Endangered and Threatened Wildlife and Plants; Determination of Endangered or Threatened Status for 24 Plants From the Island of Kauai, HI

SUMMARY: The U.S. Fish and Wildlife Service (Service) determines

endangered status pursuant to the Endangered Species Act of 1973, as

amended (Act), for 21 plant taxa and threatened status for 3 plant

taxa. All but seven of the taxa are endemic to the island of Kauai,

Hawaiian Islands. The exceptions are found on the islands of Niihau,

Oahu, Molokai, Maui, and/or Hawaii, as well as Kauai. The 24 plant taxa

and their habitats have been variously affected or are currently

threatened by 1 or more of the following: habitat degradation by

animals (e.g., goats, pigs, axis and mule deer, cattle, and red jungle

fowl); competition for space, light, water, and nutrients by

naturalized, introduced vegetation; erosion of substrate produced by

weathering, or human or animal caused disturbance; recreational and

agricultural activities; habitat loss from fires; and predation by

animals (goats and rats). Due to the small number of existing

individuals and their very narrow distributions, these taxa and most of

their populations are subject to an increased likelihood of extinction

and/or reduced reproductive vigor from stochastic events. This rule

implements the protection and recovery provisions provided by the Act

for these plant taxa.

EFFECTIVE DATE: March 28, 1994.

ADDRESSES: The complete file for this rule is available for public

inspection, by appointment, during normal business hours at the U.S.

Fish and Wildlife Service, 300 Ala Moana Boulevard, room 6307, P.O. Box

50167, Honolulu, Hawaii 96850.

FOR FURTHER INFORMATION CONTACT: Robert P. Smith, at the above address

(808/541-2749).

SUPPLEMENTARY INFORMATION:

Background

Brighamia insignis, Cyanea asarifolia, Cyrtandra limahuliensis,

Delissea rhytidosperma, Diellia pallida, Exocarpos luteolus, Hedyotis

cookiana, Hibiscus clayi, Lipochaeta fauriei, Lipochaeta micrantha,

Lipochaeta waimeaensis, Lysimachia filifolia, Melicope haupuensis,

Melicope knudsenii, Melicope pallida, Melicope quadrangularis,

Munroidendron racemosum, Nothocestrum peltatum, Peucedanum sandwicense,

Phyllostegia waimeae, Pteralyxia kauaiensis, Schiedea spergulina var.

leiopoda, Schiedea spergulina var. spergulina, and Solanum sandwicense

are endemic to or have the majority of their populations on the island

of Kauai, Hawaii. Seventeen of these taxa are endemic to the island of

Kauai, Hawaii; two additional taxa are now found only on Kauai. One of

these taxa is now or was previously also known from Niihau, four from

Oahu, two from Molokai, two from Maui, and one from the island of

Hawaii.

The island of Kauai is the northernmost and oldest of the eight

major Hawaiian Islands (Foote et al. 1972). This highly eroded island,

characterized by deeply dissected canyons and steep ridges, is 553

square miles (sq mi) (1,430 sq kilometers (km)) in area (Department of

Geography 1983). Kauai was formed about six million years ago by a

single shield volcano. Its caldera, once the largest in the Hawaiian

Islands, now extends about 10 mi (16 km) in diameter and comprises the

extremely wet, elevated tableland of Alakai Swamp (Department of

Geography 1983). Because the highest point on Kauai, at Kawaikini Peak,

is only 5,243 feet (ft) (1,598 meters (m)) in elevation (Department of

Geography 1983), it lacks the contrasting leeward montane rainfall

patterns found on other islands that have higher mountain systems.

Rainfall is, therefore, distributed throughout the upper elevations,

especially at Mount Waialeale, Kauai's second highest point at 5,148 ft

elevation (1,569 m) (Department of Geography 1983). Mount Waialeale is

one of the wettest spots on earth, where annual rainfall averages 450

inches (in) (1,140 centimeters (cm)). (Honda et al. 1967, Joesting

1984). To the west of the Alakai Swamp is the deeply dissected Waimea

Canyon, extending 10 mi (16 km) in length and up to 1 mi (1.6 km) in

width. Later volcanic activity on the southeastern flank of the volcano

formed the smaller Haupu caldera. Subsequent erosion and collapse of

its flank formed Haupu Ridge (Macdonald et al. 1983). One of the

island's most famous features is the Na Pali Coast, where stream and

wave action have cut deep valleys and eroded the northern coast to form

precipitous cliffs as high as 3,000 ft (910 m) (Joesting 1984).

Because of its age and relative isolation, levels of floristic

diversity and endemism are higher on Kauai than on any other island in

the Hawaiian archipelago. However, the vegetation of Kauai has

undergone extreme alterations because of past and present land use.

Land with rich soils was altered by the early Hawaiians and more

recently converted to agricultural use or pasture (Gagne and Cuddihy

1990). Intentional or inadvertent introduction of alien plant and

animal taxa has also contributed to the reduction of native vegetation

on the island of Kauai. Native forests are now limited to the upper

elevation mesic and wet regions within Kauai's conservation district.

The 24 taxa in this rule occur in that district, between 400 and 4,000

ft (120 and 1,200 m) elevation in the western and northwestern portions

of the island or within large State owned tracts of natural area

reserves, forest reserves, and parks. Most of the taxa included in this

rule persist on steep slopes, precipitous cliffs, valley headwalls, and

other regions where unsuitable topography has prevented agricultural

development or where inaccessibility has limited encroachment by alien

animal and plant taxa.

The 24 taxa included in this rule are distributed throughout the

island of Kauai and grow in a variety of vegetation communities

(grassland, shrubland, and forests), elevational zones (coastal to

montane), and moisture regimes (dry to wet). Six taxa are found in

various lowland dry communities. These once abundant communities are

now fragmented due to fire, development, and the ingression of alien

plants and animals. Munroidendron racemosum extends from coastal mesic

vegetation communities to higher elevations in lowland dry (Hawaii

Heritage Program (HHP) 1990a) and mesic forests. Peucedanum sandwicense

is found within a variety of vegetation communities, ranging from

coastal to lowland dry to mesic shrublands and forests. Only 1 of the

24 taxa is found in grasslands. Brighamia insignis grows within Kauai's

lowland dry grassland and shrubland communities in the Na Pali region,

where the annual rainfall is usually less than 65 in (170 cm). Three

taxa, Hibiscus clayi, Delissea rhytidosperma, and Melicope knudsenii,

are located within lowland dry forests, the latter two extending into

mesic forests. Lowland dry forests are characterized by an annual

rainfall of 20 to 80 in (50 to 200 cm), which falls between November

and March, and a well-drained, highly weathered substrate rich in

aluminum (Gagne and Cuddihy 1990).

Nineteen of the 24 taxa have all or a significant number of their

populations in lowland mesic or wet forest communities. Lowland mesic

forest communities lie between 100 and 3,000 ft (30 and 1,000 m)

elevation and are characterized by a 6.5 to 65 ft (2 to 20 m) tall

canopy and a diverse understory of shrubs, herbs, and ferns. The annual

rainfall of 45 to 150 in (120 to 380 cm) falls predominantly between

October and March (Gagne and Cuddihy 1990). This mesic community often

grades into lowland wet forests which are typically found on the

windward side of the island or in sheltered leeward situations between

330 and 3,940 ft (100 and 1,200 m) elevation. The rainfall in this

lowland wet community may exceed 200 in (500 cm) per year. These

forests were once the predominant vegetation on Kauai but now exist

only on steep rocky terrain or cliff faces. The substrate is generally

of well-drained soils that may support tree canopies up to 130 ft (40

m) in height (Cuddihy and Stone 1990, Gagne and Cuddihy 1990).

The habitat of Solanum sandwicense extends to the higher elevation

and drier portions of montane mesic forests, whereas the habitat of

Exocarpos luteolus extends into montane wet forests. Nothocestrum

peltatum and Phyllostegia waimeae are the only taxa found strictly

within these montane communities, which typically occur above 3,000 ft

(1,000 m) elevation (HHP 1991). The annual rainfall in montane

communities may exceed 280 in (700 cm) (Gagne and Cuddihy 1990).

The land that supports these 24 plant taxa is owned by various

private parties, the City and County of Honolulu, and the State of

Hawaii (including State parks, forest reserves, natural area reserves,

the Seabird Sanctuary, and land managed under a cooperative agreement

with the National Park Service).

Discussion of the 24 Plant Taxa Included in This Final Rule

Asa Gray (in Mann 1868) described Brighamia insignis based upon

alcohol-preserved flowers and fruits collected by William Tufts Brigham

on Molokai and a dried specimen collected on Kauai or Niihau by

Ezechiel Jules Remy. The specific epithet means ``outstanding,''

referring to the plant's unique appearance. Brigham's bottled material,

since lost, would today be considered to be Brighamia rockii. Other

published names that Thomas G. Lammers (1989), in the currently

accepted treatment of the genus, considers to be synonymous with B.

insignis include B. insignis f. citrina (Forbes 1917a), B. citrina (St.

John 1958), and B. citrina var. napaliensis (St. John 1969b).

Brighamia insignis, a member of the bellflower family

(Campanulaceae), is an unbranched plant 3 to 16 ft (1 to 5 m) tall with

a succulent stem that is bulbous at the bottom and tapers toward the

top. The fleshy leaves, which measure 5 to 8 in (12 to 20 cm) long and

2.5 to 4.5 in (6.5 to 11 cm) wide, are arranged in a compact rosette at

the apex of the stem. Fragrant yellow flowers are clustered in groups

of 3 to 8 in the leaf axils (the point between the leaf and the stem),

with each flower on a stalk 0.4 to 1.2 in (1 to 3 cm) long. The

hypanthium (basal portion of the flower) has 10 ribs and is topped with

5 oval or loosely triangular calyx lobes (partially fused sepals) 0.02

to 0.04 in (0.5 to 1 millimeter (mm)) long. The yellow petals are fused

into a tube 2.8 to 5.5 in (7 to 14 cm) long and 0.1 to 0.2 in (3 to 4

mm) wide, which flares into five elliptic lobes. The fruit is a capsule

0.5 to 0.7 in (13 to 19 mm) long containing numerous seeds. This

species is a member of a unique endemic Hawaiian genus with only one

other species, presently known only from Molokai, from which it differs

by the color of its petals, its shorter calyx lobes, and its longer

flower stalks (Hillebrand 1888; Johnson 1986; Lammers 1990; Rock 1919;

St. John 1958, 1969b; Takeuchi 1982).

Historically, Brighamia insignis was known from the headland

between Hoolulu and Waiahuakua Valleys along the Na Pali Coast on the

island of Kauai, and from Kaali Spring on the island of Niihau (HHP

1991a1, 1991a2, 1991a4). The Na Pali Coast populations are still extant

and additional populations are known from the same general area. The

two Na Pali Coast populations within or on the boundary of the Hono O

Na Pali Natural Area Reserve (NAR) are within 0.4 mi (0.6 km) of each

other (HHP 1991a1, 1991a3). There are also two populations in the Haupu

Range within 2.7 mi (4.3 km) of each other (HHP 1991a2, 1991a5). In

1992, Hurricane Iniki destroyed approximately half of the individuals

in the Na Pali Coast populations and 7 of the 12 individuals in the

Haupu area (Perlman 1992; Steve Perlman, Hawaii Plant Conservation

Center (HPCC), pers. comm., 1992). The 5 populations grow on State and

private land and total fewer than 40 plants. The status of the small

population on privately owned Niihau is not known, although there are

reports that it was destroyed when the supporting cliff fell away (HHP

1991a4; Wichman and St. John 1990; Charles Christensen, Hawaii

Department of Agriculture (DOA), and John Fay, U.S. Fish and Wildlife

Service (FWS), pers. comms., 1991). This species grows predominantly on

the rocky ledges with little soil or steep sea cliffs in lowland dry

grassland and shrubland from sea level to 1,300 ft (400 m) elevation

(Gagne and Cuddihy 1990, Lammers 1990). Associated plant taxa include

Canthium odoratum (alahe'e), Chamaesyce celastroides ('akoko),

Eragrostis variabilis (kawelu), and Heteropogon contortus (pili grass)

(Gagne and Cuddihy 1990; HHP 1991a1 to 1991a3).

Feral goats (Capra hircus) pose the major threat to Brighamia

insignis by causing defoliation and stem damage, restricting

populations to inaccessible cliffs, and probably causing rock slides

that degrade the plant's habitat. Alien plant taxa are another major

threat to the survival of this species, especially introduced grasses

such as Melinis minutiflora (molasses grass), Setaria gracilis (yellow

foxtail), and Sporobolus africanus (smutgrass), which prevent

establishment of seedlings. Other alien plants that potentially pose a

threat are Lantana camara (lantana), Psidium cattleianum (strawberry

guava), Psidium guajava (common guava), and Syzygium cumini (Java

plum). Hikers transport weed seeds to areas where Brighamia insignis

grows and dislodge rocks that can damage plants. Wildfire also poses a

serious threat to this species. Some plants flower but fail to set

seed, which may be due to a lack of pollinators or a reduction in

genetic variability due to the few existing individuals. Brighamia

insignis is also threatened by stochastic extinction due to low total

numbers and the frequency of disturbance events, such as the rock

slides in their cliff habitat. Carmine spider mite (Tetranychus

cinnabarinus), an introduced insect, has been observed to cause leaf

loss in both cultivated and wild individuals of Brighamia insignis

(Christensen 1979; HHP 1991a1 to 1991a4; HPCC 1990a; Perlman 1979; St.

John 1969b, 1981b; Takeuchi 1982; Wagner et al. 1990; Tim Flynn,

National Tropical Botanical Garden (NTBG), pers. comm., 1991; S.

Perlman, pers. comm., 1993).

Robert W. Hobdy collected a specimen of Cyanea asarifolia on Kauai

in 1970. Harold St. John (1975) later described and named the taxon.

The specific epithet refers to the leaves, which are similar in shape

to those in the genus Asarum. Recently, St. John (1987d, St. John and

Takeuchi 1987) placed the genus Cyanea in synonymy with Delissea,

resulting in the new combination Delissea asarifolia, but Lammers

(1990) retains both genera in the currently accepted treatment of the

family.

Cyanea asarifolia, a member of the bellflower family, is a

sparingly branched shrub 1 to 3.3 ft (0.3 to 1 m) tall. The heart-

shaped leaves are 3.3 to 4.1 in (8.5 to 10.5 cm) long and 2.8 to 3.1 in

(7 to 8 cm) wide with leaf stalks 4.7 to 5.9 in (12 to 15 cm) long.

Thirty to 40 flowers are clustered on a stalk 1 to 1.2 in (25 to 30 mm)

long, each having an individual stalk 0.3 to 0.4 in (7 to 10 mm) in

length. The slightly curved flowers are white with purple stripes, 0.8

to 0.9 in (20 to 22 mm) long, and about 0.1 in (3.5 mm) wide with

spreading lobes. The five anthers have tufts of white hairs at the

tips. The nearly spherical fruit is a dark purple berry about 0.4 in (1

cm) long. This species is distinguished from others of the genus that

grow on Kauai by the shape of the leaf base, the leaf width in

proportion to the length, and the presence of a leaf stalk (Lammers

1990, St. John 1975).

For over 20 years, Cyanea asarifolia was known only from a

population of five or six plants above the bed of Anahola Stream on

Kauai at its type locality (HHP 1991b1). Because recent attempts to

locate this population were unsuccessful, this population is now

thought to be extirpated (T. Flynn, pers. comm., 1991). In 1991, Steven

Perlman and Ken Marr discovered a population of 14 mature plants and 5

seedlings at the headwaters of the Wailua River in central Kauai on

State owned land (HHP 1991b2; S. Perlman, pers. comm., 1991). This

species typically grows in pockets of soil on sheer rock cliffs in

lowland wet forests (Ken Marr, University of British Columbia, pers.

comm., 1991) at an elevation of approximately 1,080 ft (330 m).

Associated plant taxa include ferns, Hedyotis elatior (manono),

Metrosideros polymorpha (`ohi'a), Touchardia latifolia (olona), and

Urera glabra (opuhe) (Lammers 1990; St. John 1975; Robert Hobdy, Hawaii

Department of Land and Natural Resources (DLNR), and S. Perlman, pers.

comms., 1991).

