rabies]].<ref name="Moallemi 2021">{{cite journal |last1=Moallemi |first1=Mostafa |last2=Yousofpour |first2=Mohammad |last3=Jokar |first3=Assie |title=Prevention of Rabies by Application of Lytta vesicatoria in Persian Medicine Texts in Islamic Civilization |journal=Traditional and Integrative Medicine |date=2021 |volume=6 |issue=1 |pages=70–77 |url=https://jtim.tums.ac.ir/index.php/jtim/article/download/254/244/}}</ref>

In the 19th century, Spanish fly was used externally mainly as blistering agent and local irritant; also, in chronic gonorrhoea, paralysis, lepra, ulcers therapy. ''L. vesicatoria'' was used internally as a diuretic stimulant.{{Cn|date=June 2024}}

An extract from Spanish fly was thought to be a strong [[aphrodisiac]], and various [[love potion]]s were named thusly.<ref>Joe Schwarcz, [https://www.mcgill.ca/oss/article/health-and-nutrition-you-asked/spanish-fly-really-aphrodisiac Is "Spanish Fly" really an aphrodisiac?]</ref> See {{section link|Cantharidin|Aphrodisiac preparations}} for more.

== References ==
{{Reflist|30em}}

== External links ==
{{Wikispecies|Lytta vesicatoria|Spanish fly}}
{{Commons|Lytta vesicatoria|Spanish fly}}
* {{cite EB9 |wstitle = Cantharides |volume= V | page=31 |short=1}}

{{Insects in culture}}
{{Taxonbar|from=Q624996}}

{{DEFAULTSORT:Spanish fly}}
[[Category:Meloidae]]
[[Category:Beetles described in 1758]]
[[Category:Animal taxa named by Carl Linnaeus]]
[[Category:Poisonous animals]]
[[Category:Insect products]]
[[Category:Beetles of Europe]]
[[Category:Insects in culture]]
[[Category:Parasites of bees]]