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 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 functions but also to investigate their rela- Transect) network aims to include broad tionships as a landscape unit. The latter is landscape units with three interconnected necessary to ensure that the management of biodiversity resource zones into a study pro- one resource zone does not affect the other gram. The three resource zones are an upland zones negatively and that the zonation system or inland forest system that serves as water- is managed as an integrated landscape unit. shed cover, the agroecosystems that are usu- Such is the management unit or human sup- ally below in elevation, and the coastal zone port system developed (through trial and er- ecosystems. A freshwater resource system is ror) by the early Hawaiians before contact another essential component. with Western society and named ahupua‘a. The objective is to study each of these eco- system zones individually for their internal 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 1 Manuscript accepted 20 April 2004. 2 Department of Botany, University of Hawai‘i at in the ocean water. Ahupua‘a is a Hawaiian Ma¯noa, 3190 Maile Way, Honolulu, Hawai‘i 96822. term. It refers to ahu, meaning a heap of 3 Deceased, former Director, World Wide Fund, stones, and pua‘a, the Hawaiian word for pig. Indonesia. A related, second meaning of the term ahu- pua‘a refers to an altar on which a pig was Pacific Science (2005), vol. 59, no. 2:293–314 placed as a tribute to the chief or landowner : 2005 by University of Hawai‘i Press for allowing the use of the ahupua‘a land. All rights reserved The landlord or owner and manager of an

293 294 PACIFIC SCIENCE . April 2005

Figure 1. Ahupua‘a model, after Minerbi (1999). ahupua‘a was called konohiki (Pukui and El- nahele), which may include a cloud forest bert 1986). (wao akua [akua means god]); the agricultural Luciano Minerbi (1999) described a typical zone (wao kanaka [where people work]); and ahupua‘a diagrammatically as here shown in the coastal zone (makai or kahakai [kai means Figure 1 with some slight modification em- ocean, and kahakai means beach or sea- phasizing ecotones (i.e., transition zones). As shore]). A fourth important resource zone, as seen here, an ahupua‘a consists essentially of shown on Figure 1, is the freshwater stream three vertically changing biological resource (kahawai). The stream flow is modified in zones: the upland (mauka) forest zone (wao the wao kanaka by man-made ditches (‘auwai) Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 295 for irrigating taro (Colocasia esculenta) patches elevation behind the ridge extension of Pu‘u (lo‘i) on alluvial terraces. Dryland root crops, o Kila. In front of the Pu‘u o Kila ridge is including taro and sweet potato (‘uala, Ipo- Kawa Stream, which joins Kahana Stream in moea batatas), were grown in the opened-up the midsection of the valley. uplands (kula uka). Breadfruit (‘ulu, Artocar- tenure history. The Kahana ahupua‘a pus altilis), bananas (mai‘a, Musa paradisiaca), was originally a communal property of the and coconut (niu, Cocos nucifera) were among Hawaiian people. It is estimated that from the most common tree crops planted in the 720 to 1,000 indigenous people were living lowlands and midlands. and supporting themselves from the resources Beyond crop areas, the ahupua‘a contains of the valley and the ocean bay at the time makai-mauka (toward the sea–toward the of Western contact in 1776 (Stauffer 1990). mountain) access trails as well as midelevation Over the following 100 yr, the indigenous and coastal trails, house sites (kauhale), tem- population declined to an estimated low of ples (heiau), and burial sites (pa¯ ilina). A ca. 90 people. In the late nineteenth century, most important coastal resource enhance- it rose again to ca. 250 inhabitants, primarily ment includes a fishpond (loko i‘a). This is due to Chinese immigrants. Major factors for a stonewalled enclosure in shallow coastal the rural decline of the indigenous people water that allows small fish to enter through were introduced diseases, the lure of the city, sluice gates (ma¯ka¯ha¯). After gaining in size, Honolulu, and the great mahele (land divi- the fish is trapped. Thus, a fishpond acts as a sion) of 1846–1855 with its ensuing privatiza- large fish trap. tion of native lands and the change to a The two man-made devices, the irrigated foreign cash economy. terraces (lo‘i) and fishponds, are clear evi- The great mahele resulted in a four-way dence of the agricultural engineering talents division of the land. Recipients were (1) of the early Hawaiians to make full use of the king, (2) the other high ali‘i, (3) the the productive capacity of their windward American-dominated government, and (4) ahupua‘a. the maka‘a¯inana (the common people). Stauffer (1990) documented the changes in land tenure of Kahana Valley from 1846 to The Kahana Ahupua‘a and Its Tenure History 1920. The great mahele allowed the maka‘a¯i- location. The Institute of Hawaiian nana to claim their kuleana (i.e., their house Studies, University of Hawai‘i at Ma¯noa, pre- lots and cultivated areas). This amounted to pared a map of O‘ahu Island (Figure 2), only 200 acres (ca. 81 ha). The remaining un- which shows the traditional subdivisions into cultivated land, ca. 5,000 acres (ca. 2,024 ha), five districts, called moku, and numerous ahu- became the property of the high ali‘i Annie pua‘a, all designated by names with bound- Keohoka¯lole in 1847. She was the mother of aries. Kahana stands out on windward, central King David Kala¯kaua and his sister, Queen O‘ahu in the south of Ko‘olauloa moku. Lili‘uokalani, who reigned from 1891 to A recent photo taken from a low-flying 1893. Annie Keohoka¯lole and her husband aircraft (Figure 3) shows Kahana Valley with had large land holdings but little money. To its restored fishpond at lower left (the Huilua support their lifestyle they needed money. Fishpond), the Kahana estuary next to it, its Loans were given by foreign businessmen us- coastal ironwood (Casuarina equisetifolia) for- ing land as security. Eventually, 3,000 acres est, the verdant central marsh, and forested (ca. 1,215 ha) of the ahupua‘a were sold by bottomlands. The slopes show minor land Keohoka¯lole to a Chinese merchant named scars, but are mostly forested. The peak on Ah Sing in 1857. He became the konohiki. the left is Pu‘u o Kila, 467 m (1,530 feet His son, Ah Mee, owned the property in high), and in the background, about 6 km in- 1872. Figure 4 shows Kahana Valley in a land, is the Ko‘olau mountain range with photo taken in 1888. Note absence of forest summit elevations of ca. 610 m (2,000 feet). on foothills and presence of grazing animals. Kahana Stream originates at about 300 m The maka‘a¯inana soon began to realize Figure 2. O‘ahu Island, Hawai‘i Archipelago, showing the five moku (i.e., larger island sections) and ahupua‘a, with Kahana ahupua‘a and PABITRA site shaded in the Ko‘olauloa moku on the windward side of the island (from Institute of Hawaiian Studies, University of Hawai‘i at Ma¯noa). Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 297