Cyanea asarifolia is threatened by stochastic extinction and/or

reduced reproductive vigor due to the small number of existing

individuals. Plants in the area in which the only currently known

population occurs are vulnerable to occasional hurricanes, natural rock

slides, and over-collecting for scientific purposes. In 1992, Hurricane

Iniki heavily damaged the Cyanea asarifolia population, either directly

or indirectly destroying all but four or five juvenile plants. Plants

observed after Hurricane Iniki were frequently damaged by introduced

slugs or rodents (Loyal Mehrhoff, FWS, pers. comm., 1993). Habitat

degradation by feral pigs (Sus scrofa), at least one of which has

invaded the plant's habitat, is a potential threat (T. Flynn, David

Lorence, NTBG, and S. Perlman, pers. comms., 1991).

Lawrence H. MacDaniels first collected Cyrtandra limahuliensis on

Kauai in 1926. St. John (1987a) described the species, naming it for

Limahuli Valley, where Steven Perlman collected the type specimen in

1978.

Cyrtandra limahuliensis, a member of the African violet family

(Gesneriaceae), is an unbranched or few-branched shrub up to 5 ft (1.5

m) tall. The opposite, elliptic leaves are usually 6 to 12 in (15 to 30

cm) long and 2 to 4.7 in (5 to 12 cm) wide. The upper surface of the

toothed leaves is moderately hairy and the lower surface, with deep

veins, is moderately or densely covered with yellowish brown hairs.

Single downy flowers are borne in the leaf axils. The slightly curved

corolla tube (fused petals) barely extends beyond the calyx. The calyx

encloses the approximately 0.8 in (2 cm) long berries at maturity. The

following combination of characteristics distinguish this species from

others of the genus: the leaves are usually hairy, especially on lower

surfaces; the usually symmetrical calyx is tubular or funnel-shaped and

encloses the fruit at maturity; and the flowers are borne singly (St.

John 1987a, Wagner et al. 1990).

Historically, Cyrtandra limahuliensis was known from three areas on

Kauai: Wainiha Valley; Lumahai Valley; and near Kilauea River (HHP

1991c4, 1991c5, 1991c8; C. Christensen, pers. comm., 1992). One

population remains in Wainiha Valley and 11 others exist on Kauai in

Limahuli Valley, Waipa Valley, on Mount Kahili, along the north fork of

Wahiawa Stream, along Anahola Stream, Waioli Valley, and near Powerline

Trail on private and State land (HHP 1991c1 to 1991c3, 1991c5 to

1991c7; HPCC 1991a1 to 1991a2; T. Flynn, R. Hobdy, S. Perlman, and

Warren L. Wagner, Smithsonian Institution, pers. comms., 1991; D.

Lorence et al., in litt., 1991). The 12 known populations, distributed

over a 13 by 18 mi (20 by 30 km) area, range in size from solitary

shrubs to large populations of over 1,000 plants (HHP 1991c1 to 1991c3,

1991c5 to 1991c7; D. Lorence, in litt., 1993). The largest populations

of this species occur in the upper Waioli Valley, where 3 populations

total at least 2,100 individuals (D. Lorence, in litt., 1993). Another

location with ``hundreds or perhaps thousands'' of plants (W.L. Wagner,

pers. comm., 1991) is limited to a 0.25 sq mi (0.4 sq km) area along

the north fork of the Wailua River. Other botanists familiar with this

population believe it to number no more than 500 individuals (T. Flynn

and D. Lorence, pers. comms., 1991). A total of 2,800 to 3,000 plants

are known from these 12 populations. This species typically grows along

streams in lowland wet forests at elevations between 800 and 2,850 ft

(240 and 870 m) (Wagner et al. 1990). Associated taxa include Antidesma

platyphyllum var. hillebrandii (hame), Athyrium sandwichianum (ho`i'o),

Perrottetia sandwicensis (olomea), `ohi'a, Dicranopteris linearis

(uluhe), Gunnera kauaiensis (`ape'ape), Hedyotis sp. (manono), and

Psychotria sp. (kopiko) (HHP 1991c1, 1991c7; T. Flynn, pers. comm.,

1991).

The major threat to Cyrtandra limahuliensis populations is

competition with invasive alien taxa, especially strawberry guava. Each

population has additional threats: Competition with the introduced

grass Paspalum conjugatum (Hilo grass) and Melastoma candidum (NCN) at

the Mount Kahili population; competition with common guava and habitat

degradation by feral pigs at the Anahola Stream population; and

competition with Hedychium flavescens (yellow ginger) at the Wainiha

Valley population. Individuals of the Wailua Stream population are

situated at the base of a steep cliff and are vulnerable to natural

landslides. The Waioli Valley populations are threatened by several

alien weeds: Rubus rosifolius (thimbleberry); Youngia japonica

(Oriental hawksbeard); Erechtites valerianifolia (fireweed); and

Blechnum occidentale (NCN). Hurricanes are also a potential threat, but

most of the plants have grown back vigorously since Hurricane Iniki

(HHP 1991c1; T. Flynn, R. Hobdy, D. Lorence, and W.L. Wagner, pers.

comms., 1991; D. Lorence, in litt., 1993).

Cyrtandra limahuliensis is not immediately in danger of extinction,

but if the threats outlined above are not curtailed, the species will

become endangered in the future.

Remy first collected a specimen of Delissea rhytidosperma on Kauai

between 1851 and 1855. Horace Mann, Jr. (1867) chose the specific

epithet to describe its wrinkled seeds. Heinrich Wawra (1873) later

described another species, D. kealiae, which he said was closely

related to D. rhytidosperma. In the current treatment of the family,

Lammers (1990) considers D. kealiae to be synonymous with D.

rhytidosperma.

Delissea rhytidosperma, a member of the bellflower family, is a

branched shrub 1.6 to 8.2 ft (0.5 to 2.5 m) tall. The lance-shaped or

elliptic leaves are 3.1 to 7.5 in (8 to 19 cm) long and 0.8 to 2.2 in

(2 to 5.5 cm) wide and have toothed margins. Clusters of 5 to 12

flowers are borne on stalks 0.4 to 0.8 in (1 to 2 cm) long; each flower

has a stalk 0.3 to 0.5 in (8 to 13 mm) long. The greenish white

(sometimes pale purple) corolla is 0.6 to 0.8 in (14 to 20 mm) long.

The stamens are hairless, except for a small patch of hair at the base

of the anthers. The nearly spherical dark purple fruits are 0.3 to 0.5

in (7 to 12 mm) long and contain numerous white seeds. This species

differs from other taxa of the genus by the shape, length, and margins

of the leaves and by having hairs at the base of the anthers

(Hillebrand 1888; Lammers 1990; Rock 1913, 1919; Wimmer 1953).

Historically, Delissea rhytidosperma was known from scattered

locations throughout the island of Kauai. Populations ranged as far

north as Wainiha and Limahuli Valleys, as far east as Kapaa and Kealia,

and as far south as Haupu Range between the elevations of 1,000 and

3,000 ft (300 and 1,000 m) (HHP 1991d3 to 1991d7). Today, only one

population with six individuals, located in State owned Kuia NAR, is

known to exist (HHP 1991d1; S. Perlman, pers. comm., 1993). The only

other populations seen in recent years were a single plant in Limahuli

Valley which is now dead and 20 plants in the Haupu Range (Bruegmann

1990; HHP 1991d2; Perlman 1992; S. Perlman, pers. comm., 1991). The

latter population was destroyed by Hurricane Iniki in 1992 (Perlman

1992; S. Perlman, pers. comm., 1993). This species generally grows in

diverse lowland mesic forests or Acacia koa (koa)-dominated lowland dry

forests that have well-drained soils with medium- to fine-textured

subsoil (Foote et al. 1972, Gagne and Cuddihy 1990, Lammers 1990).

Associated plant taxa include Dianella sandwicensis (`uki'uki),

Diospyros sandwicensis (lama), Nestegis sandwicensis (olopua), and

Styphelia tameiameiae (pukiawe) (HHP 1991d1, 1991d2).

Habitat degradation by mule deer or black-tailed deer (Odocoileus

hemionus), feral goats, and feral pigs is the major threat affecting

the survival of Delissea rhytidosperma. Other threats are predation by

rats (Rattus spp.), fire, over-collecting for scientific or

horticultural purposes, landslides, and competition with alien plants

such as lantana, Passiflora ligularis (sweet granadilla), and P.

mollissima (banana poka). This species, with a single extant population

of six individuals, is threatened by stochastic extinction and/or

reduced reproductive vigor due to the small number of existing

individuals. Hurricanes are an additional, and major, threat (Bruegmann

1990; HHP 1991d1; HPCC 1990b; John Obata, HPCC, and S. Perlman, pers.

comms., 1991, 1993).

About 1875, Valdemar Knudsen, a rancher on Kauai, collected a fern

at Halemanu, which Wilhelm Hillebrand (1888) named Lindsaya laciniata,

the specific epithet referring to the divided fronds. Hillebrand also

indicated two varieties: var. subpinnata, a bipinnate form, which may

actually represent another species (Wagner 1952); and an unnamed form.

Friedrick Ludwig Emil Diels (1899) transferred the species to Diellia,

resulting in Diellia laciniata, the name in use at the time the species

was proposed (Lamoureux 1988). Recent studies have recognized these

populations as a new species, Diellia pallida (W.H. Wagner 1993).

Diellia pallida, a member of the spleenwort family (Aspleniaceae),

is a plant that grows in tufts of three to four light green, lance-

shaped fronds along with a few persistent dead ones. The midrib of the

frond ranges from dark purple to brownish gray in color and has a dull

sheen. Scales on the midrib are brown, gray, or black; 0.1 to 0.2 in (3

to 5 mm) long; and rather inconspicuous. The fronds measure 12 to 22 in

(30 to 55 cm) in length and 2 to 5 in (5 to 12 cm) in width and have

short black hairs on the underside. Each frond has approximately 20 to

40 pinnae (divisions or leaflets). The largest pinnae are in the middle

section of the frond, while the lower section has triangular, somewhat

reduced pinnae, with the lowermost pair of pinnae raised above the

plane of the others. The sori (groups of spore-producing bodies), which

are frequently fused along an extended line, are encircled by a

prominent vein. This species differs from others of this endemic

Hawaiian genus by the color and sheen of the midrib, the presence and

color of scales on the midrib, and the frequent fusion of sori

(Hillebrand 1888; Wagner 1952, 1987).

Diellia pallida was known historically from Halemanu on Kauai

(Hillebrand 1888). The species had not been seen since 1949, when a

collection was made in Kuia NAR (Warren H. Wagner, University of

Michigan, pers. comm., 1991). It is currently known from two

populations on State land on the island of Kauai within Kuia NAR and

Koaie Canyon. The recently discovered population on the west side of

Waimea Canyon within Puu Ka Pele Forest Reserve is now apparently

extirpated (CPC 1989a, 1990; HHP 1991e1 to 1991e3; Wagner 1952; D.

Lorence, pers. comm., 1991; S. Perlman, pers. comm., 1993; D. Lorence

et al., in litt., 1991). The two known populations extend over a 7 by 3

mi (11 by 5 km) area. In 1987, Joel Lau of The Nature Conservancy of

Hawaii (TNCH) discovered the Koaie Canyon population of three or four

individuals (Bruegmann 1990; HHP 1991e3; Joel Lau, Hawaii Heritage

Program, and S. Perlman, pers. comms., 1991). Botanists of NTBG have

since discovered two plants in Puu Ka Pele Forest Reserve, but the

plants have since disappeared and were likely destroyed by goats (D.

Lorence et al., in litt., 1991). Recent visits to the Kuia NAR and

Koaie populations have found a total of less than 10 extant individuals

for this species (HPCC 1991c; Perlman 1992; J. Lau and D. Lorence,

pers. comms., 1991; S. Perlman, pers. comms., 1991, 1993). This species

grows on bare soil on steep, rocky, dry slopes of lowland mesic

forests, 1,700 to 2,300 ft (530 to 690 m) in elevation. Associated

plant taxa include koa, Alectryon macrococcus (mahoe), Aleurites

moluccana (kukui), Antidesma platyphyllum (hame), 'ohi'a, Myrsine

lanaiensis (kolea), and Rauvolfia sandwicensis (hao) (HHP 1991e1 to

1991e3; S. Perlman, pers. comm., 1991; D. Lorence et al., in litt.,

1991).

Competition with alien plants, especially lantana and Melia

azedarach (Chinaberry), constitutes the major threat to Diellia

pallida. Introduced grasses, such as Stenotaphrum secundatum (St.

Augustine grass) and Oplismenus hirtellus (basketgrass), and two

naturalized taxa of Polynesian introduction, kukui and Cordyline

fruticosa (ti), degrade this species' habitat. Feral goats cause

erosion near the plants and trample and possibly browse these plants.

Other threats to this species are habitat degradation by feral pigs and

mule deer, fire, over-collecting for scientific purposes, as well as

stochastic extinction and/or reduced reproductive vigor due to the

small number of existing individuals (HHP 1991e2, 1991e3; Bruegmann

1990, Wagner 1950; J. Lau, S. Perlman, and D. Lorence, pers. comms.,

1991).

Reverend John Mortimer Lydgate first collected Exocarpos luteolus

in 1908 and Charles N. Forbes (1910) described the species two years

later. The specific epithet means ``yellow'' and refers to the color of

the receptacle (base of flower) and fruit.

Exocarpos luteolus, a member of the sandalwood family

(Santalaceae), is a moderately to densely branched shrub, 1.6 to 6.6 ft

(0.5 to 2 m) tall with knobby branches. The leaves are of two kinds,

minute scales and more typical leaves. The latter, which are usually

present, are elliptical, lance-shaped or oval, usually 2 to 3.2 in (5

to 8 cm) long and 1 to 1.4 in (25 to 36 mm) wide, and lack a leaf

stalk. The green flowers have five to six petals about 0.04 in (1 mm)

long. The pale yellow fruit is a drupe (single-seeded fleshy fruit),

usually 0.4 to 0.7 in (11 to 19 mm) long, with four distinct

indentations at the apex. About 0.2 to 0.4 in (6 to 9 mm) of the drupe

is exposed above the fleshy, golden-yellow receptacle. This species is

distinguished from others of the genus by its generally larger fruit

with four indentations and by the color of the receptacle and fruit

(Degener 1932a, 1932b; Forbes 1910; Wagner et al. 1990).

Historically, Exocarpos luteolus was known from three locations on

Kauai: Wahiawa Swamp; Kaholuamanu; and Kumuwela Ridge (HHP 1991f1,

1991f5, 1991f7). This species is now known to grow on Kumuwela Ridge as

well as in Kauaikinana Valley, near Honopu Trail, Waialai, and on the

rim of Kalalau Valley within or on the boundary of Kokee State Park

(HHP 1991f3 to 1991f6; HPCC 1991c; D. Lorence et al., in litt., 1991)

in a 3 sq mi (5 sq km) area and on Kamalii Ridge in Kealia Forest

Reserve (HHP 1991f2), roughly 16 mi (26 km) away. All known populations

are on State land and are estimated at 250 individuals (HHP 1991f2,

1991f4, 1991f6; Derral Herbst, FWS, pers. comm., 1991; S. Perlman,

pers. comms., 1991, 1993; D. Lorence et al., in litt., 1991). There are

reliable, but unconfirmed, reports that this species was collected on

the slopes of Anahola Mountain about 1970 (D. Herbst, pers. comm.,

1991). Exocarpos luteolus is found at elevations between 2,000 and

3,600 ft (600 and 1,100 m) in a variety of habitats: Wet places

bordering swamps; on open, dry ridges; and lowland to montane, 'ohi'a-

dominated wet forest communities (HHP 1991f1, 1991f3, 1991f4, 1991f6;

Wagner et al. 1990). Associated taxa include koa, pukiawe, and uluhe

(HHP 1991f2 to 1991f5).

Destruction of habitat by feral goats and pigs and competition with

Erigeron karvinskianus (daisy fleabane) are major threats to Exocarpos

luteolus. Aggressive alien taxa degrading this plant's habitat include

Acacia mearnsii (black wattle), Corynocarpus laevigatus (karakanut),

Myrica faya (firetree), and Rubus argutus (prickly Florida blackberry),

all woody plants that displace native Hawaiian taxa. Other threats to

this species include: rats, that eat the fruits; goats, that browse the

plants; and fire, erosion, and over-collecting for scientific purposes

(HHP 1991f6; T. Flynn and S. Perlman, pers. comms., 1991; D. Lorence et

al., in litt., 1991).

