The Hawaiian lobelioids are a group 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 bellflower family,

Campanulaceae The family Campanulaceae (also bellflower family), of the order Asterales, contains nearly 2400 species in 84 genera of herbaceous plants, shrubs, and rarely small trees, often with milky sap. Among them are several familiar garden plants belon ...

, subfamily

Lobelioideae Lobelioideae is a subfamily of the plant family Campanulaceae. It contains 32 genera, totalling about 1200 species. Some of the larger genera are ''Lobelia'', ''Siphocampylus'', ''Centropogon'', ''Burmeistera'' and '' Cyanea''. They are perenni ...

, all of which are

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 ...

to the

Hawaiian Islands The Hawaiian Islands ( haw, Nā Mokupuni o Hawai‘i) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the North Pacific Ocean, extending some from the island of Hawaii in the south to northernmost Kur ...

. This is the largest plant

radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes: * ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...

in the Hawaiian Islands, and indeed the largest on any island archipelago, with over 125 species. The six genera involved can be broadly separated based on growth habit: ''Clermontia'' are typically branched

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...

s or small

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

s, up to tall, with fleshy fruits; ''Cyanea'' and ''Delissea'' are typically unbranched or branching only at the base, with a cluster of relatively broad leaves at the apex and fleshy fruits; ''Lobelia'' and ''Trematolobelia'' have long thin leaves down a single, non-woody stem and capsular fruits with wind-dispersed seeds; and the peculiar ''Brighamia'' have a short, thick stem with a dense cluster of broad leaves, elongate white flowers, and capsular fruits. The relationships among the genera and sections remains unsettled . Lobelia niihauensis (5762180787)

Many species have beautiful and spectacular flowers, especially those in ''Lobelia'' and ''Trematolobelia''. Unfortunately, they are also highly vulnerable to feeding by

feral A feral () animal or plant is one that lives in the wild but is descended from domesticated individuals. As with an introduced species, the introduction of feral animals or plants to non-native regions may disrupt ecosystems and has, in some ...

ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...

s such as

feral pigs The feral pig is a domestic pig which has gone feral, meaning it lives in the wild. They are found mostly in the Americas and Australia. Razorback and wild hog are Americanisms applied to feral pigs or boar-pig hybrids. Definition A feral pi ...

; the stems are only partly woody, and contain few defenses against

herbivory A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

. The bark contains a milky (but apparently non-poisonous)

latex Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latexes are found in nature, but synthetic latexes are common as well. In nature, latex is found as a milky fluid found in 10% of all flowering plants (angiosperms ...

, and is often chewed by

rat Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' ( pack rats), ''Bandicota'' (bandicoot ...

s and pigs. Seedlings are also vulnerable to disturbance by pig digging, and in areas with high densities of pigs it is not uncommon to find the only lobelioids being

epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...

on larger trees or on fallen logs.

Taxonomy

The group contains morphologically divergent species, and was long thought to have derived from at least three introductions: one for ''Lobelia'' and ''Trematolobelia'', one for ''Brighamia'', and one for ''Clermontia'', ''Cyanea'', and ''Delissea''. Based on DNA sequence evidence, it was later suggested that all are derived from a single introduction. This was likely a ''Lobelia''-like species that arrived about 13 million years ago, when

Gardner Pinnacles The Gardner Pinnacles ( haw, Pūhāhonu) are two barren rock outcrops surrounded by a reef and located in the Northwestern Hawaiian Islands at . The volcano is northwest of Honolulu and from French Frigate Shoals. The total area of the two ...

and

French Frigate Shoals The French Frigate Shoals ( Hawaiian: Kānemilohai) is the largest atoll in the Northwestern Hawaiian Islands. Its name commemorates French explorer Jean-François de La Pérouse, who nearly lost two frigates when attempting to navigate the ...

were high islands and long before the current main islands existed. A

molecular phylogenetic 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 ...

study published in 2021 found that ''Delissea'' plus ''Brighamia'', ''Trematolobelia'', ''Lobelia'' sect. ''Galeatella'', and ''Lobelia'' sect. ''Revolutella'' formed a

clade 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, ...

that was closely related to giant African and South American lobelioids. ''Clermontia'' formed a separate clade with some species of ''Cyanea'', which was not

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

. The part of their

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

that includes genera and sections from Hawaii is shown below, with added shading to show those treated as Hawaiian lobelioids, which do not form a single clade in this study. The authors of the 2021 study described the taxonomy of several genera, including ''Lobelia'', as "particularly frustrating" and called for further research.

