Alien Plants on the Northwestern Hawaiian...

ALIEN PLANTS ON THENOKIHWESTERN Hawai iAN ISLANDS

Derral R. Herbst and Warren L. Wagner

ABSTRACT

The Northwestern Hawaiìan Islands is a chain of approximately 40 emergentlow islands extending in a west-northwesterly direction for a distance ofnearly 750 mi (1,210 km). The islands are currently managed as a wildliferefuge for birds, sea turtles, and the Hawaiìan monk seal (Monachusschauinslandi). The vegetation is composed primarily of Pacific Basinstrand species, naturalized species, and, on Kure and Midway Atolls and TernIsland, cultivated plants. The present vascular flora consists of 317 taxa,of which 8 are endemic (2.5%), 42 are indigenous (13.2%), 114 are naturalized(36.0%), and 132 are cultivated (41.6%). The flora is summarized on anisland by island basis in an appendix to this paper. Some alien species havebeen deliberate or unintentional introductions by humans; others have arrivedby "natural" means. Ironwood (Casuarina equisetifolia), a species welladapted to dry conditions, is an example of an intentional introduction. Ithas been introduced on seven of the islands and has become naturalized onTern Island and Midway Atoll; it is a management problem only on Midway.Heliotropium procumbens, introduced by military flights from Oàhu to twoof the three leeward islands with runways, is an example of an accidentalintroduction that has not become a management problem. Little has been doneto manage the alien plants of the Northwestern Hawaiìan Islands other than totake precautions to prevent new ones from being introduced. There has beenlittle need for management. Proposed management needs include monitoring thevegetation of the islands, producing a manual of plants, and developing acontingency plan for the control of any alien species of plant or animalimpacting the vegetation or wildlife.

INTRODUCTION

The Northwestern Hawaiìan Islands, or Leeward Islands as they arecommonly (and hereafter) called, are a chain of islands stretching in awest-northwesterly direction from Nihoa Island to Kure Atoll, a distance ofnearly 750 mi (1,210 km). Politically, all except Midway Atoll are part ofthe City and County of Honolulu; Midway is property of the U.S. Navy and is

189

Herbst and Wagner \ Aliens on the Northwestern Hawaiìan Islands 190

not part of the State of Hawaiì. President Theodore Roosevelt signedExecutive Order 1019 on February 3, 1909, setting them aside (exceptMidway) as a preserve and breeding ground for native birds. Today, allexcept Kure and Midway Atolls are in the Northwestern Hawaiìan IslandsNational Wildlife Refuge and are managed by the U.S. Fish and WildlifeService. President Franklin Roosevelt signed Executive Order 7299 onFebruary 20, 1936, placing Kure Atoll under the jurisdiction of theSecretary of the Navy; however, there are no records of any activity on theAtoll during World War II. On November 17, 1952, President Harry Trumansigned Executive Order 10413, giving Kure Atoll to the Territory ofHawaiì. Kure presently is maintained as a wildlife refuge by the StateDivision of Forestry and Wildlife, and the Coast Guard maintains a long-range navigation (LORAN) station on Green Island, the only permanent landmass in the Atoll.

The Leeward Islands (Fig. 1) comprise approximately 40 emergent islands;numbers vary as the small sand islets are built up, fuse, separate, or washaway over time. The islands total about 3.9 mi2 (10.1 km2) ofdry land, an amount less than 1% of the total land area of the HawaiìanIslands. They vary in elevation from 10 to 910 ft (3-277 m) (Table 1).The highest is Nihoa Island, and the lowest are French Frigate Shoals andPearl and Hermes Atoll. The eastern islands (Nihoa, Necker, La PerousePinnacle of French Frigate Shoals, and Gardner Island) are eroded remnantsof shield volcanoes, the last two being residual volcanic and limestoneislands. The western islands (French Frigate Shoals except for La P6rousePinnacle, Laysan and Lisianski Islands, and Pearl and Hermes, Midway, andKure Atolls) are limestone islands resting on the summits of submergedvolcanoes. More than a dozen other reefs and shoals mark volcanoescompletely submerged below the surface of the ocean.

Compared to the main Hawaiìan Islands, the Leeward Islands are poorlyknown geologically. They are much older than the main islands; the ages offive of the islands are known and range from about 7 to 27 million years(Table 1).

VEGETATION

The vegetation of the Leeward Islands is composed chiefly of typicalstrand species and the naturalized alien plants that can tolerate strandconditions. Cultivated ornamental plants are maintained on the threeinhabited islands, and some also persist around abandoned buildings. OnNihoa, the highest island, several genera usually considered to be elementsof mid-elevation, diverse mesic forests can be found. Thus, huàhuàko(Rumex albescens), Schiedea verticillata, and perhaps Pritchardiaremota and àkoko (Chamaesyce celastroides) may be remnants of anearlier vegetation before the island eroded to its present state. Severalof the strand species, such as popplo (Solarium nelsonii), ànaunau(Lepidium bidentatum var. o-waihiense), and òhai (Sesbaniatomentosa), which are rare on the main Hawaiìan Islands, are still commonon several of the Leeward Islands. This may be due to the greaterdisturbance of strand areas on the main Hawaiìan islands.

Figure 1. The Hawai ian Archipelago.


Table 1. Area, elevation, and age of the Leeward Hawai`ian Islands.

IslandGroup

Nihoa IslandNecker IslandFrench Frigate Shoals

(13 islets)Gardner PinnaclesLaysan IslandLisianski IslandPearl and Hermes Atoll

(7 islets)Midway Atoll (4 islets)Kure Atoll (2 islets)

Areaa

(acres)

19158

563

981432

781,280

237

(hectares)

(77)(24)

(23)(1)

(397)(175)

(32)(518)

(96)

Maximumelevation3

feet (meters)

910277

1351903520

101220

(277)(84)

i

(41)b

(58)(11)(6)

(3)(4)(6)

Age(in millions of yearsby Potassium-Argon)

7.0 +10.0 +

t117b,c

—--

20.1 +27.0 +

—

0.3C

0.4C

0.5d

0.6e

Northwestern Hawai`ianIslands (Total) 3,316 (1,343)

afrom Armstrong 1983.bLa Perouse Pinnacles only.

cDalrymple et al. 1974; unreliable due to sampling problems.dClague et al. 1975; probably too young.

eDalrymple et al. 1977; probably too young.

Table 2. Summary of status information for plant taxa on Leeward Islands.

Island Group

Nihoa IslandNecker IslandFrench Frigate ShoalsGardner PinnaclesLaysan IslandLisianski IslandPearl and Hermes AtollMidway AtollKure Atoll

E

3---6-133

I

175

121

2615152616

X

51

27-1239

10141

Statusa

C

--22-223

12415

S

-122621--

p-75-

1822--

9

-

-

-

-

1

1

-

1

-

TotalTaxa

2514683

712531

25575

Totalsb 42 114 132 8 11 317

aE = endemic; I = indigenous; X = naturalized alien species; C = cultivated alien species;S = seeds that arrived via natural dispersal; P = intentionally introduced propagules;? = unknown.In calculating the totals, each species was counted only once, although it may haveoccurred on more than one island.


