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A possible explanation is as follows.Most original colonizer plants were brought to Hawai'i by water or birds which are associated with water in some way. That would mean they would first be deposited along the coast. As such plants became established, some forms would spread into neighboring areas, somewhat less harsh than the coast, such as the dry woodlands, and the moister, higher elevations more inland. The plants would eventually evolve more tree-like forms to match the more densely vegetated areas.
Below are two such examples.A. 'Akoko Chamaesyce (once in genus Euphorbia).
1. This genus of plants have many members which are worldwide, weedy, and persistent colonizers.
a. Possibly two different species arrived in Hawai'i at different times. New species developed as they spread from primarily coastal sites to the interior of the islands.
b. The trend in life form was from herbaceous weed, to shrub, and to tree forms in the rain forest. The tendency to change from herb to woody forms is called ARBORESCENCE or, literally, "becoming a tree". There are 12 plant families in Hawai'i, that are only woody or near woody, which are elsewhere in the world represented primarily by non-woody forms. Refer to the following discussion under arborescence.
2. Chamaesyce species from coastal to inland environments.
The species described below demonstrate the rends just described.
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a.C. degeneri is found primarily on the coast. Its leaves are small, round and succulent. Stems are never woody and form mats of growth. Seeds disperse readily by flotation.
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b. C. celastroides may have arrived separately (from C. degeneri) by seeds attachED to bird wings by a sticky film. It develops a low branching growth by central upright shoot which dies quickly, allowing lateral stems to take over, providing a mat-like growth adapted to dry conditions. Leaves are small and paperishy with powdery coating that protects from water loss. Leaves are readily shed in dry, hot weather and the succulent stems store needed water.
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c. C. celastroides var. loriflora is similar in appearance to the one above, but has leaves much longer, 2-4 inches, which correlates with the moister, slightly more inland location where it grows. It is slightly more upright and shrub-like and less mat-forming in growth habit.
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d. C. celastroides var. mauiensis, another variety, is definitely more upright and woody in growth form. This is achieved by some lateral shoots, growing upward rather than sideways as in the lower-growing forms discussed above.
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e. C. remyi is found in Kaua'i's rain forest. Here the leaves are strikingly different -- shiny, leathery, evergreen and larger -- which are adaptations to the moister environment. Smooth leaf surfaces discourages fungal growth. The dark green color means more chlorophyll to collect light in dim conditions. Larger leaves give more area for photosynthesis without danger of excessive water loss. This plant sprawls over other shrubs and trees in a upward direction, not having a true tree form because it can't self-support, but it appears developing in that direction.
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D. Ultimate arboresence comes with the rain forest. C. rockii grows in the dampest, rainiest high ridges of the Ko'olau. It is the tallest of 'akokos, having a trunk 20-26 feet high, with a 5-6" diameter, and 6 inch long leaves. Fruits are largest of all the species with a 1" diameter. This species lacks the sticky mucilage of the coastal species.
B. Another example of makai to mauka is the Naupaka (Scaevola), with several coastal species and many inland species. Like 'akoko, it apparently was founded by two separate colonists.
![]() Scaevola sericeaNaupaka Kahakai- - Coastal |
![]() Scaevola gaudichaudii Naupaka Kuahiwi - - dry ridges, open shrubland and forest |
![]() Scaevola gaudichaudiana Naupaka Kuahiwi - - Wet forest and open green areas |
![]() Scaevola glabra 'Ohe Naupaka - - Very wet forest |
A. Arborescence (woodiness) is found is such Hawaiian natives as the violets (Viola), 'Aweoweo(Chenopodium), Papala (Charpentiera) and Lobelioids which have the distinctive rosette growth pattern.
B. Why the trend to woodiness in the Hawaiian native flora?
1. The mild climate stimulates year-around growth
2. Weedy plants were favored as colonists but woody forms can compete best in forests and shrublands for light.
C. Tree violets, such as Viola robusta, is an upright unbranched small tree-form up to two feet high. Elsewhere in the world, violets are small herbaceous plants.
