The Kahana Valley Ahupua'a, a PABITRA Study Site on O'ahu ...

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293 The Kahana Valley Ahupua‘a, a PABITRA Study Site on O‘ahu, Hawaiian Islands 1 Dieter Mueller-Dombois 2 and Nengah Wirawan 3 Abstract: The acronym PABITRA stands for Pacific-Asia Biodiversity Tran- sect, a network of island sites and conservation professionals collaborating throughout the Pacific-Asia region. An ideal PABITRA site is a broad landscape transect from sea to summit. Such a landscape is Kahana Valley on Windward O‘ahu. Kahana Valley served during prior centuries as an ahupua‘a, a Polynesian unit of land management that integrated the three biological resource zones, the upland forests, the agriculturally used land below, and the coastal zone, into a sustainable human support system. Results of terrestrial biodiversity surveys, as begun with a vegetation/environment study and a paleoecological investigation, are presented in relation to historical land use and sea level changes. In spite of the many former human-induced modifications of the Kahana Valley landscape, the natural structure and function of its ecosystems are well preserved. The dis- tribution patterns of vegetation can be interpreted in terms of Hawaiian ecolog- ical zones in combination with the valley’s precipitation, topography, stream system, and archaeological features. Currently, efforts are under way to restore the Kahana State Park (recently renamed Ahupua‘a ‘O Kahana State Park) as a functional ahupua‘a. In addition, focused collaborative research can yield helpful information for further restoration and integrated management of the Kahana ahupua‘a as a historic Hawaiian Heritage Site. The PABITRA (Pacific-Asia Biodiversity Transect) network aims to include broad landscape units with three interconnected biodiversity resource zones into a study pro- gram. The three resource zones are an upland or inland forest system that serves as water- shed cover, the agroecosystems that are usu- ally below in elevation, and the coastal zone ecosystems. A freshwater resource system is another essential component. The objective is to study each of these eco- system zones individually for their internal functions but also to investigate their rela- tionships as a landscape unit. The latter is necessary to ensure that the management of one resource zone does not affect the other zones negatively and that the zonation system is managed as an integrated landscape unit. Such is the management unit or human sup- port system developed (through trial and er- ror) by the early Hawaiians before contact with Western society and named ahupua‘a. The Ahupua‘a Model An ahupua‘a is an island landscape represent- ing a vertical section from mountain to sea. It usually encompasses a watershed with the area to the sea as deep as a person can stand in the ocean water. Ahupua‘a is a Hawaiian term. It refers to ahu, meaning a heap of stones, and pua‘a, the Hawaiian word for pig. A related, second meaning of the term ahu- pua‘a refers to an altar on which a pig was placed as a tribute to the chief or landowner for allowing the use of the ahupua‘a land. The landlord or owner and manager of an Pacific Science (2005), vol. 59, no. 2:293–314 : 2005 by University of Hawai‘i Press All rights reserved 1 Manuscript accepted 20 April 2004. 2 Department of Botany, University of Hawai‘i at Ma ¯noa, 3190 Maile Way, Honolulu, Hawai‘i 96822. 3 Deceased, former Director, World Wide Fund, Indonesia.

Transcript of The Kahana Valley Ahupua'a, a PABITRA Study Site on O'ahu ...

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293

The Kahana Valley Ahupua‘a, a PABITRA Study Site on O‘ahu,Hawaiian Islands1

Dieter Mueller-Dombois2 and Nengah Wirawan3

Abstract: The acronym PABITRA stands for Pacific-Asia Biodiversity Tran-sect, a network of island sites and conservation professionals collaboratingthroughout the Pacific-Asia region. An ideal PABITRA site is a broad landscapetransect from sea to summit. Such a landscape is Kahana Valley on WindwardO‘ahu. Kahana Valley served during prior centuries as an ahupua‘a, a Polynesianunit of land management that integrated the three biological resource zones, theupland forests, the agriculturally used land below, and the coastal zone, into asustainable human support system. Results of terrestrial biodiversity surveys, asbegun with a vegetation/environment study and a paleoecological investigation,are presented in relation to historical land use and sea level changes. In spite ofthe many former human-induced modifications of the Kahana Valley landscape,the natural structure and function of its ecosystems are well preserved. The dis-tribution patterns of vegetation can be interpreted in terms of Hawaiian ecolog-ical zones in combination with the valley’s precipitation, topography, streamsystem, and archaeological features. Currently, efforts are under way to restorethe Kahana State Park (recently renamed Ahupua‘a ‘O Kahana State Park) as afunctional ahupua‘a. In addition, focused collaborative research can yield helpfulinformation for further restoration and integrated management of the Kahanaahupua‘a as a historic Hawaiian Heritage Site.

The PABITRA (Pacific-Asia BiodiversityTransect) network aims to include broadlandscape units with three interconnectedbiodiversity resource zones into a study pro-gram. The three resource zones are an uplandor inland forest system that serves as water-shed cover, the agroecosystems that are usu-ally below in elevation, and the coastal zoneecosystems. A freshwater resource system isanother essential component.

The objective is to study each of these eco-system zones individually for their internal

functions but also to investigate their rela-tionships as a landscape unit. The latter isnecessary to ensure that the management ofone resource zone does not affect the otherzones negatively and that the zonation systemis managed as an integrated landscape unit.Such is the management unit or human sup-port system developed (through trial and er-ror) by the early Hawaiians before contactwith Western society and named ahupua‘a.

The Ahupua‘a Model

An ahupua‘a is an island landscape represent-ing a vertical section from mountain to sea.It usually encompasses a watershed with thearea to the sea as deep as a person can standin the ocean water. Ahupua‘a is a Hawaiianterm. It refers to ahu, meaning a heap ofstones, and pua‘a, the Hawaiian word for pig.A related, second meaning of the term ahu-pua‘a refers to an altar on which a pig wasplaced as a tribute to the chief or landownerfor allowing the use of the ahupua‘a land.The landlord or owner and manager of an

Pacific Science (2005), vol. 59, no. 2:293–314: 2005 by University of Hawai‘i PressAll rights reserved

1 Manuscript accepted 20 April 2004.2 Department of Botany, University of Hawai‘i at

Manoa, 3190 Maile Way, Honolulu, Hawai‘i 96822.3 Deceased, former Director, World Wide Fund,

Indonesia.

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ahupua‘a was called konohiki (Pukui and El-bert 1986).

Luciano Minerbi (1999) described a typicalahupua‘a diagrammatically as here shown inFigure 1 with some slight modification em-phasizing ecotones (i.e., transition zones). Asseen here, an ahupua‘a consists essentially ofthree vertically changing biological resourcezones: the upland (mauka) forest zone (wao

nahele), which may include a cloud forest(wao akua [akua means god]); the agriculturalzone (wao kanaka [where people work]); andthe coastal zone (makai or kahakai [kai meansocean, and kahakai means beach or sea-shore]). A fourth important resource zone, asshown on Figure 1, is the freshwater stream(kahawai). The stream flow is modified inthe wao kanaka by man-made ditches (‘auwai)

Figure 1. Ahupua‘a model, after Minerbi (1999).

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for irrigating taro (Colocasia esculenta) patches(lo‘i) on alluvial terraces. Dryland root crops,including taro and sweet potato (‘uala, Ipo-moea batatas), were grown in the opened-upuplands (kula uka). Breadfruit (‘ulu, Artocar-pus altilis), bananas (mai‘a, Musa paradisiaca),and coconut (niu, Cocos nucifera) were amongthe most common tree crops planted in thelowlands and midlands.

Beyond crop areas, the ahupua‘a containsmakai-mauka (toward the sea–toward themountain) access trails as well as midelevationand coastal trails, house sites (kauhale), tem-ples (heiau), and burial sites (pa ilina). Amost important coastal resource enhance-ment includes a fishpond (loko i‘a). This isa stonewalled enclosure in shallow coastalwater that allows small fish to enter throughsluice gates (makaha). After gaining in size,the fish is trapped. Thus, a fishpond acts as alarge fish trap.