''Brighamia''

''Brighamia'' is quite unlike the other genera, with a

succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...

stem and long, thin, tubular

flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...

s. It was long thought to have been the result of a separate introduction, and its unique combination of characters made it difficult to place. These characters are the result of adaptation to growing on cliffs and

pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...

by the endemic Hawaiian

hawkmoth The Sphingidae are a family of moths ( Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450 species. It is best represented in the tropics, b ...

, ''

Manduca blackburni ''Manduca blackburni'', the Hawaiian tomato hornworm, Hawaiian tobacco hornworm or Blackburn's sphinx moth, is a moth in the family Sphingidae. The species was first described by Arthur Gardiner Butler in 1880. It is endemic to Hawaii. Previousl ...

''. This moth is now itself listed as endangered, surviving mainly on the southern slopes of Maui, well away from where ''Brighamia'' live. Some pollination may be done by closely related alien hawkmoths such as the five-spotted hawkmoth ('' M. quinquemaculata'') and pink-spotted hawkmoth (''

Agrius cingulata ''Agrius cingulata'', the pink-spotted hawkmoth or sweetpotato hornworm, is a moth in the family Sphingidae. The species was first described by Johan Christian Fabricius in 1775. Description The imago has a wingspan of to inches (9.5–12&n ...

''). Despite their inaccessible habitat on cliffs, ''Brighamia'' are sometimes hand-pollinated by botanists to ensure seed set. Both species are now extremely rare. The genus is named in honour of the first director of the

Bernice P. Bishop Museum The Bernice Pauahi Bishop Museum, designated the Hawaii State Museum of Natural and Cultural History, is a museum of history and science in the historic Kalihi district of Honolulu on the Hawaiian island of Oʻahu. Founded in 1889, it is the larg ...

William Tufts Brigham William Tufts Brigham (1841–1926) was an American geologist, botanist, ethnologist and the first director of the Bernice P. Bishop Museum in Honolulu. Biography William Tufts Brigham was born on May 24, 1841. After finishing the Boston Lati ...

. ''Brighamia'' species *''

Brighamia insignis ''Brighamia insignis'', commonly known as ''ʻŌlulu'' or ''Alula'' in Hawaiian, or colloquially as the vulcan palm or cabbage on a stick, is a endangered species of Hawaiian lobelioid in the bellflower family, Campanulaceae. It is native to t ...

''*

A.Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His '' Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually exc ...

– ''Ōlulu'' (

Kauai Kauai, () anglicized as Kauai ( ), is geologically the second-oldest of the main Hawaiian Islands (after Niʻihau). With an area of 562.3 square miles (1,456.4 km2), it is the fourth-largest of these islands and the 21st largest island ...

Niihau Niihau ( Hawaiian: ), anglicized as Niihau ( ), is the westernmost main and seventh largest inhabited island in Hawaii. It is southwest of Kauaʻi across the Kaulakahi Channel. Its area is . Several intermittent playa lakes provide wetland hab ...

†) *''

Brighamia rockii The Hawaiian lobelioids are a group of flowering plants in the bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are endemic to the Hawaiian Islands. This is the largest plant radiation in the Hawaiian Islands, and indeed the ...

''* H.St.John – ''Pua Ala'' (

Molokai Molokai , or Molokai (), is the fifth most populated of the eight major islands that make up the Hawaiian Islands, Hawaiian Islands archipelago in the middle of the Pacific Ocean. It is 38 by 10 miles (61 by 16 km) at its greatest length an ...

, Lānai†,

Maui The island of Maui (; Hawaiian: ) is the second-largest of the islands of the state of Hawaii at 727.2 square miles (1,883 km2) and is the 17th largest island in the United States. Maui is the largest of Maui County's four islands, which ...