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 on frontal 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. that through privatization they lost their of the hapa haole (half foreigner) Robinson ‘‘cooperative rights’’ to the ahupua‘a. These shipbuilder family. Mary Foster’s husband, rights relate to using the uncultivated land Thomas Foster, was a Canadian businessper- with the freshwater stream that fed their lo‘i son who owned interisland shipping, which via their ditch system (the ‘auwai), the use of later became the Matson Navigation Com- forest products and gathering rights as well as pany. communal fishing rights. Mary Foster was one-quarter Hawaiian, In 1874 a Kahana hui (association) under had close connections to the royal family, George William Kamakaniau bought the Ka- and spoke fluent Hawaiian. She eventually hana ahupua‘a back from Ah Mee for $6,000. became the konohiki and prime owner of At that time, a Hawaiian again became the the Kahana ahupua‘a by 1920. But she konohiki. However, the money was bor- struggled with two other haole landowners, rowed as a mortgage. This was made possible the Kaneohe Ranch Co. and the McCandless through the Mortgage Act of 1874. Eventu- brothers. The Kaneohe Ranch Co. acquired a ally the mortgage terms could not be met by lease from the Hawaiian hui for Kahana’s the Hawaiian hui, and foreclosure resulted in sloping uplands for cattle grazing. They the transfer of ahupua‘a property rights to burned much of the coastal upland and native non-Hawaiian businesspeople. Among these midland forest for conversion into pasture- were the Castle family who owned the Ka- land. The McCandless brothers acquired neohe Ranch Co., and Mary Foster, daughter rights to the mauka uplands. As professional 298 PACIFIC SCIENCE . April 2005

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

artisan well borers, they realized the poten- 1967. It is now public property and called of- tial of tapping the windward groundwater ficially the Ahupua‘a ‘O Kahana State Park. resources halfway up the Ko‘olau mountain Figure 6 shows the Kahana ahupua‘a in a range. They designed the Kahana-Waika¯ne- photo taken soon after World War II. Note Waia¯hole Ditch and mountain tunneling the barren front hills and dry terraces (kula system. The work was carried out by Asian lands). miners from 1913 to 1916. This was a great engineering accomplishment at that time, Kahana Vegetation which made the irrigated sugarcane agricul- ture at ‘Ewa basin possible. However, legal vegetation map. Nengah Wirawan battles arose over the land and water rights prepared a vegetation map in 1972 (Figure 7) with the konohiki of Kahana, Mary Foster. in conjunction with a botanical survey of She received an out-of-court settlement, and Kahana Valley (Theobald and Wirawan the McCandless brothers made a fortune by 1973). He subsequently produced a Ph.D. selling their Waia¯hole-Waika¯ne Water Co. dissertation (Wirawan 1978) on the vegeta- to Amfac, owners of the O‘ahu Sugar Co. in tion ecology of Kahana under guidance of ‘Ewa. Figure 5 shows the entrance to Kahana D.M.-D. At that time, the air photo–derived Valley from Kamehameha Highway and the vegetation map was verified with 64 vegeta- Huilua Fishpond in a photo taken in 1936. tion releve´s (sample stands of 400 m2 each) After Mary Foster’s death, the Kahana spread out from the coastal area into the ahupua‘a reverted to the State of Hawai‘i in back of the valley up to the base of the steep Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 299

Figure 5. Kahana Valley in 1936 during Mary Foster’s ownership. The valley was given to rice cultivation and cattle ranching. The Waika¯ne/Waia¯hole Ditch and tunnel system was emplaced in the back of the valley. Photo 38215 AC, courtesy of Bishop Museum Archives. wet cliff area where it rises from ca. 300 to 1972 were occasional trees of Spathodea cam- 610 m over a short distance of ca. 200 m. Fig- panulata (African tulip tree) and the shrub ure 7 is a reproduction of Wirawan’s vegeta- Clidemia hirta (Koster’s curse). tion map. It shows 13 units. Unit 3 represents an entirely alien tree Unit 1 relates to the wet, sparsely vege- planting of Paraserianthes falcataria, known tated, nearly vertical cliff area (pali) in the commonly as the tall albizia legume tree. back of the valley. There are also pali seg- This tree planting indicates the Ko‘olau ments on the eastern side, rendering Kahana Mountain ditch and tunnel system, developed an aging ‘‘amphitheater-headed valley.’’ (For between 1913 and 1916 for transferring growing on wet and dry cliffs, see some of the freshwater flow and the upper Mueller-Dombois and Fosberg 1998:488.) mountain-stored water for irrigating the for- Unit 2 is designated as Metrosideros wood- mer sugarcane fields in the ‘Ewa basin above land. The key species are clumps and scat- Pearl Harbor. The ditch system begins here tered trees of Metrosideros polymorpha (‘o¯hi‘a in the upper Kahana back valley and contin- lehua) together with the indigenous thick ues through the neighboring Waika¯ne and mat-forming uluhe , Dicranopteris linea- Waia¯hole ahupua‘a to the south. ris. Other important native trees are Euge- Unit 4 is recognized as the Acacia koa– nia sandwicensis (ha¯), Antidesma pulvinatum Pandanus (hala tree) woodland. Both koa and (hame), Psychotria (ko¯piko), small stands of hala are native trees. Most others are intro- Pritchardia palms (loulu), and some duced. The latter include the Polynesian in- tree (ha¯pu‘u). Important aliens found in troduction (‘o¯hi‘a ‘ai), 300 PACIFIC SCIENCE . April 2005

Figure 6. Kahana Valley in 1945 with sugarcane on lower left, open slopes above, smoke from a fire at right, and army installations above bottomland. Photo USA MH 5408, courtesy of Bishop Museum Archives. which is common throughout the valley and tionship has so far been ascertained (Puni- well represented by seedlings and saplings, as wai 1997). The disturbance history is most well as an occasional Artocarpus altilis (the ‘ulu likely involved. Abundant koa regeneration or breadfruit tree). Other later-introduced has been observed after fire ( Judd 1935) fruit trees include the mango tree (Mangifera or browser removal (Spatz and Mueller- indica), guava or kuawa (Psidium guajava), and Dombois 1973). After such events, koa re- the strawberry guava or waiawı¯ ‘ula‘ula (Psi- generates in cohorts (i.e., generation stands). dium cattleianum). The latter is considered a Once these cohort stands become senescent, serious weed tree; there are some large (about any temporary climatic stress can trigger 0.5-m diameter) strawberry guava trees in such tree groups into dieback because of Kahana Valley, signifying an introduction in their demographic predisposition. Insects and the late 1800s. Seedlings and saplings are not fungi often merely hasten the dieback or abundant here. Instead, a new invader, Ardisia decline because such trees have lost their ca- elliptica (the shoebutton tree), is rapidly be- pacity to regenerate foliage. If these pest or- coming abundant in forest openings of unit ganisms were constantly virulent, trees would 4. Koa saplings are sparse. Wirawan (1978) loose their foliage as soon as it forms. Of found that removal of the hala leaf litter course, younger koa trees may also become and some soil surface scarification encourages predisposed to die (for example, in this case, koa regeneration in unit 4. Some areas in this through competition with hala trees, which unit currently exhibit koa dieback or decline. have now gained dominance over koa in unit A number of insects and fungal pathogens 4). Application of some sensible silviculture have been implicated but no causative rela- could reverse the trend. Figure 7. Vegetation map of Kahana Valley after Wirawan (1978), slightly modified and updated. Solid lines labeled TR1–TR4 represent transects; TR1 is portrayed as a profile diagram in Figure 9. 302 PACIFIC SCIENCE . April 2005