Of the 317 taxa of vascular plants recorded from the Leeward Islands, 8(2.5%) are endemic and 42 (13.2%) are indigenous to these islands (althoughthe indigenous species may be endemic to the Hawaiìan Archipelago), while114 (36.0%) are naturalized alien species and 132 (41.6%) are cultivatedplants. The remaining 21 (6.6%) represent the seeds, cuttings, andseedlings deliberately introduced to the islands, or the seeds that floatedto the islands but did not become established. The flora is summarized onan island by island basis in the Appendix and in Table 2 (page 192). Ahistory of major disturbances on each of the Leeward Island groups is givenin Table 3.

Plants have been introduced by two means: deliberate or unintentionalintroductions by humans, and introductions that did not involve theintervention of humans. Naturalized species and cultivated plants fallinto the first category and native species into the last. For the purposeof this paper, alien species that moved from island to island by naturalmeans (wind, ocean currents, or birds) are still considered alien species.Examples of different categories of naturalized alien species, based upontheir means of introduction and their impacts, are given below.

CASE HISTORIES

Intentional Introduction of an Alien Plant: IronwoodCasuarinaceae is a distinctive family of a single genus of shrubs and

trees native primarily to Australia. A few species occur on Pacific Basinislands from Java to New Caledonia and in Asia as far north as Burma. Theearliest documented record of the family in Hawaiì is a collection ofironwood (Casuarina equisetifolia) made by Heller on Oàhu in 1895(Heller 1955: herbarium specimen, B.P. Bishop Museum). The speciesprobably was introduced in 1882 by P. Isenberg, who planted trees atKilohana Crater, Kauaì. In 1890, A.S. Cleghorn, the father of PrincessKaìulani, planted an avenue of ironwood trees on his estate in Waikiki.This is the avenue of ironwoods still standing in Kapiòlani Park andprobably was the source of Heller's collection.

Ironwood has several qualities that help it compete under adverseconditions (Apfelbaum et al. 1983). It is capable of fixingatmospheric nitrogen and therefore is able to colonize nutrient-poor soils,especially those lacking nitrogen. The root system takes up most of thenutrients from the soil, which, along with the thick mulch of fallenbranchlets that accumulates under the trees, prevents nearly all otherplants from growing near it (Neal 1965). Ironwoods can reproduce bothsexually and asexually, and the last, by root suckers, appears to be moreprominent in Hawaiì. The leaves are reduced to whorls of minute scaleswith stomata (minute openings) located in longitudinal grooves where theyare protected by minute cilia, reducing transpiration, an importantadaptation for dry conditions. Moreover, ironwood is a facultativehalophyte (salt lover). These qualities make ironwood a desirable, hardyornamental and shade tree on the inhabited Leeward Islands. These samequalities make this species a potential management problem.

Table 3. History of major disturbances in the Leeward Hawai`ian Islands.

ISLAND GROUP

Nihoa Islanda

Necker Islandb

French Frigate Shoalsc

Gardner Pinnaclesd

EVENT

* Prehistoric habitation; house foundations.terraces, shelters, and religious sites.

* Fire apparently destroyed much vegetation.* HIRAN projects probably introduced Cenchrus

echinatus and Paspalum sp. (since disappeared),and Portulaca oleracea (replacing, perhaps viahybridization, native P. lutea).

* Religious sites and terraces indicate muchimpact by prehistoric Hawaiians.

* Some evidence of unofficial bombing.

* Shipwreck crew of 27 on island about one month.* Vegetation partly destroyed during

military exercise on East Island.* Permanent military station on East Island.* Tern Island enlarged by dredging, naval base

built, all vegetation destroyed.* LORAN+ station on East Island.* East Island inundated by a tidal wave.* LORAN Station on Tern Island.* Tern Island inundated during heavy seas, much

vegetation damaged.* U.S. Fish & Wildlife Service refuge.

* None known.

YEAR

—1885

1961

—1940s

1867

1932-19401942-1945

1942-19461942-195219461952-1979

19691979 to present

ISLAND GROUP

Laysan Islande

fLisianski Island

Pearl and Hermes Atollg

Midway Atoll

EVENT

* Guano mining.* Various invasions by bird poachers.* Introduced rabbits virtually denuded the island.* Various plants reintroduced.* HIRAN operation may have accidentally introduced

Cenchrus echinatus and Conyza bonariensis.

* Crews from 3 shipwrecks on island for up to5 months.

* Bird poachers (at least 5 groups, largest of 77people, shacks built).

* Rabbits destroyed all vegetation, then starvedto death.

* Crews of 2 shipwrecks lived on island for 2 months.* Rabbits damaged vegetation.* Hawaiian Sea Products Co. erected

buildings and harvested oyster shell.* Temporary naval base on SE Island.* Abandoned Sea Products buildings bombed and

strafed; landing party burned them during a warexercise.

* Navy constructed a 15 ft steel tower on SE Island.

* Shipwrecked crew lived on islands for 14 monthsuntil rescued.

* Commercial Pacific Cable Co. installation built.About 9,000 tons of soil imported from Honolulu;

YEAR

1890-1910? -19151903-19231923-1966

1961-1962

1844-1872

1904-1910

after 1904-1915/16

1822before 1916-1928

1929 - 19311930s

19421961

1888-1889

Table 3, continued. 195

Table 3, continued.

ISLAND GROUP

KureAtollj

EVENT

Windbreaks and sand-binding plants, vegetables,and ornamentals planted.h

* Pan American Airways built an airport, hotel,power plant, warehouses, and other buildings;planted over 200 species of ornamentals.h,1

* Military activities, especially during WWII, .completely changed the ecology of the island.1

* Crew of shipwreck on island.* Crew of shipwreck on island (killed and ate

ca 7000 seabirds, 60 seals, and 1 dog).* Crew of shipwreck on island ca 9 weeks before rescued.* Emergency building and water tanks built,

tree seeds sown (none grew).* Navy built radar reflector.* LORAN station and runway built and maintained

by Coast Guard.

YEAR

1903-?

1935-1941?

1941 to present

1837-1838

1842-18431870-1871

18861955

1960 to present

aClapp et al. 1977.bClapp and Kridler 1977.cAmerson 1971.dClapp 1972.eEly and Clapp 1973.fClapp and Wirtz 1975.gAmerson et al. 1974.hBryan 1978.iNeff and DuMont 1955.jWoodward 1972.

+HIRAN: High-precision short-range navigation; LORAN: Long-range navigation.


