Anagyris Foetida
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''Anagyris foetida'' (English common name stinking bean trefoil), is a species of flowering plant in the family
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
, forming a malodorous, Summer-
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
Phytognosis http://regenerag.org/blog/2018/4/12/anagyris-foetida-an-underutilized-mediterranean-nitrogen-fixing-shrub Retrieved at 12.31 on Friday 30/12/22. Francisco J Valtueña, Ana Ortega-Olivencia, Tomás Rodríguez-Riaño and Josefa López "Reproductive biology in Anagyris foetida L. (Leguminosae), an autumn–winter flowering and ornithophilous Mediterranean shrub" ''Botanical Journal of the Linnean Society'', Volume 157, Issue 3, July 2008, Pages 519–532, https://doi.org/10.1111/j.1095-8339.2008.00801.x shrub or small tree 2-4m in height with green twigs bearing grey-green
trifoliate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
leaves clad beneath in silvery
hairs Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
. The red-tinged, yellow,
Laburnum ''Laburnum'', sometimes called golden chain or golden rain, is a genus of two species of small trees in the subfamily Faboideae of the pea family Fabaceae. The species are '' Laburnum anagyroides''—common laburnum and '' Laburnum alpinum''— ...
-like flowers are borne on the previous season’s growth and are pollinated mainly by birds''Encyclopedie Biologique'' - XXV (25) P. Fournier ( Professor Canon Paul-Victor Fournier ) ''Le Livre des Plantes Medicinales et Veneneuses de France'' pub. Editions Lechevalier - Paris 1947 ed. Paul Lechevalier, Tome I, pps. 81-2. and bees - notably the
buff-tailed bumblebee ''Bombus terrestris'', the buff-tailed bumblebee or large earth bumblebee, is one of the most numerous bumblebee species in Europe. It is one of the main species used in greenhouse pollination, and so can be found in many countries and areas wher ...
.The large spot of red pigmentation on the flowers is located on the exterior of the
standard Standard may refer to: Symbols * Colours, standards and guidons, kinds of military signs * Standard (emblem), a type of a large symbol or emblem used for identification Norms, conventions or requirements * Standard (metrology), an object th ...
, the interior of which bears, in turn, many small, deep red spots.


Scientific Name

The genus name ''Anagyris'' is a compound of the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
prefix ανα (''ana-'') "up / back again / backwards" and γῦρος (''gŷros''), "ring" / "round", giving the meaning "bent back into a ring" - in reference to the curved pods of the plant, while the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
specific name ''foetida'' is the feminine form of the adjective ''foetidus'', meaning "stinking" or "smelly" - in reference to the foul stench of the bruised leaves, which is said to resemble that of spoiled beans.Quattrocchi, Umberto (2012). ''CRC World dictionary of medicinal and poisonous plants: common names, scientific names, eponyms, synonyms and etymology''. Volume I, A-C, pub. CRC Press Taylor and Francis Group. page 270, heading ''Anagyris L. Fabaceae (Thermopsideae)''.


Common names

The shrub has a wide variety of vernacular names in the languages spoken around the Mediterranean, most of them alluding to its foul smell and poisonous properties. Into the former category fall the French ''bois puant'' and the Italian ''legno puzzo'' - both meaning "stinkwood" - and the Spanish '' hediondo'' meaning simply "stinker". Into the latter category (poison-related names) fall two overlapping sub-categories: the first comprising names referencing the similarity in appearance of Anagyris to the
Carob The carob ( ; ''Ceratonia siliqua'') is a flowering evergreen tree or shrub in the Caesalpinioideae sub-family of the legume family, Fabaceae. It is widely cultivated for its edible fruit pods, and as an ornamental tree in gardens and landscap ...
tree, with its similarly large and pendent pods, and contrasting it unfavourably with that edible species, and the second, pejorative names containing the names of animals perceived as being in some way noxious. A purely carob-related name is the Italian ''carrubazzo'' meaning "bad carob", the suffix ''-azzo'' providing the pejorative sense, as in ''ragazzo'' now "boy", but with the original sense of "lout", "street urchin" or "slave". Carob-and-animal names include the Arabic ''kharroub kelb'' / ''karrûb el klâb'' "carob of the dog", ''kharroub el-khinzir'' "carob of the pig" and ''kharroub el-maiz'' "carob of the goat". Animal-and-bean names include the French ''fève de loup'' "bean of the wolf" and the Maltese ''fula tal-klieb'' and Arabic ''fûl el klâb'', both meaning "bean of the dog". Of dog-related plant names it may be noted that in Europe these generally convey a perceived inferiority of some sort when compared to plants fit for humans e.g. a
dog rose ''Rosa canina'', commonly known as the dog rose, is a variable climbing, wild rose species native to Europe, northwest Africa, and western Asia. Description The dog rose is a deciduous shrub normally ranging in height from , though sometimes it ...
is scentless and dog’s mercury (as opposed to the true mercury plant) is poisonous.Mabey, Richard, ''
Flora Britannica Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''. E ...
'',
Sinclair-Stevenson Sinclair-Stevenson Ltd is a British publisher founded in 1989 by Christopher Sinclair-Stevenson. Christopher Sinclair-Stevenson became an editor at Hamish Hamilton in 1961. Thirteen years later in 1974 he became managing director, establishing ...
, London, 1996, pp256–257.
Likewise in the
Arab world The Arab world ( ar, اَلْعَالَمُ الْعَرَبِيُّ '), formally the Arab homeland ( '), also known as the Arab nation ( '), the Arabsphere, or the Arab states, refers to a vast group of countries, mainly located in Western A ...
the dog is deemed a dirty animal which feeds on
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
and other unclean things and is itself unfit for human consumption - whence the Arabic language insults ''kelb'' and ''ibn al kalb'' ("dog" and "son of a dog", respectively). Regarding the Arab conception of the "carob of the pig", it may further be noted that the pig is the only animal the meat of which is explicitly condemned as
haram ''Haram'' (; ar, حَرَام, , ) is an Arabic term meaning 'Forbidden'. This may refer to either something sacred to which access is not allowed to the people who are not in a state of purity or who are not initiated into the sacred knowle ...
in the
Quran The Quran (, ; Standard Arabic: , Classical Arabic, Quranic Arabic: , , 'the recitation'), also romanized Qur'an or Koran, is the central religious text of Islam, believed by Muslims to be a revelation in Islam, revelation from God in Islam, ...
( Quran 2:173) (see also
Islamic dietary laws Islamic dietary laws are dietary laws that Muslims follow. Islamic jurisprudence specifies which foods are '' '' (, "lawful") and which are '' '' (, "unlawful"). The dietary laws are found in the Quran, the holy book of Islam, as well as in coll ...
).