†) † species believed to be extinct
* species is listed as endangered

''Lobelia''

''Lobelia'' is a cosmopolitan genus of over 350 species, including common

ornamentals Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...

. However, many lobelioid genera are derived from it and it is highly

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

. The Hawaiian species are divided into two sections (''Galeatella'', the giant lobelias of montane

bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg; a ...

s, and ''Revolutella'', the smaller lobelias of rocky crests and interior rock walls), based on flower color and other characters. Like ''Brighamia'' and ''Trematolobelia'', the fruit of ''Lobelia'' is a dry capsule. These species are probably the closest in appearance to the original Hawaiian colonist. ''Lobelia'' species *section Galeatella: flowers red or yellow to white **'' Lobelia gaudichaudii''* A.DC (

Oahu Oahu () (Hawaiian language, Hawaiian: ''Oʻahu'' ()), also known as "The Gathering place#Island of Oʻahu as The Gathering Place, Gathering Place", is the third-largest of the Hawaiian Islands. It is home to roughly one million people—over t ...

) **'' Lobelia gloria-montis''

Rock Rock most often refers to: * Rock (geology), a naturally occurring solid aggregate of minerals or mineraloids * Rock music, a genre of popular music Rock or Rocks may also refer to: Places United Kingdom * Rock, Caerphilly, a location in Wales ...

(

?) **'' Lobelia kauaensis'' (

) A. Heller - ''Pue'' (

) **'' Lobelia villosa'' (Rock) H.St.John & Hosaka (Kauai) *section Revolutella: flowers blue or magenta **''

Lobelia dunbarii ''Lobelia'' () is a genus of flowering plants comprising 415 species, with a subcosmopolitan distribution primarily in tropical to warm temperate regions of the world, a few species extending into cooler temperate regions.Huxley, A., ed. (1992 ...

'' Rock (Molokai) **''

Lobelia grayana ''Lobelia'' () is a genus of flowering plants comprising 415 species, with a subcosmopolitan distribution primarily in tropical to warm temperate regions of the world, a few species extending into cooler temperate regions.Huxley, A., ed. (1992 ...

'' F.Wimmer (Maui) **'' Lobelia hillebrandii'' Rock (Maui) **'' Lobelia hypoleuca'' Hillebr. – ''Kuhiaikamoowahie'' (Kauai, Oahu, Molokai, Lānai, Maui,

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 ...

) **'' Lobelia monostachya''* (Rock) Lammers (Oahu) **''

Lobelia niihauensis ''Lobelia niihauensis'', commonly known as the Niihau lobelia, is a rare species of flowering plant in the bellflower family that is endemic to Hawaii. It is known only from the islands of Oahu and Kauai and is thought to be extirpated from Nii ...

''* H.St.John (

†, Kauai, Oahu) **'' Lobelia oahuensis''* Rock (Oahu) **'' Lobelia remyi'' Rock (Oahu†) **'' Lobelia yuccoides'' Hillebr. - ''Pānaunau'' (Kauai, Oahu) † species believed to be extinct
* species is listed Endangered

''Trematolobelia''

''Trematolobelia'' is distinguished from ''Lobelia'' by its unique dispersal method. Rather than drying and splitting apart, the outer (green) wall of the fruit disintegrates, revealing a perforated hard "frame" that allows the tiny wind-dispersed seeds to escape. They can be quite spectacular when in flower, with multiple flower branches and hundreds of flowers. Individual plants live for 5–10 years before flowering and dying. ''Trematobelia'' species *'' Trematolobelia grandifolia''

(

) *'' Trematolobelia kauaiensis''

– ''Kolii'' (

) *'' Trematolobelia macrostachys'' (

Hook. Sir William Jackson Hooker (6 July 178512 August 1865) was an English botanist and botanical illustrator, who became the first director of Kew when in 1841 it was recommended to be placed under state ownership as a botanic garden. At Kew h ...

Arn. George Arnott Walker Arnott of Arlary (6 February 1799 – 17 April 1868) was a Scottish botanist. Early life George Arnott Walker Arnott was born in Edinburgh in 1799, the son of David Walker Arnott of Arlary. He attended Milnathort Parish ...