Ironwood has been introduced to seven of the Leeward Islands: NeckerIsland, French Frigate Shoals, Laysan and Lisianski Islands, and Pearl andHermes, Midway, and Kure Atolls. Seeds were sown on Tern, Necker, andLaysan Islands by C.S. Judd in 1923 during the Tanager Expedition(Christophersen and Caum 1931); if any of these seeds germinated they didnot survive. Ironwood trees were planted on Pearl and Hermes by navalpersonnel in 1963 to make this low island more visible from a distance.The seedlings subsequently were removed in 1964 by the U.S. Fish andWildlife Service, as it violates Service policies to introduce alien plantson the Refuge. Ironwood trees were introduced to Laysan at least threetimes (Ely and Clapp 1973). G.P. Wilder planted a box of ironwoodseedlings on Laysan in September 1905 and additional trees in 1930, andJudd sowed seed in 1923 (Christophersen and Caum 1931). Although severalof the trees grew, only one is presently alive; it is believed to be one ofthe 1905 plantings. Coast Guard personnel planted trees around theirinstallations on Tern Island and Kure in the mid 1950s. The originalplantings are still alive, but only those on Tern Island have reproducedand become sparingly naturalized. Trees were planted on Lisianski in 1931(Clapp and Wirtz 1975). Of the original plantings, only about four or fiveare still alive.

Ironwood has become abundant enough only on Midway to cause a manage-ment problem. In 1903, the Commercial Pacific Cable Company took posses-sion of Sand Island, Midway Atoll, and erected permanent concretebuildings. Over the ensuing years, more than 9,000 tons of top soil wasbrought from Honolulu, vegetable gardens were established, and ornamentaltrees, shrubs, and flowers were planted. The island changed from a barrensand waste to an oasis around the company installations (Bryan 1978). In1907, Daniel Morrison, then Assistant Superintendent of the cable station,planted hundreds of ironwood trees to give protection against the wind andsun (Bryan 1978). Descendants of those trees now dominate the island'secosystem and have impacted the use of the island by both wildlife andhumans. Changes in bird populations have occurred with ironwoodestablishment, favoring birds associated with forested areas and selectingagainst the species of open habitats. Although black noddies (Anousminutus melanogenys) and white terns (Gygis alba rothschildi) benefitfrom the increase in their preferred nesting habitat, other species, suchas the black-footed albatross (Diomedea nigripes), which nests on theground in open areas, suffer habitat loss. Also, several of the largerseabird species need open areas because they must run in order to becomeairborne. Native vegetation also has been impacted by ironwood trees. Forexample, the beach naupaka (Scaevola sericea) is an important sandstabilizer on these islands. Since beach naupaka cannot tolerate heavyshade, the gradual encroachment and replacement of the naupaka by ironwoodhas not only changed the vegetation components of the island but hasrendered the shoreline more susceptible to erosion.

The Federal Aviation Administration requires that the vegetation alongthe sides and ends of runways be kept below certain heights for safety.The runway aprons were paved with "soil cement," an asphalt that wassprayed on the ground to prevent the encroachment of the ironwoods andreduce maintenance. According to L. Pinter (pers. comm.), however, seeds


became lodged in seams and cracks in the soil cement and germinated, theirroots threatening to break up the aprons. Graders were used to remove theunwanted vegetation but broke through the soil cement; the ironwoods theninvaded the area heavily. Tagged trees, measured at 6-month intervals bythe Navy, grew as much as 40 ft (12 m) in 18 months. When the trees werecut down, they regrew from the stumps and roots. Herbicides were not used,although the species is very susceptible to them, because the runoff of thelanding strips is the main source of water for the installation. It hasbeen necessary to keep the runway area cleared mechanically, using manuallabor to remove the seedlings. Backhoes, chain saws, and heavier equipmentare used around the buildings, antenna fields, and other areas.Additionally, the older trees often are infested with ground termites(Coptotermes formosanus), which eat out the core of the plant. Thesetrees break easily or lose large limbs during storms, damaging power linesand buildings. Ironwood control is, indeed, a major problem on Midway, butoddly, not on any of the other Leeward Islands.

Accidentally Introduced Alien Plants: Heliotropium procumbensand Golden Crown-beard

The heliotrope Heliotropium procumbens is a low, mat-forming,perennial herb. It superficially resembles, and shares the same habitatas, the indigenous nena (Heliotropium curassavicum) but can easily bedistinguished from the latter by its nonsucculent, densely pubescentleaves. Heliotropium procumbens is native from the southern UnitedStates south to Central and South America and east to the West Indies. Italso is found on Guam, where it is considered native (Stone 1970; Fosberget al. 1979).

For about three centuries, Spanish galleons regularly laid over on Guamto load fresh supplies, drop off goods, and unload passengers during theannual Acapulco to Manila trip. Because the Guam populations or thisheliotrope most closely resemble those of Mexico (Fosberg, pers. comm.),and because Heliotropium procumbens is otherwise native only inAmerica, it seems likely that it represents an early, accidentalintroduction into Guam from Mexico.

H. procumbens was first discovered in 1975 in Hawaiì, at the CampbellIndustrial Park at Barbers Point, Oàhu (Herbst and Ishikawa 5466:herbarium specimen, B.P. Bishop Museum), where it may have been broughtaccidentally by one of the frequent military flights between Hawaiì andGuam. On Oàhu, it was next seen at construction sites or other areashaving a direct connection to Barbers Point. For example, it now grows atthe Sand Island gas storage tanks that store gasoline refined at BarbersPoint (Herbst and Ishikawa 5643: herbarium specimen, B.P. Bishop Museum).This heliotrope also occurs at the ready-mix concrete plant on Sand Island(D.R. Herbst observation), which uses raw materials brought from BarbersPoint. Heliotropium procumbens has been spread from the concrete plantto various construction sites on Oàhu, such as the Entomology building atthe University of Hawaiì at Manoa, (D.R. Herbst observation) and aMariner's Ridge housing development (Nagata 1286: herbarium specimen, B.P.Bishop Museum). In recent years, it has gradually migrated to otherdisturbed sites on Oàhu and, more recently, has been collected on Maui


(Hobdy 1042: herbarium specimen, B.P. Bishop Museum). H. procumbentwas collected on Tern Island in 1978 and Midway Atoll in 1980. The Midwayand Tern Island plants probably were accidentally introduced by militaryflights from Barbers Point (Herbst 1980).

Although H. procumbens is not presently considered a managementproblem, it does represent a foreign element in the Leeward Islands andpotentially could replace native vegetation. No management plan is beingconsidered at present.

Golden crown-beard (Verbesina encelioides) has become a managementproblem. It was first reported on Kure Atoll by H.F. Clay in October 1959,growing near the radar reflector on Green Island. Bermuda grass (Cynodondactylon), ironwood, and sourbush (Pluchea symphytifolia) weregrowing with it. Clay surmised that seeds of these plants were brought onequipment from Midway Atoll in 1955 when the reflector was built (Clay1961). Golden crown-beard and sweet alyssum (Lobularia maritima) havebecome widespread on Green Island and are undergoing population explosions(Corn et al. 1981). Golden crown-beard forms a thicket that limits thespace for ground-nesting seabirds, and it has become necessary to mow thecentral plain of Green Island to restore nesting areas for the birds.