Louis Charles Adelbert von Chamisso collected a plant specimen in

1816 at Kealakekua, island of Hawaii, and named it Kadua cookiana

(Chamisso and Schlechtendal 1829). The specific epithet commemorates

Captain James Cook, the first European to anchor at Kealakekua Bay.

Ernest G. Steudel (1840) transferred the species to the genus Hedyotis,

resulting in the combination H. cookiana.

Hedyotis cookiana, a member of the coffee family (Rubiaceae), is a

small shrub with many branches 4 to 8 in (10 to 20 cm) long. The

papery-textured leaves are long and narrow, 1.5 to 3 in (4 to 8 cm)

long and about 0.2 to 0.5 in (0.5 to 1.2 cm) wide, and fused at the

base to form a sheath around the stem. The bisexual or female flowers

are arranged in clusters of threes on flower stalks about 0.3 to 0.6 in

(8 to 15 mm) long, with the central flower on the longest stalk.

Beneath the flower clusters are sharp-pointed bracts (modified leaves).

The fleshy white corolla is trumpet-shaped and about 0.3 to 0.4 in (8

to 9 mm) long, with lobes about 0.08 in (2 mm) long. Fruits are top-

shaped or spherical capsules about 0.1 in (3.0 to 3.5 mm) long and 0.1

to 0.2 in (3.5 to 4 mm) wide that open at maturity to release wedge-

shaped reddish brown seeds. This plant is distinguished from other

species in the genus that grow on Kauai by being entirely hairless

(Fosberg 1943, Hillebrand 1888, Chamisso and Schlechtendal 1829, Wagner

et al. 1990).

Historically, Hedyotis cookiana was known from only three

collections: Kealakekua on the island of Hawaii; Halawa and Kalawao on

Molokai; and at the foot of the Koolau Mountains on Oahu (Fosberg 1943,

HHP 1991g2, Hillebrand 1888). There is no evidence that it still exists

on any of those islands. This species was discovered in 1976 by Charles

Christensen on the island of Kauai in Waiahuakua Valley on State land

(HHP 1991g1). Between 50 and 100 plants are scattered along a 0.25 mi

(0.4 km) distance in the streambed and lower part of the waterfall.

Although this population has not been observed since its discovery, it

is still believed to be extant (C. Christensen, pers. comm., 1991).

Hedyotis cookiana generally grows in streambeds or on steep cliffs

close to water sources in lowland wet forest communities (C.

Christensen, pers. comm., 1991) and is believed to have formerly been

much more widespread on several of the main Hawaiian Islands at

elevations between 560 and 1,200 ft (170 and 370 m) (Wagner et al.

1990).

The major threat to Hedyotis cookiana, with only one known

population, is stochastic extinction and/or reduced reproductive vigor.

Potential threats include competition with alien plants, which are

invading the area, and habitat modification by feral pigs and goats,

which have been observed in the area. Individuals of Hedyotis cookiana

grow in a stream bed and on the side of a waterfall. These areas are

vulnerable to flooding and other natural disturbances (HHP 1991u6; C.

Christensen, pers. comm., 1991).

In 1928, Albert W. Duvel discovered several trees of Hibiscus clayi

that had been damaged by cattle (Bos taurus) and brought the species

into cultivation. Isa and Otto Degener named the species after the late

Horace F. Clay, a horticulturist and college instructor who brought the

species to their attention (Degener and Degener 1959a). Sister Margaret

James Roe, in her study of the genus in Hawaii, named H. newhousei as

another species from Kauai (Roe 1959, 1961). In the currently accepted

treatment of the Hawaiian members of the family, David M. Bates (1990)

considers H. newhousei to be a synonym of H. clayi.

Hibiscus clayi, a member of the mallow family (Malvaceae), is a

shrub or tree 13 to 26 ft (4 to 8 m) tall with stems bearing sparse

hairs at the branch tips. The oval or elliptical leaves are usually 1

to 3 in (3 to 7 cm) long and 0.6 to 1.4 in (15 to 35 mm) wide and have

a hairless upper surface and slightly hairy lower surface. The leaf

margins are entire or toothed toward the apex. The flowers are borne

singly near the ends of the branches. The flaring petals are dark red,

1.8 to 2.4 in (45 to 60 mm) long, and 0.4 to 0.7 in (10 to 18 mm) wide.

The green tubular or urn-shaped calyx is usually 0.6 to 1 in (15 to 25

mm) long with five or six shorter bracts beneath. The fruits are pale

brown capsules, 0.5 to 0.6 in (12 to 14 mm) long, containing about 10

oval, brownish-black seeds about 0.16 in (4 mm) long. This species is

distinguished from other native Hawaiian members of the genus by the

lengths of the calyx, calyx lobes, and capsule, and by the margins of

the leaves (Bates 1990, Degener and Degener 1959a).

Hibiscus clayi is known from scattered locations on private and

State land on the island of Kauai: The Kokee region on the western side

of the island; Moloaa Valley to the north; Nounou Mountain in Wailua to

the east; and as far south as Haiku near Halii Stream (HHP 1991h1 to

1991h5). At this time, only the Nounou Mountains population with four

trees, is known to still exist (HHP 1991h2, 1991h3; David Bates,

Cornell University, T. Flynn, and S. Perlman, pers. comms., 1991; D.

Lorence et al., in litt., 1991). It is unclear whether the one

individual from the Kokee region was a cultivated plant. This lowland

dry forest species generally grows on slopes at an elevation of 750 to

1,150 ft (230 to 350 m). Associated taxa include Java plum, koa, kukui,

and ti (Bates 1990; HHP 1991h1, 1991h2).

Before cattle were removed from the area, they greatly damaged the

habitat of Hibiscus clayi. Competition with alien plant taxa currently

threatens this species. Strawberry guava is the greatest threat, but

common guava, Hilo grass, Java plum, kukui, lantana, ti, and Schinus

terebinthifolius (Christmas berry) are also present. The area of the

Nounou Mountain population has been planted with Araucaria columnaris

(columnar araucaria), which is reseeding itself there and may prevent

regeneration of native plants. The close proximity of most of the

plants to a hiking trail makes them prone to disturbance. Pigs pose a

potential threat to the species. The small total number of existing

individuals poses a threat of stochastic extinction and/or reduced

reproductive vigor (Degener and Degener 1959a; HHP 1991h1 to 1991h3;

HPCC 1990c; T. Flynn, pers. comm., 1990; D. Bates, T. Flynn, D. Herbst,

and R. Hobdy, pers. comms., 1991).

Abbe Urbain Jean Faurie first collected Lipochaeta fauriei on Kauai

in 1910. During the following year, H. Leveille (1911) named the plant

in honor of him. St. John (1972) described another species from Kauai,

L. deltoidea, but the authors of the current treatment place this name

in synonymy with L. fauriei (Wagner et al. 1990).

Lipochaeta fauriei, a member of the aster family (Asteraceae), is a

perennial herb with somewhat woody, erect or climbing stems up to 16 ft

(5 m) long. The toothed leaves are narrowly triangular, slightly hairy,

3 to 5 in (7 to 13 cm) long, and about 1.2 in (3 cm) wide. Flower heads

occur in clusters of 2 to 3, each comprising 6 to 8 ray florets, 0.2 to

0.5 in (6 to 13 mm) long and about 0.1 in (2.3 mm) wide, and 30 to 35

disk florets 0.1 to 0.2 in (3.3 to 3.9 mm) long. The bracts beneath the

flower heads are purple near the base. Fruits are knobby-textured

achenes (dry, one-seeded fruits) about 0.1 in (2.5 to 3 mm) long and

0.07 in (1.5 to 2 mm) wide. The achenes of the disk florets are

sometimes thinner and shorter than those of the ray florets. This

species belongs to a genus endemic to the Hawaiian Islands and is one

of three species found only on the island of Kauai. This species

differs from the others on Kauai by having a greater number of disk and

ray flowers per flower head, typically longer leaves and leaf stalks,

and longer ray flowers (Gardner 1976, 1979; St. John 1972; Sherff

1935b; Wagner et al. 1985, 1990).

Historically, Lipochaeta fauriei was known from Olokele Canyon on

the island of Kauai (Gardner 1979, HHP 1991i5). This species is now

also known from four other areas on Kauai: Koaie Canyon; Poopooiki;

Haeleele; and lower Hikimoe Valleys (HHP 1991i1 to 1991i4; HPCC 1990d2,

1990d3; St. John 1972). All 5 populations, totalling fewer than 70

individuals, are found on State land (HHP 1991i1 to 1991i3; HPCC

1990d2, 1990d3; R. Hobdy and J. Lau, pers. comms., 1991), encompassing

a 6 by 7 mi (10 by 11 km) area. This species most often grows in

moderate shade to full sun and is usually found on the sides of steep

gulches in diverse lowland mesic forests at an elevation of about 1,570

to 2,950 ft (480 to 900 m) (Wagner et al. 1990). Associated plant taxa

include basketgrass, kukui, lama, and Hibiscus waimeae (koki'o

ke'oke'o); the major alien associate is lantana (HHP 1991i1 to 1991i3;

HPCC 1990d2, 1990d3).

The major threats to Lipochaeta fauriei are degradation of its

habitat by feral goats and competition with invasive alien plant taxa,

especially lantana. Feral pigs pose a potential threat to the species

and fire is a significant threat. The small total number of individuals

comprises a threat of stochastic extinction and/or reduced reproductive

vigor to this species (HHP 1991i1 to 1991i3; HPCC 1990d1 to 1990d3; R.

Hobdy, J. Lau, and S. Perlman, pers. comms., 1991).

Thomas Nuttall (1841) described Schizophyllum micranthum based on a

specimen collected on Kauai in 1840 during the United States Exploring

Expedition. The specific epithet refers to the small size of the

flowers. In 1843, Guilielmo Gerardo Walpers published the superfluous

name Aphanopappus nuttallii based on the same specimen described by

Nuttall (Gardner 1979). Gray (1861) transferred the species to the

genus Lipochaeta, resulting in L. micrantha. Amos Arthur Heller (1897)

transferred the species into the genus Aphanopappus, resulting in A.

micranthus. Otto Degener and Earl Edward Sherff (Sherff 1941) described

L. exigua as another Kauai taxon based on a specimen collected by Otto

Degener and Emilio Ordonez. In his monograph of the genus, Robert C.

Gardner (1979) recognized L. micrantha var. exigua along with the

typical variety, and this is accepted in the current treatment (Wagner

et al. 1990).

Lipochaeta micrantha, a member of the aster family, is a somewhat

woody perennial herb. The 1.6 to 6.6 ft (0.5 to 2 m) long stems grow

along the ground and root at the nodes, with the tip of the stem

growing upward. The roughly triangular leaves measure 0.8 to 3.8 in

(2.1 to 9.7 cm) long and 0.5 to 3.1 in (1.2 to 7.8 cm) wide. They are

sparsely hairy, with margins smooth or variously lobed. Flower heads

are in clusters of two or three. Each head contains four to five ray

florets, 0.1 to 0.2 in (2.3 to 5.8 mm) long and 0.06 to 0.14 in (1.4 to

3.5 mm) wide, and five to nine disk florets, about 0.1 in (2.7 to 3.1

mm) long. The two recognized varieties of this species, exigua and

micrantha, are distinguished by differences in leaf length and width,

degree of leaf dissection, and the length of the ray florets. The

smaller number of disk florets separates this species from the other

members of the genus on the island of Kauai (Gardner 1976, 1979;

Degener and Degener 1959b, 1962; Sherff 1935b; Wagner et al. 1990).

Only two populations of Lipochaeta micrantha var. exigua are known

from the vicinity of Haupu Range on the island of Kauai (HHP 1991j3).

The populations of this variety are distributed over a 1.5 mi (2.4 km)

distance on privately owned portions of Haupu Range and total between

100 and 500 individuals (HHP 1991j1, 1991j2; HPCC 1991d; T. Flynn,

pers. comm., 1991). Historically, Lipochaeta micrantha var. micrantha

appears to have been more widely distributed on Kauai in Olokele

Canyon, Hanapepe Valley, and in the Koloa District (HHP 1991k1, 1991k5;

HPCC 1991d; T. Flynn and S. Perlman, pers. comms., 1991). This variety

is now known only from two to four populations located on State land in

Koaie Canyon on Kauai, totalling 150 to 570 individuals (CPC 1992; HHP

1991k1, 1991k5; S. Perlman, pers. comms., 1991, 1993). The populations

encompass an area of 1.4 square miles (2.3 sq km) approximately 1.4 mi

(2.3 km) apart. Both varieties generally grow on exposed rocky slopes

in diverse lowland mesic forests and sometimes on grassy ridges at an

elevation of 1,000 to 1,300 ft (300 to 400 m) (HHP 1991j1 to 1991j3,

1991k1 to 1991k5; Wagner et al. 1990). Associated plant taxa include

alahe'e, lama, 'ohi'a, Chamaesyce celastroides var. hanapepensis

('akoko), and Neraudia kauaiensis (Gardner 1979; HHP 1991j1, 1991k1,

1991k2).

The major threats to Lipochaeta micrantha are habitat degradation

by feral ungulates and competition with alien plant taxa. Feral pigs

threaten the habitat of both varieties of Lipochaeta micrantha and

signs of damage by feral goats have been seen near individuals of var.

micrantha. Alien plant taxa, such as lantana, affect the habitats of

both varieties. Pluchea carolinensis (sourbush) is found near var.

exigua. Daisy fleabane and Stachytarpheta spp. are components of the

habitat of var. micrantha. Both varieties are threatened by stochastic

extinction and/or reduced reproductive vigor due to the small number of

existing populations (HHP 1991j1, 1991j2, 1991k1, 1991k5; HPCC 1990e,

1990f; T. Flynn, pers. comm., 1991; D. Lorence et al., in litt., 1991).

Hobdy collected the first specimen of Lipochaeta waimeaensis in

1967. Five years later, St. John (1972) described it as a new species,

naming it for the Waimea Canyon where it grows.

Lipochaeta waimeaensis, a member of the aster family, is a low

growing, somewhat woody perennial herb with stems 3 to 6.5 ft (1 to 2

m) long that root at the nodes. The linear or narrowly elliptical

leaves are 1.9 to 2 in (4.7 to 5 cm) long, 0.2 to 0.3 in (5 to 8 mm)

wide, hairy along major veins on the upper surface, and evenly hairy on

the lower surface. Flower heads are borne singly or in clusters of two

or three. The outer head bracts are lance-shaped and measure 0.1 to 0.2

in (3 to 4 mm) long and 0.06 to 0.08 in (1.5 to 2 mm) wide. The oval

ray florets number four or five per head and are about 0.13 in (3.2 to

3.5 mm) long and about 0.1 in (3 mm) wide. The disk florets number 20

to 25 per head. The fruits are knobby, winged achenes 0.1 in (2.2 to

2.5 mm) long and about 0.08 in (1.7 to 2.3 mm) wide. The ray achenes

are slightly wider and have longer wings than those of the disk. This

species differs from the two other taxa of the genus included in this

rule (L. fauriei and L. micrantha) in having a different leaf shape and

shorter leaf stalks and ray florets (Gardner 1976, 1979; St. John 1972;

Wagner et al. 1990).

Lipochaeta waimeaensis is known only from the type locality, along

the rim of Kauai's Waimea Canyon on State land (HHP 1991m1, HPCC

1991e). Fewer than 10 plants are scattered over a 2.5 acre (ac) (1-

hectare (ha)) area (Gerald Carr, University of Hawaii at Manoa, and S.

Perlman, pers. comms., 1991). This population grows on eroded soil on a

precipitous, shrub-covered gulch in a diverse lowland mesic forest at

an elevation between 1,150 and 1,300 ft (350 and 400 m) (HHP 1991m1,

Wagner et al. 1990). The vegetation at the site is predominantly alien

consisting of Grevillea robusta (silk oak), Leucaena leucocephala (koa

haole), and Rhynchelytrum repens (Natal redtop); however, native taxa

include Dodonaea viscosa ('a'ali'i) and Lipochaeta connata (nehe) (CPC

1989b, 1990; S. Perlman, pers. comm., 1991) also occur here.

