genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s in the heliotrope family,
Heliotropiaceae Heliotropiaceae is a cosmopolitan family of flowering plants with approximately 450 species worldwide, though it is concentrated especially in the tropics and subtropics.
There are the following four genera in the family:
* '' Heliotropium (''incl ...
. There are around 325 species in this almost
cosmopolitan
Cosmopolitan may refer to:
Food and drink
* Cosmopolitan (cocktail), also known as a "Cosmo"
History
* Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953
Hotels and resorts
* Cosmopoli ...
genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Etymology
The name "heliotrope" derives from the old idea that the
inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
s of these plants turned their rows of flowers to the Sun.Chittenden, Fred J. Ed., Royal Horticultural Society Dictionary of Gardening, Oxford 1951 Ἥλιος (''helios'') is
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
for "Sun", τρέπειν (''trepein'') means "to turn". The
Middle English
Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English p ...
name "turnsole" has the same meaning.
A Classical myth, told in
Ovid
Pūblius Ovidius Nāsō (; 20 March 43 BC – 17/18 AD), known in English as Ovid ( ), was a Roman poet who lived during the reign of Augustus. He was a contemporary of the older Virgil and Horace, with whom he is often ranked as one of the th ...
's ''
Metamorphoses
The ''Metamorphoses'' ( la, Metamorphōsēs, from grc, μεταμορφώσεις: "Transformations") is a Latin narrative poem from 8 CE by the Roman poet Ovid. It is considered his ''magnum opus''. The poem chronicles the history of the wo ...
'', imagines that the water nymph
Clytie
In Greek mythology, the name Clytie (Ancient Greek: Κλυτίη, Ionic) or Clytia (, Attic and other dialects) may refer to:
*Clytie (Oceanid), known for her unrequited love for Helios. Out of jealousy, Clytie arranged the death of Leucotho ...
, in love with the sun god
Helios
In ancient Greek religion and Greek mythology, mythology, Helios (; grc, , , Sun; Homeric Greek: ) is the deity, god and personification of the Sun (Solar deity). His name is also Latinized as Helius, and he is often given the epithets Hyper ...
, was betrayed by him. Wasting away, she transformed into the heliotrope, whose flowers supposedly always face the Sun.
Morphology
Like other members of the Heliotropiaceae, plants in the genus ''Heliotropium'' have 5-merous,
tetracyclic
Tetracyclics are cyclic chemical compounds that contain four interconnected rings of atoms, e.g. Tröger's base.
They have various pharmaceutical
A medication (also called medicament, medicine, pharmaceutical drug, medicinal drug or s ...
flowers and
actinomorphic
Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts.
Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
corollas. They likewise share in their characteristic terminal styles and highly modified stigmatic heads (basal stigma, infertile apex). Species in the genus are typically herbs or subshrubs exclusively and are characterized by their dry fruits that divide into two or four mericarpids.
Ecology and human use
Several heliotropes are popular garden plants, most notably
garden heliotrope
A garden is a planned space, usually outdoors, set aside for the cultivation, display, and enjoyment of plants and other forms of nature. The single feature identifying even the wildest wild garden is ''control''. The garden can incorporate bot ...
(''H. arborescens''). Some species are weeds, and many are
hepatotoxic
Hepatotoxicity (from ''hepatic toxicity'') implies chemical-driven liver damage. Drug-induced liver injury is a cause of acute and chronic liver disease caused specifically by medications and the most common reason for a drug to be withdrawn fro ...
if eaten in large quantities due to abundant pyrrolizidine alkaloids. There have been cases of canine death due to over-ingestion of this toxic plant. Some danaine
butterflies
Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
, such as male queen butterflies, visit these plants, being attracted to their pyrrolizidine alkaloids. Though it is not palatable and most animals will completely ignore it, there have been cases of horses, swine and cattle being poisoned due to contamination of hay.
Caterpillar
Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths).
As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s of the
grass jewel
''Freyeria trochylus'', the grass jewel, is a small butterfly found in Africa, Arabia (United Arab Emirates, Oman, Saudi Arabia), southern Europe (Bulgaria and Greece), India and southern Asia that belongs to the lycaenids or blues family.
Des ...
food coloring
Food coloring, or color additive, is any dye, pigment, or substance that imparts color when it is added to food or drink. They come in many forms consisting of liquids, powders, gels, and pastes. Food coloring is used in both commercial food ...
in
Middle Ages
In the history of Europe, the Middle Ages or medieval period lasted approximately from the late 5th to the late 15th centuries, similar to the post-classical period of global history. It began with the fall of the Western Roman Empire a ...
French cuisine
French cuisine () is the cooking traditions and practices from France. It has been influenced over the centuries by the many surrounding cultures of Spain, Italy, Switzerland, Germany and Belgium, in addition to the food traditions of the re ...
.
One of the most famous
ragtime
Ragtime, also spelled rag-time or rag time, is a musical style that flourished from the 1890s to 1910s. Its cardinal trait is its syncopated or "ragged" rhythm. Ragtime was popularized during the early 20th century by composers such as Scott ...
piano
The piano is a stringed keyboard instrument in which the strings are struck by wooden hammers that are coated with a softer material (modern hammers are covered with dense wool felt; some early pianos used leather). It is played using a keyboa ...
