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The Hawaiian lobelioids are a group of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
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 bel ...
, 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 peren ...
, all of which are
endemic Endemism is the state of a species being found only 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 foun ...
to the
Hawaiian Islands The Hawaiian Islands () are an archipelago of eight major volcanic islands, several atolls, and numerous smaller islets in the Pacific Ocean, North Pacific Ocean, extending some from the Hawaii (island), island of Hawaii in the south to nort ...
. 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 a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...
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 or 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 by their multiple ...
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, e.g., including only woody plants with secondary growth, only ...
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 . Many species have beautiful and spectacular flowers, especially those in ''Lobelia'' and ''Trematolobelia''. 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 som ...
ungulate Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...
s such as
feral pigs A feral pig is a domestic pig which has gone feral, meaning it lives in the wild. The term feral pig has also been applied to wild boars, which can interbreed with domestic pigs. They are found mostly in the Americas and Australia. Razorback a ...
; the stems are only partly woody, and contain few defenses against
herbivory A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
. The bark contains a milky (but apparently non-poisonous)
latex Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latices are found in nature, but synthetic latices are common as well. In nature, latex is found as a wikt:milky, milky fluid, which is present in 10% of all floweri ...
, 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'' (bandicoo ...
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 a plant or plant-like organism that grows on the surface of another 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 epiphyt ...
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 Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
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 () are two barren rock outcrops surrounded by a reef and located in the Northwestern Hawaiian Islands. The Pūhāhonu volcano responsible for the pinnacles is northwest of Honolulu and from French Frigate Shoals. The ...
and
French Frigate Shoals The French Frigate Shoals (Hawaiian language, Hawaiian: Kānemilohai) is the largest atoll in the Northwestern Hawaiian Islands, located about northwest of Honolulu, Hawaii, Honolulu. Its name commemorates France, French explorer Jean-Fran ...
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 In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
. The part of their
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
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 Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
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 (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
by the endemic Hawaiian
hawkmoth The Sphingidae are a family of moths commonly 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, but species ar ...
, ''
Manduca blackburni Lead ''Manduca blackburni'', commonly known as the Hawaiian tomato hornworm, Hawaiian tobacco hornworm, or Blackburn's sphinx moth, is a moth in the family Sphingidae. This rare moth is found in isolated locations on East Maui, Kaho'olawe, and ...
''. 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& ...
''). Despite their inaccessible habitat on cliffs, ''Brighamia'' are sometimes hand-pollinated by botanists to ensure seed set. Both species are now extremely rare in the wild, but ''B. insignis'' enjoys some small scale popularity as a house plant.National Tropical Botanical Garden
/ref> The genus is named in honour of the first director of the Bernice P. Bishop Museum,
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 La ...
. ''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 species of Hawaiian lobelioid in the bellflower family, Campanulaceae. It is native to the islands o ...
''* A.Gray – ''Ōlulu'' (
Kauai Kauai (), anglicized as Kauai ( or ), is one of the main Hawaiian Islands. It has an area of 562.3 square miles (1,456.4 km2), making it the fourth-largest of the islands and the 21st-largest island in the United States. Kauai lies 73 m ...
,
Niihau Niihau (Hawaiian language, Hawaiian: ), anglicized as Niihau ( ), is the seventh largest island in Hawaii and the westernmost of the main islands. It is southwest of Kauai, Kauai across the Channels of the Hawaiian Islands#Kaulakahi Channel, Ka ...
†) *'' Brighamia rockii''* H.St.John – ''Pua Ala'' (
Molokai Molokai or Molokai ( or ; Molokaʻi dialect: Morotaʻi ) is the fifth most populated of the eight major islands that make up the Hawaiian Islands archipelago in the middle of the Pacific Ocean. It is 38 by 10 miles (61 by 16 km) at its g ...
, Lānai†,
Maui Maui (; Hawaiian language, Hawaiian: ) is the second largest island in the Hawaiian archipelago, at 727.2 square miles (1,883 km2). It is the List of islands of the United States by area, 17th-largest in the United States. Maui is one of ...
†) † 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 th ...
. However, many lobelioid genera are derived from it and it is highly
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
. 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 musk ...
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 (, , sometimes written Oahu) is the third-largest and most populated island of the Hawaiian Islands and of the U.S. state of Hawaii. The state capital, Honolulu, is on Oahu's southeast coast. The island of Oahu and the uninhabited Northwe ...
) **'' 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 Wale ...
(
Maui Maui (; Hawaiian language, Hawaiian: ) is the second largest island in the Hawaiian archipelago, at 727.2 square miles (1,883 km2). It is the List of islands of the United States by area, 17th-largest in the United States. Maui is one of ...