D. The native lobelioids and silverswords are large shrub-tree forms coming from plants as small herbs elsewhere.
E. Rosette growth:Many herbs (non-woody plants) have very short stems and the leaves are at ground level in a rosette or whorl.
1. A number of tree lobelioids have a rosette or whorl of leaves at the top of a long stem. Investigators have found that plantsnormally producing rosette growth, when exposed to year round favorable growing conditions, sometimes start producing a longer stem with a whorl of leaves at the top.
2. This suggests one possible way tree lobelioids may have developed from annual rosette type growth.
Look here for representations of Hawaiian lobelioids and the type of plant from which they came.
F. Geranium genus, not the horticultural Pelargonium genus commonly called geraniums, have a number of shrub-like endemic species in Hawaii.
G. Chenopodium species, 'aweoweo (often called Goosefoot on the mainland, where it is a common weed) forms large semi-woody shrubs, the largest in the world for this genus, and Charpentiera, Papala, related to beets and amaranth (a garden flower and weed) forms a small tree-like growth.
'Aweoweo Papala
A. Gigantism is a feature seen especially in plants producing large fruits and seeds. Gigantism is found in many island environments and in animals too, such as the larger Galapagos Tortoises and the large Hawaiian dragonfly.
B. Ho'awa (Pittosporum) and Loulu, Hawaiian Fan Palm (Pritchardia), both have much larger fruits than other non-native species.
![]() Ho'awa Photo C. Lamoureux |
![]() Loulu Photo by P. Millen |
A.Ko'oko'olau or Spanish Needle (Bidens). The single-seeded fruits of Hawaiian native species often become larger (less dispersible), lose the barbs with hooks at the tip, and/or become curled.
In picture below, Bidens fruit on the far left is an non-native species and displays characteristics for successful dispersibility: slender, pointed ends and barbs on the top extensions, all which can anchor it into hair, fur or feathers. The native three endemic species fruits on the right, however, are broader or twisted, with little structure to hook into something.
Bidens Seeds (Photo Gerald Carr)
B. Seeds of native Wiliwili (Erythrina sandwicensis) no longer can float in sea water (likely dispersal means of dispersal to Hawai'i), unlike other non-native species.
C. One advantage of loss of dispersibility as seen in the large-seeded forest plants like Ho'awa(Pittosporum), is that the larger seeds of a forest tree or plant help to ensure survival. The abundance of stored food in the seed "fuels" the seedling's growth towards the stronger light higher up. There is great competition in the forest for light, everything else being relatively abundant (like water, favorable temperatures, and good soil).
A. Loss of Protective Chemicals.
1. Chemical poisons or strong-smelling/tasting chemicals are usually key in protecting plants from predators such as insects or grazing animals like cattle and goats. Native plants are represented by endemic species that lack that these protections. Because of lack of dispersal of many of predators to Hawaii, the native plants developed in this "safe" environment without protection.
2. Hawaiian mints, of which there are a number of genera, lack the characteristic mint odor. The chemicals associated with the odor are known to be an insect repellent, even though we find "minty" odors and flavors pleasing.
3. The genus Rhus (sumac) , neleau are known for its potent skin rash effect from the non-native species of poison ivy and oak. On the other hand, Neleau(Rhus sandwicensis) is a harmless Hawaiian native species without any chemical skin irritant.
4. The decline of the native palm, loulu ( Pritchardia), may well be due to a large extent, to the Polynesian rat that devours its tasty seeds which apparently have no chemical deterrent.
5. The native plants which are related to the nettles (Family Urticaceae, especially genus Urera, or Opuhe, with two endemic species), lack the stinging cells that protect nettles so well elsewhere in the world.