Three taxa of plants endemic to the Leeward Islands are believed to havebecome extinct within the last 20 or 25 years. These are kamanomano(Cenchrus agrimonioides var. laysanensis), Achyranthes atollensis,and Phyllostegia variabilis. On Green Island, all three grew only inthe central plain area. Competition from the golden crown-beard and sweetalyssum undoubtedly had a major role in their extinction on that island.

The Leeward Islands have had a long history of human disturbance, datingback on Nihoa and Necker to prehistoric times (Table 3). Other plants andanimals, such as Pritchardia sp., sandalwood (Santalum ellipticum),Laysan rail (Porzana palmeri), and Laysan honeycreeper (Himationesanguined freethii), on Laysan have become extinct as a direct result.

Naturally Introduced Alien Species: Nephrolepis multiflora andTree Heliotrope

Nephrolepis multiflora, a sword fern, is known from two of the LeewardIslands, Midway (Herbst, unpub. data) and Nihoa (Cpnant and Herbst 1983;Conant 1985). On Midway, the fern may have been intentionally introducedas an ornamental, or it may have been accidentally brought in with soil orother ornamental plants. Today, it persists in abandoned cultivated areasand appears to have been deliberately cultivated at one time.

The fern was first collected on Nihoa by S. Conant in 1981. A colonywas found on the western ridge of Miller Valley, a good distance away fromthe usual landing sites, and Conant believed that the spores arrived bywind dispersion rather than by accidental or intentional humanintervention. In 1983, Conant discovered two additional small coloniesthat probably were established after her 1981 sighting. In 1983 theoriginal colony had been reduced in size due to an invasion of its habitatby Eragrostis variabilis, a native grass. The fern does not appear to


be aggressive enough to impact native vegetation, and no management plan isbeing considered for its control.

Tree heliotrope, Tournefortia argentea, naturalized on the mainislands since the mid 1800s, is known from six of the Leeward Islands. Thespecies was growing on Pearl and Hermes Atoll as early as 1923 (Amerson et al. 1974), where it probably was introduced as an ornamental. It

likely was planted on Midway for the same purpose at an earlier date, butwe have no record of its original introduction. The oldest trees on TernIsland are believed to have been planted in the mid 1950s by the CoastGuard when that agency was upgrading and landscaping the old naval facilityon the island. In 1959, H. Clay found a single young tree growing on thewindward shore of Green Island, Kure Atoll (Clay 1961). Two years later asingle small tree was found growing on the northwestern beach of Laysan(Tsuda 1965). The fruits of tree heliotrope contain corky tissue and arewell adapted for water dispersal; the Kure and Laysan plants probably werethe results of natural dispersal by ocean currents.

Studies of mercury in Hawai`ian seabird eggs indicated that Midway andLaysan are influenced by the eastward-flowing Kuroshio Extension and NorthPacific Current, while the currents around the other sites sampled duringthe study were more affected by the westward-flowing California CurrentExtension and North Equatorial Current (Ohlendorf and Harrison 1968). Ifthis is correct, it would seem most probable that Midway was the source ofboth the Kure and the Laysan introductions of this species.

MANAGEMENT NEEDS

Except for the examples cited above, little has been done to manage thealien plants in the Leeward Islands. This is partly due to the logisticsinvolved, as the islands are remote and getting to them is expensive andtime consuming. Moreover, it is difficult or impossible to land on mostislands except during the few times of the year that the seas are calm.The main reason for inaction, however, is that there has been little needseen for managing alien plants, other than taking precautions that no newspecies are introduced. These precautions should include careful cleaningand inspection of all equipment and supplies used in temporary campsestablished to accommodate scientists studying the wildlife of the Islands.

Most of the naturalized species, except for those on the three islandsinhabited by humans, appear to be in balance with the rest of the flora.Some even appear to be beneficial to some of the native birds because theyprovide additional nesting sites and food and nesting material. Sandbur(Cenchrus echinatus), for example, provides additional nesting sitesfor wedge-tailed shearwaters (Puffinus pacificus chlororhynchus) andother burrowing birds. The fibrous root system binds the loose sand,allowing the birds to construct nesting burrows beneath them, while theleaves and seeds are used for nesting material and food by the Laysan finch(Telespyza cantons). Tree heliotrope, ironwoqd, and the naturalizedshrubs provide additional nesting sites for noddies and other birds thatprefer to nest off the ground.


RECOMMENDATIONS

The vegetation dynamics in the Leeward Islands should be closelymonitored to determine changes in composition and to detect newintroductions of plants or animals that may impact the existingvegetation. Also, general vegetation maps of each island are needed.Vegetation mapping methods for the Leeward Islands, using aerialphotography, have been developed. Detailed distributional maps of allalien species could be completed while ground-truthing the generalvegetation maps for all Islands.

Most of our knowledge about the vegetation of the Leeward Islands isbased upon casual observations made by ornithologists during the course oftheir primary research, or by botanists or other biologists during briefvisits. The study of ironwood by Apfelbaum et al. (1983) on Midway,although limited, constitutes one of the few detailed studies of plantsundertaken in the Leeward Islands. Permanent transects and photo stationsand the collection of baseline data are necessary for the development oflong-term management plans.

A well-illustrated, nontechnical manual of the plants of the LeewardIslands would aid in monitoring the vegetation. It could be used bynon-botanists to detect new island introductions or populations as well asdistributional changes of both native and alien plants and would be avaluable tool for individuals requiring plant identification as an aspectof their research (e.g., the host plant of an insect or the nestingsite of a bird). A complete species list of the plants of Midway Atolldoes not yet exist, but when a floristic survey of that atoll is completed,sufficient information about the Leeward Island plants will be available todevelop such a manual. The appendix provided here is intended as aninterim aid.

It is equally important that the Islands be monitored for theintroduction of alien animals, particularly rats (Rattus spp.).Shipwrecks provide the most likely means for introducing rats to theislands. Within recent years, fishing vessels have gone aground on LaysanIsland and French Frigate Atoll, but fortunately no rats were aboard. Ratshave been established on Midway Atoll since 1943 (Kramer 1971), where theyhave caused damage to both birds and vegetation, as they eat both the seedsand the vegetative parts of plants. Rat damage to beach naupaka is mostconspicuous; they chew through the branches just below the tuft of leavesto get the nutrients and moisture in the stem's pith, and the remainingtissue cannot support the weight of the leaves, causing the branches tobreak.

Mosquitoes, which are naturalized on Midway (U.S. Fish and WildlifeService, unpub. report), carry fowl pox, infecting young albatross chicks.During their inactive periods, mosquitoes rest in the shelter of the densevegetation surrounding the nests. Mowing the nesting areas has greatlydecreased the incidence of fowl pox in the albatross colony. The "Scaevoladieback" on Lisianski and Laysan Islands may have been caused by anintroduced scale insect. The insect may have weakened the shrubs to the


point where they could not withstand the stress of a prolonged period ofdrought followed by a severe winter storm.

Finally, it is important to develop a management plan for the control ofalien species, both plant and animal, impacting the vegetation and wildlifeof the islands. The plan should include a means for monitoring anddetecting new introductions and for determining population and distributionchanges of existing naturalized organisms, and it should discuss thevarious methods of containing or removing the alien species.