Alien plant taxa competing with and threatening Lipochaeta

waimeaensis include koa haole, Natal redtop, silk oak, and Opuntia

ficus-indica (prickly pear, panini). The existing soil erosion problem

is exacerbated by the presence of feral goats. The single population,

and thus the entire species, is threatened by stochastic extinction

and/or reduced reproductive vigor due to the small number of existing

individuals. Over-collecting for scientific purposes also poses a

threat (G. Carr and S. Perlman, pers. comms., 1991).

In 1912, Lydgate collected a plant specimen on Kauai that he and

Forbes named Lysimachia filifolia (Forbes 1916). They chose the

specific epithet, which means ``thread-leaved,'' in reference to the

plant's very narrow leaves. Heller (1897) created a new genus,

Lysimachiopsis, in which he placed all endemic Hawaiian taxa of

Lysimachia, and Otto and Isa Degener (1983) later published

Lysimachiopsis filifolia. The current treatment (Wagner et al. 1990)

recognizes Lysimachiopsis as a section of Lysimachia. Most recently,

St. John (1987b) published many species, varieties, and combinations of

Lysimachia, one or more of which may fit into this species (Wagner et

al. 1990).

Lysimachia filifolia, a member of the primrose family

(Primulaceae), is a small shrub 0.5 to 1.6 ft (15 to 50 cm) tall. The

linear leaves measure 0.6 to 2.1 in (15 to 54 mm) long and 0.01 to 0.07

in (0.3 to 1.8 mm) wide and are usually alternately arranged. They are

single-veined and sparsely hairy or hairless. The bell-shaped flowers

are reddish purple, 0.2 to 0.4 in (6 to 10 mm) long, and borne singly

on flower stalks about 0.7 to 1.2 in (18 to 30 mm) long that elongate

upon fruiting. Fruits are thick, hard capsules about 0.2 in (5 to 6 mm)

long that contain numerous minute, nearly black, irregularly shaped

seeds. This species is distinguished from other taxa of the genus by

its leaf shape and width, calyx lobe shape, and corolla length (Forbes

1916, Wagner et al. 1990).

Historically, Lysimachia filifolia was known only from the upper

portion of Olokele Valley on Kauai (HHP 1991n1). This species is now

known from two other areas: The headwaters of the Wailua River on

Kauai; and the slopes of Waiahole Valley in the Koolau Mountains of

Oahu (HHP 1991n2, 1991n3; HPCC 1990g1, 1990g3). Three closely situated

colonies on Kauai are located within a 0.5 sq mi (1.3 sq km) area and

total 76 individuals (K. Marr, pers. comm., 1991). The Oahu population

contains about 150 to 200 individuals (CPC 1989a; HHP 1991n3; HPCC

1990g1, 1990g3). Both populations of this species are located on State

land, totalling approximately 225 to 275 individuals. This species

typically grows on mossy banks at the base of cliff faces within the

spray zone of waterfalls or along streams in lowland wet forests at an

elevation of 800 to 2,200 ft (240 to 680 m) (HHP 1991n1 to 1991n3; HPCC

1990g1, 1990g3; Wagner et al. 1990; K. Marr, pers. comm., 1991).

Associated plant taxa include mosses, ferns, liverworts, pili grass,

Cuphea carthagenensis (tarweed), and Pilea peploides (HHP 1991n3; J.

Lau, pers. comm., 1991).

The major threat to Lysimachia filifolia is competition with alien

plant taxa. Individuals of this species on Kauai are damaged and

destroyed by natural rock slides in their habitat, which is near the

bottom of steep cliffs. Hydrocotyle sibthorpioides (marsh pennywort),

tarweed, and thimbleberry, although not invasive weeds, are present in

this near-pristine area of Wailua Stream and may degrade the native

ecosystem. At least one feral pig has made its way into this area,

indicating that this disruptive animal is a potential threat.

Individuals of Lysimachia filifolia on Oahu are vulnerable to rock

slides and compete for space with alien plants such as marsh pennywort,

tarweed, Ageratina riparia (Hamakua pamakani), and Schefflera

actinophylla (octopus tree). Because only one population of Lysimachia

filifolia occurs on each of only two islands, the species is threatened

by stochastic extinction. Hurricane Iniki caused at least some damage

to the Wailua River population (HHP 1991n3; HPCC 1990g2; D. Lorence and

S. Perlman, pers. comms., 1991; L. Mehrhoff, pers. comm., 1993).

In 1927, MacDaniels collected a plant specimen on Kauai that St.

John (1944) later named Pelea haupuensis. The specific epithet refers

to the type locality, Haupu, the only known site for this plant until

it was discovered in Waimea Canyon in 1989. Thomas G. Hartley and

Benjamin C. Stone (1989, Stone et al. 1990, Wagner et al. 1990)

synonymized the genus Pelea with Melicope, resulting in the current

name for this taxon, Melicope haupuensis.

Melicope haupuensis, a member of the citrus family (Rutaceae), is a

tree about 26 ft (8 m) tall. The oval leaves, 2 to 5.1 in (5 to 13 cm)

long and 1.1 to 2.2 in (28 to 56 mm) wide, are oppositely arranged.

Flowers grow in clusters of five to seven on stalks usually 0.1 to 2.8

in (2 to 7 mm) long, each flower on a stalk 0.04 to 0.12 in (1 to 3 mm)

long. Only female flowers are known. The flowers are about 0.14 in (3.5

mm) long, dotted with oil glands, and covered with a dense mat of

hairs. Fruits are distinct follicles (a dry fruit that splits open

lengthwise), 0.35 to 0.43 in (9 to 11 mm) long, with a hairless exocarp

and endocarp (outermost and innermost layers of the fruit wall,

respectively). Unlike other taxa of this genus on Kauai, the exocarp

and endocarp are hairless and the sepals are covered with dense hairs

(St. John 1944, Stone 1969, Stone et al. 1990).

For 62 years, Melicope haupuensis was known only from the type

locality on the north side of Haupu Ridge on Kauai (HHP 1991o3). In

1989, two plants were discovered within 1 mi (1.6 km) of each other

along the banks of Koaie Stream on State owned land in Waimea Canyon

(HHP 1991o1, 1991o2; HPCC 1991f). These plants grow on moist talus

slopes in 'ohi'a-dominated lowland mesic forests (Stone et al. 1990)

with such associated taxa as 'a'ali'i and hame, at elevations between

1,230 and 2,690 ft (375 and 820 m) (HHP 1991o1 to 1991o3).

Habitat degradation by feral goats and competition with invasive

alien plant taxa such as lantana and yellow foxtail threaten Melicope

haupuensis. A potential threat to members of this genus is their known

susceptibility to black twig borer (Xylosandrus compactus), a burrowing

beetle ubiquitous in Hawaii at elevations below 2,500 ft (670 m). The

existence of only two known trees of this species constitutes a threat

of stochastic extinction, over-collecting, and/or reduced reproductive

vigor (Hara and Beardsley 1979; HHP 1991o1, 1991o2; Medeiros et al.

1986).

Knudsen sent a plant specimen he found at Waimea to Hillebrand, who

named it Pelea knudsenii in honor of its collector (Hillebrand 1888).

In an action that was not supported by other taxonomists, Emmanuel

Drake del Castillo (1890) transferred several species from the genus

Pelea to the genus Evodia. Hartley and Stone (1989) synonymized the

genus Pelea with Melicope, resulting in the combination M. knudsenii.

Other names now included in M. knudsenii are Pelea multiflora (Rock

1911), P. knudsenii var. multiflora (Rock 1918), and P. tomentosa (St.

John 1944).

Melicope knudsenii, a member of the citrus family, is a tree

usually 10 to 33 ft (3 to 10 m) tall with smooth gray bark and

yellowish brown to olive-brown hairs on the tips of the branches.

Leaves are variable, ranging from oblong to elliptic, 3.5 to 9.8 in (9

to 25 cm) long and 1.8 to 3.9 in (4.5 to 10 cm) wide. The lower surface

of the leaves is uniformly covered with olive-brown hairs, but the

upper surface is only sparsely hairy along the midrib. The densely

hairy flowers are bisexual or may be unisexual. There are usually 20 to

200 flowers per cluster in the leaf axils. The sepals and petals are

covered with silky gray hairs and the sepals persist in fruit. The

fruits are 0.7 to 1.2 in (18 to 30 mm) wide and are comprised of

distinct follicles, 0.3 to 0.6 in (8 to 14 mm) long. The hairless

exocarp is dotted with minute glands. The endocarp also lacks hairs.

Seeds number one or two per carpel (ovule-bearing structure) and are

about 0.2 in (5 to 6 mm) long. The distinct carpels of the fruit, the

hairless endocarp, the larger number of flowers per cluster, and the

distribution of hairs on the underside of the leaves distinguish this

species from M. haupuensis and other species of the genus (Degener et

al. 1962a, 1962b; Hillebrand 1888; Rock 1913; Stone 1969; Stone et al.

1990).

Historically, Melicope knudsenii was known only from the southeast

slope of Haleakala on Maui and from Olokele Canyon on Kauai (HHP

1991p1, 1991p5). This species remains in the Auwahi and Kanaio areas of

Maui (R. Hobdy and Arthur Medeiros, Haleakala National Park, pers.

comms., 1991) on privately owned land, but its numbers have decreased

considerably from being ``very common'' in 1920 to between 20 and 30

plants when it was last observed in 1983 (CPC 1990; HHP 1991p1). On

Kauai, three populations, each consisting of one individual, remain on

State land in the Koaie drainage area of Waimea Canyon (HHP 1991p2 to

1991p4; S. Perlman, pers. comm., 1991) and are distributed across a

distance of 1.6 mi (2.6 km). This species, therefore, totals between 23

and 33 individuals at present. Melicope knudsenii grows on forested

flats or talus slopes in lowland dry to mesic forests at an elevation

of about 1,500 to 3,300 ft (450 to 1,000 m) (Stone et al. 1990). The

Auwahi population on Maui, however, grows on a substrate of 'a'a lava

in a remnant native forest, dominated by a continuous mat of Pennisetum

clandestinum (Kikuyu grass) (HHP 1991p1; Medeiros et al. 1986). Plants

associated with the Kauai populations include 'a'ali'i, hame, 'ohi'a,

and Xylosma (HHP 1991p3, 1991p4).

Competition with alien plant taxa and habitat degradation by feral

and domestic animals are the major threats affecting Melicope

knudsenii. On Kauai, this species competes with lantana and is affected

by feral goats and pigs. On Maui, Melicope knudsenii grows in an area

currently grazed by domestic cattle, where a continuous mat of Kikuyu

grass prevents seedlings from establishing. Feral goats and feral pigs

are also present in the area of the Maui population. Axis deer (Axis),

found on the south slope of Haleakala Mountain and increasing in

numbers, are a potential threat. This species is potentially threatened

by black twig borer, a ubiquitous insect that lives at elevations up to

2,500 ft (670 m) in Hawaii and is known to infest members of Melicope.

This species is also threatened by fire, stochastic extinction, and/or

reduced reproductive vigor due to the small number of existing

individuals (HHP 1991p2 to 1991p4; Hara and Beardsley 1979; Medeiros et

al. 1986; van Riper and van Riper 1982; Patrick Beil, Puu Mahoe

Arboretum, R. Hobdy, A. Medeiros, and Steve Montgomery, Bishop Museum,

pers. comms., 1991).

Hillebrand (1888) described Pelea pallida based on a specimen he

collected on Oahu. The specific epithet refers to the plant's pale leaf

veins and lower leaf surfaces. Drake del Castillo (1890) transferred

the species to the genus Evodia, a combination not accepted by other

taxonomists. Faurie described P. leveillei in 1912 based on a specimen

collected on Kauai (Stone 1969). Following the transfer of the genus

Pelea to Melicope (Hartley and Stone 1989, Wagner et al. 1990), authors

of the current treatment of the Hawaiian members of the genus (Stone et

al. 1990) now consider Evodia pallida, P. pallida, and P. leveillei to

be synonyms of Melicope pallida.

Melicope pallida, a member of the citrus family, is a 20 to 33 ft

(6 to 10 m) tree with grayish white hairs and black, resinous new

growth. The leaves, 2.4 to 8.3 in (6 to 21 cm) long and 1 to 3.1 in

(2.5 to 8 cm) wide, are grouped in threes, with each leaf loosely

folded. Fifteen to 35 pale yellowish-green flowers are also clustered

in groups of 3 along a fuzzy white stalk up to 2.4 in (6 cm) long. The

petals are usually lance-shaped and measure 0.1 to 0.2 in (3.5 to 5 mm)

long. Fruits contain two shiny black seeds about 0.1 in (3.5 mm) long

in each of the usually four distinct carpels. This species differs from

M. haupuensis, M. knudsenii, and other members of the genus by the

following combination of characteristics: Resinous new growth; leaves

folded and in clusters of three; and fruits with separate carpels

(Degener et al. 1960, Hillebrand 1888, St. John 1944, Stone et al.

1990, Wagner et al. 1990).

Historically, Melicope pallida was known from various locations in

the Waianae Mountains on Oahu and from Hanapepe on Kauai (HHP 1991q2 to

1991q4, 1991q7). This species is now known from two locations at the

base of Mount Kaala and near Palikea, within TNCH's privately owned

Honouliuli Preserve in the Waianae Mountains on Oahu, and from four

State owned locations on Kauai in Kalalau Valley, Koaie Stream in

Waimea Canyon, and Hanakapiai Valley (HHP 1991q1, 1991q6, 1991q8; HPCC

1991g1, 1991g2; T. Flynn, J. Lau, and S. Montgomery, pers. comms.,

1991). The population near Palikea was last visited in 1960 (HHP

1991q1); it is thought to contain only a few plants. Fewer than five

plants are known from the island of Oahu (S. Montgomery, pers. comm.,

1991). Populations were discovered in 1991 near the rim of Kalalau

Valley (about 65 plants) and Honopu Rim (12 plants) (Kenneth Wood,

HPCC, pers. comm., 1991), giving a total of less than 100 known plants

for this species. Melicope pallida usually grows on steep rock faces in

drier regions of lowland mesic forests at an elevation of 1,600 to over

3,000 ft (490 to 910 m) (Stone et al. 1990; J. Lau, pers. comm., 1991;

D. Lorence et al., in litt., 1991). Associated plant taxa include

Abutilon sandwicense, Alyxia oliviformis (maile), Dryopteris sp.,

'ohi'a, Pipturus albidus (mamaki), Sapindus oahuensis (lonomea),

Tetraplasandra sp. ('ohe), and Xylosma hawaiiense (mana) (HHP 1991q1,

1991q5, 1991q8; J. Lau, pers. comm., 1991).

The major threats to Melicope pallida are habitat destruction by

feral animals and competition with alien plant taxa. On Kauai, feral

goats and feral pigs destroy habitat of Melicope pallida and weeds,

such as daisy fleabane and prickly Florida blackberry, compete with the

species. The Oahu populations of Melicope pallida face strong

competition from introduced plants, especially Clidemia hirta (Koster's

curse) and Toona ciliata (Australian red cedar). A potential threat to

Melicope pallida is the black twig borer, which is known to occur in

areas where this species grows and to feed on members of the genus

Melicope. Additional threats to Melicope pallida are fire and

stochastic extinction and/or reduced reproductive vigor due to the

small number of existing individuals (Hara and Beardsley 1979; HHP

1991q6, 1991q8; Medeiros et al. 1986; T. Flynn, J. Lau, S. Montgomery,

and K. Wood, pers. comms., 1991; D. Lorence et al., in litt., 1991).

St. John and Edward P. Hume described Melicope quadrangularis,

based on a specimen collected by Forbes on Kauai in 1909 (St. John

1944). The specific epithet, meaning ``four-angled,'' describes the

cube-shaped capsule. Hartley and Stone (1989) synonymized the genus

Pelea with Melicope, resulting in the combination M. quadrangularis.

Melicope quadrangularis, a member of the citrus family, is a shrub

or small tree. Young branches are generally covered with fine yellow

fuzz but become hairless with age. The thin, leathery, elliptical

leaves, 3.5 to 6 in (9.5 to 16 cm) long and 2 to 3 in (4.5 to 7.5 cm)

wide, are oppositely arranged. The upper leaf surface is hairless and

the lower surface is sparsely hairy, especially along the veins.