Louis Chauvin
Louis Chauvin (March 13, 1881March 26, 1908) was an American ragtime pianist and composer.
Early life and education
Born in St. Louis, Missouri, to a Mexican Spanish-Indian father and an African-American mother, he widely was considered the fines ...
(the first two strains) and Scott Joplin (the last two strains).
Garden heliotrope is grown in Southern Europe as an ingredient for
perfume
Perfume (, ; french: parfum) is a mixture of fragrant essential oils or aroma compounds (fragrances), fixatives and solvents, usually in liquid form, used to give the human body, animals, food, objects, and living-spaces an agreeable scent. Th ...
.
The purplish facial rash of
dermatomyositis
Dermatomyositis (DM) is a long-term inflammatory disorder which affects skin and the muscles. Its symptoms are generally a skin rash and worsening muscle weakness over time. These may occur suddenly or develop over months. Other symptoms may inc ...
is called "heliotrope rash" because it resembles ''E. arborescens''.
Heliotrine and heliotridine
Seeds of the ''Heliotropium'' genus were discovered in the 1940s and 50s to be responsible for liver disease in populations that consumed them in large quantities, either inadvertently (as a contaminant of food crops) or deliberately (associated with the ingestion of herbal infusions for the treatment of certain ailments). The seeds contained high concentrations of
pyrrolizidine alkaloids
Pyrrolizidine alkaloids (PAs), sometimes referred to as necine bases, are a group of naturally occurring alkaloids based on the structure of pyrrolizidine. Pyrrolizidine alkaloids are produced by plants as a defense mechanism against insect he ...
, identified mainly as the N-oxide of heliotrine (74%), and one or two other compounds similar in character to lasiocarpine.(Dubrovinskii, 1947, 1952; Khanin, 1948; Bras et al., 1954, 1961; Bras & Hill, 1956; cited in World Health Organization (1988) PYRROLIZIDINE ALKALOIDS INTERNATIONAL PROGRAMME ON CHEMICAL SAFETY, ENVIRONMENTAL HEALTH CRITERIA 80.
Taxonomy
Taxonomic revision supported through
molecular phylogenetics
Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
led to the recognition of ''
Euploca
''Euploca'' is an almost cosmopolitan genus of plants with around 100 species. It was first described by Thomas Nuttall in 1837. While part of the broadly defined Boraginaceae in the APG IV system from 2016, a revision of the order Boraginales ...
'' as genus separate from ''Heliotropium''. In contrast, the genus ''
Tournefortia
''Tournefortia'', commonly known as soldierbush, is a genus of flowering plants in the borage family, Boraginaceae.
It was first published under the name ''Pittonia'' by Charles Plumier in 1703, in honour of Joseph Pitton de Tournefort. Later, ...
'' was included in ''Heliotropium'' in a 2016 revision.
Within ''Heliotropium'', there are four major
clades
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, t ...
:
*''Heliotropium'' sect. ''Heliothamnus''
*Old World ''Heliotropium'' clade
*''Heliotropium'' sect. ''Cochranea''
*''Tournefortia'' clade, comprising ''Tournefortia'' sect. ''Tournefortia'' and all remaining New World ''Heliotropium species''
Origins of diversification
Three of the four major clades within ''Heliotropium'' have their centers of diversity in South America. The origins of the remaining
Old World
The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
''Heliotropium'' clade can be traced back to a single colonization event from the
New World
The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. 3 ...
.
ITS1
Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript.
I ...
data shows there is a single characteristic long deletion between positions 61 and 111 in the genome of the Old World species, which defines the Old World ''Heliotropium'' species and separates them from their New World counterparts. Researchers concluded this is a single autapomorphic character from a single deletion event in the past. This most reasonably explains how the whole group may have come to share this characteristic deletion when comparing the genomes of Old World and New World ''Heliotropium''.
The most likely driver of ''Heliotropium'' diversification across the three New World clades is early Andean uplift. Researchers identified three independent diversification events in the phylogeny of Andean ''Heliotropium'', whose timings correspond to late
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
Andean uplift as well as the development of arid environments in South America during the
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58
''Heliothamnus'' diversification is estimated to have taken place in the late Miocene. The age of ''Heliothamnus'' suggest that its diversification could have been triggered directly by the uplift of the Andes, something that would have promoted speciation in inner-Andean valleys and the Andean scrub. The majority of
endemic
Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
''Heliothamnus''
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
in the region are restricted to these sorts of environments, further supporting this theory as the current leading theory explaining ''Heliothamnus'' diversification.
Before the main rise of the Andes, ''Cochranea'' and ''Tournefortia'' coinhabited the Andean region at the same time and significant
speciation
Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
had not yet occurred. Once the Andes began to rise, ''Cochranea'' became isolated on the western side of the Andes while ''Tournefortia'' grew on the eastern side. This east-west division is still true of each group’s present distributions. The rise of the Andes affected the climate of the region and is believed to have contributed to the hyperaridity of the
Atacama Desert
The Atacama Desert ( es, Desierto de Atacama) is a desert plateau in South America covering a 1,600 km (990 mi) strip of land on the Pacific coast, west of the Andes Mountains. The Atacama Desert is the driest nonpolar desert in th ...