Distribution

The plant has a circum-Mediterranean distribution in Europe and North Africa, with extensions into Turkey, Iran and the Arabian peninsula.


Habitat

A. foetida is a tropical or subtropical
relict species In biogeography and paleontology, a relict is a population or taxon of organisms that was more widespread or more diverse in the past. A relictual population is a population currently inhabiting a restricted area whose range was far wider during a ...
from the
Tertiary Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
( a status suggested (i.a.) by its unusually large and heavy seeds),Francisco J Valtueña, Ana Ortega-Olivencia and Rodríguez-Riaño Tomás "Germination and seed bank biology in some Iberian populations of Anagyris foetida L. (Leguminosae)" April 2008 ''Plant Systematics and Evolution'' 275(3):231-243 DOI:10.1007/s00606-008-0067-2 https://www.researchgate.net/profile/Rodriguez-Riano-Tomas/publication/225357180_Germination_and_seed_bank_biology_in_some_Iberian_populations_of_Anagyris_foetida_L_Leguminosae/links/0fcfd50a21249dccc2000000/Germination-and-seed-bank-biology-in-some-Iberian-populations-of-Anagyris-foetida-L-Leguminosae.pdf?origin=publication_detail Retrieved at 10.23 on Friday 16/12/22. which suggests that its original homeland may have lain somewhere in the refugium encompassing the
Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
, Türkiye and the Black Sea coastal region of the Caucasus. The picture is complicated by the fact that it is also an
archaeophyte An archaeophyte is a plant species which is non-native to a geographical region, but which was an introduced species in "ancient" times, rather than being a modern introduction. Those arriving after are called neophytes. The cut-off date is usua ...
(ancient introduction - original homeland uncertain). In France it is a rare and protected species now found only in coastal
départements A department (, ) is an administrative or political division in several countries. Departments are the first-level divisions of 11 countries, nine in the Americas and two in Africa. An additional 10 countries use departments as second-level div ...
, although it once grew also in
Alpes-Maritimes Alpes-Maritimes (; oc, Aups Maritims; it, Alpi Marittime, "Maritime Alps") is a department of France located in the country's southeast corner, on the Italian border and Mediterranean coast. Part of the Provence-Alpes-Côte d'Azur region, it ...
where it is now extinct. It grows in
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
soils in seasonally
arid A region is arid when it severely lacks available water, to the extent of hindering or preventing the growth and development of plant and animal life. Regions with arid climates tend to lack vegetation and are called xeric or desertic. Most ar ...
areas exposed to the hottest summer sun, favouring rocky
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
escarpment An escarpment is a steep slope or long cliff that forms as a result of faulting or erosion and separates two relatively level areas having different elevations. The terms ''scarp'' and ''scarp face'' are often used interchangeably with ''escar ...
s (as currently at
Mont Faron Mont Faron is a mountain overlooking the city and roadstead of Toulon, France. It is 584m high. At its peak is a memorial dedicated to the 1944 Allied landings in Provence (Operation Dragoon), and to the liberation of Toulon. The top can be rea ...
), often among the ruins of
castle A castle is a type of fortified structure built during the Middle Ages predominantly by the nobility or royalty and by military orders. Scholars debate the scope of the word ''castle'', but usually consider it to be the private fortified r ...
s and
abbey An abbey is a type of monastery used by members of a religious order under the governance of an abbot or abbess. Abbeys provide a complex of buildings and land for religious activities, work, and housing of Christian monks and nuns. The conce ...
s (see also
monastic garden A monastic garden was used by many people and for multiple purposes. Gardening was the chief source of food for households, but also encompassed orchards, cemeteries and pleasure gardens, as well as providing plants for medicinal and cultural uses. ...
), suggesting that it may be a relic of cultivation for
medicinal purposes ''Medicinal Purposes'' is a Big Finish Productions audio drama based on the long-running British science fiction television series ''Doctor Who''. Plot Edinburgh, 1827. Body snatchers William Burke and William Hare are on the loose while the ...
or even to furnish an
arrow poison Arrow poisons are used to poison arrow heads or darts for the purposes of hunting and warfare. They have been used by indigenous peoples worldwide and are still in use in areas of South America, Africa and Asia. Notable examples are the poisons se ...
for military use.