The two man-made devices, the irrigatedterraces (lo‘i) and fishponds, are clear evi-dence of the agricultural engineering talentsof the early Hawaiians to make full use ofthe productive capacity of their windwardahupua‘a.

The Kahana Ahupua‘a and Its Tenure History

location. The Institute of HawaiianStudies, University of Hawai‘i at Manoa, pre-pared a map of O‘ahu Island (Figure 2),which shows the traditional subdivisions intofive districts, called moku, and numerous ahu-pua‘a, all designated by names with bound-aries. Kahana stands out on windward, centralO‘ahu in the south of Ko‘olauloa moku.

A recent photo taken from a low-flyingaircraft (Figure 3) shows Kahana Valley withits restored fishpond at lower left (the HuiluaFishpond), the Kahana estuary next to it, itscoastal ironwood (Casuarina equisetifolia) for-est, the verdant central marsh, and forestedbottomlands. The slopes show minor landscars, but are mostly forested. The peak onthe left is Pu‘u o Kila, 467 m (1,530 feethigh), and in the background, about 6 km in-land, is the Ko‘olau mountain range withsummit elevations of ca. 610 m (2,000 feet).Kahana Stream originates at about 300 m

elevation behind the ridge extension of Pu‘uo Kila. In front of the Pu‘u o Kila ridge isKawa Stream, which joins Kahana Stream inthe midsection of the valley.

tenure history. The Kahana ahupua‘awas originally a communal property of theHawaiian people. It is estimated that from720 to 1,000 indigenous people were livingand supporting themselves from the resourcesof the valley and the ocean bay at the timeof Western contact in 1776 (Stauffer 1990).Over the following 100 yr, the indigenouspopulation declined to an estimated low ofca. 90 people. In the late nineteenth century,it rose again to ca. 250 inhabitants, primarilydue to Chinese immigrants. Major factors forthe rural decline of the indigenous peoplewere introduced diseases, the lure of the city,Honolulu, and the great mahele (land divi-sion) of 1846–1855 with its ensuing privatiza-tion of native lands and the change to aforeign cash economy.

The great mahele resulted in a four-waydivision of the land. Recipients were (1)the king, (2) the other high ali‘i, (3) theAmerican-dominated government, and (4)the maka‘ainana (the common people).Stauffer (1990) documented the changes inland tenure of Kahana Valley from 1846 to1920. The great mahele allowed the maka‘ai-nana to claim their kuleana (i.e., their houselots and cultivated areas). This amounted toonly 200 acres (ca. 81 ha). The remaining un-cultivated land, ca. 5,000 acres (ca. 2,024 ha),became the property of the high ali‘i AnnieKeohokalole in 1847. She was the mother ofKing David Kalakaua and his sister, QueenLili‘uokalani, who reigned from 1891 to1893. Annie Keohokalole and her husbandhad large land holdings but little money. Tosupport their lifestyle they needed money.Loans were given by foreign businessmen us-ing land as security. Eventually, 3,000 acres(ca. 1,215 ha) of the ahupua‘a were sold byKeohokalole to a Chinese merchant namedAh Sing in 1857. He became the konohiki.His son, Ah Mee, owned the property in1872. Figure 4 shows Kahana Valley in aphoto taken in 1888. Note absence of foreston foothills and presence of grazing animals.

The maka‘ainana soon began to realize

Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 295

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that through privatization they lost their‘‘cooperative rights’’ to the ahupua‘a. Theserights relate to using the uncultivated landwith the freshwater stream that fed their lo‘ivia their ditch system (the ‘auwai), the use offorest products and gathering rights as well ascommunal fishing rights.

In 1874 a Kahana hui (association) underGeorge William Kamakaniau bought the Ka-hana ahupua‘a back from Ah Mee for $6,000.At that time, a Hawaiian again became thekonohiki. However, the money was bor-rowed as a mortgage. This was made possiblethrough the Mortgage Act of 1874. Eventu-ally the mortgage terms could not be met bythe Hawaiian hui, and foreclosure resulted inthe transfer of ahupua‘a property rights tonon-Hawaiian businesspeople. Among thesewere the Castle family who owned the Ka-neohe Ranch Co., and Mary Foster, daughter

of the hapa haole (half foreigner) Robinsonshipbuilder family. Mary Foster’s husband,Thomas Foster, was a Canadian businessper-son who owned interisland shipping, whichlater became the Matson Navigation Com-pany.

Mary Foster was one-quarter Hawaiian,had close connections to the royal family,and spoke fluent Hawaiian. She eventuallybecame the konohiki and prime owner ofthe Kahana ahupua‘a by 1920. But shestruggled with two other haole landowners,the Kaneohe Ranch Co. and the McCandlessbrothers. The Kaneohe Ranch Co. acquired alease from the Hawaiian hui for Kahana’ssloping uplands for cattle grazing. Theyburned much of the coastal upland and nativemidland forest for conversion into pasture-land. The McCandless brothers acquiredrights to the mauka uplands. As professional

Figure 3. Kahana Valley as seen recently from a low-flying aircraft, with its restored fishpond at lower left (Huilua),the Kahana estuary next to it, the coastal ironwood (Casuarina equisetifolia) forest, the lush central marsh, land scars onfrontal slopes, forested bottomland, and back slopes. Pu‘u o Kila (467 m) stands out as the peak on the left horizon.Photo courtesy of Douglas Peebles.

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artisan well borers, they realized the poten-tial of tapping the windward groundwaterresources halfway up the Ko‘olau mountainrange. They designed the Kahana-Waikane-Waiahole Ditch and mountain tunnelingsystem. The work was carried out by Asianminers from 1913 to 1916. This was a greatengineering accomplishment at that time,which made the irrigated sugarcane agricul-ture at ‘Ewa basin possible. However, legalbattles arose over the land and water rightswith the konohiki of Kahana, Mary Foster.She received an out-of-court settlement, andthe McCandless brothers made a fortune byselling their Waiahole-Waikane Water Co.to Amfac, owners of the O‘ahu Sugar Co. in‘Ewa. Figure 5 shows the entrance to KahanaValley from Kamehameha Highway and theHuilua Fishpond in a photo taken in 1936.

After Mary Foster’s death, the Kahanaahupua‘a reverted to the State of Hawai‘i in

1967. It is now public property and called of-ficially the Ahupua‘a ‘O Kahana State Park.Figure 6 shows the Kahana ahupua‘a in aphoto taken soon after World War II. Notethe barren front hills and dry terraces (kulalands).

Kahana Vegetation

vegetation map. Nengah Wirawanprepared a vegetation map in 1972 (Figure 7)in conjunction with a botanical survey ofKahana Valley (Theobald and Wirawan1973). He subsequently produced a Ph.D.dissertation (Wirawan 1978) on the vegeta-tion ecology of Kahana under guidance ofD.M.-D. At that time, the air photo–derivedvegetation map was verified with 64 vegeta-tion releves (sample stands of 400 m2 each)spread out from the coastal area into theback of the valley up to the base of the steep

Figure 4. Frontal area of Kahana Valley in 1888 during time of ranching use. Grass and low scrub in foreground withgrazing horses and almost barren slopes on opposite side of bay. Photo CP 96720, courtesy of Bishop MuseumArchives.

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wet cliff area where it rises from ca. 300 to610 m over a short distance of ca. 200 m. Fig-ure 7 is a reproduction of Wirawan’s vegeta-tion map. It shows 13 units.

Unit 1 relates to the wet, sparsely vege-tated, nearly vertical cliff area (pali) in theback of the valley. There are also pali seg-ments on the eastern side, rendering Kahanaan aging ‘‘amphitheater-headed valley.’’ (Forplants growing on wet and dry cliffs, seeMueller-Dombois and Fosberg 1998:488.)