ACKNOWLEDGMENTS

The contribution by Warren L. Wagner was supported by a grant from theIrwin Charity Foundation of San Francisco to S.H. Sohmer. We thank S.W.Mill for assisting with the manuscript preparation and editing. Weappreciate comments on an earlier draft of the paper by S. Conant andS. Fefer.


APPENDIX

Checklist of the Vascular Plants of the Leeward Hawaiìan Islands

This list is based on the collections in the herbaria of the B.P. Bishop Museum and theUniversity of Hawaiì, literature citations, and personal collections and observations. Itshould be considered a preliminary report, as the identification of many of the specimenshas not been verified. Each entry includes the scientific name, the common name, thedistribution, and the status of the species within the Leeward Islands.

The symbols used for status are:E- endemic to the Leeward Islands; i.e., occurring naturally nowhere else in the

world, including the main Hawaiìan Islands.I - indigenous; i.e., native to the Leeward Islands but occurring naturally (without

the aid of humans) elsewhere; however, the plant may be endemic to the Hawaiìanarchipelago.

X- non-cultivated alien species; i.e., plants of accidental or deliberate introductionthat have become naturalized on the islands.

C- cultivated species which have not become naturalized.S - seeds that arrived through natural dispersal means. These are most commonly found in

the debris line.P - intentionally introduced propagules, as seeds or cuttings, that did not germinate or

live.? - status unknown.

- an underlined symbol denotes a non-vouchered species reported in the literature as anobservation.

In the case of multiple entries, only the symbol which best describes the presentstatus of the species is used. For example, Tern Island, French Frigate Shoals, has bothcultivated ironwood trees (C) and wild plants that have seeded from these planted specimens(X); only the "X" symbol is used in the list, as it gives the most dynamic statement ofthat plant's present status on that island.

Symbols for the island groups are: N = Nihoa; Ne = Necker; FF = French Frigate Shoals;GP = Gardner Pinnacles; La = Laysan Island; Li = Lisianski Island; PH = Pearl andHermes Atoll; Mi = Midway Atoll; Ku = Kure Atoll.

Appendix, continued.


Taxon

PTERIDOPHYTA

DAVALLIACEAE (Sword fern family)Nephrolepis multiflora (Roxb.) Morton

DICKSONIACEAE (Dicksonia family)Cibotium sp.

POLYPODIACEAE (Common fern family)Phymatodes scolopendria (J. Burm.) Clung

PSILOTACEAE (Psilotum family)Psilotum nudum (L.) Griseb.

GYMNOSPERMAE

ARAUCARIACEAE (Araucaria family)Araucaria heterophylla (Salisb.) Franco

CUPRESSACEAE (Cypress family)Cupressus sp.

CYCADACEAE (Cycas family)Cycas cirdnalis L.Cycas revoluta Thunb.

Common Name

sword fern

Hawaiian tree fern

laua`e

moa

Norfolk Island pine

sago palmsago palm

Island GroupN

X

Ne FF GP La Li PH Mi

C

C

C

?

C

C

CC

Ku

C


Taxon

MONOCOTYLEDONAE

AGAVACEAE (Agave family)Agave sisalana PerrineCordyline fruticosa (L.) A. Chev.Cordyline sp.Dracaena sp.Sanseviera sp.

ALOEACEAE (Aloe family)Aloe sp.

ARACEAE (Taro family)Alocasia cucullata (Lour.) G. DonAnthuriwn andraeanum Lindl.Caladium bicolor (Alton) Vent.Colocasia esculenta (L.) SchottDieffenbachia sp.Monstera delidosa Liebm.Philodendron sp.Rhaphidophora aurea (Andre) BirdseySyngonium podophyllum SchottXanthosoma sp.

ARECACEAE (Palm family)Cocos nudfera L.Livistona australis (R. Br.) Mart.

Common Name

sisalti

bowstring hemp

aloe

Chinese taroanthuriumcaladiumtarodumb canemonsteraphilodendrontaro vinesyngonium`ape

coconutcabbage palm

Island GroupN Ne

P

FF

CP

GP La

X

Li

C

PH Mi

CCCC

CCCCCCCC

C

C

Ku

C

C


Taxon

Phoenix dactylifera L.Phoenix sp.Pritchardia martii (Gaud.) HA. Wendl.Pritchardia pacifica Seem. & HA. Wendl.Pritchardia remota (Kuntze) Becc.Pritchardia sp.Roystonea sp.

CANNACEAE (Canna family)Canna indica L.

COMMELINACEAE (Spiderwort family)Commelina diffusa N.L. Burm.Dichorisandra thyrsiflora Mikan.Tradescantia spathacea Sw.Tradescantia zebrina Bosse

CYPERACEAE (Sedge family)Cyperus altemifolius

subsp. flabelliformis (Rottb.) Kukenth.Cyperus laevigatus L.Cyperus papyrus L.Cyperus rotundas L.Fimbristylis cymosa

subsp. spathacea (Roth) T. KoyamaFimbristylis cymosa subsp.

umbellato-capitata (Hillebr.) T. Koyama

Common Name

date palm

louluvinloulu

canna

honohonoblue gingeroyster plantwandering Jew

umbrella plantmakaloapapyrusnutgrass

button sedge

button sedge

Island GroupN

E

Ne

P

P

FF

PP

X

I

GP La

P

P

?

I

I

I

Li PH

P

Mi

CC

C

XCCC

X

CX

I

I

Ku

X

I


Taxon

Mariscus javanicus (Houtt.) Merr. & MetcalfeMariscus pennatiformis

subsp. bryanii (Kukenth.) T. KoyamaPycreus polystachyos (Rottb.) P. Beauv.

subsp. pofystachyos

LILIACEAE (Lily family)Allium cepa L.Allium fistulosum L.Asparagus setaceus (Kunth) JessopChlorophytum capense (L.) VossCrinum asiaticum L.Hemerocallis sp.Hippeastrum sp.Pancratium littorale Jacq.

MUSACEAE (Banana family)Heliconia psittacorum L. ffl.Musa sp.Strelitzia reginae Banks

ORCHIDACEAE (Orchid family)Vanda sp.Various orchids

PANDANACEAE (Screwpine family)Pandanus tectorius S. Parldnson ex Z.

Common Name

`ahu`awa

onionbunch onionasparagus fernbracket plantspider lilyday lily

spider lily

bananabird of paradise

hala

Island GroupN Ne FF

C

C

C

GP La

E

X

Li PH

X

Mi

I

I

CCCC

CCC

CC

C

Ku

C

C


Taxon

POACEAE (Grass family)Ammophila arenaria (L.) LinkBrachiaria mutica (Forssk.) StapfBromus willdenowii KunthCenchrus agrimonioides var. laysanensis F. Br.Cenchrus echinatus L.Chloris barbata (L.) Sw.Chloris virgata Sw.Cymbopogon citratus (Nees) StapfCynodon dactylon (L.) Pers.Digitaria ciliaris (Retz.) KoelerDigitaria insularis (L.) EkmanEleusine indica (L.) Gaertn.Eragrostis tenella (L.) Roem. & Schult.Eragrostis variabilis (Gaud.) Steud.Eragrostis paupera Jedwabn.Eustachys petraea Desf.Hordeum leporinum LinkLepturus repens (G. Forst.) R. Br. var. repensLepturus repens var. cinereus (Burch.) Fosb.Lepturus repens var. occidentalis Fosb.Lepturus repens var. subulatus Fosb.Melinis minutiflora P. Beauv.Panicum torridum Gaud.Paspalum fimbriatum KunthPaspalum urvillei Steud.Paspalum sp.