Flowers are solitary or in clusters of two. The specific floral details

are not known. The fruits are somewhat cube-shaped, flattened capsules,

about 0.5 in (13 mm) long and about 0.8 in (19 to 22 mm) wide with a

conspicuous central depression at the top of the fruit. The capsules

are four-lobed and completely fused. The exocarp is sparsely hairy and

the endocarp is hairless. This species differs from others in the genus

in having the following combination of characters: Oppositely arranged

leaves; only one or two flowers per cluster; cube-shaped capsules with

fused lobes; and a deep central depression at the top of the fruit (St.

John 1944, Stone 1969, Stone et al. 1990).

Melicope quadrangularis is known from the type locality in the

Wahiawa Bog region of Kauai (HHP 1991r1; Stone et al. 1990). One adult

plant and two seedlings were discovered in 1991 in that area by Ken

Wood of HPCC on an east-facing slope of Wahiawa Ridge at 2,800 ft (850

m) elevation on privately owned land. Subsequent exploration has

resulted in the location of 13 individuals of this species. The plants

are growing in a diverse lowland forest that ranges from mesic to wet

conditions with other plants, such as 'ohi'a, opuhe, uluhe, Broussaisia

arguta (kanawao), Cyrtandra pickeringii (ha'iwale), other Melicope

species (alani), Metrosideros waialealae, and abundant ferns and mosses

(K. Wood, pers. comm., 1991, D. Lorence et al., in litt., 1991).

The existence of only 13 known plants of this species causes the

species to be threatened by over-collecting for scientific purposes,

stochastic extinction, and/or reduced reproductive vigor. The alien

strawberry guava grows in the area and is a potential threat (Hara and

Beardsley 1979; K. Wood, pers. comm., 1991; D. Lorence et al., in

litt., 1991).

Forbes collected specimens of a tree on Kauai in 1916 that he

described the following year (1917b) as Tetraplasandra racemosa. The

specific epithet describes the inflorescence that Forbes considered a

raceme. Sherff (1952) transferred the species to the new endemic,

monotypic genus Munroidendron, named in honor of George C. Munro, who

was apparently the first to recognize the plant as a new taxon. Sherff

(1952) also published two varieties, Munroidendron racemosum var.

forbesii and M. racemosum var. macdanielsii. In the current treatment

of the species, Porter P. Lowrey II (1990) recognizes no subspecific

taxa.

Munroidendron racemosum, a member of the ginseng family

(Araliaceae), is a tree up to about 23 ft (7 m) in height with a

straight gray trunk crowned with spreading branches. The leaves are 6

to 12 in (15 to 30 cm) long and comprise five to nine oval or

elliptical leaflets with clasping leaf stalks. Each leaflet is 3.1 to

6.7 in (8 to 17 cm) long and usually 1.6 to 3.9 in (4 to 10 cm) wide.

About 250 pale yellow flowers are borne along a stout hanging stalk 10

to 24 in (25 to 60 cm) long. Each flower has five or six lance-shaped

petals 0.3 to 0.4 in (8 to 10 mm) long emerging from a cup-shaped or

ellipsoid calyx tube. Both the lower surface of the petals and the

calyx tube are covered with whitish scaly hairs. The fruit is an egg-

shaped drupe 0.3 to 0.5 in (8 to 12 mm) long and nearly as wide,

situated atop a flat, dark red disk (stylopodium). This species is the

only member of a genus endemic to Hawaii, differing from other closely

related Hawaiian genera of the family primarily in its distinct flower

clusters and corolla (Forbes 1917b, Lamoureux 1982, Lowrey 1990, St.

John 1981b, Sherff 1952).

Historically, Munroidendron racemosum was known from scattered

locations throughout the island of Kauai (HHP 1991s1, 1991s3, 1991s6,

1991s13). Fifteen populations are now found at elevations of 390 to

1,310 ft (120 to 400 m) on private and State land in the following

areas: Along the Na Pali Coast within Na Pali Coast State Park and Hono

O Na Pali NAR; in the Poomau and Koaie branches of Waimea Canyon; in

the Haupu Range area; and on Nounou Mountain (HHP 1991s1 to 1991s12,

1991s14, 1991s15; Lamoureux 1982). Although widely distributed, the

largest population contains fewer than 50 individuals, with most

populations numbering only 1 or 2 individuals. Estimates of the total

number of individuals range from 57 to 100 (HHP 1991s1 to 1991s15).

Most populations are found on steep exposed cliffs or on ridge slopes

in coastal to lowland mesic forests (Lowrey 1990), but a few

populations are in mesic Pandanus tectorius (hala) forests, lantana-

dominated shrubland, or Eragrostis grassland. Other associated plant

taxa include common guava, kopiko, kukui, and lama (Gagne and Cuddihy

1990; HHP 1991s1, 1991s3 to 1991s5, 1991s8 to 1991s11, 1991s15;

Lamoureux 1982).

Competition with introduced plants is the major threat to

Munroidendron racemosum. Kukui and ti, plants introduced by Polynesian

immigrants to the Hawaiian Islands, compete with this species for space

in the forests of Kauai. Other introduced plants threatening this

species' habitat include Chinaberry, common guava, firetree, koa haole,

lantana, and Triumfetta semitriloba (Sacramento bur). Feral goats

degrade the habitat of Munroidendron and cattle were formerly present

in areas where the trees grow. Fire is a threat to the habitat.

Predation of the fruit by rats is probable. An introduced insect of the

longhorned beetle family (Cerambycidae) that killed a mature,

cultivated tree has the potential of affecting wild trees. Because each

population of this species contains only one or a few trees, the total

number of individuals is small, threatening the species through over-

collecting for scientific or horticultural purposes, stochastic

extinction, and/or reduced reproductive vigor (HHP 1991s1, 1991s3 to

1991s5, 1991s8 to 1991s11, 1991s15; HPCC 1990h; Lamoureux 1982).

First collected on Kauai before 1900, Nothocestrum peltatum was

described by Carl J. F. Skottsberg in 1944, based on a specimen

collected by Olof H. Selling in 1938. The specific epithet refers to

the peltate leaves, attached to the stalk by the lower surface, inside

the leaf margin rather than at its edge. St. John (1986) later

described N. inconcinnum, but David E. Symon (1990), in the currently

accepted treatment of the genus, regards that name as a synonym of N.

peltatum.

Nothocestrum peltatum, a member of the nightshade family

(Solanaceae), is a small tree up to 26 ft (8 m) tall with ash-brown

bark and woolly stems. The leathery leaves are usually peltate, measure

2.4 to 9.1 in (6 to 23 cm) long and 1.4 to 3 in (3.5 to 7.5 cm) wide

and vary in shape from oval or elliptic to oblong. The densely hairy

flowers number up to 10 per cluster. The corolla is greenish yellow

fading to yellow orange and 0.5 to 0.6 in (12 to 14 mm) long. The

orange berries are 0.5 to 0.6 in (13 to 14 mm) long and contain

numerous irregularly shaped seeds about 0.1 in (2.5 mm) in diameter.

The usually peltate leaves and shorter leaf stalks separate this

species from others in the genus (St. John 1986, Selling 1947,

Skottsberg 1944, Symon 1990).

Historically, Nothocestrum peltatum was known from Kauai at

Kumuwela, Kaholuamanu, and the region of Nualolo (HHP 1991t3, 1991t5,

1991t6). This species is now known from five populations on Kauai

located near the Kalalau Lookout area, in Awaawapuhi and Makaha

Valleys, and in Waimea Canyon (HHP 1991t1, 1991t2, 1991t4, 1991t7; HPCC

1990i1, 1990i2, 1990i4; S. Perlman, pers. comm., 1991, 1993), scattered

over a 5.5 by 2.5 mi (8.9 by 4 km) area. These populations, totalling

about 15 individuals (CPC 1989b, 1990; S. Perlman, pers. comm., 1993),

are on State owned land between 3,000 and 4,000 ft (915 and 1,220 m)

elevation (Symon 1990). This species generally grows in rich soil on

steep slopes in montane mesic forests dominated by koa or a mixture of

'ohi'a and koa, with associates such as hame, uluhe, Bobea brevipes

('ahakea lau li'i), Elaeocarpus bifidus (kalia), and more common

Melicope species (alani) (HHP 1991t1, 1991t7; Sohmer and Gustafson

1987; J. Lau, pers. comm., 1991).

Competition with alien plants and habitat degradation by introduced

animals constitute the major threats to Nothocestrum peltatum.

Introduced plants competing with this species include banana poka,

daisy fleabane, lantana, prickly Florida blackberry, and Passiflora

edulis (passion fruit). Animals disturbing the habitat of this species

include feral goats, feral pigs, mule deer, and red jungle fowl (Gallus

gallus). Although plants of this species flower, they rarely set fruit;

this could be the result of a loss of pollinators, reduced genetic

variability, or self-incompatibility (S. Perlman, pers. comm., 1991; D.

Lorence et al., in litt., 1991). This species is threatened by fire,

over-collecting for scientific or horticultural purposes, stochastic

extinction, and/or reduced reproductive vigor due to the small number

of existing individuals (HHP 1991t7; HPCC 1990i3, 1990i4).

Hillebrand (1888) described Peucedanum sandwicense based on a

specimen collected on Molokai and P. kauaiense based on a specimen

collected on Kauai. He also referred to an unnamed variety of P.

sandwicense from Maui. Otto and Isa Degener (1960) later named the Maui

plant P. sandwicense var. hiroi. In their current treatment, Lincoln

Constance and James Affolter (1990) recognize only P. sandwicense for

all populations of the genus in the Hawaiian Islands.

Peucedanum sandwicense, a member of the parsley family (Apiaceae),

is a parsley-scented, sprawling herb usually 20 to 40 in (0.5 to 1 m)

tall. Hollow stems arise from a short, vertical, perennial stem with

several fleshy roots. The compound leaves are generally three-parted

with stalkless leaflets, each egg- or lance-shaped and toothed. The

larger terminal leaflet is usually one- to three-lobed and 2.8 to 5.1

in (7 to 13 cm) long. The other leaflets have leaf stalks 4 to 20 in

(10 to 50 cm) long or are stalkless. Flowers are clustered in a

compound umbel of 10 to 20 flowers. The round petals are white and bent

inward at the tips. The flat, dry, oval fruits are 0.4 to 0.5 in (10 to

13 mm) long and 0.2 to 0.3 in (5 to 8 mm) wide, splitting in half to

release a single flat seed. This species is the only member of the

genus in the Hawaiian Islands, one of three genera of the family with

taxa endemic to the island of Kauai. This species differs from the

other Kauai members of the parsley family in having larger fruit and

pinnately compound leaves with broad leaflets (Constance and Affolter

1990, Degener and Constance 1959, Degener and Degener 1960, Hillebrand

1888).

Historically, Peucedanum sandwicense was known from three islands:

Kalaupapa, Pauonuakea Kui, Waikolu, and Wailau Valley on Molokai;

Wailuku and Waiehu on Maui; and various locations in the Waimea Canyon

and Olokele regions of Kauai (HHP 1991u1, 1991u2, 1991u4, 1991u7,

1991u9 to 1991u12; HPCC 1991h1, 1991h2). Discoveries in 1990 extended

the known distribution of this species to the island of Oahu, where 2

populations totalling about 85 individuals exist in the Waianae

Mountains on County and State land (J. Lau, in litt. and pers. comm.,

1991; J. Obata, pers. comm., 1990). One population of 20 to 30

individuals is known from State owned Keopuka Rock, an islet off the

coast of Maui (HHP 1991u8; Hobdy 1982; R. Hobdy, pers. comm., 1991). On

Molokai, 3 populations totalling fewer than 30 individuals are found on

private and State owned land in Pelekunu Preserve, Kalaupapa National

Historical Park, and Huelo, an islet off the coast of Molokai (HHP

1991u7, 1991u16, 1991u20; S. Perlman, pers. comm., 1991). The 10 Kauai

populations of 130 to 190 individuals are distributed in Waimea Canyon

and along the Na Pali Coast within 1.5 mi (2.4 km) of the ocean (HHP

1991u1, 1991u3, 1991u5, 1991u6, 1991u13 to 1991u15, 1991u17 to 1991u19;

T. Flynn, pers. comm., 1991). These populations are found within a 7 by

8 mi (11 by 13 km) area on private and State land. The total number of

plants in the known populations of this species is estimated to exceed

1,000 and possibly 5,000 individuals (CPC 1992; S. Perlman, pers.

comm., 1993). This species grows in cliff habitats from sea level to

above 3,000 ft (900 m) (Constance and Affolter 1990) with such plant

associates as 'akoko, kawelu, lama, 'ohi'a, Artemisia australis

('ahinahina), and alien species such as common guava and lantana (HHP

1991u1 to 1991u3, 1991u5 to 1991u8, 1991u14 to 1991u18, 1991u20; J.

Lau, in litt. and pers. comm., 1991; D. Lorence et al., in litt.,

1991).

Competition with introduced plants and habitat degradation and

browsing by feral goats are the major threats to Peucedanum

sandwicense. Kauai populations are affected by alien plant species such

as Kalanchoe pinnata (air plant), banana poka, common guava, daisy

fleabane, firetree, introduced grasses, Java plum, and lantana, as well

as by feral goats. The Hanakapiai population on Kauai is close enough

to the trail that it is potentially affected by hikers and trail

clearing. Oahu populations are threatened by alien plants such as

Christmas berry, common guava, daisy fleabane, Hamakua pamakani, silk

oak, and Stachytarpheta; feral goats; fire; and landslides. The

Kalaupapa, Molokai, population of P. sandwicense competes with

Christmas berry, common guava, and molasses grass. The Pelekunu,

Molokai, population is threatened by common guava, Hamakua pamakani,

Ageratina adenophora (Maui pamakani), and potentially by axis deer.

Plants of this species on Huelo are vulnerable to natural rock slides.

The population on Keopuka Rock is threatened by alien grasses, lantana,

and sourbush (Clarke and Cuddihy 1980; HHP 1991u1, 1991u3, 1991u5,

1991u15, 1991u16; HPCC 1990j1 to 1990j3; R. Hobdy, J. Lau, J. Obata,

and S. Perlman, pers. comms., 1991; D. Lorence et al., in litt., 1991).

Peucedanum sandwicense is not in immediate danger of extinction,

but if these threats are not curtailed, this species will become

endangered in the future.

Wawra collected a specimen of Phyllostegia waimeae on Kauai in 1870

while he was a member of the Austrian East Asiatic Exploring

Expedition. In 1872, he described the species, naming it for Waimea

Canyon where he collected it. St. John (1987c) recently published many

species, varieties, and combinations in Phyllostegia, one or more of

which may fit into this species (Wagner et al. 1990).

Phyllostegia waimeae, a nonaromatic member of the mint family

(Lamiaceae), is a climbing perennial plant with hairy four-angled stems

that are woody at the base. The oval leaves are 2 to 5 in (5 to 13 cm)

long, 1 to 2.4 in (2.5 to 6 cm) wide, and have rounded, toothed

margins. They are wrinkled and sparsely dotted with oil glands. Flowers

grow in groups of six along an unbranched leafy stalk usually 3.9 to

5.9 in (10 to 15 cm) long. The bracts below each flower stalk are broad

and partially overlap the flowers. The calyx resembles an inverted cone

with broad lobes. The corolla, 0.3 to 0.5 in (8 to 12 mm) long, is

pinkish or may be white. The fruits, probably nutlets, have not been

observed. Characteristics that distinguish this species from others in

the genus are the nearly stalkless bracts that partially overlap and

cover the flowers and relatively fewer oil glands on the leaves

(Hillebrand 1888, Sherff 1935a, Wagner et al. 1990, Wawra 1872).

Historically, Phyllostegia waimeae was known from Kaholuamanu and

Kaaha on Kauai (HHP 1991v2, 1991v3). In recent years, it is known from

State land on Kauai in the Halemanu and Waimea Canyon areas (HHP

1991v1, 1991v4). Because the Halemanu population has not been seen for

almost 40 years (HHP 1991v1), the number of extant individuals is

unknown. The Waimea Canyon population consists of a single plant which

has not been observed recently (R. Hobdy, pers. comm., 1991; S.