Unit 2 is designated as Metrosideros wood-land. The key species are clumps and scat-tered trees of Metrosideros polymorpha (‘ohi‘alehua) together with the indigenous thickmat-forming uluhe fern, Dicranopteris linea-ris. Other important native trees are Euge-nia sandwicensis (ha), Antidesma pulvinatum(hame), Psychotria (kopiko), small stands ofPritchardia palms (loulu), and some Cibotiumtree ferns (hapu‘u). Important aliens found in

1972 were occasional trees of Spathodea cam-panulata (African tulip tree) and the shrubClidemia hirta (Koster’s curse).

Unit 3 represents an entirely alien treeplanting of Paraserianthes falcataria, knowncommonly as the tall albizia legume tree.This tree planting indicates the Ko‘olauMountain ditch and tunnel system, developedbetween 1913 and 1916 for transferringsome of the freshwater flow and the uppermountain-stored water for irrigating the for-mer sugarcane fields in the ‘Ewa basin abovePearl Harbor. The ditch system begins herein the upper Kahana back valley and contin-ues through the neighboring Waikane andWaiahole ahupua‘a to the south.

Unit 4 is recognized as the Acacia koa–Pandanus (hala tree) woodland. Both koa andhala are native trees. Most others are intro-duced. The latter include the Polynesian in-troduction Syzygium malaccense (‘ohi‘a ‘ai),

Figure 5. Kahana Valley in 1936 during Mary Foster’s ownership. The valley was given to rice cultivation and cattleranching. The Waikane/Waiahole Ditch and tunnel system was emplaced in the back of the valley. Photo 38215 AC,courtesy of Bishop Museum Archives.

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which is common throughout the valley andwell represented by seedlings and saplings, aswell as an occasional Artocarpus altilis (the ‘uluor breadfruit tree). Other later-introducedfruit trees include the mango tree (Mangiferaindica), guava or kuawa (Psidium guajava), andthe strawberry guava or waiawı ‘ula‘ula (Psi-dium cattleianum). The latter is considered aserious weed tree; there are some large (about0.5-m diameter) strawberry guava trees inKahana Valley, signifying an introduction inthe late 1800s. Seedlings and saplings are notabundant here. Instead, a new invader, Ardisiaelliptica (the shoebutton tree), is rapidly be-coming abundant in forest openings of unit4. Koa saplings are sparse. Wirawan (1978)found that removal of the hala leaf litterand some soil surface scarification encourageskoa regeneration in unit 4. Some areas in thisunit currently exhibit koa dieback or decline.A number of insects and fungal pathogenshave been implicated but no causative rela-

tionship has so far been ascertained (Puni-wai 1997). The disturbance history is mostlikely involved. Abundant koa regenerationhas been observed after fire ( Judd 1935)or browser removal (Spatz and Mueller-Dombois 1973). After such events, koa re-generates in cohorts (i.e., generation stands).Once these cohort stands become senescent,any temporary climatic stress can triggersuch tree groups into dieback because oftheir demographic predisposition. Insects andfungi often merely hasten the dieback ordecline because such trees have lost their ca-pacity to regenerate foliage. If these pest or-ganisms were constantly virulent, trees wouldloose their foliage as soon as it forms. Ofcourse, younger koa trees may also becomepredisposed to die (for example, in this case,through competition with hala trees, whichhave now gained dominance over koa in unit4). Application of some sensible silviculturecould reverse the trend.

Figure 6. Kahana Valley in 1945 with sugarcane on lower left, open slopes above, smoke from a fire at right, and armyinstallations above bottomland. Photo USA MH 5408, courtesy of Bishop Museum Archives.

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Unit 5 (crosshatched on the map), desig-nated as forest scrub, consists primarily ofsmall, pure-stand patches of Syzygium cumini( Java plum), Syzygium jambos (rose apple or‘ohi‘a loke), and stands of Phyllostachis nigra(black bamboo), and Schyzostachium glaucifo-lium (‘ohe, possibly of Polynesian introduc-tion). This unit grades into the riparian Unit8.

Unit 6 is the Aleurites moluccana (kukui)tree community of Polynesian introduction.It fills the gulches on both the north andsouth slopes from near the coast to the mid-dle of the valley. It is most prevalent in themesic part of the valley and drops out wherethe mean annual rainfall exceeds 4,000 mm.Wirawan (1978) found that kukui tree regen-eration is related to storm surges, when tum-bling rocks in the gulches happen to crack theaccumulated kukui nuts on the forest floor.

Unit 7 is the Acacia koa–Pandanus forest, avariation of unit 4, recognized by taller anddenser tree growth indicating either a moreproductive habitat or less disturbed vegeta-tion.

Unit 8 is dominated by the hau tree, Hi-biscus tiliaceus, which is either native or an-other Polynesian introduction (Wagner et al.1990). It forms a jungle or krummholz andcovers many former irrigated terraces (lo‘i).It grows over sections of Kahana Stream andforms a dense cover all along its estuary.There it grows intermixed with the intro-duced American mangrove (Rhizophora man-gle).

Unit 9 is occupied dominantly by the octo-pus tree (Schefflera actinophylla). It is a fast-growing secondary rain-forest tree, probablyintroduced from Australia as an ornamental.Wagner et al. (1990) cited Degener as havingsaid that the octopus plant was grown in potsand Hawaiian gardens as an ornamental in1900. In Kahana, this tree has rapidly invadedthe formerly open grass-covered slopes (kulakai) on both sides of the lower valley fromthe coast to about 1.5 km inland. It is nowalso invading the marsh (unit 11).

Unit 10, designated as mixed grass-scrubon the map, has now mostly disappeared.In 1972, this was dominantly occupied byAndropogon virginicus (the broomsedge), a py-

rophytic grass introduced from the southeast-ern United States in 1924. Its spread waspromoted by fire, and it contributed sub-stantially to soil avalanching on slopes withdeeply oxidized soil (ultisol) here in KahanaValley and in many other windward valleys(Mueller-Dombois 1973). The grass-scrubcover is now mostly invaded by the octopustree. However, a native shrub, Osteomelesanthyllidifolia (‘ulei), still prevails in the un-dergrowth on sun-exposed slopes, whereAndropogon has become very sparse. The oc-topus tree is still on the move and currentlyis invading unit 4 farther inland as well asunit 11.

Unit 11 is primarily marsh-grassland ofBrachiaria mutica (California grass). Like thebroomsedge it was first collected in Hawai‘iin 1924 (Wagner et al. 1990). At the time ofmapping the marsh, which gets less swampyinland, this unit served as cow pasture, andin the wetter parts rice was still cultivated inthe mid 1970s.

Unit 12 represents the dry coastal cliffcommunity, sparsely vegetated with wisps ofgrasses including Heteropogon contortus (piligrass) and Rhynchelytrum repens (Natal red-top).

Unit 13 relates to the strand community,which at Kahana Bay is dominated by anironwood (Casuarina equisetifolia) forest witha row of Terminalia catappa (kamani haole)trees along Kamehameha Highway. An up-dated plant checklist is included in the Ap-pendix.

physiographic environment. Impor-tant geomorphic features of Kahana Valleyare displayed on the physiographic map (Fig-ure 8). The figure shows the physiographyand topographical diversity by 200-ft (61-m)contour lines. Steep slopes are indicated bycrowding of the contour lines. The back val-ley rises steeply to the Ko‘olau summit ridge.Another ridge, topped by the 1,530 ft (467 m)Pu‘u o Kila, juts into the valley along the5,000-mm precipitation line. A third ridgewith a steep slope down into the valley isfound on the east side near the coast. Wig-gling of the contour lines indicates numeroussteep gulches along each of the slopes, whichfrom near the coast to beyond the midvalley

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are dominated by kukui, unit 6 on the vegeta-tion map.