Unit 5 (crosshatched on the map), desig- rophytic grass introduced from the southeast- nated as forest scrub, consists primarily of ern United States in 1924. Its spread was small, pure-stand patches of Syzygium cumini promoted by fire, and it contributed sub- ( Java plum), Syzygium jambos (rose apple or stantially to soil avalanching on slopes with ‘o¯hi‘a loke), and stands of Phyllostachis nigra deeply oxidized soil (ultisol) here in Kahana (black bamboo), and Schyzostachium glaucifo- Valley and in many other windward valleys lium (‘ohe, possibly of Polynesian introduc- (Mueller-Dombois 1973). The grass-scrub tion). This unit grades into the riparian Unit cover is now mostly invaded by the octopus 8. tree. However, a native shrub, Osteomeles Unit 6 is the Aleurites moluccana (kukui) anthyllidifolia (‘u¯lei), still prevails in the un- tree community of Polynesian introduction. dergrowth on sun-exposed slopes, where It fills the gulches on both the north and Andropogon has become very sparse. The oc- south slopes from near the coast to the mid- topus tree is still on the move and currently dle of the valley. It is most prevalent in the is invading unit 4 farther inland as well as mesic part of the valley and drops out where unit 11. the mean annual rainfall exceeds 4,000 mm. Unit 11 is primarily marsh-grassland of Wirawan (1978) found that kukui tree regen- Brachiaria mutica (California grass). Like the eration is related to storm surges, when tum- broomsedge it was first collected in Hawai‘i bling rocks in the gulches happen to crack the in 1924 (Wagner et al. 1990). At the time of accumulated kukui nuts on the forest floor. mapping the marsh, which gets less swampy Unit 7 is the Acacia koa–Pandanus forest, a inland, this unit served as cow pasture, and variation of unit 4, recognized by taller and in the wetter parts rice was still cultivated in denser tree growth indicating either a more the mid 1970s. productive habitat or less disturbed vegeta- Unit 12 represents the dry coastal cliff tion. community, sparsely vegetated with wisps of Unit 8 is dominated by the hau tree, Hi- grasses including Heteropogon contortus (pili biscus tiliaceus, which is either native or an- grass) and Rhynchelytrum repens (Natal red- other Polynesian introduction (Wagner et al. top). 1990). It forms a jungle or krummholz and Unit 13 relates to the strand community, covers many former irrigated terraces (lo‘i). which at Kahana Bay is dominated by an It grows over sections of Kahana Stream and ironwood (Casuarina equisetifolia) forest with forms a dense cover all along its estuary. a row of Terminalia catappa (kamani haole) There it grows intermixed with the intro- trees along Kamehameha Highway. An up- duced American mangrove (Rhizophora man- dated checklist is included in the Ap- gle). pendix. Unit 9 is occupied dominantly by the octo- physiographic environment. Impor- pus tree (Schefflera actinophylla). It is a fast- tant geomorphic features of Kahana Valley growing secondary rain-forest tree, probably are displayed on the physiographic map (Fig- introduced from Australia as an ornamental. ure 8). The figure shows the physiography Wagner et al. (1990) cited Degener as having and topographical diversity by 200-ft (61-m) said that the octopus plant was grown in pots contour lines. Steep slopes are indicated by and Hawaiian gardens as an ornamental in crowding of the contour lines. The back val- 1900. In Kahana, this tree has rapidly invaded ley rises steeply to the Ko‘olau summit ridge. the formerly open grass-covered slopes (kula Another ridge, topped by the 1,530 ft (467 m) kai) on both sides of the lower valley from Pu‘u o Kila, juts into the valley along the the coast to about 1.5 km inland. It is now 5,000-mm precipitation line. A third ridge also invading the marsh (unit 11). with a steep slope down into the valley is Unit 10, designated as mixed grass-scrub found on the east side near the coast. Wig- on the map, has now mostly disappeared. gling of the contour lines indicates numerous In 1972, this was dominantly occupied by steep gulches along each of the slopes, which Andropogon virginicus (the broomsedge), a py- from near the coast to beyond the midvalley Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 303 are dominated by kukui, unit 6 on the vegeta- Zone C, wao koa, relates to the Acacia koa– tion map. Pandanus area. Zone D, kula kai, refers to stream system and precipitation. the formerly more open grassy slopes, now The Kahana Stream system is shown in Fig- occupied mostly by introduced octopus trees. ure 8 as originating in the back of the valley Zone E, kahakai, relates to the coastal strand, at about 1,000 ft (305 m) elevation. The with Casuarina forest, mixed grass-scrub, and dashed line running between 800 (244 m) fishpond as well as into the sea up to about and 1,000 ft (305 m) elevation indicates the 1.5 m depth. ditch where fresh water is extracted for trans- water yield in kahana valley. Taka- fer to leeward O‘ahu. The freshwater stream saki et al. (1969) estimated the water yield for system follows a course that relates directly Kahana Valley. They calculated a total input to the topographic pattern of the valley. from precipitation amounting to 60 million Another important feature is the precipitation gallons (240 million liters) per day. This esti- gradient, which starts at the coast with a mean mate appears reasonable. A relationship of annual rainfall of 1,875 mm (74 in.). In the height to area, which is easy to remember, is midvalley where Kawa Stream joins Kahana that 1 mm/m2 yields 1 liter. If one considers Stream, the mean annual precipitation is the effective catchment area to be 2,000 ha 2,500 mm (98 in.). From there rainfall dou- (the Kahana ahupua‘a is estimated to include bles toward the back valley, where rainfall ex- 5,200 acres) and the average daily precipita- ceeds 7 m (275 in.) per year. Thus, there is a tion to be 12 mm, the incoming yield would steep rainfall gradient. From the coast to the be 12 mm 2,000 ha ¼ 240 million liters or back valley rainfall increases from ca. 2 m to 7 ca. 60 million gallons. m over a distance of 6 km. The four physical Takasaki et al. (1969) considered the fol- environmental features of topography, geo- lowing fractionation of their estimated 60 morphology, stream system, and precipitation million gallons (240 million liters) input per pattern explain much of the observed vegeta- day: 12 million gallons (48 million liters) are tion pattern. lost as evapotranspiration (20%), 15 million topographic profile with climate gallons (60 million liters) are lost as runoff diagram. In addition, there is a gentle slope (25%), 19 million gallons (76 million liters) in the center of the valley, descending in step- enter the stream system (32%), 4 million gal- like intervals from mauka to makai, as seen on lons (16 million liters) are removed by the the profile map along Transect 1 (Figure 9), Kahana-Waia¯hole Ditch system (6%), and which explains the direction of stream flow. 10 million gallons (40 million liters) leave The climate diagram substantiates the great the area as groundwater (17%). This gives differences in rainfall from mauka to makai. some idea of the valley’s water yield poten- It also shows how rainfall is distributed tial. Further direct measurements at different throughout the year, indicating a short dry rainfall events including storm surges and dry period in June/July at the coast versus periods would be of considerable interest and monthly rainfall in excess of 300 mm inland practical value because this is known to be a at the Kahana Valley ditch and tunnel system. highly dynamic relationship. Here, one can see that it is also cooler by archaeological map. Hommon and 1.5C throughout the year. Barrera (1971) found 119 archaeological sites hawaiian ecological zones. The pro- in Kahana Valley. They produced a foldout file also shows an attempt to recognize the map that shows all of the site locations with ecological zonation system developed by an outline of their spatial extent. This map is the early Hawaiians. It is here related to here reproduced as Figure 10 and superim- topography, precipitation, stream flow, and posed on the vegetation map units drawn vegetation in five zones from mauka to makai. at the same scale. The archaeological map Zone A, kua mauna, refers to the wet cliff shows that former human habitation sites (pali) area in the back valley. Zone B, wao extend about 5 km into the valley along the kele, refers to the wet Metrosideros forest. streams. The authors recorded 120 wet ter- Figure 8. Physiographic map of Kahana Valley. Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 305