,
Molokai Molokai or Molokai ( or ; Molokaʻi dialect: Morotaʻi ) is the fifth most populated of the eight major islands that make up the Hawaiian Islands archipelago in the middle of the Pacific Ocean. It is 38 by 10 miles (61 by 16 km) at its g ...
?) **'' Lobelia kauaensis'' ( A.Gray) A. Heller - ''Pue'' (
Kauai Kauai (), anglicized as Kauai ( or ), is one of the main Hawaiian Islands. It has an area of 562.3 square miles (1,456.4 km2), making it the fourth-largest of the islands and the 21st-largest island in the United States. Kauai lies 73 m ...
) **'' Lobelia villosa'' (Rock) H.St.John & Hosaka (Kauai) *section Revolutella: flowers blue or magenta **'' Lobelia dunbarii'' Rock (Molokai) **'' Lobelia grayana'' F.Wimmer (Maui) **''
Lobelia hillebrandii ''Lobelia'' () is a genus of flowering plants in the family Campanulaceae comprising 415 species, with a subcosmopolitan distribution primarily in tropical to warm temperate regions of the world, a few species extending into cooler temperate r ...
'' Rock (Maui) **'' Lobelia hypoleuca'' Hillebr. – ''Kuhiaikamoowahie'' (Kauai, Oahu, Molokai, Lānai, Maui,
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
) **'' 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 Niiha ...
''* H.St.John (
Niihau Niihau (Hawaiian language, Hawaiian: ), anglicized as Niihau ( ), is the seventh largest island in Hawaii and the westernmost of the main islands. It is southwest of Kauai, Kauai across the Channels of the Hawaiian Islands#Kaulakahi Channel, Ka ...
†, 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''
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 Wale ...
(
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
) *'' Trematolobelia kauaiensis''
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 Wale ...
– ''Kolii'' (
Kauai Kauai (), anglicized as Kauai ( or ), is one of the main Hawaiian Islands. It has an area of 562.3 square miles (1,456.4 km2), making it the fourth-largest of the islands and the 21st-largest island in the United States. Kauai lies 73 m ...
) *'' 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 he ...
&
Arn. George Arnott Walker Arnott of Arlary (6 February 1799 – 17 April 1868) was a Scottish botanist. He collaborated with botanists from around the world and served as a regius professor of botany at the University of Glasgow. An orchid genus ' ...
) A. Zahlbr.
– ''Kolii'' (
Oahu Oahu (, , sometimes written Oahu) is the third-largest and most populated island of the Hawaiian Islands and of the U.S. state of Hawaii. The state capital, Honolulu, is on Oahu's southeast coast. The island of Oahu and the uninhabited Northwe ...
,
Molokai Molokai or Molokai ( or ; Molokaʻi dialect: Morotaʻi ) is the fifth most populated of the eight major islands that make up the Hawaiian Islands archipelago in the middle of the Pacific Ocean. It is 38 by 10 miles (61 by 16 km) at its g ...
†, Lānai†,
Maui Maui (; Hawaiian language, Hawaiian: ) is the second largest island in the Hawaiian archipelago, at 727.2 square miles (1,883 km2). It is the List of islands of the United States by area, 17th-largest in the United States. Maui is one of ...
, 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 CALYX, Inc. is a non-profit publisher of art and literature by women founded in 1976 based in Corvallis, Oregon. CALYX publishes both '' CALYX, A Journal of Art and Literature by Women'' twice a year and CALYX Books, which publishes one to three ...
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, such as ''
Drosophila silvestris ''Drosophila silvestris'' is a large species of fly in the Family (biology), family Drosophilidae that are primarily black with yellow spots. As a rare species of Drosophila, fruit fly endemism in the Hawaiian Islands, endemic to Hawaii (“the B ...
''. 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
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another 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 epiphyt ...
. ''Clermontia'' species *section ''Clermontia'': calyx lobes similar to petals **'' Clermontia calophylla''* F.Wimmer - ''Ōhā wai nui'' ( Kohala,
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
) **'' Clermontia drepanomorpha''*
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 Wale ...