stream system and precipitation.The Kahana Stream system is shown in Fig-ure 8 as originating in the back of the valleyat about 1,000 ft (305 m) elevation. Thedashed line running between 800 (244 m)and 1,000 ft (305 m) elevation indicates theditch where fresh water is extracted for trans-fer to leeward O‘ahu. The freshwater streamsystem follows a course that relates directlyto the topographic pattern of the valley.Another important feature is the precipitationgradient, which starts at the coast with a meanannual rainfall of 1,875 mm (74 in.). In themidvalley where Kawa Stream joins KahanaStream, the mean annual precipitation is2,500 mm (98 in.). From there rainfall dou-bles toward the back valley, where rainfall ex-ceeds 7 m (275 in.) per year. Thus, there is asteep rainfall gradient. From the coast to theback valley rainfall increases from ca. 2 m to 7m over a distance of 6 km. The four physicalenvironmental features of topography, geo-morphology, stream system, and precipitationpattern explain much of the observed vegeta-tion pattern.

topographic profile with climate

diagram. In addition, there is a gentle slopein the center of the valley, descending in step-like intervals from mauka to makai, as seen onthe profile map along Transect 1 (Figure 9),which explains the direction of stream flow.The climate diagram substantiates the greatdifferences in rainfall from mauka to makai.It also shows how rainfall is distributedthroughout the year, indicating a short dryperiod in June/July at the coast versusmonthly rainfall in excess of 300 mm inlandat the Kahana Valley ditch and tunnel system.Here, one can see that it is also cooler by1.5�C throughout the year.

hawaiian ecological zones. The pro-file also shows an attempt to recognize theecological zonation system developed bythe early Hawaiians. It is here related totopography, precipitation, stream flow, andvegetation in five zones from mauka to makai.Zone A, kua mauna, refers to the wet cliff(pali) area in the back valley. Zone B, waokele, refers to the wet Metrosideros forest.

Zone C, wao koa, relates to the Acacia koa–Pandanus area. Zone D, kula kai, refers tothe formerly more open grassy slopes, nowoccupied mostly by introduced octopus trees.Zone E, kahakai, relates to the coastal strand,with Casuarina forest, mixed grass-scrub, andfishpond as well as into the sea up to about1.5 m depth.

water yield in kahana valley. Taka-saki et al. (1969) estimated the water yield forKahana Valley. They calculated a total inputfrom precipitation amounting to 60 milliongallons (240 million liters) per day. This esti-mate appears reasonable. A relationship ofheight to area, which is easy to remember, isthat 1 mm/m2 yields 1 liter. If one considersthe effective catchment area to be 2,000 ha(the Kahana ahupua‘a is estimated to include5,200 acres) and the average daily precipita-tion to be 12 mm, the incoming yield wouldbe 12 mm� 2,000 ha ¼ 240 million liters orca. 60 million gallons.

Takasaki et al. (1969) considered the fol-lowing fractionation of their estimated 60million gallons (240 million liters) input perday: 12 million gallons (48 million liters) arelost as evapotranspiration (20%), 15 milliongallons (60 million liters) are lost as runoff(25%), 19 million gallons (76 million liters)enter the stream system (32%), 4 million gal-lons (16 million liters) are removed by theKahana-Waiahole Ditch system (6%), and10 million gallons (40 million liters) leavethe area as groundwater (17%). This givessome idea of the valley’s water yield poten-tial. Further direct measurements at differentrainfall events including storm surges and dryperiods would be of considerable interest andpractical value because this is known to be ahighly dynamic relationship.

archaeological map. Hommon andBarrera (1971) found 119 archaeological sitesin Kahana Valley. They produced a foldoutmap that shows all of the site locations withan outline of their spatial extent. This map ishere reproduced as Figure 10 and superim-posed on the vegetation map units drawnat the same scale. The archaeological mapshows that former human habitation sitesextend about 5 km into the valley along thestreams. The authors recorded 120 wet ter-

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races (lo‘i) connected by man-made irrigationchannels (‘auwai). Most of these terraces werebuilt on gently sloping alluvial areas with re-taining walls. The total growing surface ofthe wet terraces was calculated as 10,800 m2

(1.08 ha). The authors suggested that manylower terraces were probably buried underalluvial material. A second archaeological fea-ture was clearings for the cultivation of non-irrigated crops (probably dry taro and sweetpotato). These dry terraces were recognized

by removal of stones. Terraces marked bystone walls were often rather small, only 2 by10 m. In addition, the authors identifiedanimal enclosures, house sites, men’s eatinghouses (mua), and scarcely legible petro-glyphs. The archaeological sites coinciderather well with three current vegetationtypes, the mixed forest scrub (unit 5), the Hi-biscus (hau) forest (unit 8), and the pasture andmarsh (unit 11) as shown on the vegetationmap.

Figure 9. Topographic profile from mountain to sea (mauka-makai) in central Kahana Valley following the courseof Transect 1 (TR1) on the physiographic map (Figure 8). A Walter-type climate diagram (top right) shows meanmonthly temperature and rainfall curves with a short dry season at the coast in June/July and year-round high rainfallin the back of the valley (top left) with a mean annual rainfall gradient from 1.9 to 5.9 m over 6 km distance from thecoast to the back of the valley. Along this profile one can distinguish five Hawaiian ecological zones as indicated.

Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 305

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Kahana Valley, a Geomorphic Artifact?

a former inland lagoon. Patricia RiceBeggerly (1990) presented a very thoroughand thought-provoking Ph.D. dissertation onher paleoecological research in Kahana Val-ley. She considered Kahana Valley to be a‘‘geomorphic artifact’’ based on past humanimpacts. She dug deep soil trenches in eightareas at the borders of the marsh and in thepasture up to 1.2 km inland. She used opentrenches and auger borings to distinguish soillayers with inclusions of organic remains suchas snail shells, charcoal, wood, pollen, andfern spores. For charcoal, shells, and somewoody remains, she obtained radiometricdates. She distinguished seven soil stratafrom about 3 m depths upward to the currentsoil surface and clarified that the valley bot-tom was once an ocean inlet, a finding veri-fied also for Kawainui marsh in the Kailuaahupua‘a, farther south on windward O‘ahu(see Kirch 1985). This was not surprising be-cause there was a higher sea level followingthe Pleistocene. A shallow-water phase suc-ceeded in the form of a lagoon. As evi-dence, Beggerly found shells of brackish-water snails, such as Tellima macoma dispar.In the lagoon phase, at the bottom of themarsh, she found charcoal. Dates for theseorganic remains turned out to be 430–30 b.c.Beggerly suggested that the charcoal remainsargue for a first invasion of human settlersinto Kahana Valley by at least 30 b.c. She be-lieved that wood fires could not have startedin this rain-forest valley because of its year-round high humidity. On the shallow lagoonbottom she found colluvial material as thenext deposit upward. This material, she sug-gested, was deposited from the slopes byaccelerated erosion, when the first human set-tlers used slash-and-burn swidden cultivation.Following the colluvial layer upward, mineralsoil and organic muck mixtures indicate con-tinued terrestrification. However, there wasno charcoal inclusion until she came nearerto the surface (between 2.2 and 2.5 m). Thesecond find of charcoal was dated 1425G80 yr a.d.

pandanus versus pritchardia pollen.Based on this second charcoal date, Beggerly

(1990) proposed that Kahana Valley wasabandoned for at least 1,200 yr and was rein-vaded by a new wave of indigenous settlersat about 1200 a.d. (i.e., 800 b.p.) at the ear-liest. She proposed that the earlier settlers,who came into the valley around 230G 200b.c., experienced an ‘‘ecological crunch’’ dueto their overexploitation of the valley’s re-sources, with the ecological crunch havingresulted primarily from their traditionalslash-and-burn practice. This would have in-volved accelerated soil erosion and nutrientloss with slow growth of vegetation, thusshifting activity across the sloping landscape.This also implied exhaustion of readily avail-able food resources, such as starch from tree-fern trunks and fish from the inland lagoonas it continued to decline from acceleratedsedimentation. At that time, the first settlerscould still migrate to another empty valley.Beggerly argued that the first settlers usedtree-fern starch as a food resource becausetree fern spores showed an early ebb in hersoil core samples.