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

races (lo‘i) connected by man-made irrigation by removal of stones. Terraces marked by channels (‘auwai). Most of these terraces were stone walls were often rather small, only 2 by built on gently sloping alluvial areas with re- 10 m. In addition, the authors identified taining walls. The total growing surface of animal enclosures, house sites, men’s eating the wet terraces was calculated as 10,800 m2 houses (mua), and scarcely legible petro- (1.08 ha). The authors suggested that many glyphs. The archaeological sites coincide lower terraces were probably buried under rather well with three current vegetation alluvial material. A second archaeological fea- types, the mixed forest scrub (unit 5), the Hi- ture was clearings for the cultivation of non- biscus (hau) forest (unit 8), and the pasture and irrigated crops (probably dry taro and sweet marsh (unit 11) as shown on the vegetation potato). These dry terraces were recognized map. Figure 10. Archaeological map outlining the realm of the known archaeological sites discovered by Hommon and Barrera (1971). Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 307

Kahana Valley, a Geomorphic Artifact? (1990) proposed that Kahana Valley was abandoned for at least 1,200 yr and was rein- a former inland lagoon. Patricia Rice vaded by a new wave of indigenous settlers Beggerly (1990) presented a very thorough at about 1200 a.d. (i.e., 800 b.p.) at the ear- and thought-provoking Ph.D. dissertation on liest. She proposed that the earlier settlers, her paleoecological research in Kahana Val- who came into the valley around 230 G 200 ley. She considered Kahana Valley to be a b.c., experienced an ‘‘ecological crunch’’ due ‘‘geomorphic artifact’’ based on past human to their overexploitation of the valley’s re- impacts. She dug deep soil trenches in eight sources, with the ecological crunch having areas at the borders of the marsh and in the resulted primarily from their traditional pasture up to 1.2 km inland. She used open slash-and-burn practice. This would have in- trenches and auger borings to distinguish soil volved accelerated soil erosion and nutrient layers with inclusions of organic remains such loss with slow growth of vegetation, thus as snail shells, charcoal, wood, pollen, and shifting activity across the sloping landscape. fern spores. For charcoal, shells, and some This also implied exhaustion of readily avail- woody remains, she obtained radiometric able food resources, such as starch from tree- dates. She distinguished seven soil strata fern trunks and fish from the inland lagoon from about 3 m depths upward to the current as it continued to decline from accelerated soil surface and clarified that the valley bot- sedimentation. At that time, the first settlers tom was once an ocean inlet, a finding veri- could still migrate to another empty valley. fied also for Kawainui marsh in the Kailua Beggerly argued that the first settlers used ahupua‘a, farther south on windward O‘ahu tree-fern starch as a food resource because (see Kirch 1985). This was not surprising be- tree fern spores showed an early ebb in her cause there was a higher sea level following soil core samples. the Pleistocene. A shallow-water phase suc- She found Pandanus pollen in the lagoon ceeded in the form of a lagoon. As evi- phase, which may explain today’s presence dence, Beggerly found shells of brackish- of the many hala trees on the midslopes of water snails, such as Tellima macoma dispar. both sides of the valley. Pandanus is usually In the lagoon phase, at the bottom of the found in coastal settings near the ocean shore. marsh, she found charcoal. Dates for these She also found ebbs in Metrosideros pollen organic remains turned out to be 430–30 b.c. and tree-fern spores alternating with ebbs of Beggerly suggested that the charcoal remains Pandanus pollen and concluded that these al- argue for a first invasion of human settlers ternations may be the result of human inter- into Kahana Valley by at least 30 b.c. She be- vention with Kahana’s rain forest. Athens lieved that wood fires could not have started and Ward (1993), doing similar coring studies in this rain-forest valley because of its year- in Kawainui marsh, showed the startling re- round high humidity. On the shallow lagoon sult of an abundance of Pritchardia (loulu) bottom she found colluvial material as the palm pollen before the first centuries of Poly- next deposit upward. This material, she sug- nesian land occupation. They found that the gested, was deposited from the slopes by loulu palm pollen began to disappear around accelerated erosion, when the first human set- 1200 a.d. and suggested disease rather than tlers used slash-and-burn swidden cultivation. human-induced fire as the cause. They also Following the colluvial layer upward, mineral hold that loulu palm forest was the dominant soil and organic muck mixtures indicate con- lowland vegetation type on windward O‘ahu tinued terrestrification. However, there was when the first human settlers arrived. Because no charcoal inclusion until she came nearer Patricia Beggerly did not find loulu palm pol- to the surface (between 2.2 and 2.5 m). The len in her corings of Kahana marsh, it seems second find of charcoal was dated 1425 G worthwhile to reinvestigate this area, particu- 80 yr a.d. larly because loulu palms are still present in pandanus versus pritchardia pollen. Kahana’s back valley today. The same applies Based on this second charcoal date, Beggerly to koa tree pollen, which was not found by 308 PACIFIC SCIENCE . April 2005