) A. Zahlbr. – ''Kolii'' (

†, Lānai†,

, Hawaii†) *'' Trematolobelia singularis''* H.St.John (Oahu) † species believed to be extinct
* species is listed Endangered

''Clermontia''

''Clermontia'', with 22 species, are the most common of Hawaiian lobelioids. Unlike ''Cyanea'', which are typically found in dense forest, ''Clermontia'' are frequently found in more open areas and edges, and therefore persist better when forests become fragmented. Nevertheless, there are still many endangered species. The flowers are often large and spectacular; in section ''Clermontia'', the calyx lobes are similar in color and size to the corolla, giving the appearance of a flower with twice the normal number of petals. ''Clermontia'' is a very important host plant for many species of

Hawaiian Drosophilidae The Hawaiian Drosophilidae are a lineage of flies within the genus ''Drosophila.'' This monophyletic clade includes all of the endemic Hawaiian ''Drosophila'' and all members of the genus '' Scaptomyza'', which contains both Hawaiian and non-Hawa ...

, such as ''

Drosophila silvestris ''Drosophila silvestris'' is a large species of fly in the family Drosophilidae that are primarily black with yellow spots. As a rare species of fruit fly endemic to Hawaii (“the Big Island”), the fly often experiences reproductive isolatio ...

''. The larvae of Drosophilidae flies breed in the rotting bark, leaves, flowers, and fruit of all lobelioids, but primarily ''Clermontia'' since it is largest and most common. Several species of these native plants, especially on the Big Island, are

''Clermontia'' species *section ''Clermontia'': calyx lobes similar to petals **''

Clermontia calophylla The Hawaiian lobelioids are a group of flowering plants in the bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are endemic to the Hawaiian Islands. This is the largest plant radiation in the Hawaiian Islands, and indeed the ...

''* F.Wimmer - ''Ōhā wai nui'' ( Kohala,

) **''

Clermontia drepanomorpha ''Clermontia drepanomorpha'' is a rare species of flowering plant in the Campanulaceae, bellflower family known by the common name Kohala Mountain clermontia. It is one of several Hawaiian lobelioids in genus ''Clermontia'' that are known as ''`o ...

''*

- ''Ōhā wai'' (Kohala, Hawaii) **'' Clermontia grandiflora'' Gaudich. - ''Ōhā wai'' (Molokai, Lānai, Maui) **'' Clermontia hawaiiensis'' (Hillebr.) Rock - ''Ōhā kēpau'' ( Puna and Kaū on Hawaii) **'' Clermontia kakeana'' Meyen - ''Ōhā wai'' (Oahu, Molokai, Maui) **'' Clermontia kohalae'' Rock - ''Ōhā wai'' (Kohala and Hāmākua on Hawaii) **''

Clermontia lindseyana ''Clermontia lindseyana'' is a rare species of flowering plant in the bellflower family known by the common name hillside clermontia. It is one of several Hawaiian lobelioids in genus ''Clermontia'' that are known as ''`oha wai''. This plant is ...

''* Rock - ''Ōhā wai'' (Hawaii, east Maui) **'' Clermontia micrantha'' (Hillebr.) Rock - ''Ōhā wai'' (Lānai, west Maui) **''

Clermontia montis-loa ''Clermontia montis-loa'', or Mauna Loa clermontia, is a species of Hawaiian lobelioid endemic to the eastern windward slopes of Mauna Loa on the island of Hawai'i. Like other Clermontia, it is referred to as ʻŌhā wai in Hawaiian. Descripti ...

'' Rock - ''Ōhā wai'' (

Hilo Hilo () is a census-designated place (CDP) and the largest settlement in Hawaii County, Hawaii, Hawaii County, Hawaii, United States, which encompasses the Hawaii (island), Island of Hawaii. The population was 44,186 according to the 2020 United ...

, Puna, and Kaū on Hawaii) **'' Clermontia multiflora'' Hillebr. - ''Ōhā wai'' (Oahu†, west Maui†) **'' Clermontia oblongifolia''* Gaudich. - ''Ōhā wai'' (Oahu, Molokai, Lānai, Maui) **''

Clermontia pallida The Hawaiian lobelioids are a group of flowering plants in the bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are endemic to the Hawaiian Islands. This is the largest plant radiation in the Hawaiian Islands, and indeed the ...