Common Name

European beachgrassparagrassprairie grass

sandburswollen fingergrassfeather fingergrasslemon grassBermuda grasskukaepua`asourgrassgoosegrassJapanese lovegrass`emoloa

molasses grasskakonakonaColumbia grassvasey grass

Island GroupN

X

I

I

Ne

I

X

FF

X

CXX

X

IX

I

I

GP La

EX

X

I

IP

Li

X

I

I

PH

X

II

II

Mi

XXXEXX

XXXXXIIXXI

I

X

Ku

EXXX

XX

XXII

I

X

X

Taxon

Poa annua L.Polypogon interruptus KunthPolypogon monspeliensis (L.) Desf.Rhynchelytrum repens (Willd.) Hubb.Setaria verticillata (L.) P. Beauv.Sporobolus africanus (Poir.) Robyns&TournaySporobolus indicus (L.) R. Br.Sporobolus pyramidatus (Lam) Hitchc.Sporobolus virginicus (L.) KunthStenotaphrum secundatum (Walter) KuntzeVulpia myuros (L.) GmelinZea mays L.

ZINGIBERACEAE (Ginger family)Catimbium speciosum (Wendl.) HolttumHedychium gardnerianum Ker-Gawl.

DICOTYLEDONAE

ACANTHACEAE (Acanthus family)Asystasia gangetica (L.) T. AndersonOdontonema stictum (Nees) KuntzeRuellia brittoniana E. Leonard

AIZOACEAE (Carpetweed family)Sesuvium portulacastrum (L.) L.Tetragonia tetragonioides (Pall.) Kuntze

Common Name

annual bluegrassditch polypogonrabbitfoot grassNatal redtopbristly foxtailAfrican dropseedWest Indian dropseed

beach dropseedbuffalo grassfoxtail fescuecorn

shell gingerkahili ginger

asystasiaodontonema

`akulikuliNew Zealand spinach

Island GroupN

X

X

Ne

I

FF

X

X

C

GP La

I

I

Li

I

PH

X

I

Mi

XXXXXXXXIXX

CC

XCX

Ku

XX

X

X

Appendix, continued. 209


Taxon

AMARANTHACEAE (Amaranth family)Achyranthes aspera L. var. asperaAchyranthes atollensis St. JohnAltemanthera tenella CollaAmaranthus brownii Christoph. & GaumAmaranthus dubius Mart.Amaranthus hybridus L.Amaranthus spinosus L.Amaranthus viridis L.

ANACARDIACEAE (Mango family)Mangifera indica L.Schinus terebinthifolius Raddi

APIACEAE (Carrot family)Ciclospermum leptophyllum (Pers.) Sprague

APOCYNACEAE (Periwinkle family)Allamanda cathartica L.Carissa macrocarpa (Eckl.) DC.Cascabela thevetia (L.) LippoldCatharanthus roseus (L.) G. DonErvatamia sp.Nerium oleander L.Plumeria obtusa L.Plumeria sp.

Common Name

joyweed

pakaigreen amaranthspiny pigweedslender amaranth

mangoChristmas berry

fine-leaved celery

allamandaNatal plumbe-still treeperiwinklecrape jasmineoleanderSingapore plumeriaplumeria

Island GroupN

E

Ne FF GP La

XE

X

Li PH

E

Mi

EC

XXXX

CX

X

CCCC

Ku

E

XX

X


Taxon

ARALIACEAE (Ginseng family)Polyscias guilfoylei (Bull) BaileySchefflera actinophylla (Endl.) Harms

ASTERACEAE (Daisy family)Arctium lappa L.Bidens alba var. radiata

(Schultz-Bip.) MelchertBidens pilosa L.Conyza bonariensis (L.) Cronq.Emilia fosbergii NicolsonGaillardia pulchella Foug.Gnaphalium purpureum L.Gnaphalium sandwicensium Gaud.var. sandwicensium

Helianthus annuusvar. macrocarpus (DC.) Cockerell

Lactuca sativa L.Lipochaeta integrifolia (Nutt.) A. GrayPluchea x fosbergii Cooperr. & GalangPluchea indica (L.) Less.Pluchea symphytifolia (Mill.) GillisSonchus oleraceus L.Tridaxprocumbens L.Verbesina encelioides (Cav.) Benth. & Hook.Wedelia trilobata (L.) Hitchc.Xanthium strumarium

var. canadense (Mill.) Torr. & A. Gray

Common Name

panaxoctopus tree

gobo

Spanish needlesSpanish needlesiliohared pualelewaikahulicudweed

`ena`ena

sunflowerlettucenehe

Indian plucheasour bushsow thistlecoat buttonsgolden crown-beardwedelia

cocklebur

Island GroupN Ne FF

X

C

X

GP La

X

I

X

Li PH

X

Mi

CC

C

XXX

X

I

C

XXXXXXX

X

Ku

X

XXX

I

CI

XX

X


Taxon

BIGNONIACEAE (Bignonia family)Spathodea campanulata P. Beauv.Tabebuia sp.

BORAGINACEAE (Heliotrope family)Cordia sebestena L.Cordia subcordata Lam.Heliotropium curassavicum L.Heliotropium procumbens var. depressum (Cham.) Fosb.Tournefortia argentea L. fil.

BRASSICACEAE (Mustard family)Brassica campestris L.Brassica nigra (L.) W. KochCapsella rubella Reut.Coronopus didymus (L.) Sm.Lepidium bidentatum var. o-waihiense

(Cham. & Schlecht.) Fosb.Lepidium virginicum L.Lobularia maritima (L.) Desv.Raphanus sativus L.

CACTACEAE (Cactus family)Epiphyllum oxypetalum (DC.) Haw.Hylocereus undatus (Haw.) Britton & Rose

CAPPARACEAE (Caper family)Capparis sandwichiana DC.

Common Name

African tulip tree

kou-haolekounena

tree heliotrope

field mustardblack mustardshepherd's purseswine cress

`anaunaupepper grasssweet alyssumradish

gooseneck cactusnight blooming cereus

puapilo

Island GrouoN

I

Ne FF

IXX

X

GP La

PI

X

I

I

Li

X

PH

X

X

X

I

I

Mi

CC

C

XX

XXX

IXXC

XC

I

Ku

X

X

IXX

to


Taxon

CARICACEAE (Papaya family)Carica papaya L.