- ''Ōhā wai'' (Kohala, Hawaii) **'' Clermontia grandiflora''† Gaudich. - ''Ōhā wai'' (Molokai, Lānai, Maui) **'' Clermontia hawaiiensis'' (Hillebr.) Rock - ''Ōhā kēpau'' ( Puna and Kaū on Hawaii) **'' Clermontia hanaulaensis''* - ''Ōhā wai'' (west Maui) **'' Clermontia kakeana'' Meyen - ''Ōhā wai'' (Oahu, Molokai, Maui) **'' Clermontia kohalae'' Rock - ''Ōhā wai'' (Kohala and
Hāmākua Hāmākua is a district on the northeast coast of Hawaii's Big Island, administered by the County of Hawaii in the state of Hawaii. It is also the name given for the coastline in the region, the "Hāmākua Coast". Description Hāmākua's coast ...
on Hawaii) **'' Clermontia lindseyana''* Rock - ''Ōhā wai'' (Hawaii, east Maui) **'' Clermontia micrantha'' (Hillebr.) Rock - ''Ōhā wai'' (Lānai, west Maui) **'' Clermontia montis-loa'' Rock - ''Ōhā wai'' (
Hilo Hilo () is the largest settlement in and the county seat of Hawaii County, Hawaiʻi, United States, which encompasses the Island of Hawaiʻi, and is a census-designated place (CDP). The population was 44,186 according to the 2020 census. I ...
, Puna, and Kaū on Hawaii) **'' Clermontia multiflora'' Hillebr. - ''Ōhā wai'' (Oahu†, west Maui†) **'' Clermontia oblongifolia''* Gaudich. - ''Ōhā wai'' (Oahu, Molokai, Lānai, Maui) **'' Clermontia pallida'' Hillebr. - ''Ōhā wai'' (Molokai) **'' Clermontia parviflora'' Gaudich. ex A.Gray - ''Ōhā wai'' (windward Hawaii) **'' Clermontia persicifolia'' Gaudich. - ''Ōhā wai'' (Oahu) **'' Clermontia samuelii''* F.B.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'' H.Lév - ''Hāhāaiakamanu'' (Kauai, Oahu) **''
Clermontia peleana ''Clermontia peleana'' is a rare species of flowering plant in the bellflower family known by the common name Pele clermontia. It is one of several Hawaiian lobelioids in genus ''Clermontia'' that are known as ''`oha wai''. This plant is endemic ...
''* Rock (Hawaii, Maui?) **'' Clermontia pyrularia''* - ''Ōhā wai'' (Hawaii) **'' Clermontia tuberculata''* - ''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 transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
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. In context of humans, it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing open sh ...
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 (island), Hawaii itself, in the Pacific. They were the major herbivores on most of these islands until they became extinct a ...
, giant browsing
geese A goose (: geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera '' Anser'' (grey geese and white geese) and ''Branta'' (black geese). Some members of the Tadorninae subfamily (e.g., Egyp ...
and goose-like
duck Duck is the common name for numerous species of waterfowl in the family (biology), family Anatidae. Ducks are generally smaller and shorter-necked than swans and goose, geese, which are members of the same family. Divided among several subfam ...
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 In geography and seamanship, windward () and leeward () are directions relative to 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 o ...
East Maui where almost no native habitat remains. Because they are particularly sensitive to disturbance by
pigs The pig (''Sus domesticus''), also called swine (: swine) or hog, is an omnivorous, domesticated, even-toed, hoofed mammal. It is named the domestic pig when distinguishing it from other members of the genus '' Sus''. Some authorities cons ...
, ''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
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
(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''
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 Wale ...
(east
Maui Maui (; Hawaiian language, Hawaiian: ) is the second largest island in the Hawaiian archipelago, at 727.2 square miles (1,883 km2). It is the List of islands of the United States by area, 17th-largest in the United States. Maui is one of ...
) *'' Cyanea acuminata''* ( Gaudich.) Hillebr. (
Oahu Oahu (, , sometimes written Oahu) is the third-largest and most populated island of the Hawaiian Islands and of the U.S. state of Hawaii. The state capital, Honolulu, is on Oahu's southeast coast. The island of Oahu and the uninhabited Northwe ...
) *'' Cyanea angustifolia''* (
Cham. Adelbert von Chamisso (; 30 January 1781 – 21 August 1838) was a German poet, writer and botanist. He was commonly known in French as Adelbert de Chamisso (or Chamissot) de Boncourt, a name referring to the family estate at Boncourt. Life ...
) Hillebr.
(Oahu,
Molokai Molokai or Molokai ( or ; Molokaʻi dialect: Morotaʻi ) is the fifth most populated of the eight major islands that make up the Hawaiian Islands archipelago in the middle of the Pacific Ocean. It is 38 by 10 miles (61 by 16 km) at its g ...
, Lānai,
Maui Maui (; Hawaiian language, Hawaiian: ) is the second largest island in the Hawaiian archipelago, at 727.2 square miles (1,883 km2). It is the List of islands of the United States by area, 17th-largest in the United States. Maui is one of ...