She found Pandanus pollen in the lagoonphase, which may explain today’s presenceof the many hala trees on the midslopes ofboth sides of the valley. Pandanus is usuallyfound in coastal settings near the ocean shore.She also found ebbs in Metrosideros pollenand tree-fern spores alternating with ebbs ofPandanus pollen and concluded that these al-ternations may be the result of human inter-vention with Kahana’s rain forest. Athensand Ward (1993), doing similar coring studiesin Kawainui marsh, showed the startling re-sult of an abundance of Pritchardia (loulu)palm pollen before the first centuries of Poly-nesian land occupation. They found that theloulu palm pollen began to disappear around1200 a.d. and suggested disease rather thanhuman-induced fire as the cause. They alsohold that loulu palm forest was the dominantlowland vegetation type on windward O‘ahuwhen the first human settlers arrived. BecausePatricia Beggerly did not find loulu palm pol-len in her corings of Kahana marsh, it seemsworthwhile to reinvestigate this area, particu-larly because loulu palms are still present inKahana’s back valley today. The same appliesto koa tree pollen, which was not found by

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Beggerly in her soil cores. However, koa pol-len was found by Athens et al. (1992), even inareas that lack Acacia koa today. Athens andWard (1993) did not dispute Beggerly’s earlyhuman invasion date but suggested resam-pling her charcoal findings for comparabilitywith other areas of O‘ahu.

Two Paleoecological Land Use Models

continuous succession of human

occupation. Patrick Kirch (2000) providedan interesting theory of a successional se-quence of the Hawaiian cultural develop-ment. He divides it into four phases: phase 1:from 300 to ca. 600 a.d., the colonization pe-riod; phase 2: from 600 to ca. 1100 a.d., thedevelopment period; phase 3: from 1100 toca. 1650, the expansion period; and phase 4:from 1650 to Western contact (1776), theprotohistoric period.

During phase 1, the prevailing agriculturaltechnology was shifting cultivation, as stillpracticed today, for example, in parts of Va-nuatu, Solomon Islands, and New Guinea.Taro irrigation in windward valleys had al-ready begun in the colonization phase. It didnot develop into a more widely used agricul-tural technology until the beginning of theexpansion phase, around 1100 a.d. Its greatestsophistication was reached during the proto-historic phase from 1600 to 1800 a.d.

discontinuous succession of human

occupation. Patricia Beggerly’s (1990)model, based directly on her research in Ka-hana Valley, suggested four different phases,as follows.

Phase 1: Between 400 and 30 b.c., the firsthuman settlement occurred. She suggestedthat it involved ca. 100 people. They initiallyused tree-fern starch as a food source whilethey began swidden cultivation with ti (Cor-dyline terminalis) and taro (Colocasia esculenta)through opening the forest by cutting itdown with subsequent burning of the slash.In so removing the forest on slopes, theycaused soil avalanching, which resulted in theaccumulation of colluvium in the lagoonphase at the valley bottom. That also de-stroyed the fish populations in the lagoon.Overexploitation forced the first colonizers

to migrate into neighboring valleys. At thattime, such valleys may have been unoc-cupied.

Phase 2: From 30 b.c. to 1200 a.d., KahanaValley was abandoned. The indigenous forestregenerated. Fewer soil avalanches occurred.The marsh formed by further, mostly or-ganic, terrestrification processes and herba-ceous cover dominated on the valley floor.

Phase 3: From 1200 to 1800 the secondwave of settlers had arrived. They againstarted with slash-and-burn agriculture. Asecond phase of environmental degradationensued but not for long. This time, the set-tlers did not move out of Kahana Valley.Because the population had expanded in allwindward valleys, space had become limited.In conjunction with swidden cultivation byremoving forest for growing crops on dryland, the settlers began irrigation agricultureby constructing wet terraces (the lo‘i) andconverting stream water through ditch sys-tems (the ‘auwai).

Phase 4: From 1800 to 1900, after West-ern contact, a third phase of forest degra-dation began, combined with acceleratederosion. Early in this phase, ca. 1825, as toldin written historical accounts, slopes werebrown and bare and forest was confined tothe inner valley and the upland region. Thewestern slopes were still cultivated in spotswith dryland taro, banana, and sweet potato,and the riparian lowlands with irrigated taroin spotty wet terraces. The traditional usechanged with overseas immigrants: Easternagricultural practices of rice growing andWestern practices of cattle grazing were in-troduced. In addition to the free-roaming cat-tle, other hoofed animals such as goats, sheep,and pigs were using the valley’s vegetation.This was followed by sugarcane growing inthe pasture area with the Castle-owned Ka-huku Railroad reaching south across KahanaStream into the eastern valley in the lateeighteenth to early nineteenth century. Dur-ing World War II Kahana Valley was usedas a military training area. Remnant bunkerstructures are still found on the loop trail inthe back part of the valley. Related to this isthe tragic story of land tenure, the changesin land ownership as discussed earlier.

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sea level changes. Patrick Nunn’s(1999) diagram of sea level changes over thelast 1,200 yr (Figure 11) provides further in-sights. The first human colonization phasecertainly had occurred before 800 a.d. (i.e.,1200 b.p.) when the sea level was perhaps2 m higher than today. The bottomland ofKahana Valley was then a marine embaymentup to the inland margin of the current pas-ture, or meadow (ca. 1.2 km inland of thecurrent Kahana beach), as confirmed by Pa-tricia Beggerly (1990). Kahana’s inland marinebay changed into a lagoon most likely whenthe sea level dropped thereafter to about 1 mabove the current level. According to Beg-gerly’s interpretation, the first settlers hadalready abandoned Kahana Valley due tooverexploitation, and the valley was experi-encing a fallow period of about 12 centuriesuntil ca. 800 b.p. (i.e., 1200 a.d.). Then thesecond wave of settlers arrived eight centuriesago, during the ‘‘Little Climatic Optimum.’’They again began to use Kahana Valley bypracticing slash-and-burn agriculture. This

second group of settlers remained in thevalley. They soon changed their land-usepractices from degrading the environment toenhancing its natural resources. They accom-plished this by developing irrigation schemesand removing stones from dryland terracesfor growing newly introduced crop plants,such as sweet potato (Ipomoea batatas, ‘uala),as well as by constructing fish traps culminat-ing in the Huilua Fishpond. The latter is saidto have been developed about four centuriesago.

According to Patrick Kirch’s (2000) suc-cessional interpretation of human impact onwindward O‘ahu, there was only one phaseof settlement. Colonization with slash-and-burn practices occurred only once. This wasca. five centuries before the Little ClimaticOptimum. During the warmer and calmerclimatic phase, when ocean canoe travel wascommon practice, Kirch’s development phasegave way to his expansion phase ca. nine cen-turies ago. From that time onward, the Ha-waiian land-use practices started to enhance

Figure 11. Changes in emergence over the last 1,200 yr as portrayed by Patrick D. Nunn (1999). Reproduced withsome additions for clarification and permission by the author.

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the natural ecosystem services through con-servation practices. This then became thetraditional Hawaiian ahupua‘a managementsystem. It existed through the ‘‘Little IceAge,’’ when European sea travel began in thePacific. At the end of the Little Ice Age,the Hawaiian land-care system collapsed,around 1860. The reason for that was not cli-mate change. It was primarily the Western-introduced privatization of land and the casheconomy that corrupted the traditional lifestyle and culture of the indigenous Hawaiianpeople. Jane Allen (1997), who reviewed pre-historic development on windward O‘ahu,also related landscape changes to the LittleClimatic Optimum and the Little Ice Age.She says that windward valleys were alreadysettled by about 600 a.d., and that human im-pact upslope was strong during the Little Cli-matic Optimum. During this period of highersea level, much erosive material was depositeddownhill on marine platforms. Thereafter,terracing began during the Little Ice Ageand the irrigation of lo‘i began. She con-cluded that both natural forces and humanactivity interacted in prehistoric landscapechange.