'' Hillebr. - ''Ōhā wai'' (Molokai) **'' Clermontia parviflora'' Gaudich. ex A.Gray - ''Ōhā wai'' (windward Hawaii) **'' Clermontia persicifolia'' Gaudich. - ''Ōhā wai'' (Oahu) **''

Clermontia samuelii ''Clermontia samuelii'' is a rare species of flowering plant in the bellflower family known by the common name Hana clermontia. It is one of several Hawaiian lobelioids in genus ''Clermontia'' that are known as ''`oha wai''. This plant is endemi ...

''* C.N.Forbes - ''Ōhā wai'' (east Maui) *section ''Clermontioideae'': calyx lobes short, green **'' Clermontia arborescens'' (H.Mann) Hillebr. - ''Ōhā wai nui'' (Molokai, Lānai, Maui) **'' Clermontia clermontioides'' (Gaudich.) A.Heller (Kaū and Kona Districts on Hawaii) **''

Clermontia fauriei The Hawaiian lobelioids are a group of flowering plants in the bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are endemic to the Hawaiian Islands. This is the largest plant radiation in the Hawaiian Islands, and indeed the ...

'' H.Lév - ''Hāhāaiakamanu'' (Kauai, Oahu) **'' Clermontia peleana''* Rock (Hawaii, Maui?) **''

Clermontia pyrularia ''Clermontia pyrularia'' is a rare species of flowering plant in the bellflower family known by the common names Hamakua clermontia and pear clermontia. It is one of several Hawaiian lobelioids in genus ''Clermontia'' that are known as ''oha wa ...

''* - ''Ōhā wai'' (Hawaii) **''

Clermontia tuberculata The Hawaiian lobelioids are a group of flowering plants in the Campanula, bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are Endemism, endemic to the Hawaiian Islands. This is the largest plant Adaptive radiation, radiation ...

''* - ''Ōhā wai'' (Maui) **'' Clermontia waimeae''* - ''Ōhā wai'' (Hawaii) † species believed to be extinct
* species is listed Endangered

''Cyanea''

'' Cyanea'' is the largest and most morphologically diverse group of Hawaiian lobelioids, with more than 70 species. Most grow as a single stem or as a cluster branching near the ground, but a few, such as ''C. stictophylla'', grow as multi-branched shrubs. Some, such as ''C. leptostegia'' of Kauai, can grow to over tall - something that is especially notable given the relative thinness of the stem and soft wood. Part of the reason ''Cyanea'' are able to grow tall stalks is that they tend to grow in deep forest, often in narrow gulches on the older islands, where there is little wind. This characteristic of growing under dense cover also makes them more sensitive to disturbance of the forest. An interesting character of many ''Cyanea'' is their tendency to grow spines or thorns on the stem and leaves (see the photo of ''Cyanea platyphylla''). This is most pronounced in younger plants, and some species undergo a kind of

metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...

as they mature, to the extent that different growth stages were described as separate species, due in part to the presence or absence of spines. The purpose of the spines was puzzling, since in most island situations there is a tendency for plants to lose defenses - Hawaii is noted for its nettle-less nettles, mintless mints, and (not quite) thornless raspberries - and no native

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 ...

animals were known. However, it is now believed that the spines were a defense against the

moa-nalo The moa-nalo are a group of extinct aberrant, goose-like ducks that lived on the larger Hawaiian Islands, except Hawaii itself, in the Pacific. They were the major herbivores on most of these islands until they became extinct after human settle ...

, giant browsing

geese A goose (plural, : geese) is a bird of any of several waterfowl species in the family (biology), family Anatidae. This group comprises the genera ''Anser (bird), Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some o ...

and goose-like

duck Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...

s that formerly inhabited the islands (Givnish et al. 1995). These birds were apparently driven extinct by the Hawaiians before Europeans reached the islands, but their evolutionary effects live on. Many species are now extinct or have not been seen in decades. These include ''C. arborea'', ''C. comata'', and ''C. pohaku'', a cluster of species that formerly inhabited the drier, mesic areas of

leeward Windward () and leeward () are terms used to describe the direction of the wind. Windward is ''upwind'' from the point of reference, i.e. towards the direction from which the wind is coming; leeward is ''downwind'' from the point of reference ...