) *'' Cyanea arborea'' Hillebr. (east Maui†) *'' Cyanea asarifolia''* H.St.John (
Kauai Kauai (), anglicized as Kauai ( or ), is one of the main Hawaiian Islands. It has an area of 562.3 square miles (1,456.4 km2), making it the fourth-largest of the islands and the 21st-largest island in the United States. Kauai lies 73 m ...
) *'' Cyanea asplenifolia''* (H.Mann) Hillebr. (Maui) *'' Cyanea calycina''* Lammers (Oahu) *'' Cyanea comata'' Hillebr. (east Maui†) *'' Cyanea copelandii''* Rock (east Maui,
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
) *'' Cyanea coriacea'' ( A.Gray) Hillebr. (Kauai) *'' Cyanea crispa''* Gaudich. (Oahu) *'' Cyanea degeneriana'' F. Wimmer (Hawaii) *'' Cyanea dolichopoda'' Lammers & Lorence (Kauai†) *'' Cyanea dunbariae''* Rock - Molokai†) *'' Cyanea eleeleensis'' (H.St.John) Lammers (Kauai†) *'' Cyanea elliptica'' (Rock) Lammers (Lānai, Maui) *'' Cyanea fauriei'' H.Lév (Kauai) *'' Cyanea fissa'' (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 Cyanea may refer to: * Cyanea (jellyfish), ''Cyanea'' (jellyfish), a genus of jellyfish in the family Cyaneidae * Cyanea (plant), ''Cyanea'' (plant), a genus of Hawaiian plants in the family Campanulaceae * Cyanea, a Naiad * ''Cyanea'', a List of ...
'' (H.Mann) Hillebr. (Kauai) *'' Cyanea horrida'' (Rock) O.Deg. & Hosaka - ''Hāhā nui'' (east Maui) *''
Cyanea humboldtiana ''Cyanea humboldtiana'' (formerly ''Rollandia humboldtiana'') is a rare species of flowering plant in the Campanulaceae, bellflower family known by the common name Oʻahu rollandia. It is native to Oahu, Oʻahu, where it is known only from the Ko ...
''* Gaudich. (Oahu) *'' Cyanea kahiliensis'' (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'' Hillebr. (Maui) *'' Cyanea lanceolata'' Gaudich. (Oahu) *'' Cyanea leptostegia'' A.Gray - ''Hāhā lua'' (Kauai) *'' Cyanea linearifolia'' Rock (Kauai†) *'' Cyanea lobata''* H.Mann (Lānai, west 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 ''Cyanea mceldowneyi'' is a rare species of flowering plant in the bellflower family known by the common name McEldowney cyanea. It is endemic to Hawaii, where it is known only from the island of Maui.Cyanea membranacea'' Rock (Oahu) *'' Cyanea obtusa'' (A.Gray) Hillebr. (Maui†) *'' Cyanea parvifolia'' C.N.Forbes (Kauai†) *'' Cyanea pilosa'' A.Gray (Hawaii) *''
Cyanea pinnatifida ''Cyanea pinnatifida'' is a rare species of flowering plant in the Campanulaceae, bellflower family known by the common name sharktail cyanea. It is Endemism, endemic to Oahu, but it is now extinct in the wild and only exists in cultivation.Brueg ...
''* (Cham.) F.Wimmer (Oahu) *'' Cyanea platyphylla''* (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'' (Hillebr.) F.Wimmer (Hawaii†) *'' Cyanea quercifolia'' (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'' Hillebr. - ''Pua kala'' (Molokai) *'' Cyanea spathulata'' (Hillebr.) A.Heller (Kauai) *'' Cyanea st.-johnii''* Hosaka (Oahu) *'' Cyanea stictophylla''* Rock (Kona and Kaū on Hawaii) *'' Cyanea superba''* (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†) *'' Cyanea leptostegia''* A.Gray - hanalau (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 of Germany, capital and largest city of Germany, by both area and List of cities in Germany by population, population. With 3.7 million inhabitants, it has the List of cities in the European Union by population withi ...
during
World War II World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
. 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'' (E.Wimm.) H.St.John **'' Delissea fallax'' Hillebr. (Hawaii†) **'' Delissea kauaiensis'' (Lammers) Lammers (Kauai) **'' Delissea lauliiana'' Lammers (Oahu†) **''
Delissea parviflora 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 th ...
'' 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 ''Delissea subcordata'' is a rare species of flowering plant in the family Campanulaceae known by the common names Koʻolau Range delissea and ''oha''. It is endemic to Hawaii, where it is known only from the island of Oahu. It is now only found ...
''† 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 t ...
Lists of flora of Hawaii . Natural history of Hawaii Trees of Hawaii Flora without expected TNC conservation status