Conclusions

By synthesizing the former study of Kahana’svegetation ecology with its archaeological andpaleoecological investigations, it has becomeabundantly clear that human impact in thevalley has been substantial for at least the past12 centuries. However, characterization ofKahana Valley as a ‘‘geomorphic artifact,’’ asproposed by Beggerly (1990), does not appearto be warranted. The valley’s geomorphologycertainly has undergone changes. The con-tinued drop of sea level following the post-Pleistocene warming may be the prime rea-son for the change of the inland marine bayinto a lagoon. Moreover, the terrestrificationof the inland lagoon may have become accel-erated with the transition period to the LittleIce Age seven centuries ago, when the sealevel dropped another meter in roughly 100yr. Human invasion of the valley during orbefore the Little Climatic Optimum certainlycontributed to landscape change, but it wasnot the only force. Natural forces were

equally apparent, if not of greater impor-tance.

Moreover, Beggerly’s (1990) argumentthat her charcoal find, dated 230G 200 yrb.c., indicates human presence is tenuous.Her argument is based on the assumptionthat natural fires could not have occurred inthis wet rain-forest valley. That argumentoverlooks the fact that the widespread uluhefern dries up periodically and surely can serveas an easily combustible material during a dryspell. As Vogl (1969) documented, fire causedby lightning has occurred repeatedly in wind-ward valleys. Also, Mueller-Dombois and La-moureux (1967) documented the occurrenceof past natural fire in Hawai‘i before Polyne-sian land occupation.

The back-valley slopes, occupied by ‘ohi‘alehua (Metrosideros polymorpha) woodland withuluhe fern (Dicranopteris linearis), representthe outcome of a prolonged (ca. 2 million yr)primary succession. As fine soil is formedthere from weathering, it also wastes con-stantly or abruptly away by erosion, therebykeeping the substrate primarily as rockland.Both the ‘ohi‘a lehua tree and the uluhe ferninvade land scars after soil avalanches. Theoccurrence of rockland and avalanche scarson slopes maintains a vegetation whose dy-namics is arrested in primary succession.This vegetation never reaches a climax statein the sense of Clements (1916, 1928). Like-wise, the koa and hala tree are native pioneerspecies that are highly resilient. They regen-erate after fire. Thus, native colonizer speciesdominate the two larger vegetation types inthe mid- and back valley. The Metrosideros-Dicranopteris back-valley type is resilientto geomorphic instability. The Acacia koa–Pandanus midvalley type is resilient to humanimpacts, including an occasional fire. Themarsh area in the valley represents an ad-vanced successional stage of a typical hy-drarch succession in the Clementsian sense.If swamp tree species had evolved in theHawaiian flora, as are prevalent in the west-ern Pacific islands (Stemmermann 1981), themarsh would have long become a swamp for-est. Such process is currently ongoing withinvasion of the octopus tree. Another intro-duced tree species capable of invading themarsh is the paperbark tree (Melaleuca quin-

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quenervia). It has been planted in Kahana Val-ley but is not yet documented as naturalized.The current management goal of an arrestedwetland succession can still be upheld by treeremoval. Currently, efforts are under way torestore the Kahana State Park (recently re-named Ahupua‘a ‘O Kahana State Park) as afunctional ahupua‘a. Results of follow-up his-torical and more complete biodiversity re-search with PABITRA methodologies willcontinue to yield helpful information. Suchinformation can be used to support restora-tion of ecosystem functions and biodiversitycomponents. In addition, new research canbe directed to determine if the restored Ka-hana ahupua‘a will be able to function as asustainable system in the future.

acknowledgments

We thank Mark Nakamura for computerdrafting the line drawings and AnnetteMueller-Dombois for word processing andeditorial counseling.

Literature Cited

Allen, J. 1997. Pre-contact landscape trans-formation and cultural change in wind-ward O‘ahu. Pages 230–247 in P. V.Kirch and T. L. Hunt, eds. Historicalecology in the Pacific islands: Prehistoricenvironmental and landscape change. YaleUniversity Press, New Haven, Connecti-cut.

Athens, J. S., and J. V. Ward. 1993. Environ-mental change and prehistoric Polynesiansettlement in Hawai‘i. Asian Perspect. 32(2): 205–222.

Athens, J. S., J. V. Ward, and S. Whickler.1992. Late Holocene lowland vegetation,O‘ahu, Hawai‘i. N. Z. J. Archaeol. 14:9–34.

Beggerly, P. R. 1990. Kahana Valley, Hawai‘i,a geomorphic artifact: A study of the in-terrelationships among geomorphic struc-tures, natural processes, and ancientHawaiian technology, land use, and settle-ment patterns. Ph.D. diss., University ofHawai‘i at Manoa, Honolulu.

Clements, F. E. 1916. Plant succession: An

analysis of the development of vegetation.Carnegie Institution, Washington.

———. 1928. Plant succession and indica-tors. Republished, 1963. Hafner Publ.Co., New York.

Hommon, R. J., and W. M. Barrera Jr. 1971.Archeological survey of Kahana Valley,Ko‘olau District, Island of O‘ahu. Depart-ment of Anthropology, B. P. Bishop Mu-seum, Honolulu, Hawai‘i (prepared forDivision of State Parks). Department ofLand and Natural Recources Report 71-3.Available from Division of State Parks,State of Hawai‘i, Honolulu.

Judd, C. S. 1935. Koa reproduction after fire.J. For. 33:176.

Kirch, P. V. 1985. Feathered gods and fish-hooks: An introduction to Hawaiian ar-chaeology and prehistory. University ofHawai‘i Press, Honolulu.

———. 2000. On the roads of the winds: Anarchaeological history of the Pacific islandsbefore European contact. University ofCalifornia Press, Berkeley.

Minerbi, L. 1999. Indigenous managementmodels and protection of the ahupua‘a.Soc. Process Hawai‘i 39:208–225.

Mueller-Dombois, D. 1973. A non-adaptedvegetation interferes with soil water re-moval in a tropical rain forest area in Ha-wai‘i. Trop. Ecol. 24:1–18.

Mueller-Dombois, D., and F. R. Fosberg.1998. Vegetation of the tropical Pacific is-lands. Ecol. Stud. 132. Springer-Verlag.

Mueller-Dombois, D., and C. H. Lamoureux.1967. Soil-vegetation relationships in Ha-waiian kipukas. Pac. Sci. 21:286–299.

Nunn, P. D. 1999. Environmental change inthe Pacific Basin. Wiley & Sons, Ltd.,Chichester, England.

Palmer, D. D. 2003. Hawai‘i’s ferns and fernallies. University of Hawai‘i Press, Hono-lulu.

Pukui, M. K., and S. H. Elbert. 1986. Hawai-ian dictionary. University of Hawai‘i Press,Honolulu.

Puniwai, N. 1997. Decline of Acacia koaon windward O‘ahu. Final report. USDAForest Service, Institute of Pacific IslandsForestry, Honolulu. Available from Divi-sion of State Parks, State of Hawai‘i, Ho-nolulu.

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Spatz, G., and D. Mueller-Dombois. 1973.The influence of feral goats on koa tree re-production in Hawai‘i Volcanoes NationalPark. Ecology 54:870–876.

Stauffer, R. H. 1990. Land tenure in Kahana,Hawai‘i, 1846–1920. Ph.D. diss., Univer-sity of Hawai‘i, Honolulu.

Stemmermann, L. 1981. A guide to Pacificwetland plants. U.S. Army Corps of Engi-neers, Honolulu District, Honolulu.

Takasaki, K. J., G. T. Hirashima, and E. R.Lubke. 1969. Water resources of Wind-ward O‘ahu, Hawai‘i. Geological SurveyWater-Supply Paper 1894. Prepared incooperation with Division of Water andLand Development, Department of Landand Natural Resources, State of Hawai‘i.U.S. Government Printing Office, Wash-ington, D.C.