East Maui where almost no native habitat remains. Because they are particularly sensitive to disturbance by

pigs The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus s ...

, ''Cyanea'' are often the first plants to disappear, even when the forest as a whole appears relatively healthy. Extinct species tended to have longer, more highly specialized flowers and to have narrower elevational and geographic ranges than the species that survived (Givnish et al. 1995). Species of ''Cyanea'' on each major island tend to differ in flower tube length and mean elevation, apparently reflecting a partitioning of ecological and reproductive resources. The total number of species of ''Cyanea'' can be predicted rather precisely from the height and area of each of the major islands except

(the Big Island), suggesting that the assembly of ''Cyanea'' communities requires more than 0.6 million years (the age of Hawaii) and less than 1.5 million years (the age of Maui) to run to ecological saturation (Givnish et al. 2008). ''Cyanea'' species *''

Cyanea aculeatiflora Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

(east

) *''

Cyanea acuminata ''Cyanea acuminata'' is a rare species of flowering plant known by the common names Honolulu cyanea. It is endemic to Oahu, where there are no more than 250 individuals remaining.Bruegmann, M. M. & V. Caraway. (2003)''Cyanea acuminata'' IUCN Red ...

''* ( Gaudich.) Hillebr. (

) *''

Cyanea angustifolia ''Cyanea angustifolia'' is a plant in the genus '' Cyanea'' that is found in Hawaii.Cyanea angustifolia ...

''* (

Cham. Adelbert von Chamisso (; 30 January 178121 August 1838) was a Germans, German poet and botanist, author of ''Peter Schlemihl'', a famous story about a man who sold his shadow. He was commonly known in French as Adelbert de Chamisso (or Chamissot ...

) Hillebr. (Oahu,

, Lānai,

) *''

Cyanea arborea Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' Hillebr. (east Maui†) *''

Cyanea asarifolia ''Cyanea asarifolia'' is a rare species of flowering plant in the bellflower family known by the common name gingerleaf cyanea. It is endemic to Kauai, where there were no more than 30 individuals in a single population as of 2005.USFWS''Cyanea ...

''* H.St.John (

) *'' Cyanea asplenifolia''* (H.Mann) Hillebr. (Maui) *'' Cyanea calycina''* Lammers (Oahu) *'' Cyanea comata'' Hillebr. (east Maui†) *''

Cyanea copelandii ''Cyanea copelandii'' is a rare species of flowering plant in the bellflower family known by the common name treetrunk cyanea. It is endemic to Maui, where there are no more than 250 individuals remaining in the wild. It is a federally listed en ...

''* Rock (east Maui,

) *''

Cyanea coriacea Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' (

) Hillebr. (Kauai) *''

Cyanea crispa ''Cyanea crispa'' is a rare species of flowering plant known by the common names crimped rollandia and Koolau Range rollandia. It is endemic to Oahu, where there are no more than fifty individuals remaining in the Koʻolau Range.Bruegmann, M. M. ...

''* Gaudich. (Oahu) *'' Cyanea degeneriana'' F. Wimmer (Hawaii) *''

Cyanea dolichopoda ''Cyanea dolichopoda'' was a species of shrub in the bellflower family that was endemic to Kauai. It was discovered in 1990 and has not been located in the wild since 1992. Like other '' Cyanea'' it is known as ''haha'' in Hawaiian. This Hawa ...

'' Lammers & Lorence (Kauai†) *'' Cyanea dunbariae''* Rock - Molokai†) *''

Cyanea eleeleensis ''Cyanea eleeleensis'' was a rare species of flowering plant in the bellflower family known by the common name Eleele cyanea. It was endemic to Kauai, where it has been declared extinct.USFWSDetermination of endangered status for 48 species on K ...