B. Loss of Physical Protective Features:
Neleau Photo by G. Carr
Loulu Photo P. Millen
1. Thorns, prickles and spines are absent from many native plants.
a. The Hawaiian raspberry, 'Akala(Rubus hawaiensis), is a thornless raspberry, unlike most other related species world-wide. This is because of the absence of large, grazing predators.
b. The Hawaiian poppy, Pua kala ( Argemone glauca), still has prickles. This may be explained by its status as a relative "newcomer" to the islands - maybe in the last two thousand years. The theory of its relatively recent arrival is supported by its unusual biennial growth habit, living only 2 years, whereas most native flora are perennial.
2. Loss of deep-rootedness which protects against the trampling of large herbivores:
a. Many natives are shallow-rooted. They are very vulnerable to the walking around of hoofed animals which destroys their root systems. When cattle are released on lands with native species, both the grazing and trampling kills them.
Shallow roots result in wind damage (left) and cattle damage (right). Forests are reduced to pasture in many area where cattle graze. Photos M. Bauer.
b. A'ali'i, or Dodonaea viscosa, is one member of the native flora which is less imcted by the presence of hoofed animals because of its deep roots. It is clearly the exception.
A. Adaptive radiation is a pattern of evolutionary development in which many different forms or species develop from a single founder organism. This is demonstrated in many Hawaiian groups such as Naupaka and Chamaesyce as well as in the Silversword Alliance.
B. The Silversword alliance is a group of closely related species (28) belonging to 3 different genera: Ili'au(Wilkesia), Na'ena'e(Dibautia) and="../../audio/ahinahina.aiff">'Ahinahina
or the Silverswords (Argyroxiphium).
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1. The genus Wilkesia is
composed of the two of the oldest species of the alliance and is found only
on Kaua'i. It is the most ancient of the three genera which is reflected
by its location on the oldest existing island. (Photo by Dave Boynton). |
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2. Dubautia with 21 species,
has many species on the Big Island as well as on other islands. The species
can be shrubs, semi-shrubs and even a vine. It is the youngest genus found
in greatest diversity on the youngest island. Dubautia
is still in process of active speciation, a young genus, unlike Wilkesia,
which is considered a relict genus (one which is a remnant). (Photo by Priscilla Millen) |
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3. World famous Argyroxiphium genus has five species members, called green and silver swords 'Ahinahina, for the Haleakala species. Many form large rosette (a plant-growth form where all the leaves grow close together in a big whorl usually at the ground level or at the end of a long stem). They are found from the alpine deserts to the bogs of Hawai'i. (Photo by Charles Lamoureux) |
C. Through the work of Dr. Gerald Carr at University of Hawaii and Dr. Donald Kyhos of University of California at Davis, there is evidence that all the silversword alliance members evolved from a single seed arriving about 5 million years ago. Their original colonist source is related to the 99 species of "tarweeds" found in California and Mexican deserts today. The adaptive radiation of these Hawaiian species in the Silversword Alliance from a single colonist plant is quite a spectacular evolutionary event.
1. The proposed ancestors of the colonist plant are found in North America in the dry areas of California and Mexico. They are shrubby, sprawling plants, not as striking as the silverswords but have important structural similarities. They are called "tarweeds" because of sticky substances found on the flowers and fruits. You can guess what way they probably got to Hawai'i! They hitched a ride on a bird feathers!
Tar Weed
Silversword
The relationship between the Hawaiian silversword alliance and these "tarweeds" was determined by "screening" the present-day tarweeds, mostly by genetic tests, to find closely-related descendants.
a. When two were found that seemed to be likely relatives, they were crossed and, produced healthy hybrids with their Hawaiian "cousins." This means that they still have a great deal of genetic material in common and are not that "distantly" related, though they are far apart in space and in time -- by hundreds of miles and millions of years!
b. A California tarweed called Raillardiopsis muirii, found in the Southern Sierra Mountains on sandy soils of a granite outcrops, was crossed with Dubautia laevigata,, from the wet forests of Kaua'i. A hybrid formed between these two unlikely plants and was dubbed the "Transoceanic Hybrid."
c. The creation of this artificial hybrid supports the genetic closeness of the two groups of plants found on opposite sides of the Pacific. Chromosomes sets from two different parents can form a successful hybrid only when they are closely related. Thus chromosome compatibility can tell us with a great deal of certainty that these two plants must be related. We do not have to be standing on the shore million of years ago to watch the first colonist arrive to come to this conclusion.
d. Another important outcome of this study was to inform the scientific world that long-distance dispersal is more significant than previously suspected.