Adaptations to growth in Winter and early Spring


Summer leaf-fall

Anagyris foetida is an ultra
drought-tolerant Drought tolerance is the ability to which a plant maintains its biomass production during arid or drought conditions. Some plants are naturally adapted to dry conditions'','' surviving with protection mechanisms such as desiccation tolerance, deto ...
member of the Fabaceae family. The large shrub originates in the
Mediterranean Basin In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and w ...
. The plant illustrates one of the best drought-resisting strategies of
mediterranean flora Mediterranean forests, woodlands, and scrub is a biome defined by the World Wide Fund for Nature. The biome is generally characterized by dry summers and rainy winters, although in some areas rainfall may be uniform. Summers are typically hot in ...
: the plant keeps its leaves through autumn, winter and spring, then it goes completely
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
in summer, becoming dormant and highly resistant to
drought A drought is defined as drier than normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, an ...
.
This pattern of aestivation (summer dormancy) is more characteristic of many Mediterranean bulbs than it is of the majority of Mediterranean shrubs, which have often adopted the different strategy of
sclerophyll Sclerophyll is a type of vegetation that is adapted to long periods of dryness and heat. The plants feature hard leaf, leaves, short Internode (botany), internodes (the distance between leaves along the stem) and leaf orientation which is paral ...
y to cope with high Summer temperatures and drought.


Pollination by birds

Canon Fournier noted as early as 1947 that ''A. foetida'' was primarily bird-pollinated. This was substantiated by Ortega-Olivencia et al. who carried out a study of the pollinators visiting two populations of the plant in Southwest Spain over a three-year period and recorded their findings in a paper of 2005."First Confirmation of a Native Bird-Pollinated Plant in Europe" Ana Ortega-Olivencia, Tomás Rodríguez-Riaño, Francisco J. Valtueña, Josefa López, Juan A. Devesa and Jane Memmott, ''Oikos'' Vol. 110, No. 3 (Sep., 2005), pp. 578-590 (13 pages) Abstract viewable at https://onlinelibrary.wiley.com/doi/abs/10.1111/j.0030-1299.2005.13877.x Retrieved at 8.52 on Friday 9/12/22. Such bird-pollination (
ornithophily Ornithophily or bird pollination is the pollination of flowering plants by birds. This sometimes (but not always) coevolutionary association is derived from insect pollination (entomophily) and is particularly well developed in some parts of the w ...
) is, so far, unique among the thousands of plant species comprising the
flora of 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 subcontinent of Eurasia and it is located entire ...
. The avian pollinators recorded in the study were three
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
species: the
Common chiffchaff The common chiffchaff (''Phylloscopus collybita''), or simply the chiffchaff, is a common and widespread leaf warbler which breeds in open woodlands throughout northern and temperate Europe and the Palearctic. It is a migratory passerine which ...
, the
Eurasian blackcap The Eurasian blackcap (''Sylvia atricapilla''), usually known simply as the blackcap, is a common and widespread typical warbler. It has mainly olive-grey upperparts and pale grey underparts, and differences between the five subspecies are sm ...
and the
Sardinian warbler The Sardinian warbler (''Curruca melanocephala'') is a common and widespread typical warbler from the Mediterranean region. Like most ''Curruca'' species, it has distinct male and female plumages. The adult male has a grey back, whitish underpart ...
. Relevant in this context are the red markings on the flowers of A. foetida, red being a colour particularly noticeable by (and therefore attractive to) birds, with their
tetrachromat Tetrachromacy (from Greek ''tetra'', meaning "four" and ''chromo'', meaning "color") is the condition of possessing four independent channels for conveying color information, or possessing four types of cone cell in the eye. Organisms with tet ...
vision. The authors offer the following plausible explanation for the unusual form of pollination which has evolved in the case of ''A. foetida'':
The negligible existence of insect visitors and pollinators of ''A. foetida'' flowers could be explained by the early flowering of this species during the autumn–winter, coinciding with the coolest and wettest months of the year (cold, high winds, almost daily mists that persist until nightfall, and above all, abundant rainfall).
A complementary study of a similar kind undertaken by Haran et al. in Israel in 2018 revealed yet further bird species involved in the pollination of ''A. foetida'', along with a single species which steals nectar from the flowers while seldom acting as a pollen vector."Specialist nectarivorous birds (''Cinnyris osea'') steal nectar whereas omnivorous birds are pollen transfer vectors of ''Anagyris foetida''" Ron Haran, Ido Izhaki, Amots Dafni ''Journal of Pollination Ecology'', 23(9), 2018, pp 82-89 https://www.researchgate.net/profile/Ido-Izhaki/publication/325527432_Specialist_nectarivorous_birds_Cinnyris_osea_steal_nectar_whereas_omnivorous_birds_are_pollen_transfer_vectors_of_Anagyris_foetida/links/5b122a874585150a0a608d51/Specialist-nectarivorous-birds-Cinnyris-osea-steal-nectar-whereas-omnivorous-birds-are-pollen-transfer-vectors-of-Anagyris-foetida.pdf?origin=publication_detail Retrieved at 19.05 on Friday 9/12/22 In addition to the Common chiffchaff, the Eurasian Blackcap and the Sardinian Warbler the Israeli study documents another six pollinating species: the
White-spectacled bulbul The white-spectacled bulbul (''Pycnonotus xanthopygos'') is a member of the bulbul family. It is in length with a wingspan of . These birds live in fruit plantations, gardens, and cities. It is the most common member of the bulbul family in Isr ...
, the
Spanish Sparrow The Spanish sparrow or willow sparrow (''Passer hispaniolensis'') is a passerine bird of the sparrow family Passeridae. It is found in the Mediterranean region and south-west and central Asia. It is very similar to the closely related house spar ...
, the
Common whitethroat The common whitethroat or greater whitethroat (''Curruca communis'') is a common and widespread typical warbler which breeds throughout Europe and across much of temperate western Asia. This small passerine bird is strongly migratory, and winter ...
, the
Lesser whitethroat The lesser whitethroat (''Curruca curruca'') is a common and widespread typical warbler which breeds in temperate Europe, except the southwest, and in the western and central Palearctic. This small passerine bird is strongly migratory, winteri ...
, Rüppell's warbler and the
Eastern Orphean warbler The eastern Orphean warbler (''Curruca crassirostris'') is a typical warbler of the genus ''Curruca''. This species occurs in summer around the Mediterranean, through the Balkans via Turkey, the Caucasus and surrounding regions to Central Asia. I ...
. The species found generally to operate as a nectar thief was the
Palestine sunbird The Palestine sunbird (''Cinnyris osea'') is a small passerine bird of the sunbird family, Nectariniidae. Found in parts of the Middle East and sub-Saharan Africa, it is also known as the orange-tufted sunbird, a name also used for the similar ...
(in consequence of its longer bill), although even it could act as a pollinator, on occasion. As in Spain, bird pollination is explicable by the plant’s flowering at a time when weather and temperatures are uncongenial to insect activity: the plant is almost the only shrub in the eastern Mediterranean region that flowers in winter.