'' (H.St.John) Lammers (Kauai†) *'' Cyanea elliptica'' (Rock) Lammers (Lānai, Maui) *''

Cyanea fauriei Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' H.Lév (Kauai) *''

Cyanea fissa Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' (H.Mann) Hillebr. (Kauai) *'' Cyanea giffardii'' Rock (Hawaii†) *'' Cyanea glabra''* (F.Wimmer) H.St.John (Maui†) *'' Cyanea grimesiana''* Gaudich. (Oahu, Molokai, Lānai, Maui, Hawaii†) *'' Cyanea habenata'' (H.St.John) Lammers (Kauai) *'' Cyanea hamatiflora''* Rock (east Maui, Hawaii) *'' Cyanea hardyi'' Rock (Kauai) *'' Cyanea hirtella'' (H.Mann) Hillebr. (Kauai) *'' Cyanea horrida'' (Rock) O.Deg. & Hosaka - ''Hāhā nui'' (east Maui) *'' Cyanea humboldtiana''* Gaudich. (Oahu) *''

Cyanea kahiliensis Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' (H.St.John) Lammers (Kauai) *'' Cyanea kolekoleensis'' (H.St.John) Lammers (Kauai) *'' Cyanea koolauensis''* Lammers Givnish & Sytsma (new name for ''Rollandia angustifolia'') (Oahu) *''

Cyanea kunthiana Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' Hillebr. (Maui) *''

Cyanea lanceolata Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' Gaudich. (Oahu) *''

Cyanea leptostegia Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' A.Gray - ''Hāhā lua'' (Kauai) *''

Cyanea linearifolia Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' Rock (Kauai†) *''

Cyanea lobata ''Cyanea lobata'' is a rare species of flowering plant in the bellflower family known by the common name Waihee Valley cyanea. It is endemic to Hawaii, where it is known from Lanai and Maui.Cyanea longissima'' (Rock) H.St.John (east Maui†) *'' Cyanea longiflora''* Wawra (Oahu) *'' Cyanea macrostegia''* Hillebr. (Lānai, Maui) *'' Cyanea mannii''* ( Brigham) Hillebr. (Molokai) *'' Cyanea marksii'' Rock ( Kona, Hawaii) *'' Cyanea mceldowneyi''* Rock (east Maui) *'' Cyanea membranacea'' Rock (Oahu) *''

Cyanea obtusa Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' (A.Gray) Hillebr. (Maui†) *''

Cyanea parvifolia Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' C.N.Forbes (Kauai†) *''

Cyanea pilosa Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' A.Gray (Hawaii) *''

Cyanea pinnatifida ''Cyanea pinnatifida'' is a rare species of flowering plant in the bellflower family known by the common name sharktail cyanea. It is endemic to Oahu, but it is now extinct in the wild and only exists in cultivation.Bruegmann, M. M. & V. Caraway ...

''* (Cham.) F.Wimmer (Oahu) *''

Cyanea platyphylla ''Cyanea platyphylla'' is a rare species of flowering plant in the Campanulaceae, bellflower family known by the common names puna cyanea and flatleaf cyanea. It is Endemism, endemic to the island of Hawaii (island), Hawaii, where there are fewer ...

''* (A.Gray) Hillebr. - ''Akūakū'' (Hawaii) *'' Cyanea pohaku'' Lammers (east Maui†) *'' Cyanea procera''* Hillebr. (Molokai†) *'' Cyanea profuga'' C.N.Forbes (Molokai†) *'' Cyanea purpurellifolia'' Rock (Oahu) *''

Cyanea pycnocarpa Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' (Hillebr.) F.Wimmer (Hawaii†) *''

Cyanea quercifolia ''Cyanea quercifolia'', known as oakleaf cyanea, was a species of plant native to the Hawaiian island of Maui. The plant is now considered extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a ...

'' (Hillebr.) F.Wimmer (east Maui†) *'' Cyanea recta''* (Wawra) Hillebr. (Kauai†) *'' Cyanea rivularis''* (Rock) (Kauai†) *'' Cyanea scabra'' Hillebr. (west Maui) *'' Cyanea shipmanii''* Rock (Hawaii) *'' Cyanea solenacea'' Hillebr. - ''Pōpolo'' (Molokai) *''

Cyanea solenocalyx Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' Hillebr. - ''Pua kala'' (Molokai) *''

Cyanea spathulata Cyanea may refer to: * ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * An ancient Greek name that means "blue sky" and in the Greek alphabet it' ...