CARYOPHYLLACEAE (Pink family)Cerastium fontanumvar. triviale (Link) Jalas

Schiedea verticillata F. Br.Spergularia marina (L.) Griseb.Stellaria media (L.) Vill.

CASUARINACEAE (Casuarina family)Casuarina equisetifolia L.

CHENOPODIACEAE (Goosefoot family)Atriplex suberecta Verd.Beta vulgaris var. cicla L.Chenopodium murale L.Chenopodium oahuense (Meyen) AellenSalicomia virginica L.

CLUSIACEAE (Mangosteen family)Calophyllum inophyllum L.Clusia rosea Jacq.

COMBRETACEAE (Indian almond family)Conocarpus erectus L.Terminalia catappa L.Terminalia myriocarpa Heurck & Muell.-Arg.

Common Name

papaya

common mouse ear

mimi iliocommon duckweed

ironwood

salt bushSwiss chardnettle-leaved goosefoot`aweoweosamphire

kamanicopey

buttonwoodfalse kamani

Island GroupN

E

I

Ne

P

I

FF

X

X

XCXIX

S

CC

GP La

C

P

I

P

PPP

Li

C

I

PH

C

Mi

C

X

XX

X

X

CC

CC

Ku

XX

C

X

C


Taxon

CONVOLVULACEAE (Morning-glory family)Convolvulus sp.Ipomoea batatas (L.) Lam.Ipomoea indica f. albiflora StoneIpomoea indica (J. Burm.) Merr.

f. indicaIpomoea pes-caprae

subsp. brasiliensis (L.) Ooststr.Ipomoea sp.Ipomoea triloba L.Merremia tuberosa (L.) Rendle

CRASSULACEAE (Stonecrop family)Crassula sp.Kalanchoe daigremontiana x tubifloraKalanchoe pinnata (Lam.) Pers.Kalanchoe tubiflora (Harv.) Raym.-Hamet

CUCURBITACEAE (Gourd family)Cucumis sativus L.Cucurbita pepo L.Sicyos maximowiczii Cogn.Sicyos pachycarpus Hook. & ArnottSicyos semitonsus St. John

EUPHORBIACEAE (Spurge family)Acafypha wilkesiana Muell.-Arg.Aleurites moluccana (L.) Willd.

Common Name

sweet potato

koali`awahia

beach morning glory

little bellwood rose

air plantchandelier plant

cucumbersquash, zucchini

beefsteak plantkukui

Island GroupN

I

I

I

Ne

S

FF

I

CC

S

GP La

P

I

I

PIIE

S

Li

I

I

IS

I

S

PH

I

S

Mi

C

I

ICXC

CCXC

C

C

Ku

I

I

I

214

Taxon

Breynia disticha var. rosi-picta Hort.Chamaesyce celastroides (Boiss.) Croizat & Degenervar. celastroides

Chamaesyce hirta (L.) Millsp.Chamaesyce hypericifolia (L.) Millsp.Chamaesyce maculata (L.) SmallChamaesyce prostata (Alton) SmallCodiaeum variegatumvar.pictum (Lodd.) Muell.-Arg.

Euphorbia cyathopora MurrayEuphorbia heterophylla L.Euphorbia peplus L.Euphorbia pulcherrima Klotz.Pedilanthus tithymaloides (L.) PoitRicinus communis L.

FABACEAE (Pea family)Acacia famesiana (L.) Willd.Albizia lebbeck (L.) Benth.Caesalpinia bonduc (L.) Roxb.Canavalia ensiformis (L.) DC.Cassia sp.Crotalaria incana L.Crotalaria pallida AitonDelonix regia (Hook.) Raf .Desmanthus virgatus (L.) Willd.Desmodium sandwicense E. Mey.

Common Name

snow bush

`akokohairy spurge

small ground fig

crotonwild poinsettia

petty spurgepoinsettiaslipper flowercastor bean

kmsiris treekakalaioa, kinikinijackbean

fuzzy rattle-podrattle-podroyal poincianavirgate mimosaiclover

Island GroupN

I

Ne FF

XX

X

GP La

SP

Li PH Mi

C

XXXX

CXXXCCX

XC

CXXCXX

Ku

XXX

C

X



Taxon

Dioclea wilsonii Standl.Entada scandens (Rosb.) Benth.Eiythrina variegatavar. orientalis (L.) Merr.

Erythrina sp.Glydne soja (L.) Sieb. & Zucc.Haematoxylum campechianum L.Leucaena leucocephala (Lam.) de WitMedicago lupulina L.Medicago polymorpha L.Medicago sativa L.Melilotus alba Medik.Melilotus indica (L.) All.Mucuna gigantea (Willd.) Medik.Mucuna urens (L.) Medik.Mucuna sp.Phaseolus vulgaris L.Prosopis pallida (Willd.) KunthSamanea saman (Jacq.) Merr.Senna surattensis (N. L. Burm.)

H. Irwin & BarnebySesbania tomentosa Hook & ArnottSeedling legume

FRANKENIACEAE (Frankenia family)Frankenia salina (Molina) I.M. Johnst.

Common Name

sea bean

tiger's claw

soy beanlogwood treekoa haoleblack medicbur cloveralfalfawhite sweet cloversweet cloverkaèèsea bean

green beankiawemonkeypod

kolomonaòhai

Island GroupN

I

Ne

P

I

FF

PX

C

X

GP

S

La

SS

PP

SS

Li

?

PH Mi

C

C

XX

XXX

CCC

C

Ku

C

X

216

Taxon

GENTIANACEAE (Gentian family)Centaurium erythraea Raf.subsp. erythraea

GERANIACEAE (Geranium family)Pelargonium hortorum Bailey

GOODENIACEAE (Goodenia family)Scaevola sericea Vahl

HYDROPHYLLACEAE (Water-leaf family)Nama sandwicensis A. Gray

LAMIACEAE (Mint family)Phyllostegia variabilis BitterPlectranthus scutellarioides (L.) R. Br.Stachys arvensis L.

LECYTHIDACEAE (Barringtonia family)Barringtonia asiatica (L.) Kurz

MALPIGHIACEAE (Malphigia family)Tristellateia australasiae Rich.

MALVACEAE (Mallow family)Abutilon grandifolium (Willd.) SweetHibiscus esculentus L.

Common Name

bitter herb

fish geranium

naupaka kahakai

Hawaiian nama

coleusstaggerweed

barringtonia

bagnit

hairy abutilonokra

Island GroupN Ne FF

I

C

GP La

I

I

E

P

Li

I

I

P

PH

I

Mi

X

C

I

ECX

C

XC

Ku

I

E



Taxon

Hibiscus rosa-sinensis L.Hibiscus tiliaceus L.Hibiscus sp.Malva parviflora L.Malvastrum coromandelianum (L.) Garcke

subsp. coromandelianumMalvaviscus arboreus Cav.Sidafallax Walp.Thespesia populnea (L.) Correa

MORACEAE (Mulberry family)Ficus benghalensis L.Ficus elastica Hornem.Ficus microcarpa L. fil.Ficus sp.Morns alba L.