Beggerly in her soil cores. However, koa pol- to migrate into neighboring valleys. At that len was found by Athens et al. (1992), even in time, such valleys may have been unoc- areas that lack Acacia koa today. Athens and cupied. Ward (1993) did not dispute Beggerly’s early Phase 2: From 30 b.c. to 1200 a.d., Kahana human invasion date but suggested resam- Valley was abandoned. The indigenous forest pling her charcoal findings for comparability regenerated. Fewer soil avalanches occurred. with other areas of O‘ahu. The marsh formed by further, mostly or- ganic, terrestrification processes and herba- ceous cover dominated on the valley floor. Two Paleoecological Land Use Models Phase 3: From 1200 to 1800 the second continuous succession of human wave of settlers had arrived. They again occupation. Patrick Kirch (2000) provided started with slash-and-burn agriculture. A an interesting theory of a successional se- second phase of environmental degradation quence of the Hawaiian cultural develop- ensued but not for long. This time, the set- ment. He divides it into four phases: phase 1: tlers did not move out of Kahana Valley. from 300 to ca. 600 a.d., the colonization pe- Because the population had expanded in all riod; phase 2: from 600 to ca. 1100 a.d., the windward valleys, space had become limited. development period; phase 3: from 1100 to In conjunction with swidden cultivation by ca. 1650, the expansion period; and phase 4: removing forest for growing crops on dry from 1650 to Western contact (1776), the land, the settlers began irrigation agriculture protohistoric period. by constructing wet terraces (the lo‘i) and During phase 1, the prevailing agricultural converting stream water through ditch sys- technology was shifting cultivation, as still tems (the ‘auwai). practiced today, for example, in parts of Va- Phase 4: From 1800 to 1900, after West- nuatu, Solomon Islands, and New Guinea. ern contact, a third phase of forest degra- Taro irrigation in windward valleys had al- dation began, combined with accelerated ready begun in the colonization phase. It did erosion. Early in this phase, ca. 1825, as told not develop into a more widely used agricul- in written historical accounts, slopes were tural technology until the beginning of the brown and bare and forest was confined to expansion phase, around 1100 a.d. Its greatest the inner valley and the upland region. The sophistication was reached during the proto- western slopes were still cultivated in spots historic phase from 1600 to 1800 a.d. with dryland taro, banana, and sweet potato, discontinuous succession of human and the riparian lowlands with irrigated taro occupation. Patricia Beggerly’s (1990) in spotty wet terraces. The traditional use model, based directly on her research in Ka- changed with overseas immigrants: Eastern hana Valley, suggested four different phases, agricultural practices of rice growing and as follows. Western practices of cattle grazing were in- Phase 1: Between 400 and 30 b.c., the first troduced. In addition to the free-roaming cat- human settlement occurred. She suggested tle, other hoofed animals such as goats, sheep, that it involved ca. 100 people. They initially and pigs were using the valley’s vegetation. used tree-fern starch as a food source while This was followed by sugarcane growing in they began swidden cultivation with ti (Cor- the pasture area with the Castle-owned Ka- dyline terminalis) and taro (Colocasia esculenta) huku Railroad reaching south across Kahana through opening the forest by cutting it Stream into the eastern valley in the late down with subsequent burning of the slash. eighteenth to early nineteenth century. Dur- In so removing the forest on slopes, they ing World War II Kahana Valley was used caused soil avalanching, which resulted in the as a military training area. Remnant bunker accumulation of colluvium in the lagoon structures are still found on the loop trail in phase at the valley bottom. That also de- the back part of the valley. Related to this is stroyed the fish populations in the lagoon. the tragic story of land tenure, the changes Overexploitation forced the first colonizers in land ownership as discussed earlier. Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 309

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

sea level changes. Patrick Nunn’s second group of settlers remained in the (1999) diagram of sea level changes over the valley. They soon changed their land-use last 1,200 yr (Figure 11) provides further in- practices from degrading the environment to sights. The first human colonization phase enhancing its natural resources. They accom- certainly had occurred before 800 a.d. (i.e., plished this by developing irrigation schemes 1200 b.p.) when the sea level was perhaps and removing stones from dryland terraces 2 m higher than today. The bottomland of for growing newly introduced crop plants, Kahana Valley was then a marine embayment such as sweet potato (Ipomoea batatas, ‘uala), up to the inland margin of the current pas- as well as by constructing fish traps culminat- ture, or meadow (ca. 1.2 km inland of the ing in the Huilua Fishpond. The latter is said current Kahana beach), as confirmed by Pa- to have been developed about four centuries tricia Beggerly (1990). Kahana’s inland marine ago. bay changed into a lagoon most likely when According to Patrick Kirch’s (2000) suc- the sea level dropped thereafter to about 1 m cessional interpretation of human impact on above the current level. According to Beg- windward O‘ahu, there was only one phase gerly’s interpretation, the first settlers had of settlement. Colonization with slash-and- already abandoned Kahana Valley due to burn practices occurred only once. This was overexploitation, and the valley was experi- ca. five centuries before the Little Climatic encing a fallow period of about 12 centuries Optimum. During the warmer and calmer until ca. 800 b.p. (i.e., 1200 a.d.). Then the climatic phase, when ocean canoe travel was second wave of settlers arrived eight centuries common practice, Kirch’s development phase ago, during the ‘‘Little Climatic Optimum.’’ gave way to his expansion phase ca. nine cen- They again began to use Kahana Valley by turies ago. From that time onward, the Ha- practicing slash-and-burn agriculture. This waiian land-use practices started to enhance 310 PACIFIC SCIENCE . April 2005 the natural ecosystem services through con- equally apparent, if not of greater impor- servation practices. This then became the tance. traditional Hawaiian ahupua‘a management Moreover, Beggerly’s (1990) argument system. It existed through the ‘‘Little Ice that her charcoal find, dated 230 G 200 yr Age,’’ when European sea travel began in the b.c., indicates human presence is tenuous. Pacific. At the end of the Little Ice Age, Her argument is based on the assumption the Hawaiian land-care system collapsed, that natural fires could not have occurred in around 1860. The reason for that was not cli- this wet rain-forest valley. That argument mate change. It was primarily the Western- overlooks the fact that the widespread uluhe introduced privatization of land and the cash fern dries up periodically and surely can serve economy that corrupted the traditional life as an easily combustible material during a dry style and culture of the indigenous Hawaiian spell. As Vogl (1969) documented, fire caused people. Jane Allen (1997), who reviewed pre- by lightning has occurred repeatedly in wind- historic development on windward O‘ahu, ward valleys. Also, Mueller-Dombois and La- also related landscape changes to the Little moureux (1967) documented the occurrence Climatic Optimum and the Little Ice Age. of past natural fire in Hawai‘i before Polyne- She says that windward valleys were already sian land occupation. settled by about 600 a.d., and that human im- The back-valley slopes, occupied by ‘o¯hi‘a pact upslope was strong during the Little Cli- lehua (Metrosideros polymorpha) woodland with matic Optimum. During this period of higher uluhe fern (Dicranopteris linearis), represent sea level, much erosive material was deposited the outcome of a prolonged (ca. 2 million yr) downhill on marine platforms. Thereafter, primary succession. As fine soil is formed terracing began during the Little Ice Age there from weathering, it also wastes con- and the irrigation of lo‘i began. She con- stantly or abruptly away by erosion, thereby cluded that both natural forces and human keeping the substrate primarily as rockland. activity interacted in prehistoric landscape Both the ‘o¯hi‘a lehua tree and the uluhe fern change. invade land scars after soil avalanches. The occurrence of rockland and avalanche scars on slopes maintains a vegetation whose dy- Conclusions namics is arrested in primary succession. By synthesizing the former study of Kahana’s This vegetation never reaches a climax state vegetation ecology with its archaeological and in the sense of Clements (1916, 1928). Like- paleoecological investigations, it has become wise, the koa and hala tree are native pioneer abundantly clear that human impact in the species that are highly resilient. They regen- valley has been substantial for at least the past erate after fire. Thus, native colonizer species 12 centuries. However, characterization of dominate the two larger vegetation types in Kahana Valley as a ‘‘geomorphic artifact,’’ as the mid- and back valley. The Metrosideros- proposed by Beggerly (1990), does not appear Dicranopteris back-valley type is resilient to be warranted. The valley’s geomorphology to geomorphic instability. The Acacia koa– certainly has undergone changes. The con- Pandanus midvalley type is resilient to human tinued drop of sea level following the post- impacts, including an occasional fire. The Pleistocene warming may be the prime rea- marsh area in the valley represents an ad- son for the change of the inland marine bay vanced successional stage of a typical hy- into a lagoon. Moreover, the terrestrification drarch succession in the Clementsian sense. of the inland lagoon may have become accel- If swamp tree species had evolved in the erated with the transition period to the Little Hawaiian flora, as are prevalent in the west- Ice Age seven centuries ago, when the sea ern Pacific islands (Stemmermann 1981), the level dropped another meter in roughly 100 marsh would have long become a swamp for- yr. Human invasion of the valley during or est. Such process is currently ongoing with before the Little Climatic Optimum certainly invasion of the octopus tree. Another intro- contributed to landscape change, but it was duced tree species capable of invading the not the only force. Natural forces were marsh is the paperbark tree (Melaleuca quin- Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 311 quenervia). It has been planted in Kahana Val- analysis of the development of vegetation. ley but is not yet documented as naturalized. Carnegie Institution, Washington. The current management goal of an arrested ———. 1928. Plant succession and indica- wetland succession can still be upheld by tree tors. Republished, 1963. Hafner Publ. removal. Currently, efforts are under way to Co., New York. restore the Kahana State Park (recently re- Hommon, R. J., and W. M. Barrera Jr. 1971. named Ahupua‘a ‘O Kahana State Park) as a Archeological survey of Kahana Valley, functional ahupua‘a. Results of follow-up his- Ko‘olau District, Island of O‘ahu. Depart- torical and more complete biodiversity re- ment of Anthropology, B. P. Bishop Mu- search with PABITRA methodologies will seum, Honolulu, Hawai‘i (prepared for continue to yield helpful information. Such Division of State Parks). Department of information can be used to support restora- Land and Natural Recources Report 71-3. tion of ecosystem functions and biodiversity Available from Division of State Parks, components. In addition, new research can State of Hawai‘i, Honolulu. be directed to determine if the restored Ka- Judd, C. S. 1935. Koa reproduction after fire. hana ahupua‘a will be able to function as a J. For. 33:176. sustainable system in the future. Kirch, P. V. 1985. Feathered gods and fish- hooks: An introduction to Hawaiian ar- chaeology and prehistory. University of acknowledgments Hawai‘i Press, Honolulu. ———. 2000. On the roads of the winds: An We thank Mark Nakamura for computer archaeological history of the Pacific islands drafting the line drawings and Annette before European contact. University of Mueller-Dombois for word processing and California Press, Berkeley. editorial counseling. Minerbi, L. 1999. Indigenous management models and protection of the ahupua‘a. Literature Cited Soc. Process Hawai‘i 39:208–225. Mueller-Dombois, D. 1973. A non-adapted Allen, J. 1997. Pre-contact landscape trans- vegetation interferes with soil water re- formation and cultural change in wind- moval in a tropical rain forest area in Ha- ward O‘ahu. Pages 230–247 in P. V. wai‘i. Trop. Ecol. 24:1–18. Kirch and T. L. Hunt, eds. Historical Mueller-Dombois, D., and F. R. Fosberg. ecology in the Pacific islands: Prehistoric 1998. Vegetation of the tropical Pacific is- environmental and landscape change. Yale lands. Ecol. Stud. 132. Springer-Verlag. University Press, New Haven, Connecti- Mueller-Dombois, D., and C. H. Lamoureux. cut. 1967. Soil-vegetation relationships in Ha- Athens, J. S., and J. V. Ward. 1993. Environ- waiian kipukas. Pac. Sci. 21:286–299. mental change and prehistoric Polynesian Nunn, P. D. 1999. Environmental change in settlement in Hawai‘i. Asian Perspect. 32 the Pacific Basin. Wiley & Sons, Ltd., (2): 205–222. Chichester, England. Athens, J. S., J. V. Ward, and S. Whickler. Palmer, D. D. 2003. Hawai‘i’s ferns and fern 1992. Late Holocene lowland vegetation, allies. University of Hawai‘i Press, Hono- O‘ahu, Hawai‘i. N. Z. J. Archaeol. 14:9– lulu. 34. Pukui, M. K., and S. H. Elbert. 1986. Hawai- Beggerly, P. R. 1990. Kahana Valley, Hawai‘i, ian dictionary. University of Hawai‘i Press, a geomorphic artifact: A study of the in- Honolulu. terrelationships among geomorphic struc- Puniwai, N. 1997. Decline of Acacia koa tures, natural processes, and ancient on windward O‘ahu. Final report. USDA Hawaiian technology, land use, and settle- Forest Service, Institute of Pacific Islands ment patterns. Ph.D. diss., University of Forestry, Honolulu. Available from Divi- Hawai‘i at Ma¯noa, Honolulu. sion of State Parks, State of Hawai‘i, Ho- Clements, F. E. 1916. Plant succession: An nolulu. 312 PACIFIC SCIENCE . April 2005