Seed dispersal

The fruits of this species mature at the beginning of Summer, and are dispersed by simple fall under gravity. The process of fruit
abscission Abscission () is the shedding of various parts of an organism, such as a plant dropping a leaf, fruit, flower, or seed. In zoology, abscission is the intentional shedding of a body part, such as the shedding of a claw, husk, or the autotomy of a ...
lasts about a month, although really most of the fruits fall in the first two weeks. The fruits are
indehiscent Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that op ...
and, if they are not eaten by
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animals ...
, remain on the ground. After the heat of summer and the arrival of autumnal rains, the papyraceous–
coriaceous This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...
pericarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Aggre ...
(papery, corky pod) begins to rot, passing from brown to blackish in colour, and leaving the seeds exposed. These are large in size, very hard, and of an attractive bluish or violet-bluish colour. At least at present and in the populations studied, A. foetida shows no specialized dispersal mechanism, as is the case in many species of the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
- thus Ortega-Olivencia, Valtueña and Rodríguez-Riaño in their data-rich paper of 2008 attempting to account for the relative rarity and decline of ''Anagyris foetida'' in Spain - the study focussing on populations of the plant in Andalusia and Valencia. The authors are cautious in applying conclusions drawn from data gathered solely in Spain, but nonetheless offer pointers for future research in other countries where the plant is native.


Soil seed banks

The authors sought to determine whether the plant could form a
soil seed bank The soil seed bank is the natural storage of seeds, often dormant, within the soil of most ecosystems. The study of soil seed banks started in 1859 when Charles Darwin observed the emergence of seedlings using soil samples from the bottom of a lak ...
, while doubting that this could be possible, given the large size and high surface-to-volume ratio of the somewhat flattened, reniform seeds, which would be likely to hinder their slipping through gaps in the soil and their sinking in soil through the action of earthworms and other soil organisms. Their doubts, however, proved unfounded: since the species could indeed form seed banks through two mechanisms: first the seed could sink during the rainy season, when the soil became waterlogged and
colloidal A colloid is a mixture in which one substance consisting of microscopically dispersed insoluble particles is suspended throughout another substance. Some definitions specify that the particles must be dispersed in a liquid, while others extend ...
, and secondly there emerged evidence that
moles Moles can refer to: * Moles de Xert, a mountain range in the Baix Maestrat comarca, Valencian Community, Spain * The Moles (Australian band) *The Moles, alter ego of Scottish band Simon Dupree and the Big Sound People *Abraham Moles, French engin ...
were involved in seed burial when they raised molehills while digging their burrows during the winter months, although this occurred only in areas where the soil had not been overly compacted by the trampling of
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animals ...
.


Olive mimicry

While the authors were unable to discover a seed dispersal vector for the plant in Spain, they did discover an intriguing strategy encouraging
endozoochory In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
(consumption of seeds followed by
defecation Defecation (or defaecation) follows digestion, and is a necessary process by which organisms eliminate a solid, semisolid, or liquid waste material known as feces from the digestive tract via the anus. The act has a variety of names ranging f ...
at a distance) that suggested that a species of bird might be involved. The large blue-black seeds of ''Anagyris foetida'' bear a strong resemblance (particularly when seen from a distance) to the fruits of ''Olea europaea'' var. ''sylvestris'', the wild
olive The olive, botanical name ''Olea europaea'', meaning 'European olive' in Latin, is a species of small tree or shrub in the family Oleaceae, found traditionally in the Mediterranean Basin. When in shrub form, it is known as ''Olea europaea'' ...
, which, being fleshy, brightly coloured and
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
-rich, have typical characteristics of fruits encouraging dispersal by birds.
The similarity between these fruits and the seeds of Anagyris could lead to dispersal of the seeds of Anagyris by birds in habitats where Olive and Anagyris coexist and are abundant (they are frequently found in the same habitat), because of the birds confusing them with the fruits of Olea. Nevertheless, the test of this hypothesis did not give a positive result in the population studied…
While the authors failed to observe any birds which are habitual consumers of black olives in Spain consuming Anagyris seeds in error, they do not rule out the role of olive
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
in seed dispersal in other Mediterranean countries where both Anagyris and Olea are present. (See also
Vavilovian mimicry In plant biology, Vavilovian mimicry (also crop mimicry or weed mimicry) is a form of mimicry in plants where a weed evolves to share one or more characteristics with a domesticated plant through generations of artificial selection. It is name ...
).