'' (Hillebr.) A.Heller (Kauai) *'' Cyanea st.-johnii''* Hosaka (Oahu) *''

Cyanea stictophylla ''Cyanea stictophylla'' is a rare species of flowering plant in the bellflower family known by the common name Kaiholena cyanea. It is endemic to the island of Hawaii, where it is known only from the rainforests of Mauna Loa.World Conservation M ...

''* Rock (Kona and Kaū on Hawaii) *''

Cyanea superba ''Cyanea superba'' is a rare species of flowering plant in the bellflower family known by the common names Mt. Kaala cyanea and superb cyanea. It is endemic to the island of Oahu, but it is now extinct in the wild. It exists in cultivation and s ...

''* (Cham.) A.Gray (Oahu) *'' Cyanea sylvestris'' A.Heller (Kauai) *'' Cyanea tritomantha'' A.Gray - ''Akūakū'' (Hawaii) *'' Cyanea truncata''* (Rock) Rock (Oahu) *'' Cyanea undulata''* C.N.Forbes (Kauai†) † species believed to be extinct
* species is listed Endangered

''Delissea''

''Delissea'' is similar to ''Cyanea'' in many ways, differing primarily in the flower (with a small knob on the dorsal side) and fruit (dark purple; most ''Cyanea'' fruit are orange, though some are also purple or blue). It is notable in part because it has suffered so much: only three of the nine species known to science are still extant, and one of these (''D. undulata'') is extinct in the wild. Several species are known only from type specimens collected in the late 1800s. Several species are very poorly known, and their status as species is questionable. For example, ''D. fallax'' and ''D. parviflora'' are both from Hawaii and their flowers are identical; it is possible that they represent different growth forms of the same species (both ''Delissea'' and ''Cyanea'' are known to undergo changes in vegetative morphology during the lifetime of the plant). ''Delissea lauliiana'' was known only from the type, which was destroyed in

Berlin Berlin ( , ) is the capital and largest city of Germany by both area and population. Its 3.7 million inhabitants make it the European Union's most populous city, according to population within city limits. One of Germany's sixteen constitue ...

during

World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...

. All three of these are believed to be extinct, and unless new specimens turn up there is no way to resolve questions about them. ''Delissea'' species *section Delissea: nearly straight flowers long **''

Delissea argutidentata ''Delissea argutidentata'' is a species of flowering plant in the Hawaiian lobelioids section of the Campanulaceae family, native to Hawaii (island). The species was long thought to be extinct, but in March 2021 three individuals were discovered ...

'' (E.Wimm.) H.St.John **'' Delissea fallax'' Hillebr. (Hawaii†) **''

Delissea kauaiensis ''Delissea kauaiensis'' is a critically endangered species of the bellflower family. It is found on Mt. Ha'upu near Hanapepe Hanapepe or Hanapēpē is a historic, unincorporated community in Kauai County, Hawaii, United States. The name means ...

'' (Lammers) Lammers (Kauai) **'' Delissea lauliiana'' Lammers (Oahu†) **'' Delissea parviflora'' Hillebr. (Hawaii†) **'' Delissea rhytidosperma''* H.Mann (Kauai) **'' Delissea undulata''* Gaudich. (Niihau†, Kauai†, Maui†, Hawaii) *section Macranthae: curved flowers long **'' Delissea laciniata'' Hillebr (Oahu†) **'' Delissea sinuata'' Hillebr. (Oahu†, Lānai†) **'' Delissea subcordata''† Gaudich. (Oahu) **'' Delissea waianaeensis''* Lammers (Oahu) † species believed to be extinct
* species is listed Endangered

References

* :The standard reference for Hawaiian plant taxonomy. * {{DEFAULTSORT:Hawaiian Lobelioids

Hawaiian lobelioids The Hawaiian lobelioids are a group of flowering plants in the bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are endemic to the Hawaiian Islands. This is the largest plant radiation in the Hawaiian Islands, and indeed the ...

Lists of flora of Hawaii . Natural history of Hawaii Trees of Hawaii Flora without expected TNC conservation status