Perlman, pers. comm., 1993). This species typically grows on shallow to

deep, well-drained soils in clearings (HHP 1991v1) or along the banks

of streams of diverse montane mesic to wet forests at elevations from

3,000 to 3,600 ft (915 m to 1,100 m) (Wagner et al. 1990). Associated

taxa include 'ohi'a and Pritchardia minor (loulu) (HHP 1991v4).

Habitat destruction by feral goats, erosion, and competition with

introduced grasses are the major threats to Phyllostegia waimeae. The

species is also threatened by over-collecting for scientific purposes,

stochastic extinction, and/or reduced reproductive vigor due to the

small number of existing individuals (R. Hobdy, pers. comm., 1991).

Based on a specimen collected by Duvel and Harold L. Lyon in 1925,

Edward L. Caum (1933) described Pteralyxia kauaiensis, named for the

island where it grows. St. John (1981a) later published P. elliptica,

but the authors of the current treatment of the genus (Wagner et al.

1990) regard that name to be synonymous with P. kauaiensis.

Pteralyxia kauaiensis, a member of the dogbane family

(Apocynaceae), is a tree 10 to 26 ft (3 to 8 m) tall. The leaves are

dark green and shiny on the upper surfaces but pale and dull on the

lower surfaces. They are generally egg-shaped and usually 4.3 to 8.7 in

(11 to 22 cm) long and 1.6 to 2.6 in (40 to 65 mm) wide. The pale

yellow flowers are trumpet-shaped, 0.3 to 0.5 in (8 to 12 mm) long,

with each of the five lobes 0.1 to 0.2 in (3 to 4 mm) long. The paired

fruits, of which usually only one matures, are drupe-like, bright red,

and fleshy. The woody endocarp that encloses the single seed has two

prominent central wings and two reduced lateral wings. This species

differs from the only other taxa in this endemic Hawaiian genus in

having reduced lateral wings on the seed (Caum 1933; Degener 1933,

1936; Lamb 1981; St. John 1981a; Wagner et al. 1990).

Historically, Pteralyxia kauaiensis was known from the Wahiawa

Mountains in the southern portion of Kauai (HHP 1991w8). This species

is now known from the following scattered locations on private and

State land on Kauai at elevations between 820 and 2,000 ft (250 and 610

m) (Wagner et al. 1990): Mahanaloa-Kuia Valley in Kuia NAR; Haeleele

Valley; Na Pali Coast State Park; Limahuli Valley; the Koaie branch of

Waimea Canyon; Haupu Range; Wailua River; and Moloaa Forest Reserve

(HHP 1991w1 to 1991w7, 1991w9, 1991w10, 1991w11; HPCC 1990k1; HPCC

1991j1, 1991j2; T. Flynn and S. Montgomery, pers. comms., 1991). There

is an undocumented sighting of one individual at Makaleha, above the

town of Kapaa (T. Flynn, pers. comm., 1991). The 13 known populations,

totaling 170 to 300 individuals, typically grow on the sides of gulches

in diverse lowland mesic forests and sometimes lowland wet forests

(Wagner et al. 1990). Associated plant taxa include hame, lama,

lantana, 'ohi'a, and Pouteria sandwicensis ('ala'a) (Degener 1936; HHP

1991w1 to 1991w7, 1991w10; D. Herbst, pers. comm., 1991).

The major threats to Pteralyxia kauaiensis are habitat destruction

by feral animals and competition with introduced plants. Animals

affecting the survival of this species include feral goats, feral pigs,

and possibly rats, which may eat the fruits. Fire and over-collecting

for scientific purposes could threaten some populations. Introduced

plants competing with this species include common guava, daisy

fleabane, kukui, lantana, strawberry guava, and ti (HHP 1991w1, 1991w4,

1991w5, 1991w7; HPCC 1990k1, 1990k2; T. Flynn and S. Perlman, pers.

comms., 1991).

Gray (1854) described Schiedea spergulina based on a specimen

collected in 1840 on Kauai during the United States Exploring

Expedition. The specific epithet means ``resembling Spergula,'' another

genus in the same plant family. Two varieties of S. spergulina are

recognized in the current treatment of the genus (Wagner et al. 1990).

The typical variety, which includes var. degeneriana, was named by

Sherff (1956) and var. leiopoda (Sherff 1944), which includes var.

major, was also named by Sherff (1944).

Schiedea spergulina, a member of the pink family (Caryophyllaceae),

is a 1 to 2 ft (30 to 60 cm) tall subshrub. The opposite leaves are

very narrow, usually 1.2 to 2.6 in (30 to 65 mm) long and about 0.04 in

(1.4 mm) wide, one-veined, and attached directly to the stem. The

flowers are unisexual, with male and female flowers on different

plants. Flowers occur in compact clusters of three. The sepals usually

number five and are green and purple-tinged, 0.08 to 0.13 in (2 to 3.3

mm) long. The capsular fruits are about 0.08 to 0.12 in (2 to 3 mm)

long and contain nearly smooth, kidney-shaped seeds. Of the 22 species

in this endemic genus, only 2 other species have smooth seeds. This

species differs from those two in having very compact flower clusters.

The two weakly defined varieties differ primarily in the degree of

hairiness (Heller 1897; Hillebrand 1888; Sherff 1944, 1945; Wagner et

al. 1990).

Historically, Schiedea spergulina var. leiopoda was found on a

ridge on the east side of Hanapepe on Kauai (HHP 1991x1). One

population of 50 to 100 individuals of this variety is now known to

grow in Lawai Valley on Kauai on privately owned land (HHP 1991x2; HPCC

1991k; T. Flynn, J. Lau, and S. Perlman, pers. comms., 1991). Schiedea

spergulina var. spergulina is more numerous, once found in Olokele

Canyon but now known from Kalalau Rim and four locations in Waimea

Canyon on State land (HHP 1991y1 to 1991y5). One population contains

only five plants, whereas others number in the thousands. However,

these populations are estimated to total no more than 5,000 individuals

(HHP 1991y1 to 1991y5; T. Flynn, pers. comm., 1991; S. Perlman, pers.

comm., 1993). This taxon is usually found on bare rock outcrops or

sparsely vegetated portions of rocky cliff faces or cliff bases in

diverse lowland mesic forests at elevations between 590 and 3,000 ft

(180 and 800 m) (Wagner et al. 1990). Plants associated with the Lawai

population of S. spergulina var. leiopoda are Bidens sandvicensis

(ko'oko'olau), Doryopteris (kumuniu), Peperomia leptostachya, and

Plectranthus parviflorus ('ala'ala wai nui) (T. Flynn and J. Lau, pers.

comms., 1991; D. Lorence et al., in litt., 1991). Plant taxa associated

with S. spergulina var. spergulina include 'ahinahina, Chinaberry,

lantana, Sacramento bur, and Nototrichium sandwicense (kulu'i) (HHP

1991y5, Sherff 1956).

The major threats to Schiedea spergulina are habitat destruction by

feral goats and competition with introduced plants. Variety leiopoda is

threatened by competition with alien plant taxa such as koa haole,

lantana, and Furcraea foetida (Mauritius hemp), and individuals are

also damaged and destroyed by rock slides. This variety is potentially

threatened by pesticide use in nearby sugarcane fields. This variety is

threatened by stochastic extinction and/or reduced reproductive vigor

due to the small number of existing individuals (CPC 1990; D. Lorence,

T. Flynn, pers. comms., 1991). Variety spergulina is threatened by

competition with alien taxa, including daisy fleabane and lantana. The

area in which this variety grows is used heavily by feral goats and

there is evidence that plants are being browsed and trampled (HHP

1991y2, 1991y5; T. Flynn, J. Lau, and S. Perlman, pers. comms., 1991;

D. Lorence et al., in litt., 1991).

The intensity of threats and small number of populations known for

Schiedea spergulina var. leiopoda indicate that this taxon is in

serious danger of extinction. Schiedea spergulina var. spergulina has

significantly more individuals and populations and is facing less

intense threats since it is found on more protected cliffs. However,

while the latter taxon is not now in danger of extinction, if these

threats are not curtailed, the taxon will become endangered in the

future.

William Jackson Hooker and G.A.W. Arnott (1830-1841) described

Solanum sandwicense based on a specimen collected in 1826 or 1827 on

Oahu during the voyage of H.M.S. Blossom. The plant was named for the

Sandwich Islands, an older name for the Hawaiian Islands. Other names

by which portions of this species have been known include S.

hillebrandii (St. John 1969a), S. kauaiense (Hillebrand 1888), S.

sandwicense var.? kavaiense (Gray 1862), S. woahense (Symon 1990), and

S. woahense var. eroso-crenulatum (Symon 1990). In the current

treatment of this genus, Symon (1990) considers the Oahu and Kauai

populations as Solanum sandwicense and recognizes no subspecific taxa.

Solanum sandwicense, a member of the nightshade family, is a large

sprawling shrub that grows up to 13 ft (4 m) tall. The younger branches

are more densely hairy than older branches. The oval leaves are usually

4 to 6 in (10 to 15 cm) long and 2 to 5.5 in (5 to 14 cm) wide and have

up to four lobes along the margins. Leaf stalks are 0.8 to 1.6 in (2 to

4 cm) long. On the flowering stem, a few to as many as 40 flowers are

grouped in threes, with each flower on a stalk about 0.6 in (15 mm)

long, bent at the end so that the flower faces downward. The corolla is

white with a faint purplish stripe, each lobe is curved somewhat

backward. Stamens are attached low on the corolla tube, with anthers

curved inward. The fruit is a berry 0.5 to 0.6 in (13 to 15 mm) in

diameter, black when ripe. This species differs from others of the

genus in having dense hairs on young plant parts, a greater height, and

its lack of prickles (Gray 1862, St. John 1969a, Sohmer and Gustafson

1987, Symon 1990).

Historically, Solanum sandwicense was known from widely scattered

populations throughout the Waianae Mountains and southern portions of

the Koolau Mountains on Oahu (HHP 1991z1 to 1991z5, 1991z7 to 1991z10).

On Kauai, this species was known from locations in the Kokee region

bounded by Kalalau Valley to the north, Milolii Ridge to the west, and

Kawaikoi to the east, extending southward to the Hanapepe River (HHP

1991z13 to 1991z17, 1991z21, 1991z22, 1991z24). On Oahu, this species

is known from a single population on privately owned land in what is

now Honouliuli Preserve (HPCC 1991l). One other recent population was

destroyed by a landslide in 1986 (HHP 1991z6; J. Obata, pers. comm.,

1991; D. Lorence et al., in litt., 1991). The Kauai populations are on

private and State land and most are from Kokee and Na Pali Coast State

Parks. Of the 12 known populations, only 4 are currently extant; they

total about 20 plants (Bruegmann 1990; CPC 1990; HHP 1991z11, 1991z12,

1991z19, 1991z20, 1991z26; D. Herbst, pers. comm., 1991; S. Perlman,

pers. comm., 1993). This species is typically found in open, sunny

areas at elevations between 2,500 and 4,000 ft (760 and 1,220 m) in

diverse lowland to montane mesic forests and occasionally in wet

forests (HHP 1991z1, 1991z11, 1991z16, 1991z19 to 1991z26; Symon 1990).

Associated plant taxa include koa, 'ohi'a, uluhe, and wet forest plants

such as kopiko, ho'i'o, and more common Melicope species (alani) (HHP

1991z11, 1991z18, 1991z20, 1991z26).

The major threats to populations of Solanum sandwicense on Kauai

are habitat degradation by feral pigs and competition with alien plant

taxa. Alien taxa that have heavily invaded this species' habitat on

Kauai include banana poka, prickly Florida blackberry, strawberry

guava, Hedychium gardnerianum (kahili ginger), and Lonicera japonica

(Japanese honeysuckle). This species is also threatened by fire, over-

collecting for scientific purposes, stochastic extinction, and/or

reduced reproductive vigor due to the small number of existing

individuals. All Oahu populations of Solanum sandwicense except one are

now apparently extinct, the result of its habitat being destroyed by

urbanization, landslides, feral pigs, and weedy alien taxa (Bruegmann

1990; HHP 1991z1 to 1991z7, 1991z18, 1991z25; HPCC 1990m; R. Hobdy, J.

Lau, J. Obata, and S. Perlman, pers. comms., 1991; D. Lorence et al.,

in litt., 1991).

Previous Federal Action

Federal action on these plants began as a result of section 12 of

the Endangered Species Act (16 U.S.C. 1531 et seq.), which directed the

Secretary of the Smithsonian Institution to prepare a report on plants

considered to be endangered, threatened, or extinct in the United

States. This report, designated as House Document No. 94-51, was

presented to Congress on January 9, 1975. In that document, Brighamia

insignis (as B. insignis and B. citrina var. napaliensis), Delissea

rhytidosperma, Exocarpos luteolus, Hibiscus clayi (as H. clayi and H.

newhousei), Lipochaeta fauriei, Lipochaeta micrantha (as L. exigua),

Melicope haupuensis (as Pelea haupuensis), Melicope knudsenii (as Pelea

multiflora), Melicope pallida (as Pelea pallida and P. leveillei),

Melicope quadrangularis (as Pelea quadrangularis), Nothocestrum

peltatum, Peucedanum sandwicense (as P. kauaiense), Pteralyxia

kauaiensis, and Solanum sandwicense were considered to be endangered.

Diellia pallida (as D. laciniata), Lipochaeta fauriei, Lipochaeta

micrantha, Lipochaeta waimeaensis, Lysimachia filifolia, and Solanum

sandwicense (as S. kauaiense) were considered to be threatened.

Hedyotis cookiana, Melicope knudsenii (as Pelea knudsenii and P.

tomentosa), Munroidendron racemosum (as M. racemosum var.

macdanielsii), and Solanum sandwicense (as S. hillebrandii) were

considered to be extinct.

On July 1, 1975, the Service published a notice in the Federal

Register (40 FR 27823) of its acceptance of the Smithsonian report as a

petition within the context of section 4(c)(2) (now section 4(b)(3)) of

the Act, and giving notice of its intention to review the status of the

plant species named therein. As a result of that review, on June 16,

1976, the Service published a proposed rule in the Federal Register (41

FR 24523) to determine endangered status pursuant to section 4 of the

Act for approximately 1,700 vascular plant species, including all of

the above species considered to be endangered or thought to be extinct.

The list of 1,700 plant species was assembled on the basis of comments

and data received by the Smithsonian Institution and the Service in

response to House Document No. 94-51 and the July 1, 1975, Federal

Register publication.

General comments received in response to the 1976 proposal are

summarized in an April 26, 1978, Federal Register publication (43 FR

17909). In 1978, amendments to the Act required that all proposals over

two years old be withdrawn. A one-year grace period was given to

proposals already over two years old. On December 10, 1979, the Service

published a notice in the Federal Register (44 FR 70796) withdrawing

the portion of the June 16, 1976, proposal that had not been made

final, along with four other proposals that had expired. The Service

published updated notices of review for plants on December 15, 1980 (45

FR 82479), September 27, 1985 (50 FR 39525), and February 21, 1990 (55

FR 6183). In these notices, 10 of the species (including synonymous

species) that had been in the 1976 proposed rule were treated as

Category 1 candidates for Federal listing. Category 1 species are those

for which the Service has on file substantial information on biological

vulnerability and threats to support preparation of listing proposals.

Other than Diellia pallida (as D. laciniata), Hedyotis cookiana,

Lipochaeta fauriei, Lipochaeta micrantha (as L. exigua), Lysimachia

filifolia, Melicope knudsenii (as Pelea knudsenii), Melicope pallida,

M. quadrangularis, Peucedanum sandwicense, and Solanum sandwicense (as

S. hillebrandii), all the aforementioned species that were either

proposed as endangered or thought to be extinct in the June 16, 1976,

proposed rule were considered Category 1 candidates on all three

notices of review.

In the 1980 and 1985 notices, Lipochaeta fauriei, Melicope

knudsenii (as Pelea knudsenii), and Solanum sandwicense (as S.

hillebrandii) were considered Category 1* species. Category 1* species

are those that are possibly extinct. Lysimachia filifolia appeared as a

Category 2 species and Hedyotis cookiana as a Category 3A species in

the 1980 and 1985 notices. Category 2 species are those for which there

is some evidence of vulnerability, but for which there are not enough

data to support listing proposals at the time. Category 3A species are

those for which the Service has persuasive evidence of extinction.