2. Why is there much diversity in the Hawaiian silverswords and less in the California tarweeds?This can be answered in part by the fact that relatively few different kinds of plants (disharmonic flora concept) were able to establish in Hawai'i. This means that the ones that did had far greater opportunities, with all the available habitats in which to live. Many different forms developed in response to the different habitats, the specific locations where they live.
3 In further studies, it has been shown that all three genera can hybridize and a number do in the wild where their habitats overlap. In the crater of Haleakala, the silverswords and a DUBAUTIA species form hybrids in certain intermediate locations.
(See Sohmer and Gustafson,pp. 34-35.)
This means that all three genera of the Silversword alliance share the same gene pool, thus making the pool larger than if each species is restricted to its own genus, the usual case. This clearly allows for greater opportunity to survive because of the greater genetic diversity within the alliance.
D. The silversword alliance story fits the theories of dispersal and colonist characteristics in at least five major ways:
1. Dispersal from North America
2. Adaptive radiation influenced by the varied life zones of Hawai'i.
3. Survival with small gene pool by forming hybrid "bridges" between genera which can allow gene flow between groups.
4. From weedy annual or low growing semi-shrubs to massive rosettes (like the Silversword) and other larger woody plants.
5. Bird dispersal by sticky fruits.
For further information on adaptive radiation in the silversword alliance, visit Dr. Gerald Carr's Web site.
E. The Silversword has many features which allow it to live successfully in the harsh environment of alpine desert of Haleakala.
1. The long thin curved leaves allow for sufficient surface for light absorption without excessive water loss. The gray hairs on the leaves are even curved in such a way as to re-radiate the excessive light and heat of the sun away from the plant. It is called 'Ahinahina by the Hawaiians, which means very gray (gray-gray).
2. The spaces between leaf cells are filled with a gelatinous substance which helps to prevent water loss and, perhaps, to protect from extreme cold, also part of the alpine desert environment. The Silversword can also absorb great amounts of water, especially when the large inflorescence is being formed.
3. The plants live about 5 to 15 years and bloom only once, in July. The mature plant can be six feet high with a massive inflorescence covered with maroon flowers. Goats once threatened silverswords with extinction. While goats have been fenced out of Haleakala, now the Argentine ants threaten by their predation on the silversword pollinator.
A. Large colorful red-purple twisted flowers are bird-pollinated and small white inconspicuous ones are pollinated by small crawling beetles and other insects. In Hawai'i there are few native bees and only one native butterfly.
B. Many native plants have small, inconspicuous flowers:
1. There are few native bees to Hawai'i and only one native butterfly. Few large pollinators, such as social honey bees, larger butterflies and moths, made it to the islands for obvious reasons.
2. As a result, a number Hawaiian flowers are small, colored greenish/white, and shaped flattish or cupped slightly since the pollinators are small insects and beetles, which are not generally attracted by bright colors or large flowers as are bees and butterflies.
C. Naupaka species are adapted to different pollinators.
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1. In the Scaevola genus (Naupaka) the familiar white, half-flower is found in many of the lowland species which reflect pollination by small insects.
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2. On the other hand, a flower of a rain forest species, 'ohe naupaka (Scaevola glabra) has curved tubular yellow flower which is clearly bird-pollinated.
D. Bird-pollinated flowers found in the native flora are usually bright colored, larger, and frequently with a curved corolla (the structure made up of all the petals).
1. Some good examples are the native Koki'o(Hibiscus), a related genus such as hau kuahiwi
(Hibiscadelphus) and 'ohi'a (Metrosideros), all which have flowers of bright colors or produce copious amounts of nectar. The HONEY CREEPER native birds are important pollinators of many native plants.