MYRTACEAE (Myrtle family)Eugenia uniflora L.Psidium guajava L.

NYCTAGINACEAE (Four o'clock family)Boerhavia herbstii Fosb.Boerhavia repens L.Bougainvillea spectabilis Willd.Mirabilis jalapa L.Pisonia grandis R. Br.

Common Name

red hibiscushau

cheese weed

false mallowturks cap`ilimamilo

Indian banyanIndian rubber treeChinese banyan

white mulberry

Surinam cherryguava

alenabougainvilleafour-o'clock

N

I

I

Ne

P

FF

C

X

C

I

IslanGP

d GrLa

C

I

oupLi

II

I

PH

P

X

II

Mi

CCCX

XCIC

CCC

CC

cC

ICX

Ku

C

C

I

218

Appendix, continued.219

Taxon

OLEACEAE (Olive family)Jasminum sambac (L.) AltonNoronhia emarginata (Lam.) Stadm.

ONAGRACEAE (Evening primrose family)Oenothera ladniata J. Hill

OXALIDACEAE (Wood sorrel family)Oxalis comiculata L.Oxalis corymbosa DC.

PASSIFLORACEAE (Passion flower family)Passiflora edulis Sims

PLANTAGINACEAE (Plantain family)Plantago lanceolata L.Plantago major L.

PLUMBAGINACEAE (Leadwort family)Plumbago auriculata Lam.

POLYGONACEAE (Buckwheat family)Coccoloba uvifera (L.) L.Rumex albescens Hillebr.

PORTULACACEAE (Purslane family)Portulaca lutea G. ForsterPortulaca oleracea L.

Common Name

pikakeMadagascar olive

yellow wood sorrelpink wood sorrel

liliko i

English plantaincommon plantain

plumbago

sea grape

`ihicommon purslane

Island GroupN

I

IX

Ne

I

FF

X

C

IX

GP

I

La

P

IX

Li

I

PH

IX

Mi

CC

X

XX

C

XX

C

C

IX

Ku

C

X

X

C

X


Taxon

Portulaca villosa Cham.Portulacaria afra Jacq.

PRIMULACEAE (Primrose family)Anagallis arvensis L.

ROSACEAE (Rose family)Rosa sp.

RUBIACEAE (Coffee family)Gardenia sp.

RUTACEAE (Citrus family)Citrus sp.Murraya paniculata (L.) Jack

SANTALACEAE (Sandalwood family)Santalum ellipticum Gaud.

SCROPHULARIACEAE (Figwort family)Bacopa monnieri (L.) Wettst.

SOLANACEAE (Nightshade family)Capsicum annuum L.Lycopersicon esculentum Mill.Nicotiana glauca R.C. GrahamNicotiana tabacum L.

Common Name

jade tree

scarlet pimpernel

rose

mock orange

iliahialoè

àeàe

nioitomatotree tobaccotobacco

Island GroupN

I

Ne

P

FF

C

GP La

I

PX

Li

X

PH Mi

X

C

C

CC

I

C

Ku

X

Taxon

Solatium americanum Mill.Solanum melongena L.Solanum nelsonii DunalSolanum tuberosum L.

STERCULIACEAE (Cocoa family)Waltheria indica L.

TROPAEOLACEAE (Nasturtium family)Tropaeolum majus L.

URTICACEAE (Nettle family)Pilea microphylla (L.) Liebm.

VERBENACEAE (Verbena family)Lantana camara L.Phyla nodiflora (L.) GreeneStachytarpheta dichotoma (Ruiz & Pav.) VahlStachytarpheta jamaicensis (L.) VahlVerbena litoralis KunthVitex trifolia var. subtrisecta (Kuntze) Mold,

f. subtrisectaVitex trifolia var. subtrisecta

f. variegata Mold.

VIOLACEAE (Violet family)Viola odorata L.

Common Name

popoloeggplant

potato

`uhaloa

nasturtium

artillery plant

lantana

oiowlha`uowl

polinalina

polinalina

violet

Island GrouoN

I

I

Ne FF

C

GP La

I

IX

Li

I

PH

I

I

Mi

I

I

I

X

X

XXXXX

C

C

C

Ku

I

I



Taxon

VITACEAE (Grape family)Vitis sp.

ZYGOPHYLLACEAE (Tribulus family)Tributes cistoides L.

Common Name

grape

nohu

Island GroupN

I

Ne FF

I

GP

S

La

I

Li

I

PH

I

Mi

C

I

Ku

I


Literature Cited

Amerson, A.B., Jr. 1971. The natural history of French Frigate Shoals, NorthwesternHawaiìan Islands. Atoll Res. Bull. 150.

Amerson, A.B., Jr., R.B. Clapp, and W.O. Wirtz II. 1974. The natural history of Pearl andHermes Reef, Northwestern Hawaiìan Islands. Atoll Res. Bull. 174.

Apfelbaum, S.I., J.P. Ludwig, and C.E. Ludwig. 1983. Ecological problems associated withdisruption of dune vegetation dynamics by Casuarina equisetifolia L. at Sand Island,Midway Atoll. Atoll Res. Bull. 261.

Armstrong, R.W., ed. 1983. Atlas ofHawaiì, 2nd ed. Honolulu: Univ. Hawaiì Pr.

Bryan, E.H., Jr. 1978. The Northwestern Hawaiìan Islands: an annotatedbibliography. Honolulu: U.S. Fish and Wildl. Serv.

Christophersen, E., and E.L. Caum. 1931. Vascular plants of the Leeward Islands,Hawaiì. B.P. Bishop Mus. Bull. 81. Honolulu: Bishop Mus. Pr.

Clague, D., G.B. Dalrymple, and R. Moberly. 1975. Petrography and K-Ar ages of dredgedvolcanic rocks from the western Hawaiìan Ridge and the southern Emperor Seamount Chain.Bull. Geol. Soc. Amer. 86: 991-998.

Clapp, R.B. 1972. The natural history of Gardner Pinnacles, Northwestern HawaiìanIslands. Atoll Res. Bull. 163.

Clapp, R.B., and E. Kridler. 1977. The natural history of Necker Island, NorthwesternHawaiìan Islands. Atoll Res. Bull. 206.

Clapp, R.B., E. Kridler, and R.R. Fleet. 1977. The natural history of Nihoa Island,Northwestern Hawaiìan Islands. Atoll Res. Bull. 207.

Clapp, R.B., and W.O. Wirtz II. 1975. The natural history of Lisianski Island,Northwestern Hawaiìan Islands. Atoll Res. Bull. 186.

Clay, H.F. 1961. Narrative report of botanical field work on Kure Island. Atoll Res.Bull. 78.

Conant, S. 1985. Recent observations on the plants of Nihoa Island, Northwestern HawaiìanIslands. Pac. Sci. 39(2):135-149.

Conant, S., and D.R. Herbst. 1983. A record of Nephrolepis multiflora from NihoaIsland, Northwestern Hawaiìan Islands. Newsletter, Hawn. Bot. Soc. 22:17-19.

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