Spatz, G., and D. Mueller-Dombois. 1973. District, island of O‘ahu. Department of The influence of feral goats on koa tree re- Botany, University of Hawai‘i, Honolulu, production in Hawai‘i Volcanoes National Technical Report prepared for Division Park. Ecology 54:870–876. of State Parks, Outdoor and Historical Stauffer, R. H. 1990. Land tenure in Kahana, Sites, Department of Land and Natural Hawai‘i, 1846–1920. Ph.D. diss., Univer- Resources, State of Hawai‘i, Honolulu. sity of Hawai‘i, Honolulu. Available from Division of State Parks. Stemmermann, L. 1981. A guide to Pacific Vogl, R. J. 1969. The role of fire in the evo- wetland plants. U.S. Army Corps of Engi- lution of the Hawaiian flora and vegeta- neers, Honolulu District, Honolulu. tion. Proc. Tall Timbers Fire Ecol. Conf., Takasaki, K. J., G. T. Hirashima, and E. R. 1969: 5–60. Lubke. 1969. Water resources of Wind- Wagner, W. L., D. R. Herbst, and S. H. ward O‘ahu, Hawai‘i. Geological Survey Sohmer. 1990. Manual of the flowering Water-Supply Paper 1894. Prepared in plants of Hawai‘i. 2 vols. University of cooperation with Division of Water and Hawai‘i Press and Bishop Museum Press, Land Development, Department of Land Honolulu. and Natural Resources, State of Hawai‘i. Wirawan, N. 1978. Vegetation and soil-water U.S. Government Printing Office, Wash- regimes in a tropical rain forest valley ington, D.C. on O‘ahu, Hawaiian Islands. Ph.D. diss., Theobald, W. L., and N. Wirawan. 1973. University of Hawai‘i at Ma¯noa, Hono- Kahana Valley botanical survey, Ko‘olau lulu.