Theobald, W. L., and N. Wirawan. 1973.Kahana Valley botanical survey, Ko‘olau

District, island of O‘ahu. Department ofBotany, University of Hawai‘i, Honolulu,Technical Report prepared for Divisionof State Parks, Outdoor and HistoricalSites, Department of Land and NaturalResources, State of Hawai‘i, Honolulu.Available from Division of State Parks.

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Wagner, W. L., D. R. Herbst, and S. H.Sohmer. 1990. Manual of the floweringplants of Hawai‘i. 2 vols. University ofHawai‘i Press and Bishop Museum Press,Honolulu.

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Appendix

Kahana Valley Plant Checklist

Abutilon grandifolium* (Willd.) Sweet (Malvaceae), ma‘o,hairy abutilon

Acacia koa A. Gray (Fabaceae), koaAdenophorus pinnatifidus Gaudich. (Grammitidaceae), kihiAdenophorus tamariscinus (Kaulf.) Hook. & Grev.

(Grammitidaceae), wahine noho maunaAdiantum raddianum C. Presl (Pteridaceae), maidenhair

fernAgeratina riparia* (Reget) R. King & H. Robinson

(Asteraceae), pamakaniAleurites moluccana* (L.) Willd. (Euphorbiaceae), kukuiAlocasia macrorrhiza* (L.) Schott (Arecaceae), ‘apeAlyxia oliviformis Gaud. (Apocynaceae), maile, endemicAmaranthus viridis* L. (Amaranthaceae), pakaiAndropogon virginicus* L. (Poaceae), broomsedgeAntidesma pulvinatum Hillebr. (Euphorbiaceae), hameArdisia crenata* Sims (Myrsinaceae), Hilo hollyArdisia elliptica* Thunb. (Myrsinaceae), shoebutton

ardisiaArtocarpus altilis* (S. Parkinson ex Z) Fosb. (Moraceae),

‘ulu, breadfruitArundina graminifolia* (D. Don) Hochr. (Orchidaceae),

bamboo orchidBidens pilosa* L. (Asteraceae), kı, kı nehe, Spanish needleBidens populifolia Sherff (Asteraceae)Bixa orellana* L. (Bixaceae), ‘alaea, annatto, lipstick treeBlechnum appendiculatum Willd. (Blechnaceae)Brachiaria mutica* (Forssk.) Stapf (Poaceae), Calfornia

grassBroussonetia papyrifera (L.) Vonton. (Moraceae), waukeBulbostylis capillaris (L.) C. B. Clarke (Cyperaceae)

Calophyllum inophyllum* L. (Clusiaceae), kamaniCanavalia cathartica* Thouars (Fabaceae), maunaloaCarex wahuensis C. A. Mey (Cyperaceae)Casuarina equisetifolia* L. (Casuarinaceae), paina,

ironwood treeCenchrus enchinatus* L. (Poaceae), ‘ume‘alu, common

sandburCentaurium erythraea* Raf. (Gentianaceae), European

centauryCentella asiatica* (L.) Urb. (Apiaceae), pohe kula, Asiatic

pennywortChamaecrista nictitans* (L.) Moench (Fabaceae), laukı,

partridge peaChamaesyce hypericifolia* (L.) Millsp. (Euphorbiaceae),

graceful spurgeCharpentiera obovata Gaud. (Amaranthaceae), papalaChristella dentata (Forssk.) Brownsey & Jermy

(Thelypteridaceae), pai‘i‘ihaChrysopogon aciculatus* (Rets.) Trin. (Poaceae), manienie

‘ulaCibotium chamissoi Kaulf. (Dicksoniaceae), hapu‘uCibotium menziesii Hook. (Dicksoniaceae), hapu‘u ‘i‘iClerodendrum philippinum* Schauer (Verbenaceae),

pıkake hohonoClidemia hirta* (L.) D. Don (Melastomataceae), Koster’s

curseCocculus trilobus (Thunb.) DC (Menispermaceae),

huehueCoix lachryma-jobi* L. (Poaceae), pu‘ohe‘ohe, kukaekolea,

Job’s tearsColocasia esculenta* (L.) Schott. (Araceae), kalo, taro

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Appendix (continued)

Commelina diffusa* N. L. Burm. (Commelinaceae),honohono, honohono wai

Convolvulus arvensis* L. (Convolvulaceae), field bindweedConyza canadensis* (L.) Cronq. (Asteraceae), lani wela,

Canada fleabaneCordyline fruticosa* (L.) A. Chev. (Agavaceae), tiCoronopus didymus* (L.) Sm. (Brassicaceae), swinecressCrotalaria pallida* Aiton (Fabaceae), pikakani, smooth

rattleboxCuphea carthagenensis* ( Jacq.) Macbr. (Lythraceae),

tarweedCuscuta sandwichiana Choisy (Cuscutaceae), kauna‘oa,

kauna‘oa leiCyclosorus interruptus (Willd.) H. Ito (Thelypteridaceae),

nekeCyperus rotundus* L. (Cyperaceae), kili‘o‘opu, nutgrassCyrtandra hawaiiensis C. B. Clark (Gesneriaceae),

ha‘iwaleCyrtandra laxiflora H. Mann (Gesneriaceae)Cyrtandra spp., endemicDicranopteris linearis (Burm. f.) Underw.

(Gleicheniaceae), uluheDioscorea bulbifera* L. (Dioscoreaceae), hoi, bitter yamDioscorea pentaphylla* L. (Dioscoreaceae), pi‘a, yamDiospyros sandwicensis (A. DC) Fosb. (Ebenaceae), lamaDiplazium sandwichianum (C. Presl) Diels (Athyriaceae),

ho‘i‘oDiplopterygium pinnatum (Kunze) Nakai

(Gleicheniaceae), uluhe lau nuiDoryopteris decipiens (Hook.) J. Sm. (Pteridaceae),

‘iwa‘iwaDryopteris spp. (Dryopteridaceae), oluaElaeocarpus bifidus Hook. & Arnott (Elaeocarpaceae),

kaliaElaphoglossum aemulum (Kaulf.) Brack.

(Lomariopsidaceae), ‘ekahaElaphoglossum crassifolium (Gaudich.) W. R. Anderson &

Crosby (Lomariopsidaceae), ‘ekahaEmilia fosbergii* Nicolson (Asteraceae), pualeleEmilia sonchifolia* (L.) DC (Asteraceae), Flora’s

paintbrushErechtites valerianifolia* (Wolf ) DC (Asteraceae)Eucalyptus* sp. (Myrtaceae)Fimbristylis dichotoma (L.) Vahl (Cyperaceae)Freycinetia arborea Gaud. (Pandanaceae), ‘ie‘ieGardenia mannii St. John & Kuykendall (Rubiaceae),

nanuGonocormus minutus (Blume) Bosch (Hymenophyllaceae)Grammites tenella Kaulf. (Grammitidaceae), kolokoloHedychium coronarium* J. Konig (Zingiberaceae),

‘awapuhi ke‘oke‘o, white gingerHedychium flavescens* N. Carey ex Roscoe

(Zingiberaceae), ‘awapuhi melemele, yellow gingerHibiscus arnottianus A. Gray (Malvaceae), koki‘o ke‘oke‘oHibiscus tiliaceus L. (Malvaceae), hauHuperzia phyllantha (Hook. & Arn.) Holub

(Lycopodiaceae), wawae‘iole, rat’s footIpomoea alba* L. (Convolvulaceae), koali pehu, moon

flower

Ipomoea indica ( J. Burm.) Merr. (Convolvulaceae), koali‘awa

Ipomoea pes-caprae (L.) R. Br. (convolvulaceae) pohuehueKorthalsella latissima (Tiegh.) Degener (Viscaceae),

kaumahana, mistletoeKyllingia nemoralis* ( J. R. Forster & G. Forster) Dandy

(Cyperaceae), kili‘o‘opuLantana camara* L. (Verbenaceae), lakanaLepisorus thunbergianus (Kaulf.) Ching (Polypodiaceae),

pakahakahaLeucaena leucocephala* (Lam.) de Wit (Fabaceae), koa

haoleLudwigia octivalvis* ( Jacq.) Raven (Onagraceae), kamole,

primrose willowLycopodiella cernua (L.) Pic. Serm. (Lycopodiaceae),

wawae‘ioleMachaerina angustifolia (Gaud.) T. Koyama (Cyperaceae),

‘ukiMangifera indica* L. (Anacardiaceae), manako, mangoMecodium recurvum (Gaudich.) Copel.