Because new information indicated their current existence or provided

support for listing, the above five species were conferred Category 1

status in the 1990 notice. Lipochaeta exigua appeared as a Category 3B

species in the 1980 and 1985 notices; in the 1990 notice, it was

considered synonymous with L. micrantha, a Category 1 species. Category

3B species are those which, on the basis of current taxonomic

understanding, do not represent distinct taxa meeting the Act's

definition of ``species.'' Diellia pallida (as D. laciniata), Melicope

pallida, and M. quadrangularis were accorded Category 1* status in the

1990 notice, but because new information regarding their existence has

become available, they are included herein for listing. In 1980,

Peucedanum sandwicense appeared as a Category 2 species and retained

that status in the 1985 and 1990 notices. Information obtained since

the 1990 notice suggests that its numbers and distribution are

sufficiently restricted to warrant listing. Schiedea spergulina first

appeared on the 1985 notice of review as a Category 1 species. In the

1990 notice, two varieties were recognized: Variety spergulina as a

Category 1 species; and variety leiopoda as a Category 1* species for

which recently obtained information indicates that it is extant.

Cyrtandra limahuliensis first appeared in the 1990 notice of review as

Category 1 species after it was described in 1987. The 1990 notice also

recognized Cyanea asarifolia and Phyllostegia waimeae as Category 1

species for the first time.

Section 4(b)(3)(B) of the Act requires the Secretary to make

findings on certain pending petitions within 12 months of their

receipt. Section 2(b)(1) of the 1982 amendments further requires all

petitions pending on October 13, 1982, be treated as having been newly

submitted on that date. On October 13, 1983, the Service found that the

petitioned listing of these species was warranted, but precluded by

other pending listing actions, in accordance with section

4(b)(3)(B)(iii) of the Act; notification of this finding was published

on January 20, 1984 (49 FR 2485). Such a finding requires the petition

to be recycled, pursuant to section 4(b)(3)(C)(i) of the Act. The

finding was reviewed in October of 1984, 1985, 1986, 1987, 1988, 1989,

and 1990. The proposed rule published on October 30, 1991 (56 FR 55862)

to list 23 plant species primarily from the island of Kauai as

endangered species constituted the final 1-year finding that was

required for the species discussed above.

Based on comments and recommendations received in response to the

proposal (see Comments and Recommendations, below), the Service now

determines 21 plant species to be endangered and 3 plant species to be

threatened with the publication of this rule. One of the species

proposed in 1991 is now being listed as two separate entities: Schiedea

spergulina var. spergulina (as threatened); and Schiedea spergulina

var. leiopoda (as endangered). This results in the apparent increase

from 23 species in the proposed rule to 24 species in the final rule.

Summary of Comments and Recommendations

In the October 30, 1991, proposed rule and associated

notifications, all interested parties were requested to submit factual

reports or information that might contribute to the development of a

final listing decision. The public comment period ended on December 30,

1991. Appropriate State agencies, county governments, Federal agencies,

scientific organizations, and other interested parties were contacted

and requested to comment. A newspaper notice inviting public comment

was published in ``The Garden Island'' on November 10, 1991. Fifteen

letters of comment, including 2 from State agencies, 11 from

organizations, and 2 from individuals, were received and are discussed

below. A public hearing was requested by Kamehameha Schools/Bernice

Pauahi Bishop Estate on December 1, 1992. On January 13, 1993, the

Service published a notice (58 FR 4145) reopening the comment period

until February 20, 1993, to accommodate the requested public hearing

which was held in Kapaa, Kauai, on February 10, 1993. A newspaper

notice announcing the public hearing was published in ``The Garden

Island'' on February 10, 1993. Eleven people presented oral comments;

their testimony is included in the following summary.

Two respondents acknowledged receipt of the proposed rule but had

no comments. Of the remaining 13 letters, 7 supported the listing of

these taxa from the island of Kauai and 6 opposed 1 or more of the

listings. Additional information included in the letters has been

incorporated into this final rule. Comments of similar content were

grouped into a number of general issues for discussion. These issues

and the Service's response to each are discussed below.

Issue 1: Status of Cyrtandra limahuliensis: Two respondents stated

that more populations of Cyrtandra limahuliensis exist than were

discussed in the proposed rule and that there is a good possibility

that more populations await discovery. One respondent asked whether

this species should still be considered endangered, while the second

stated that this species does not warrant listing as endangered.

Response: At the time the proposed rule was written, only 9

populations of Cyrtandra limahuliensis were known; however, 3

additional populations were subsequently discovered in Waioli Valley,

bringing the total number of plants to 2,800 to 3,000 (D. Lorence, in

litt., 1993). This information has been included in the final rule.

Based on additional information, the Service determines that Cyrtandra

limahuliensis is not now in danger of extinction, but that Cyrtandra

limahuliensis is likely to become endangered in the foreseeable future

if the threats posed by competition from alien species are not curbed.

Thus, Cyrtandra limahuliensis is designated a threatened species.

Issue 2: Status of Peucedanum sandwicense: One respondent stated

that more populations of Peucedanum sandwicense exist than were

discussed in the proposed rule and that this species is not immediately

threatened with extinction. The proposed rule indicated that only 16

populations of the species were known, 2 on Oahu, 3 on Molokai, 10 on

Kauai, and 1 on Maui. Additional populations are now known from the

Wailau sea cliffs on Molokai and the Kalalau Valley rim on Kauai (D.

Lorence et al., in litt., 1991).

Response: At the time the proposed rule was written, 250 to 350

individuals of Peucedanum sandwicense were known; however, several

additional colonies were subsequently brought to the Service's

attention, bringing the totals to between 1,000 to 5,000 individuals

(CPC 1992; D. Lorence et al., in litt., 1991). This information has

been included in the final rule. Based on the above information, the

Service determines that Peucedanum sandwicense is not now in danger of

extinction, but that it is likely to become endangered in the

foreseeable future if the threats posed by competition from alien plant

species and feral goats are not curbed. Thus, Peucedanum sandwicense is

designated a threatened species.

Issue 3: Status of Schiedea spergulina: One respondent stated that

one variety of Schiedea spergulina, var. spergulina, is not immediately

threatened with extinction. The proposed rule treated the species as

one entity in terms of listing.

Response: Schiedea spergulina var. leiopoda is known from only 1

population of 50 to 100 individuals (HHP 1991x2; T. Flynn, J. Lau, and

S. Perlman, pers. comms., 1991). The numbers of plants and populations

of this variety are sufficiently small that, given its threats, it is

in immediate danger of extinction and meets the definition of an

endangered species as defined in the Act. Schiedea spergulina var.

spergulina is known from 5 locations with from 1,000 to 5,000

individuals total. Based on the above information, the Service

determines that Schiedea spergulina var. spergulina is not now in

danger of extinction, but that it is likely to become endangered in the

foreseeable future if the threats posed by competition from alien taxa

and feral goats are not curbed. Thus, Schiedea spergulina var.

spergulina is designated a threatened species and S. spergulina var.

leiopoda is designated as an endangered species.

Issue 4: Changes to the Act: Several concerns were expressed in

letters of comment and in testimony at the public hearing about the

Endangered Species Act and its effect on private citizens and

landowners. One individual asked both during testimony at the hearing

and in multiple letters of comment that the ability of the Federal

government to seize land with endangered species be removed from the

Act. Concern was expressed by four individuals during their testimony

and in three letters of comment that the Endangered Species Act does

not allow private citizens to grow and propagate endangered species.

Many respondents asked that the permit process be streamlined for

propagation and scientific research purposes.

Response: The Act does not require the seizure or acquisition of

private lands containing endangered species. The Act also does not

restrict the growth and propagation of endangered plant taxa by private

citizens as long as the plants are not collected from Federal lands,

are not transported between states or outside of the U.S., and that

these actions do not violate State laws. A Federal permit for

collecting is only required if the collection involves Federal actions,

Federal funding, or activities on Federal lands.

Issue 5: Changes to State regulations: Concern was expressed by

four individuals during their testimony and in three letters of comment

that State regulations might make the propagation or cultivation of

these 24 taxa more difficult if they were listed as endangered, since

State regulations prohibit possessing endangered plants or collecting

their propagules. Permits to collect or possess endangered plants may

be issued by the State to enhance the propagation or survival of the

species, but the respondents believe that it is difficult and

burdensome to obtain these permits. One individual was also concerned

about the seizure of private lands by the State to protect endangered

species. Another individual stated that rezoning for conservation

districts should be limited to public lands.

Response: Hawaii Administration Rule 13-124-4(a) allows for

``Permits to * * * possess * * * any endangered or threatened species

of wildlife or plants'' to be issued ``to enhance the propagation or

survival of the species.'' To date, the State has worked with private

landowners to reach a cooperative agreement for management of

endangered species habitat or to reach a fair purchase price. Concerns

about the State's endangered species law and how it affects private

citizens, and any requests to change State requirements, should be

addressed to State authorities. The Service believes that existing

State policies should not delay or prevent the Federal protection that

listing would afford the species.

Issue 6: Threat of game mammals: One individual at the hearing was

concerned that there is inadequate information on the effects of feral

ungulates to indicate that their eradication would truly benefit the

proposed plants.

Response: The Service is not proposing the eradication of feral

ungulates from the island of Kauai. However, several studies verify

that feral ungulates damage native plants and habitats. Feral goats

have been implicated in the damage of native vegetation ranging from

lowland to subalpine areas (Mueller-Dombois and Spatz 1972, Spatz and

Mueller-Dombois 1973, Scowcroft and Sakai 1983). Goat browsing damage

has been observed on individuals of Exocarpus luteolus, Lipochaeta

micrantha, and Schiedea spergulina var. spergulina (HHP 1991k5; T.

Flynn, pers. comm., 1991; S. Perlman, pers. comm., 1993). Goats

threaten 12 other species through habitat degradation.

Pigs have been similarly implicated in damaging native habitats.

Specific studies have demonstrated that native vegetation recovers

significantly after the exclusion of pigs in rainforests and subalpine

grasslands (Spatz and Mueller-Dombois 1975, Stone 1985). Eleven species

in this rule are threatened by habitat degradation caused by pigs.

Elimination or reduction of habitat damage by feral ungulates will be

addressed during the recovery process for these species.

Issue 7: Lack of adequate management: Four respondents and three

individuals in their testimony stated that inadequate management of

State and Federal lands in the past has caused the decline of many of

the 24 taxa.

Response: Listing a species as either endangered or threatened does

enable States, such as Hawaii, to apply for Federal funds to undertake

programs aimed at conserving endangered species. Additional concerns

about the management of State lands should be addressed to State

authorities. None of the 24 taxa are found on federally owned lands.

Populations of one species, Peucedanum sandwicense, are found on State

land managed under a cooperative agreement with the National Park

Service. The National Park Service assumed management of the area in

1980. A natural resource management plan will be developed for the area

soon, but must be approved by the State before it can be implemented

(Gary Barbano, National Park Service, pers. comm., 1993).

Summary of Factors Affecting the Species

After a thorough review and consideration of all information

available, the Service has determined that Brighamia insignis ('olulu),

Cyanea asarifolia (haha), Delissea rhytidosperma (no common name

(NCN)), Diellia pallida (NCN), Exocarpos luteolus (heau), Hedyotis

cookiana ('awiwi), Hibiscus clayi (Clay's hibiscus), Lipochaeta fauriei

(nehe), Lipochaeta micrantha (nehe), Lipochaeta waimeaensis (nehe),

Lysimachia filifolia (NCN), Melicope haupuensis (alani), Melicope

knudsenii (alani), Melicope pallida (alani), Melicope quadrangularis

(alani), Munroidendron racemosum (NCN), Nothocestrum peltatum ('aiea),

Phyllostegia waimeae (NCN), Pteralyxia kauaiensis (kaulu), Schiedea

spergulina var. leiopoda (NCN), and Solanum sandwicense

(popolo'aiakeakua) should be classified as endangered species; and that

Cyrtandra limahuliensis (ha'iwale), Peucedanum sandwicense (makou), and

Schiedea spergulina var. spergulina (NCN) should be classified as

threatened species. The provisions of section 4 of the Endangered

Species Act (16 U.S.C 1533) and regulations (50 CFR part 424)

promulgated to implement the listing provisions of the Act were

followed in making these determinations. A species may be determined to

be an endangered or threatened species due to one or more of the five

factors described in section 4(a)(1). The threats facing these 24 taxa

are summarized in Table 1.

Table1.--Summary of Threats

--------------------------------------------------------------------------------------------------------------------------------------------------------

Alien mammals

Species -------------------------------------------------- Insects Alien Fire Natural Human Limited

Cattle Deer Goats Pigs Rats plants disaster* impacts numbers**

--------------------------------------------------------------------------------------------------------------------------------------------------------

Brighamia insignis.................... ........ ........ X ........ ........ X X X X X X1,3

Cyanea asarifolia..................... ........ ........ ........ P P ........ ........ ........ X X X1,2

Cyrtandra limahuliensis............... ........ ........ ........ X ........ ........ X ........ P P

Delissea rhytidosperma................ ........ X X X X ........ X X X X X1,2

Diellia pallida....................... ........ X X X ........ ........ X ........ X X X1,2

Exocarpos luteolus.................... ........ ........ X X X ........ X X X P

Hedyotis cookiana..................... ........ ........ P P ........ ........ P ........ P P X1,3

Hibiscus clayi........................ ........ ........ ........ P ........ ........ X ........ .......... X X1,2

Lipochaeta fauriei.................... ........ ........ X P ........ ........ X X .......... P X1,3

Lipochaeta micrantha var. exigua...... ........ ........ ........ X ........ ........ X ........ .......... P X1

Lipochaeta micrantha vir. micrantha... ........ ........ X X ........ ........ X ........ .......... P X1

Lipochaeta waimeaensis................ ........ ........ X ........ ........ ........ X ........ X X X1,2

Lysimachia filifolia.................. ........ ........ ........ P ........ ........ X ........ X P X1

Melicope haupuensis................... ........ ........ X ........ ........ P X ........ .......... X X1,2

Melicope knudsenii.................... X P X X ........ P X P .......... P X1,3

Melicope pallida...................... ........ ........ X X ........ P X X .......... P X3

Melicope quadrangularis............... ........ ........ ........ ........ ........ P P ........ .......... P X1,3

Munroidendron racemosum............... ........ ........ X ........ P P X X X P X3

Nothocestrum peltatum................. ........ X X X ........ ........ X X .......... X X1,3

Peucedanum sandwicense................ ........ P X ........ ........ ........ X X X X

Phyllostegia waimeae.................. ........ ........ X ........ ........ ........ X ........ X X X1,2

Pteralyxia kauaiensis................. ........ ........ X X P ........ X X .......... P

Schiedea spergulina var. leiopoda..... ........ ........ X ........ ........ ........ X ........ X P X1,3

Schiedea spergulina var. spergulina... ........ ........ X ........ ........ ........ X ........ .......... P X1

Solanum sandwicense................... ........ ........ ........ X ........ ........ X X X X X1,3

--------------------------------------------------------------------------------------------------------------------------------------------------------

X=Immediate and significant threat.

P=Potential threat.

*=Natural disasters include hurricanes, erosion, rock slides, and landslides.

**=No more than 100 individuals and/or no more than 5 populations.

1=No more than 5 populations.

2=No more than 10 individuals.

3=No more than 100 individuals.

These factors and their application to Brighamia insignis A. Gray

('olulu), Cyanea asarifolia St. John (haha), Cyrtandra limahuliensis

St. John (ha'iwale), Delissea rhytidosperma H. Mann (no common name

(NCN)), Diellia pallida W.H. Wagner (NCN), Exocarpos luteolus C. Forbes

(heau), Hedyotis cookiana (Cham. and Schlechtend.) Steud. ('awiwi),

Hibiscus clayi Degener and I. Degener (Clay's hibiscus), Lipochaeta

fauriei H. Levl. (nehe), Lipochaeta micrantha (Nutt.) A. Gray (nehe),

Lipochaeta waimeaensis St. John (nehe), Lysimachia filifolia C. Forbes

and Lydgate (NCN), Melicope haupuensis (St. John) Hartley and Stone

(alani), Melicope knudsenii (Hillebr.) Hartley and Stone (alani),

Melicope pallida (Hillebr.) Hartley and Stone (alani), Melicope

quadrangularis (St. John and E. Hume) T. Hartley and B. Stone (alani),

Munroidendron racemosum (C. Forbes) Sherff (NCN), Nothocestrum peltatum

Skottsb. ('aiea), Peucedanum sandwicense Hillebr. (makou), Phyllostegia

waimeae Wawra (NCN), Pteralyxia kauaiensis Caum (kaulu), Schiedea

spergulina var. leiopoda Sherff (NCN), Schiedea spergulina A. Gray var.

spergulina (NCN), and Solanum sandwicense Hook. and Arnott.