Role of sheep

Few seeds can sink to the seed bank in the soil due to the intense pressure of predation by sheep. This is due principally to the shrubby habit of the plants and constant presence of sheep (always present during the period of ripening of the fruits, when predation has the most negative effect).
In Spain, sheep proved to have both negative and positive effects upon seed production and germination. On the negative side, the animals showed a predilection for
browsing Browsing is a kind of orienting strategy. It is supposed to identify something of relevance for the browsing organism. When used about human beings it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing o ...
on unripe pods (at which stage of development the seeds have not yet acquired a hard testa) and thus destroyed, by
mastication Chewing or mastication is the process by which food is crushed and ground by teeth. It is the first step of digestion, and it increases the surface area of foods to allow a more efficient break down by enzymes. During the mastication process, th ...
, a great many seeds before these had had a chance to ripen. On the positive side, in years of drought, when there was no grass upon which they could graze, sheep would resort to eating fallen pods containing ripe seeds. This suggested to the authors that sheep might be involved in endozoochoric seed dispersal, but examination of their
droppings Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
failed to reveal any ripe seeds that had passed through their guts (which would have involved their being exposed to acid scarification in the animals’ stomachs). However, the authors discovered that, in consuming the ripe, fallen pods, the sheep had swallowed the pericarps, but spat out the ripe seeds, having (sometimes) chewed them enough to rupture their leathery testae (seed coats) but not enough to kill the embryos in them - thus improving their rate of germination by up to 48%.


Other possible mammalian and avian vectors

The authors note that horses and cattle play no role in seed dispersal, because they do not browse on the fruits of A. foetida. They speculate that, in habitats modified less by
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
than those of the Spanish locales in which they carried out their research, other mammals, such as deer, foxes and hares might be involved (indeed, they do not even rule out sheep completely as vectors elsewhere in the plant’s range). Olive mimicry by the seeds might suggest the existence of an avian vector of some kind, but in that instance, scarification would be purely chemical (by stomach acid),not involving mastication, as the seeds would be swallowed whole, without any preliminary chewing. Such chemical scarification by a bird would be less effective than mechanical scarification by the teeth of a mammal in rupturing the seed coat to facilitate imbibition.


Unknown seed dispersal vector

In Spain, the dispersers of A. foetida fruit remain unknown, although this does not mean that the species does not possess them in other populations in its area of distribution. ''If it still exists'' talics added this disperser is likely to be a mammal since the latency of the seeds of this species, which are hard and impermeable, is broken physically through biting and by the action of stomach acid.
As the above makes plain, the principal disperser or dispersers of A. foetida may actually be extinct, which conclusion would go some way toward accounting for its relative rarity and would fit with its status as an archaeophyte possibly no longer to be found growing in a truly wild state.


Tough, waterproof seed coats

As suggested by their accumulation in soil seed banks, the seeds of ''A. foetida'' have tough, waterproof testae (seed coats) which enable them to remain viable for a long time. The authors demonstrated this water-resistance clearly by immersing a batch of unscarified seeds (i.e. seeds with undamaged seed coats) in a beaker of water for no fewer than four years - without the least ensuing sign of
imbibition Imbibition is a special type of diffusion that takes place when liquid is absorbed by solids-colloids causing an increase in volume. Water surface potential movement takes place along a concentration gradient; some dry materials absorb water. A ...
(water uptake necessary for the initiation of germination).
Scarification Scarification involves scratching, etching, burning/branding, or superficially cutting designs, pictures, or words into the skin as a permanent body modification or body art. The body modification can take roughly 6–12 months to heal. In the p ...
of the seeds before their immersion (whether chemically by acids or mechanically by nicking, filing or similar damage from the teeth of sheep) on the other hand, could result in complete imbibition within 48 hours - this being indicated by colour change. The fully-imbibed seeds swelled and lost their purple-black, olive-like tones as the
anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical compo ...
pigments responsible for this colouration leached away into the water, leaving the swollen seeds cream-coloured - and thus no longer potentially attractive to birds fooled by olive mimicry. These anthocyanin pigments not only mimic the dark colours of black olives, but also confer resistance to pathogens, since they posess
antibacterial An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention of ...
properties providing a measure of protection of the seed coats from breakdown by soil
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
.


Value as a nitrogen-fixing pioneer and companion plant

A. foetida will tolerate a wide variety of soils and has been grown as a fast-growing, nitrogen-fixing
pioneer species Pioneer species are hardy species that are the first to colonize barren environments or previously biodiverse steady-state ecosystems that have been disrupted, such as by wildfire. Pioneer flora Some lichens grow on rocks without soil, so m ...
in the large-scale
restoration Restoration is the act of restoring something to its original state and may refer to: * Conservation and restoration of cultural heritage ** Audio restoration ** Film restoration ** Image restoration ** Textile restoration * Restoration ecology ...
of fire-damaged areas in Southern Europe, using native species. It has also been planted at high densities as a
nitrogen-fixing Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmos ...
companion plant (albeit a very toxic one) among fruit and nut trees.