(Hymenophyllaceae), ‘ohi‘a ku, filmy fernMelaleuca quinquenervia* (Car.) S. T. Blake (Myrtaceae),

paperbark treeMetrosideros polymorpha Gaud. (Myrtaceae), ‘ohi‘a lehua,

endemicMetrosideros tremuloides (A. Heller) P. Knuth

(Myrtaceae), lehua ‘ahihiMimosa pudica* L. (Fabaceae), pua hilahila, sensitive

plantMorinda citrifolia* L. (Rubiaceae), noni, Indian mulberryMyrsine lessertiana A. DC (Myrsinaceae), kolea lau nuiNephrolepis multiflora (Roxb.) F. M. Jarrett ex C. V.

Morton (Nephrolepidaceae)Nestegis sandwicensis (A. Gray) Degener (Oleaceae),

olopuaOchrosia compta K. Schum. (Apocynaceae)Ophioglossum polyphyllum A. Braun (Ophioglossaceae),

pololei, adder’s tongueOplismenus hirtellus* (L.) P. Bear. (Poaceae), honohono,

basketgrassOsteomeles anthyllidifolia (Sm.) Lindl. (Rosaceae), ‘uleiPandanus tectorius S. Parkinson ex Z (Pandanaceae), hala

treePanicum maximum* Jacq. (Poaceae), Guinea grassParaserianthes falcataria* (Fabaceae), albiziaPaspalum conjugatum* Bergius (Poaceae), Hilo grassPaspalum scrobiculatum* L. (Poaceae), mau‘u laiki,

ricegrassPaspalum urvillei* Steud. (Poaceae), Vasey grassPassiflora edulis* Sims (Passifloraceae), passion fruit,

liliko‘iPassiflora foetida* L. (Passifloraceae) pohapoha, wild

water lemonPassiflora mollisima* (Kunth) L. H. Bailey

(Passifloraceae), banana pokaPassiflora pulchella* Kunth (Passifloraceae), two-lobed

passion flower, naturalizedPassiflora suberosa* L. (Passifloraceae), huehue haole,

introduced; wild passion flower

Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 313

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Appendix (continued)

Peperomia oahuensis C. DC (Piperaceae)Phaius tankarvilleae* (Banks ex L’Her.) Blume

(Orchidaceae), Chinese ground orchidPhyllostachys nigra* (Lodd.) Munro (Poaceae), black

bambooPhymatosorus grossus (Langsd. & Fisch.) Brownlie

(Polypodiaceae), laua‘ePiper methysticum* G. Forster (Piperaceae), ‘awa, kavaPipturus albidus (Hook. & Arnott.) A. Gray (Urticaceae),

mamakiPisonia umbellifera (G. Forster) Seem. (Nyctaginaceae),

papala kepauPityrogramma austroamericana* Domin (Pteridaceae),

goldback fernPityrogramma calomelanos* (L.) Link (Pteridaceae),

silverback fernPluchea symphytifolia* (Mill.) Gillis (Asteraceae), sourbushPolypodium pellucidum Kaulf. (Polypodiaceae), ‘aePritchardia martii (Gaud.) H. A. Wendl. (Arecaceae),

loulu hiwaPsidium cattleianum* Sabine (Myrtaceae), waiawı,

strawberry guavaPsidium guajava* L. (Myrtaceae), kuawa, common guavaPsilotum complanatum* Sw. (Psilotaceae), moa, moa haolePsilotum nudum (L.) P. Beauv. (Psilotaceae), moaPsychotria kaduana (Cham. &. Schlechtend.) Fosb.

(Rubiaceae), kopikoRhizophora mangle* L. (Rhizophoraceae), American or

red mangroveRhynchospora sclerioides (Hook. & Arnott) (Cyperaceae)Ricinus communis* L. (Euphorbiaceae), kolıRubus rosifolius* Sm. (Rosaceae), ‘akala, thimbleberrySacciolepis indica* (L.) Chase (Poaceae), Glenwood grassSadleria pallida Hook. & Arn. (Blechnaceae), ‘ama‘uSalvia coccinea* Etl. (Lamiaceae), lılılehua, scarlet sage or

Texas sageSantalum ellipticum Gaud. (Santalaceae), ‘iliahialo‘e, coast

sandalwoodScaevola gaudichaudiana Cham. (Goodeniaceae), naupaka

kuahiwiScaevola sericea Vahl. (Goodeniaceae), naupaka kahakaiSchefflera actinophylla* (Endl.) Harms (Araliaceae),

octopus treeSchinus terebinthifolius* Raddi (Anacardiaceae), wilelaiki,

Christmas berry treeSchizostachyum glaucifolium* (Rupr.) Munro (Poaceae),

‘oheScirpus sp. (Cyperaceae)

Selaginella arbuscula (Kaulf.) Spring (Sellaginellaceae),lepelepe a moa

Setaria gracilis* Kunth (Poaceae), mau‘u Kaleponi,yellow foxtail

Setaria verticillata* (L.) P. Beauv. (Poaceae), mau‘upilipili, bristly foxtail

Spathodea campanulata* P. Beauv. (Bignoniaceae), Africantulip tree

Spathoglottis plicata* Blume (Orchidaceae), Malayan orPhilippine ground orchid

Sphaerocionium obtusum (Hook. & Arn.) Copel.(Hymenophyllaceae), palai lau li‘i, small-leaved fern

Sphenomeris chinensis (L.) Maxon (Lindsaeaceae), pala‘a,lace fern

Stachytarpheta dichotoma* (Ruiz & Par.) Vahl(Verbenaceae), owı, oı weed

Stachytarpheta jamaicensis* (L.) Vahl (Verbenaceae), owı,oı, Jamaican vervain

Styphelia tameiameiae (Cham. & Schlechtend.) F. v.Muell. (Epacridaceae), pukiawe

Syzygium cumini* (L.) Skeels (Myrtaceae), Java plumSyzygium jambos* (L.) Alston (Myrtaceae), ‘ohi‘a loke,

rose appleSyzygium malaccense* (L.) Merr. & Perry (Myrtaceae),

‘ohi‘a ‘ai, mountain appleSyzygium sandwicensis (A. Gray) Nied. (Myrtaceae), ‘ohi‘a

haTectaria gaudichaudii (Mett.) Maxon (Dryopteridaceae),

‘iwa‘iwa lau nui, large-leaved ‘iwa‘iwaTerminalia catappa* L. (Combretaceae), kamani haole,

Indian almondTetraplasandra gymnocarpa (Hillebr.) Sherff (Araliaceae),

‘ohe‘oheTouchardia latifolia Gaud. (Urticaceae), olonaTrema orientalis* (L.) Blume (Ulmaceae), gunpowder

treeUrena lobata L. (Malvaceae), araminaUrera glabra (Hook. & Arnott) Wedd. (Urticaceae),

opuheVandenboschia davallioides (Gaudich.) Copel.

(Hymenophyllaceae), palai hihi, filmy fernVerbena litoralis* Kunth (Verbenaceae), owıVittaria elongata Sw. (Vittariaceae), ‘ohe‘oheWedelia trilobata* (L.) Hitchc. (Asteraceae)Wikstroemia oahuensis (A. Gray) Rock (Thymelaeaceae)

‘akiaZingiber zerumbet* (L.) Sm. (Zingiberaceae), ‘awapuhi

kuahiwi, shampoo ginger

Note: After Wirawan (1978); updated by D.M.-D. in 2003. Names are from Wagner et al. (1990) and Palmer (2003).* Nonnative.

PACIFIC SCIENCE . April 2005314