Koki'o ula Hau kuahiwi 'Ohi'a lehua
>2. The many extinctions and very low numbers of the honey creepers today threaten the survival of the many native plants that depend on their pollinating activities.
A. Why are Hawaiian native plants not as competitive as many introduced plants?
1. They are narrowly adapted to very specific climatic conditions which exist in small regions on the islands.
2. Originally they had little competition for space because competition was low in the developing flora. That is related to few new plants arriving over a long time.
3. Competitive plants develop where there is a lot of disturbance in the environment and are often found where humans have been intensively active for thousands of years. Hawai'i was free from humans for millions of years!
B. What is the impact of introduction of non-native plants?
1. Introduction of "weedy", non-native plants from other parts of the world has greatly impacted native plants. Wherever a disturbance is created in the Hawaiian environment -- a road, clearing for farming, or rooting by wild pigs -- the introduced plants often can move in and become established more quickly than the native.
2. For example, Banana poka (Passiflora mollissima) doesn't even wait for an opening, but grows right over the natives, smothering them! A more recent threat is the ivy or scarlet-fruited gourd (Coccinia grandis), introduced in the mid '80's and already a serious vining pest on many islands. The Christmas berry tree (Schinus terebinthifolius) arches over and shades out native plants. Strawberry guavas (Psidium cattleianum) grow so thickly that nothing else can grow between them.
Banana Poka
C. Exceptions.
1. Some native Hawaiian plants are adapted to harsh conditions such as the coastal zone and recent lava flows, and are able to compete fairly successfully with introduced plants.
2. Uluhe fern (Dicranopteris) has adapted well to road cuts and other human-made disturbances, but its original adaptation is to landslide areas, a natural phenomenon of the Hawaiian environment.
3. The tree fern, 'Ama'u, (Sadleria), an endemic genus, and 'ohi'a lehua (Metrosideros polymorpha) are other native plants which can aggressively colonize new lava flows.
![]() Uluhe Photo by R. Saito |
![]() 'Ama'u Photo by E. Petteys |
A. In conclusion, the adaptations of colonist plants to their new island home did not prepare them for the future, when humans physically altered the landscape and brought introduced plants and animals from which native plants lacked protection. Furthermore, it would probably take thousands of years for them to adapt to the altered, human-impacted environment. A massive change has occurred in a few hundred years and the native plants don't have the luxury of time.
B. As a result, the Hawaiian flora (along with much of the fauna) is one of the most endangered in the world.
1. There have been more animal and plant species extinction in the Hawaiian Islands than in all of the rest of the United States.
2. As a result, Hawaii - with 0.2% of land area in the United States - has almost 50% of endangered plant species for the country
3. If conditions are not reversed, another significant percentage of native plants is predicted to become extinct in the next 20 years.
4. Dr. Gary Ray, representing the Center for Plant Conservation, an organization dedicated to preserving endangered plants, has found that there are 120 taxa (groups of plants at the species and subspecies level) with 20 or fewer individuals left in the wild. He estimates it would take a million dollars just to save them from immediate extinction.
5. Reasons given for saving native plants and animals often stress the practical side of their value and what's "in it" for humans. Unknown chemical properties of medicinal value, genetic content that could become a lifeline to humans by saving crop or other valuable plants, their stabilizing role in the ecosystem, and even their inherent beauty or appeal, have all been given as "good reasons" for saving any kind of plant or animal.
Maybe we should look at the problem from an ethical and moral stance. Do we need to justify their value only in terms of their use to us? What is it about us humans, an integral part of the web of life, which permits us to take over much of the livable surface of the planet to the exclusion and destruction of numerous plant and animal forms, many of which have been on the planet much longer than we? In respect and in valuing all forms of life, I hope that we can create safe and sustainable havens for many of the special plant and animal communities around us. Certainly we can co-exist! To do less would be less than human.
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