Appendix Kahana Valley Plant Checklist

Abutilon grandifolium* (Willd.) Sweet (Malvaceae), ma‘o, Calophyllum inophyllum* L. (Clusiaceae), kamani hairy abutilon Canavalia cathartica* Thouars (Fabaceae), maunaloa Acacia koa A. Gray (Fabaceae), koa Carex wahuensis C. A. Mey (Cyperaceae) Adenophorus pinnatifidus Gaudich. (Grammitidaceae), kihi Casuarina equisetifolia* L. (Casuarinaceae), paina, Adenophorus tamariscinus (Kaulf.) Hook. & Grev. ironwood tree (Grammitidaceae), wahine noho mauna Cenchrus enchinatus* L. (Poaceae), ‘ume‘alu, common Adiantum raddianum C. Presl (Pteridaceae), maidenhair sandbur fern Centaurium erythraea* Raf. (Gentianaceae), European Ageratina riparia* (Reget) R. King & H. Robinson centaury (Asteraceae), pa¯makani Centella asiatica* (L.) Urb. (Apiaceae), pohe kula, Asiatic Aleurites moluccana* (L.) Willd. (Euphorbiaceae), kukui pennywort Alocasia macrorrhiza* (L.) Schott (Arecaceae), ‘ape Chamaecrista nictitans* (L.) Moench (Fabaceae), laukı¯, Alyxia oliviformis Gaud. (Apocynaceae), maile, endemic partridge pea Amaranthus viridis* L. (Amaranthaceae), pakai Chamaesyce hypericifolia* (L.) Millsp. (Euphorbiaceae), Andropogon virginicus* L. (Poaceae), broomsedge graceful spurge Antidesma pulvinatum Hillebr. (Euphorbiaceae), hame Charpentiera obovata Gaud. (Amaranthaceae), pa¯pala Ardisia crenata* Sims (Myrsinaceae), Hilo holly Christella dentata (Forssk.) Brownsey & Jermy Ardisia elliptica* Thunb. (Myrsinaceae), shoebutton (Thelypteridaceae), pai‘i‘iha¯ ardisia Chrysopogon aciculatus* (Rets.) Trin. (Poaceae), ma¯nienie Artocarpus altilis* (S. Parkinson ex Z) Fosb. (Moraceae), ‘ula ‘ulu, breadfruit Cibotium chamissoi Kaulf. (Dicksoniaceae), ha¯pu‘u Arundina graminifolia* (D. Don) Hochr. (Orchidaceae), Cibotium menziesii Hook. (Dicksoniaceae), ha¯pu‘u ‘i‘i bamboo orchid Clerodendrum philippinum* Schauer (Verbenaceae), Bidens pilosa* L. (Asteraceae), kı¯,kı¯ nehe, Spanish needle pı¯kake hohono Bidens populifolia Sherff (Asteraceae) Clidemia hirta* (L.) D. Don (Melastomataceae), Koster’s Bixa orellana* L. (Bixaceae), ‘alaea, annatto, lipstick tree curse Blechnum appendiculatum Willd. (Blechnaceae) Cocculus trilobus (Thunb.) DC (Menispermaceae), Brachiaria mutica* (Forssk.) Stapf (Poaceae), Calfornia huehue grass Coix lachryma-jobi* L. (Poaceae), pu¯‘ohe‘ohe, ku¯kaeko¯lea, Broussonetia papyrifera (L.) Vonton. (Moraceae), wauke Job’s tears Bulbostylis capillaris (L.) C. B. Clarke (Cyperaceae) Colocasia esculenta* (L.) Schott. (Araceae), kalo, taro Kahana Valley Ahupua‘a, PABITRA Study Site on O‘ahu . Mueller-Dombois et al. 313

Appendix (continued)

Commelina diffusa* N. L. Burm. (Commelinaceae), Ipomoea indica ( J. Burm.) Merr. (Convolvulaceae), koali honohono, honohono wai ‘awa Convolvulus arvensis* L. (Convolvulaceae), field bindweed Ipomoea pes-caprae (L.) R. Br. (convolvulaceae) po¯huehue Conyza canadensis* (L.) Cronq. (Asteraceae), lani wela, Korthalsella latissima (Tiegh.) Degener (Viscaceae), Canada fleabane kaumahana, mistletoe Cordyline fruticosa* (L.) A. Chev. (Agavaceae), ti Kyllingia nemoralis* ( J. R. Forster & G. Forster) Dandy Coronopus didymus* (L.) Sm. (Brassicaceae), swinecress (Cyperaceae), kili‘o‘opu Crotalaria pallida* Aiton (Fabaceae), pikakani, smooth Lantana camara* L. (Verbenaceae), la¯kana rattlebox Lepisorus thunbergianus (Kaulf.) Ching (Polypodiaceae), Cuphea carthagenensis* ( Jacq.) Macbr. (Lythraceae), pa¯kahakaha tarweed Leucaena leucocephala* (Lam.) de Wit (Fabaceae), koa Cuscuta sandwichiana Choisy (Cuscutaceae), kauna‘oa, haole kauna‘oa lei Ludwigia octivalvis* ( Jacq.) Raven (Onagraceae), ka¯mole, Cyclosorus interruptus (Willd.) H. Ito (Thelypteridaceae), primrose willow neke Lycopodiella cernua (L.) Pic. Serm. (Lycopodiaceae), Cyperus rotundus* L. (Cyperaceae), kili‘o‘opu, nutgrass wa¯wae‘iole Cyrtandra hawaiiensis C. B. Clark (Gesneriaceae), Machaerina angustifolia (Gaud.) T. Koyama (Cyperaceae), ha‘iwale ‘uki Cyrtandra laxiflora H. Mann (Gesneriaceae) Mangifera indica* L. (Anacardiaceae), manako¯, mango Cyrtandra spp., endemic Mecodium recurvum (Gaudich.) Copel. Dicranopteris linearis (Burm. f.) Underw. (Hymenophyllaceae), ‘o¯hi‘a ku¯, filmy fern (Gleicheniaceae), uluhe Melaleuca quinquenervia* (Car.) S. T. Blake (Myrtaceae), Dioscorea bulbifera* L. (Dioscoreaceae), hoi, bitter yam paperbark tree Dioscorea pentaphylla* L. (Dioscoreaceae), pi‘a, yam Metrosideros polymorpha Gaud. (Myrtaceae), ‘o¯hi‘a lehua, Diospyros sandwicensis (A. DC) Fosb. (Ebenaceae), lama endemic Diplazium sandwichianum (C. Presl) Diels (Athyriaceae), Metrosideros tremuloides (A. Heller) P. Knuth ho¯‘i‘o (Myrtaceae), lehua ‘a¯hihi Diplopterygium pinnatum (Kunze) Nakai Mimosa pudica* L. (Fabaceae), pua hilahila, sensitive (Gleicheniaceae), uluhe lau nui plant Doryopteris decipiens (Hook.) J. Sm. (Pteridaceae), * L. (Rubiaceae), noni, Indian mulberry ‘iwa‘iwa Myrsine lessertiana A. DC (Myrsinaceae), ko¯lea lau nui Dryopteris spp. (Dryopteridaceae), olua Nephrolepis multiflora (Roxb.) F. M. Jarrett ex C. V. Elaeocarpus bifidus Hook. & Arnott (Elaeocarpaceae), Morton (Nephrolepidaceae) kalia Nestegis sandwicensis (A. Gray) Degener (Oleaceae), Elaphoglossum aemulum (Kaulf.) Brack. olopua (Lomariopsidaceae), ‘e¯kaha Ochrosia compta K. Schum. (Apocynaceae) Elaphoglossum crassifolium (Gaudich.) W. R. Anderson & Ophioglossum polyphyllum A. Braun (Ophioglossaceae), Crosby (Lomariopsidaceae), ‘e¯kaha pololei, adder’s tongue Emilia fosbergii* Nicolson (Asteraceae), pualele Oplismenus hirtellus* (L.) P. Bear. (Poaceae), honohono, Emilia sonchifolia* (L.) DC (Asteraceae), Flora’s basketgrass paintbrush Osteomeles anthyllidifolia (Sm.) Lindl. (Rosaceae), ‘u¯lei Erechtites valerianifolia* (Wolf ) DC (Asteraceae) Pandanus tectorius S. Parkinson ex Z (Pandanaceae), hala Eucalyptus* sp. (Myrtaceae) tree Fimbristylis dichotoma (L.) Vahl (Cyperaceae) Panicum maximum* Jacq. (Poaceae), Guinea grass Freycinetia arborea Gaud. (Pandanaceae), ‘ie‘ie Paraserianthes falcataria* (Fabaceae), albizia Gardenia mannii St. John & Kuykendall (Rubiaceae), Paspalum conjugatum* Bergius (Poaceae), Hilo grass na¯nu¯ Paspalum scrobiculatum* L. (Poaceae), mau‘u laiki, Gonocormus minutus (Blume) Bosch (Hymenophyllaceae) ricegrass Grammites tenella Kaulf. (Grammitidaceae), kolokolo Paspalum urvillei* Steud. (Poaceae), Vasey grass Hedychium coronarium* J. Ko¨nig (Zingiberaceae), Passiflora edulis* Sims (Passifloraceae), passion fruit, ‘awapuhi ke‘oke‘o, white ginger liliko‘i Hedychium flavescens* N. Carey ex Roscoe Passiflora foetida* L. (Passifloraceae) poha¯poha¯, wild (Zingiberaceae), ‘awapuhi melemele, yellow ginger water lemon Hibiscus arnottianus A. Gray (Malvaceae), koki‘o ke‘oke‘o Passiflora mollisima* (Kunth) L. H. Bailey Hibiscus tiliaceus L. (Malvaceae), hau (Passifloraceae), banana poka Huperzia phyllantha (Hook. & Arn.) Holub Passiflora pulchella* Kunth (Passifloraceae), two-lobed (Lycopodiaceae), wa¯wae‘iole, rat’s foot passion flower, naturalized Ipomoea alba* L. (Convolvulaceae), koali pehu, moon Passiflora suberosa* L. (Passifloraceae), huehue haole, flower introduced; wild passion flower 314 PACIFIC SCIENCE . April 2005