Medicinal use

The blossom smells strongly of
cabbage Cabbage, comprising several cultivars of ''Brassica oleracea'', is a leafy green, red (purple), or white (pale green) biennial plant grown as an annual vegetable crop for its dense-leaved heads. It is descended from the wild cabbage ( ''B.&nb ...
, and all parts of the plant were used medicinally, particularly in cases surrounding childbirth problems.
The first to write an account of the folk medicinal properties of ''A. foetida'' Guérin, Paul Émile Alexis, ''Recherches sur la localisation de l'anagyrine & de la cytisine'' (thèse), École supérieure de pharmacie de Paris, 1895 (translation: Research upon the sources of the alkaloids anagyrine and cytisine : thesis submitted to the Higher School of Pharmacy, Paris 1885) https://archive.org/details/BIUSante_pharma_p5293x1895x01/page/n9/mode/2up Retrieved at 12.39 on 29/11/22. was Ancient Greek herbalist and physician
Pedanius Dioscorides Pedanius Dioscorides ( grc-gre, Πεδάνιος Διοσκουρίδης, ; 40–90 AD), “the father of pharmacognosy”, was a Greek physician, pharmacologist, botanist, and author of ''De materia medica'' (, On Medical Material) —a 5-vol ...
in his monumental work ''
De materia medica (Latin name for the Greek work , , both meaning "On Medical Material") is a pharmacopoeia of medicinal plants and the medicines that can be obtained from them. The five-volume work was written between 50 and 70 CE by Pedanius Dioscorides, a ...
''. He lists the names ''onaguris'', ''anaguris'', ''anagyros'', ''acopon'', and ''agnacopum'' for the shrub and records distinct uses for the leaves, root and seeds. Concerning the leaves, he recommends a
poultice A poultice, also called a cataplasm, is a soft moist mass, often heated and medicated, that is spread on cloth and placed over the skin to treat an aching, inflamed, or painful part of the body. It can be used on wounds, such as cuts. 'Poultice' ...
of the young foliage to treat
oedema Edema, also spelled oedema, and also known as fluid retention, dropsy, hydropsy and swelling, is the build-up of fluid in the body's tissue. Most commonly, the legs or arms are affected. Symptoms may include skin which feels tight, the area ma ...
and an infusion of a small quantity of the leaves in
raisin wine Straw wine, or raisin wine, is a wine made from grapes that have been dried to concentrate their juice. The result is similar to that of the ice wine process, but is a much older process and suitable for warm climates. The technique dates back ...
to treat
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, cou ...
,
headache Headache is the symptom of pain in the face, head, or neck. It can occur as a migraine, tension-type headache, or cluster headache. There is an increased risk of depression in those with severe headaches. Headaches can occur as a result ...
and delayed menstruation. He further records two uses of the leaves suggesting uterine stimulant properties: an aid to
placental expulsion Placental expulsion (also called afterbirth) occurs when the placenta comes out of the birth canal after childbirth. The period from just after the baby is expelled until just after the placenta is expelled is called the third stage of labor. The ...
and an
abortifacient An abortifacient ("that which will cause a miscarriage" from Latin: ''abortus'' "miscarriage" and '' faciens'' "making") is a substance that induces abortion. This is a nonspecific term which may refer to any number of substances or medications, ...
. Of the root, he writes that it "dissolves and ripens" (presumably in reference to
tumour A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
s and
boil A boil, also called a furuncle, is a deep folliculitis, which is an infection of the hair follicle. It is most commonly caused by infection by the bacterium ''Staphylococcus aureus'', resulting in a painful swollen area on the skin caused by an ...
s, although this is not specifically stated). His statement "It is hung as an
amulet An amulet, also known as a good luck charm or phylactery, is an object believed to confer protection upon its possessor. The word "amulet" comes from the Latin word amuletum, which Pliny's ''Natural History'' describes as "an object that protects ...
on those who have hard
labour Labour or labor may refer to: * Childbirth, the delivery of a baby * Labour (human activity), or work ** Manual labour, physical work ** Wage labour, a socioeconomic relationship between a worker and an employer ** Organized labour and the labour ...
, yet one must at once (after the woman’s delivery) take off the amulet and put it away" presumably also refers to the root, although it is possible that other plant parts are also intended (contact between plant parts and bare skin could conceivably lead to some degree of absorption of active constituents). Concerning the seeds, he says only that their consumption causes excessive vomiting. Eminent French botanist Professor Canon Paul-Victor Fournier (1877-1964) devotes two pages of his three-volume work on the medicinal and poisonous plants of France (published in 1947) to the uses and toxicity of the plant. The seeds of the plant were formerly employed as an
emetic Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteri ...
(which
Renaissance The Renaissance ( , ) , from , with the same meanings. is a period in European history marking the transition from the Middle Ages to modernity and covering the 15th and 16th centuries, characterized by an effort to revive and surpass ideas ...
doctor and botanist Matthiolus observed was so violent in its action that it could cause
internal bleeding Internal bleeding (also called internal hemorrhage) is a loss of blood from a blood vessel that collects inside the body. Internal bleeding is usually not visible from the outside. It is a serious medical emergency but the extent of severity depen ...
), while an infusion of the leaves was used as a
purgative Laxatives, purgatives, or aperients are substances that loosen stools and increase bowel movements. They are used to treat and prevent constipation. Laxatives vary as to how they work and the side effects they may have. Certain stimulant, lubri ...
, but neither use should be considered safe, given the extreme toxicity of the species.


Toxicity

The alkaloid anagyrine, present in the seeds, causes, in warm-blooded animals, first slowing of the breathing and heartbeat and ultimately cessation of respiration and
cardiac arrest Cardiac arrest is when the heart suddenly and unexpectedly stops beating. It is a medical emergency that, without immediate medical intervention, will result in sudden cardiac death within minutes. Cardiopulmonary resuscitation (CPR) and possib ...
.


Cause of birth defects

As the specific name foetida (as do various common names in European languages) indicates, the foliage emits an unpleasant smell when brushed against or handled. The plant is very poisonous, containing a variety of
quinolizidine alkaloids Quinolizidine alkaloids are natural products that have a quinolizidine structure; this includes the lupine alkaloids. Occurrence Quinolizidine alkaloids can be found in the plant family of legumes, especially in papilionaceous plants. While t ...
, including the
teratogenic Teratology is the study of abnormalities of physiological development in organisms during their life span. It is a sub-discipline in medical genetics which focuses on the classification of congenital abnormalities in dysmorphology. The related t ...
anagyrine (named for the genus Anagyris and occurring also in certain toxic species belonging to the genus
Lupinus ''Lupinus'', commonly known as lupin, lupine, or regionally bluebonnet etc., is a genus of plants in the legume family Fabaceae. The genus includes over 199 species, with centers of diversity in North and South America. Smaller centers occur ...
Couch, James Fitton. "Lupine Studies. XIV.1 The Isolation of Anagyrine From ''Lupinus laxiflorus var.silvicola'' C. P. Smith." ''Journal of the American Chemical Society'' 61.12 (1939): 3327-328. https://pubs.acs.org/doi/pdf/10.1021/ja01267a027 Retrieved at 12.18 on 28/11/22. ). The highest concentration of the toxins occurs in the small, greenish-purple, bean-like seeds.