, something that could have acted as an additional barrier to filter out other ''Heliotropium'' species into the range of ''Cochranea'', thus promoting ''Cochranea'' speciation. Elevation differences would have also acted as barriers that helped promote speciation in ''Tournefortia'' species as many large groups within ''Tournefortia'' became well-adapted to high-elevation environments while other ''Heliotropium'' clades did not and thus could not coinhabit the same environments as ''Tournefortia''.
Heliotropium anomalum
''Heliotropium anomalum'' is a species of flowering shrub in the borage family, Boraginaceae, that is native to the Hawaiian Islands, Guam, Christmas Island, Saipan, Tinian, Wake Island and New Caledonia. Common names include Polynesian heliotrop ...
Hawaii
Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
)
* ''
Heliotropium arborescens
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...
Benth.
George Bentham (22 September 1800 – 10 September 1884) was an English botanist, described by the weed botanist Duane Isely as "the premier systematic botanist of the nineteenth century". Born into a distinguished family, he initially studie ...
Heliotropium chenopodiaceum
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...
Heliotropium curassavicum
''Heliotropium curassavicum'', commonly called salt heliotrope (among other names), a species of flowering plant in the borage family (Boraginaceae). It is native to much of the Americas, from Canada to Argentina, including the West Indies and Ha ...
'' L. – seaside heliotrope, salt heliotrope, monkey tail, quail plant, Chinese parsley; ''cola de mico'' (
Spanish
Spanish might refer to:
* Items from or related to Spain:
**Spaniards are a nation and ethnic group indigenous to Spain
**Spanish language, spoken in Spain and many Latin American countries
**Spanish cuisine
Other places
* Spanish, Ontario, Can ...
Heliotropium epacrideum
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...
'' F.Muell. ex Benth.
* ''
Heliotropium europaeum
''Heliotropium europaeum'' is a species of heliotrope known by the common names European heliotrope and European turn-sole. It is native to Europe, Asia, and North Africa, but it is widely naturalized elsewhere, such as in Australia and North Am ...
'' L. – European heliotrope, European turnsole (Europe, Asia, and North Africa)
* '' Heliotropium fasciculatum'' R.Br.
* ''
Heliotropium flintii
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...
'' F.Muell. ex A.S.Mitch.
* ''
Heliotropium foertherianum
''Heliotropium arboreum'' is a species of flowering plant in the borage family, Boraginaceae. It is native to tropical Asia including southern China, Madagascar, northern Australia, and most of the atolls and high islands of Micronesia and Polyn ...
'' Diane & Hilger – tree heliotrope, velvet soldierbush, octopus bush (
South Asia
South Asia is the southern subregion of Asia, which is defined in both geographical and ethno-cultural terms. The region consists of the countries of Afghanistan, Bangladesh, Bhutan, India, Maldives, Nepal, Pakistan, and Sri Lanka.;;;;;;;; ...
,
East Asia
East Asia is the eastern region of Asia, which is defined in both geographical and ethno-cultural terms. The modern states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. China, North Korea, South Korea and ...
,
Melanesia
Melanesia (, ) is a subregion of Oceania in the southwestern Pacific Ocean. It extends from Indonesia's New Guinea in the west to Fiji in the east, and includes the Arafura Sea.
The region includes the four independent countries of Fiji, Va ...
, western
Polynesia
Polynesia () "many" and νῆσος () "island"), to, Polinisia; mi, Porinihia; haw, Polenekia; fj, Polinisia; sm, Polenisia; rar, Porinetia; ty, Pōrīnetia; tvl, Polenisia; tkl, Polenihia (, ) is a subregion of Oceania, made up of ...
, northern
Australia
Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
Heliotropium glabellum
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...
Heliotropium megalanthumn
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...
Heliotropium pannifolium
''Heliotropium pannifolium'', the Saint Helena heliotrope, is now extinct but was formerly a hairy-leaved small shrub up to 1 m in height. it was only seen once, by the explorer W. Burchell in Broad Gut, Saint Helena (ca. 1808) and has never be ...
'' – St. Helena heliotrope (
Saint Helena
Saint Helena () is a British overseas territory located in the South Atlantic Ocean. It is a remote volcanic tropical island west of the coast of south-western Africa, and east of Rio de Janeiro in South America. It is one of three constitu ...
) (
extinct
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
Heliotropium paulayanum
''Heliotropium paulayanum'' is a species of plant in the family Boraginaceae. It is Endemism, endemic to Yemen. Its natural habitat is subtropical or tropical dry shrubland.
References
Endemic flora of Socotra
Heliotropium, paulayanum
V ...
Heliotropium popovii
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...
Heliotropium shoabense
''Heliotropium shoabense'' is a species of plant in the family Boraginaceae. It is endemic to Yemen
Yemen (; ar, ٱلْيَمَن, al-Yaman), officially the Republic of Yemen,, ) is a country in Western Asia. It is situated on the souther ...
Heliotropium strigosum
''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats.
Et ...