Appendix (continued)

Peperomia oahuensis C. DC (Piperaceae) Selaginella arbuscula (Kaulf.) Spring (Sellaginellaceae), Phaius tankarvilleae* (Banks ex L’He´r.) Blume lepelepe a moa (Orchidaceae), Chinese ground orchid Setaria gracilis* Kunth (Poaceae), mau‘u Kaleponi, Phyllostachys nigra* (Lodd.) Munro (Poaceae), black yellow foxtail bamboo Setaria verticillata* (L.) P. Beauv. (Poaceae), mau‘u Phymatosorus grossus (Langsd. & Fisch.) Brownlie pilipili, bristly foxtail (Polypodiaceae), laua‘e Spathodea campanulata* P. Beauv. (Bignoniaceae), African Piper methysticum* G. Forster (Piperaceae), ‘awa, kava tulip tree Pipturus albidus (Hook. & Arnott.) A. Gray (Urticaceae), Spathoglottis plicata* Blume (Orchidaceae), Malayan or ma¯maki Philippine ground orchid Pisonia umbellifera (G. Forster) Seem. (Nyctaginaceae), Sphaerocionium obtusum (Hook. & Arn.) Copel. pa¯pala ke¯pau (Hymenophyllaceae), palai lau li‘i, small-leaved fern Pityrogramma austroamericana* Domin (Pteridaceae), Sphenomeris chinensis (L.) Maxon (Lindsaeaceae), pala‘a¯, goldback fern lace fern Pityrogramma calomelanos* (L.) Link (Pteridaceae), Stachytarpheta dichotoma* (Ruiz & Par.) Vahl silverback fern (Verbenaceae), o¯wı¯,oı¯ weed Pluchea symphytifolia* (Mill.) Gillis (Asteraceae), sourbush Stachytarpheta jamaicensis* (L.) Vahl (Verbenaceae), o¯wı¯, Polypodium pellucidum Kaulf. (Polypodiaceae), ‘ae oı¯, Jamaican vervain Pritchardia martii (Gaud.) H. A. Wendl. (Arecaceae), Styphelia tameiameiae (Cham. & Schlechtend.) F. v. loulu hiwa Muell. (Epacridaceae), pu¯kiawe Psidium cattleianum* Sabine (Myrtaceae), waiawı¯, Syzygium cumini* (L.) Skeels (Myrtaceae), Java plum strawberry guava Syzygium jambos* (L.) Alston (Myrtaceae), ‘o¯hi‘a loke, Psidium guajava* L. (Myrtaceae), kuawa, common guava rose apple Psilotum complanatum* Sw. (Psilotaceae), moa, moa haole Syzygium malaccense* (L.) Merr. & Perry (Myrtaceae), Psilotum nudum (L.) P. Beauv. (Psilotaceae), moa ‘o¯hi‘a ‘ai, mountain apple Psychotria kaduana (Cham. &. Schlechtend.) Fosb. Syzygium sandwicensis (A. Gray) Nied. (Myrtaceae), ‘o¯hi‘a (Rubiaceae), ko¯piko ha¯ Rhizophora mangle* L. (Rhizophoraceae), American or Tectaria gaudichaudii (Mett.) Maxon (Dryopteridaceae), red mangrove ‘iwa‘iwa lau nui, large-leaved ‘iwa‘iwa Rhynchospora sclerioides (Hook. & Arnott) (Cyperaceae) Terminalia catappa* L. (Combretaceae), kamani haole, Ricinus communis* L. (Euphorbiaceae), kolı¯ Indian almond Rubus rosifolius* Sm. (Rosaceae), ‘a¯kala, thimbleberry Tetraplasandra gymnocarpa (Hillebr.) Sherff (Araliaceae), Sacciolepis indica* (L.) Chase (Poaceae), Glenwood grass ‘ohe‘ohe Sadleria pallida Hook. & Arn. (Blechnaceae), ‘ama‘u Touchardia latifolia Gaud. (Urticaceae), olona¯ Salvia coccinea* Etl. (Lamiaceae), lı¯lı¯lehua, scarlet sage or Trema orientalis* (L.) Blume (Ulmaceae), gunpowder Texas sage tree Santalum ellipticum Gaud. (Santalaceae), ‘iliahialo‘e, coast Urena lobata L. (Malvaceae), aramina sandalwood Urera glabra (Hook. & Arnott) Wedd. (Urticaceae), Scaevola gaudichaudiana Cham. (Goodeniaceae), naupaka o¯puhe kuahiwi Vandenboschia davallioides (Gaudich.) Copel. Scaevola sericea Vahl. (Goodeniaceae), naupaka kahakai (Hymenophyllaceae), palai hihi, filmy fern Schefflera actinophylla* (Endl.) Harms (Araliaceae), Verbena litoralis* Kunth (Verbenaceae), o¯wı¯ octopus tree Vittaria elongata Sw. (Vittariaceae), ‘ohe‘ohe Schinus terebinthifolius* Raddi (Anacardiaceae), wilelaiki, Wedelia trilobata* (L.) Hitchc. (Asteraceae) Christmas berry tree Wikstroemia oahuensis (A. Gray) Rock (Thymelaeaceae) Schizostachyum glaucifolium* (Rupr.) Munro (Poaceae), ‘a¯kia ‘ohe Zingiber zerumbet* (L.) Sm. (Zingiberaceae), ‘awapuhi Scirpus sp. (Cyperaceae) 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.