Danger to children

The shrub should not be grown in areas where children play because they may be tempted by the bean-like appearance of the seed pods to nibble the attractive seeds, with potentially fatal consequences (- the most dangerous poisonous plants are those combining high toxicity with fruits resembling those of edible species).


Hazardous tainting of milk products

As canon Fournier points out, stock generally avoid browsing upon the plant because of its unpleasant smell and taste, this being fortunate for humans, for, on the rare occasions when ewes have been forced, by hunger and the unavailability of other fodder, to consume the plant in quantity, their milk has become not merely tainted, but poisonous. Persons who have eaten
cheese Cheese is a dairy product produced in wide ranges of flavors, textures, and forms by coagulation of the milk protein casein. It comprises proteins and fat from milk, usually the milk of cows, buffalo, goats, or sheep. During production, ...
prepared from such milk have suffered violent vomiting, and, on occasion, (unspecified) symptoms of an even more severe nature.


Case of accidental poisoning in Algeria

Trotter mentions, in passing (and without further elaboration), a case of accidental poisoning by the plant recorded in a work of Vesque’s: a group of hungry soldiers,
foraging Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...
for edible wild plants while stationed in Algeria, mistook the seeds of ''A. foetida'' for
bean A bean is the seed of several plants in the family Fabaceae, which are used as vegetables for human or animal food. They can be cooked in many different ways, including boiling, frying, and baking, and are used in many traditional dishes th ...
s, with grave consequences.


Cultural references: Anagyris and Anagyrous

That the plant was deemed proverbially smelly in
Ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cult ...
is demonstrated by a
punning A pun, also known as paronomasia, is a form of word play that exploits multiple meanings of a term, or of similar-sounding words, for an intended humorous or rhetorical effect. These ambiguities can arise from the intentional use of homophon ...
exchange near the beginning of the celebrated comedy of circa 411BCE
Lysistrata ''Lysistrata'' ( or ; Attic Greek: , ''Lysistrátē'', "Army Disbander") is an ancient Greek comedy by Aristophanes, originally performed in classical Athens in 411 BC. It is a comic account of a woman's extraordinary mission to end the Peloponne ...
, by the playwright
Aristophanes Aristophanes (; grc, Ἀριστοφάνης, ; c. 446 – c. 386 BC), son of Philippus, of the deme In Ancient Greece, a deme or ( grc, δῆμος, plural: demoi, δημοι) was a suburb or a subdivision of Athens and other city-states ...
:
everal_women_enter,_headed_by_MYRRHINA,_from_the_deme_(=suburb)_of_Anagyrous.html" ;"title="suburb.html" ;"title="everal women enter, headed by MYRRHINA, from the deme (=suburb">everal women enter, headed by MYRRHINA, from the deme (=suburb) of Anagyrous">suburb.html" ;"title="everal women enter, headed by MYRRHINA, from the deme (=suburb">everal women enter, headed by MYRRHINA, from the deme (=suburb) of Anagyrous. Others soon follow.] CALONICE: Hi ! but they're coming now : here they all are : First one,and then another - hoity-toity! Where are this lot from? LYSISTRATA: From Anagyrous. CALONICE: Aha! Well, at any rate we’ve "stirred up Anagyrous" omophonic pun upon the plant name "Anagyris" * * Note: κινεῖν τὸν ἀνάγυρον (kinein tòn anáguron) "To stir up
hake The term hake refers to fish in the: * Family Merlucciidae of northern and southern oceans * Family Phycidae (sometimes considered the subfamily Phycinae in the family Gadidae) of the northern oceans Hake Hake is in the same taxonomic order (Gad ...
the Anagyris" (meaning the nauseous-smelling shrub ''Anagyris foetida'') was a proverb, used of persons who brought some unpleasantness on themselves ompare "Let sleeping dogs lie" Calonice applies the proverb to the imilarly nameddeme
suburb A suburb (more broadly suburban area) is an area within a metropolitan area, which may include commercial and mixed-use, that is primarily a residential area. A suburb can exist either as part of a larger city/urban area or as a separate ...
meaning that the influx of Anagyrasian women proved that the deme uburb of ancient AthensAnagyrous was thoroughly stirred up .e. in uproar/out on the street
Further layers of meaning can be guessed at in the pun: some commentators have inferred that the Athenian suburb of Anagyrus may have been so named because the rank-smelling Anagyris grew plentifully there: others that a playful (possibly traditional) jibe at the inhabitants of Anagyrus may have been intended - to the effect that the Anagyrasians were notably smelly, because their
personal hygiene Hygiene is a series of practices performed to preserve health. According to the World Health Organization (WHO), "Hygiene refers to conditions and practices that help to maintain health and prevent the spread of diseases." Personal hygiene refer ...
was poor. ristophanes_may_here_be_referencing_an_in-joke_that_his_Athenian_audience_would_have_appreciated.html" ;"title="in-joke.html" ;"title="ristophanes may here be referencing an
ristophanes_may_here_be_referencing_an_in-joke_that_his_Athenian_audience_would_have_appreciated">in-joke.html"_;"title="ristophanes_may_here_be_referencing_an_in-joke">ristophanes_may_here_be_referencing_an_in-joke_that_his_Athenian_audience_would_have_appreciated There_are_two_theories_concerning_the_etymology_of_the_suburb_name_Anagyrous_punned_upon_in_Lysistrata_(as_quoted_above):_first_that_it_was_named_for_an_Eponym.html" "title="in-joke">ristophanes may here be referencing an in-joke that his Athenian audience would have appreciated">in-joke.html" ;"title="ristophanes may here be referencing an in-joke">ristophanes may here be referencing an in-joke that his Athenian audience would have appreciated There are two theories concerning the etymology of the suburb name Anagyrous punned upon in Lysistrata (as quoted above): first that it was named for an Eponym">eponymous An eponym is a person, a place, or a thing after whom or which someone or something is, or is believed to be, named. The adjectives which are derived from the word eponym include ''eponymous'' and ''eponymic''. Usage of the word The term ''epon ...
hero Anagyrous, famed for exacting a terrible revenge upon those who had cut branches from the sacred grove dedicated to him, and secondly that it was actually named for the foul-smelling shrub Anagyris that grew there in such profusion. More light might have been cast on the matter by a lost play by Aristophanes entitled Anagyrous and devoted to the deeds of the vengeful hero. However a measure of confusion over the title of a second play possibly devoted to the same hero makes plain the availability to Aristophanes of a second pun at the expense of the Anagyrasians: the playwright
Diphilus Diphilus (Greek: Δίφιλος), of Sinope, was a poet of the new Attic comedy and a contemporary of Menander (342–291 BC). He is frequently listed together with Menander and Philemon, considered the three greatest poets of New Comedy. He wa ...
also wrote a play entitled Anagyrous, but it remains uncertain whether the title was actually Anargyrous - the insertion of a second 'r' changing the meaning to 'without silver' (prefix αν 'without' + ἁργυρυσ 'silver') i.e. 'penniless' or 'impoverished'. Such wordplay could lend itself both to a comic characterisation of the hero Anagyrous as a ne'er-do-well and/or of the inhabitants of the suburb named for him as paupers.Loeb Classics https://www.loebclassics.com/view/aristophanes-attributed_fragments/2008/pb_LCL502.129.xml Retrieved at 23.51 on Thursday 15/12/22.


Gallery


''Anagyris foetida''

File:PikiWiki Israel 83195 mediterranean stinkbush.jpg, Closeup of flower cluster, showing red markings on
standard Standard may refer to: Symbols * Colours, standards and guidons, kinds of military signs * Standard (emblem), a type of a large symbol or emblem used for identification Norms, conventions or requirements * Standard (metrology), an object th ...
attractive to avian pollinators File:Anagyris foetida063.JPG, Closeup of flowers showing stamens and a young, protruding pod File:Anagyris foetidus (15427825069).jpg, Containerised sapling, showing young foliage File:Anagyris.jpg, Mature shrubby specimen festooned with ripening pods File:Fesols moros (Anagyris foetida) de la poblacion de Canèssa Lorcha-l`Orxa Vall de Perputxent.jpg, Low-growing, wild specimen, showing spreading canopy of grey-green foliage File:Anagyris foetida.jpg, Canopy viewed from beneath, showing bark, foliage and ripening pods 4150-Anagyris foetida-Gargano-5.06.JPG, Unripe, paired pods, hanging among proverbially unpleasant-smelling foliage. File:Anagyris foetida pods and seeds.JPG, Ripe pods juxtaposed with ripe, bean-like seeds File:Semillas de fesols moros (Anagyris foetida) Canèssia i Castell de Perputxent Lorcha.jpg, Twigs, showing ash-grey bark and single pod split open to reveal toxic, bean-like seeds File:Anagyris-foetida seeds.jpg, Highly toxic and
emetic Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteri ...
bean-like seeds, rich in the
teratogenic Teratology is the study of abnormalities of physiological development in organisms during their life span. It is a sub-discipline in medical genetics which focuses on the classification of congenital abnormalities in dysmorphology. The related t ...
alkaloid anagyrine (note superficial similarity to olive fruit) File:Anagyris foetida Taub104a.png, Engraving showing anatomical detail from the ''Natürliche Pflanzenfamilien'' of Paul Hermann Wilhelm Taubert (1891)


Avian pollinators

Generalist species A generalist species is able to thrive in a wide variety of environmental conditions and can make use of a variety of different natural resource, resources (for example, a heterotroph with a varied diet (nutrition), diet). A specialist species can ...
whose short beaks increase the likelihood of transporting ''A. foetida'' pollen on their foreheads and throats File:Common Chiffchaff.jpg, Common chiffchaff ''Phylloscopus collybita'' File:Blackcap (Sylvia atricapilla) male.jpg, Eurasian blackcap ''Sylvia atricapilla'' File:Sardinian Warbler.jpg, Sardinian warbler ''Curruca melanocephala'' File:White-spectacled bulbul.jpg, White-spectacled bulbul
''Pycnonotus xanthopygos'' File:Passer Hispaniolensis Male.JPG, Spanish sparrow ''Passer hispaniolensis'' File:Common Whitethroat.jpg, Common whitethroat
''Curruca communis'' File:Sylvia curruca 1 (Martin Mecnarowski).jpg, Lesser whitethroat ''Curruca curruca'' File:Ruppell's warbler.jpg, Rüppell's warbler ''Curruca ruppeli'' File:Eastern Orphean Warbler - Uzbekistan S4E8419 (18675147404)-cropped.jpg, Eastern orphean warbler
''Curruca crassirostris''


Avian nectar thief

File:Nectarinia osea2.jpg, Palestine sunbird ''Cinnyris osea'' showing
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for food ...
specially adapted for feeding on nectar File:Palestine sunbird (Cinnyris osea osea) male.jpg, Palestine sunbird feeding


Insect pollinators

File:Bombus terrestris.jpg, Buff-tailed bumblebee
''Bombus terrestris'' File:Apis mellifera Western honey bee.jpg, Western honey bee ''Apis mellifera''


References

{{Taxonbar, from=Q2630262 foetida Flora of France Flora of Spain Flora of Greece Flora of Malta Poisonous plants Medicinal plants