(popolo'aiakeakua) are as follows:

A. The Present or Threatened Destruction, Modification, or Curtailment

of Its Habitat or Range

The habitats of the plants included in this rule have undergone

extreme alteration because of past and present land management

practices, including deliberate alien animal and plant introductions,

agricultural development, and recreational use. Natural disturbances

such as storms and landslides also destroy habitat and can have a

significant effect on small populations of plants. Destruction and

modification of habitat by introduced animals and competition with

alien plants are the primary threats facing the 24 plant taxa in this

rule (see Table 1).

When Polynesian immigrants settled in the Hawaiian Islands, they

brought with them water-control and slash-and-burn systems of

agriculture and encouraged plants that they introduced to grow in

valleys. Their use of the land resulted in erosion, changes in the

composition of native communities, and a reduction of biodiversity

(Cuddihy and Stone 1990; HHP 1990b; Kirch 1982; Wagner et al. 1985).

Hawaiians settled and altered many areas of Kauai including areas in

which some of the taxa in this rule grew (DLNR 1981a; HHP 1990a,

1990b). Many forested slopes were denuded in the mid-1800s to supply

firewood to whaling ships, plantations, and Honolulu residents. Native

plants, such as the historic population of Lipochaeta micrantha var.

micrantha in Koloa District (HHP 1991k4), were undoubtedly affected by

this practice. Also, sandalwood and tree fern harvesting occurred in

many areas, changing forest composition and affecting native taxa

(Cuddihy and Stone 1990).

Beginning with Captain James Cook in 1792, early European explorers

introduced livestock, which became feral, increased in number and

range, and caused significant changes to the natural environment of

Hawaii. The 1848 provision for land sales to individuals allowed large-

scale agricultural and ranching ventures to begin. So much land was

cleared for these enterprises that climatic conditions began to change

and the amount and distribution of rainfall were altered (Wenkam 1969).

Plantation owners supported reforestation programs that resulted in

many alien trees being introduced in the hope that the watershed could

be conserved. Beginning in the 1920s, water collection and diversion

systems were constructed in upland areas to irrigate lowland fields. It

is probable that individuals and populations of native plants were

destroyed. Some of the taxa, such as a Kokee population of Exocarpos

luteolus and a Waimea Canyon population of Schiedea spergulina var.

spergulina, which now occur near ditches of the irrigation system, may

have been affected (HHP 1991f4, 1991y2). The irrigation system also

opened new routes for the invasion of alien plants and animals into

native forests (Cuddihy and Stone 1990, Culliney 1988, Wagner et al.

1990, Wenkam 1969).

Past and present activities of introduced alien mammals are the

primary factor in altering and degrading vegetation and habitats on

Kauai, Niihau, Oahu, Molokai, and Maui. Feral ungulates trample and eat

native vegetation and disturb and open new areas. This causes erosion

and allows the entry of alien plant taxa (Cuddihy and Stone 1990,

Wagner et al. 1990). Nineteen taxa in this proposal are directly

threatened by habitat degradation resulting from introduced ungulates:

17 taxa are threatened by goats; 10 by pigs; 3 by deer; and 1 by

cattle. In addition, an introduced ground-nesting bird potentially

threatens one taxon by disturbing its habitat.

Goat (Capra hircus), a species originally native to the Middle East

and India, was successfully introduced to the Hawaiian Islands in 1792

and there currently are populations on Kauai, Oahu, Molokai, Maui, and

Hawaii. All feral goats were removed from Niihau about 1910, but by

that time they had caused considerable damage to the island's dry and

mesic forests. On Kauai, feral goats have been present in drier, more

rugged areas since the 1820s. They still occur in Waimea Canyon and

along the Na Pali Coast, as well as in the drier perimeter of Alakai

Swamp and even in its wetter areas during periods with low rainfall.

Goats have been on Oahu since about 1820 and they currently occur in

the northern Waianae Mountains. On Molokai, goats degrade dry forests

at low elevations and they are expanding their range (J. Lau, pers.

comm., 1991). On Maui, goats have been widespread for 100 to 150 years

and are common throughout the south slope of Haleakala (Medeiros et al.

1986). Goats are managed in Hawaii as a game animal, but many herds

populate inaccessible areas where hunting has little effect on their

numbers (HHP 1990c). Goat hunting is allowed year-round or during

certain months, depending on the area (DLNR n.d.-a, n.d.-b, n.d.-c,

1990). Goats browse on introduced grasses and native plants, especially

in drier and more open ecosystems. Feral goats eat native vegetation,

trample roots and seedlings, cause erosion, and promote the invasion of

alien plants. They are able to forage in extremely rugged terrain and

have a high reproductive capacity (Clarke and Cuddihy 1980, Cuddihy and

Stone 1990, Culliney 1988, Scott et al. 1986, Tomich 1986, van Riper

and van Riper 1982).

Although many of the plants survive on steep cliffs inaccessible to

goats, their original range was probably much larger. Plants are

vulnerable to the long-term, indirect effects of goats, such as large-

scale erosion (Corn et al. 1979). The habitats of many of the 24 plant

taxa in this rule were damaged in the past by goats and these effects

are still apparent today in the form of alien vegetation and erosion.

One or more populations of 17 of the following taxa are currently

threatened by direct damage from feral goats, such as trampling of

plants and seedlings and erosion of substrate: Brighamia insignis;

Delissea rhytidosperma; Diellia pallida; Exocarpos luteolus; Lipochaeta

fauriei; Lipochaeta micrantha var. micrantha; Lipochaeta waimeaensis;

Melicope haupuensis; Melicope knudsenii; Melicope pallida;

Munroidendron racemosum; Nothocestrum peltatum; Peucedanum sandwicense;

Phyllostegia waimeae; Pteralyxia kauaiensis, and both varieties of

Schiedea spergulina. In addition, it is probable that goats have

invaded the area in which the only known population of Hedyotis

cookiana occurs (Bruegmann 1990; Clarke and Cuddihy 1980; Culliney

1988; HHP 1991a1, 1991e3, 1991f6, 1991i3, 1991k5, 1991o1, 1991o2,

1991p1 to 1991p4, 1991q6, 1991q8, 1991s1, 1991s8 to 1991s10, 1991s15,

1991t7, 1991u14, 1991w5, 1991y5; HPCC 1990a, 1990i4, 1990j2, 1990j3,

1990k1, 1990k2; Lammers 1990; Lamoureux 1982; Medeiros et al. 1986;

Perlman 1979; St. John 1981b; Scott et al. 1986; Takeuchi 1982; van

Riper and van Riper 1982; C. Christensen, T. Flynn, R. Hobdy, J. Lau,

D. Lorence, S. Montgomery, S. Perlman, and K. Wood, pers. comms.,

1991).

Pig (Sus scrofa) is a species originally native to Europe, northern

Africa, Asia Minor, and Asia. European pigs, introduced to Hawaii by

Captain James Cook in 1778, became feral and invaded forested areas,

especially wet and mesic forests and dry areas at high elevations. They

are currently present on Kauai, Oahu, Molokai, Maui, and Hawaii, and

inhabit rain forests and grasslands. Pig hunting is allowed on all

islands either year-round or during certain months, depending on the

area (DLNR n.d.-a, n.d.-b, n.d.-c, 1990). While rooting in the ground

in search of the invertebrates and plant material they eat, feral pigs

disturb and destroy vegetative cover, trample plants and seedlings, and

threaten forest regeneration by damaging seeds and seedlings. They

disturb soil substrates and cause erosion, especially on slopes. Alien

plant seeds are dispersed on their hooves and coats as well as through

their digestive tracts. The disturbed soil is fertilized by their

feces, helping these plants to establish. Pigs are a major vector in

the spread of banana poka, firetree, and strawberry guava and enhance

populations of common guava, kahili ginger, Hamakua pamakani, prickly

Florida blackberry, sweet granadilla, and yellow ginger, all of which

threaten one or more of the taxa (Cuddihy and Stone 1990, Medeiros et

al. 1986, Scott et al. 1986, Smith 1985, Stone 1985, Tomich 1986,

Wagner et al. 1990).

Feral pigs pose an immediate threat to 1 or more populations of 11

of the taxa. At least one population of each of the following taxa is

threatened by feral pigs: Pteralyxia kauaiensis; Solanum sandwicense;

both varieties of Lipochaeta micrantha; Cyrtandra limahuliensis;

Delissea rhytidosperma; Diellia pallida; Exocarpos luteolus; Lipochaeta

fauriei; Melicope knudsenii; Melicope pallida; and Nothocestrum

peltatum. Pigs also constitute a potential threat to Cyanea asarifolia,

Hedyotis cookiana, Hibiscus clayi, Lipochaeta fauriei, and Lysimachia

filifolia (Bruegmann 1990; HHP 1991f6, 1991p1, 1991p3; HPCC 1990i3,

1990i4; J. Obata, pers. comm., 1990; C. Christensen, T. Flynn, R.

Hobdy, J. Lau, D. Lorence, and S. Perlman, pers. comms., 1991).

Cattle (Bos taurus), the wild progenitor of which was native to

Europe, north Africa, and southwestern Asia, were introduced to the

Hawaiian Islands in 1793. Large feral herds developed as a result of

restrictions on killing cattle decreed by King Kamehameha I. Feral

cattle formerly occurred on Niihau and, along with goats and sheep

(Ovis aries), caused much damage on the island (Stone 1985). On Kauai,

parts of Kokee were leased for cattle grazing in the 1850s and both

sides of Waimea Canyon were supporting large cattle ranching operations

by the 1870s (Joesting 1984, Ryan and Chang 1985). Cattle grazing began

about 1920 in the Na Pali region (DLNR 1981a). Cattle roamed lowland

areas and eventually began invading wet forests from adjacent mesic

areas. Around 1900, Augustus Knudsen, the district forester of Kauai

and a rancher, realizing the amount of destruction being caused to the

forests by cattle, initiated some fencing (Daehler 1973). Sugar company

interests funded additional fencing as well as feral cattle removal to

protect the forest from further degradation and to safeguard water

reserves for their crops (Wenkam 1969). On Kauai, feral cattle were

still present in Kokee as late as 1960 and in the Puu Ka Pele area in

the 1980s. Feral cattle roamed Oahu, but most were removed by the early

1960s. Today, only a few can be found in the northwestern part of the

island (J. Lau, pers. comm., 1990). Feral cattle were formerly found on

Molokai and Maui and damaged the forests there. Hunting of feral cattle

is no longer allowed in Hawaii (DLNR 1985). Cattle eat native

vegetation, trample roots and seedlings, cause erosion, create

disturbed areas into which alien plants invade, and spread seeds of

alien plants in their feces and on their bodies. The forest, in areas

grazed by cattle, becomes degraded to grassland pasture. Plant cover is

reduced for many years following removal of cattle from an area.

Several alien grasses and legumes purposely introduced for cattle

forage have become noxious weeds (Cuddihy and Stone 1990, Scott et al.

1986, Tomich 1986).

The habitats of many of the plants included in this rule were

degraded in the past by feral cattle. This has had effects which still

persist. Examples of plant taxa whose habitats have been altered by

feral cattle include Hibiscus clayi and Munroidendron racemosum. The

Maui population of Melicope knudsenii, growing in an area currently

used as a domestic cattle pasture, is directly threatened by trampling

by this animal (Degener and Degener 1959a; HHP 1991h3, 1991p1;

Lamoureux 1982).

Individuals of mule deer or black-tailed deer (Odocoileus

hemionus), native from western North America to central Mexico, were

brought to Kauai from Oregon in the 1960s for game hunting and have not

been introduced to any other Hawaiian island. In part, mule deer were

introduced to provide another animal for hunting, since the State had

planned to reduce the number of goats on Kauai because they were so

destructive to the landscape (Kramer 1971). There are about 400 animals

in and near Waimea Canyon, with some invasion into Alakai Swamp in

drier periods. Mule deer, legally hunted during only one month each

year, trample native vegetation and cause erosion by creating trails

and removing vegetation (Cuddihy and Stone 1990, DLNR 1985, Tomich

1986). They are a threat to Delissea rhytidosperma, Diellia pallida,

and Nothocestrum peltatum (Bruegmann 1990; HPCC 1990b, 1990i3, 1990i4;

S. Perlman, pers. comm., 1991).

Axis deer (Axis), native to Sri Lanka and India, were first

introduced to the Hawaiian Islands in 1868 as a game animal on Molokai,

later to Oahu and Lanai, and finally to East Maui in 1960. Hunting of

axis deer is allowed only on Molokai and Lanai during two months of the

year. Considerable damage has been done to the forests on Molokai and

Lanai by this animal, especially through browsing of vegetation and

compaction of the soil (Cuddihy and Stone 1990, Culliney 1988, DLNR

1985, Scott et al. 1986, Tomich 1986). With a population of about 100

animals on the lower southwest slope of Haleakala, the range of the

axis deer is expanding on East Maui and constitutes a potential threat

to Melicope knudsenii (Medeiros et al. 1986). On Molokai, axis deer are

encroaching on Pelekunu Valley and are already present in Kalaupapa,

thus posing a potential threat to populations of Peucedanum sandwicense

in these areas (HHP 1991u7, 1991u16; J. Lau, pers. comm., 1990).

Red jungle fowl (Gallus), ground-nesting chickens native to India

and southeast Asia, were introduced to Hawaii by the Polynesian

immigrants and became feral in the forests. A current threat to the

Makaha Valley, Kauai, population of Nothocestrum peltatum, red jungle

fowl disturb the ground cover while searching for seeds, fruits, and

small invertebrates, thus disrupting seedling establishment (Cuddihy

and Stone 1990, HPCC 1990i3, Scott et al. 1986).

Substrate loss due to agriculture, grazing animals (especially

goats), hikers, and vegetation change results in habitat degradation

and loss. This particularly affects plant populations located on cliffs

or steep slopes, including Lipochaeta waimeaensis, Brighamia insignis,

Diellia pallida, Exocarpos luteolus, Peucedanum sandwicense, and

Phyllostegia waimeae (Bruegmann 1990; Christensen 1979; HHP 1991f6;

Takeuchi 1982; G. Carr, R. Hobdy, and J. Obata, pers. comms., 1991).

B. Overutilization for Commercial, Recreational, Scientific, or

Educational Purposes.

Unrestricted collecting for scientific or horticultural purposes

and excessive visits by individuals interested in seeing rare plants

could result from increased publicity. This is a potential threat to

all 24 of the taxa, but especially to Cyanea asarifolia, Delissea

rhytidosperma, Diellia pallida, Hibiscus clayi, Lipochaeta waimeaensis,

Melicope haupuensis, and Phyllostegia waimeae, each of which has only 1

or 2 populations and a total of 10 or fewer individuals. Collection of

whole plants or reproductive parts of any of these seven taxa could

cause an adverse impact on the gene pool and threaten the survival of

the taxa. Some taxa, such as Brighamia insignis, Exocarpos luteolus,

Hibiscus clayi, Nothocestrum peltatum, Peucedanum sandwicense, and

Solanum sandwicense, have populations close to trails or roads and are,

thus, easily accessible to collectors (HHP 1991a3, 1991f6, 1991h1,

1991t1, 1991t2, 1991t4, 1991t7, 1991u1, 1991u3, 1991u5, 1991u7,

1991u15, 1991z11, 1991z12, 1991z18, 1991z20, 1991z23, 1991z25, 1991z26;

HPCC 1990c, 1990i1 to 1990i4, 1990m).

Many of the plants occur in recreational areas used for hiking,

camping, and hunting. Tourism is a growing industry in Hawaii and as

more people seek recreational activities, they are more likely to come

into contact with rare native plants. People can transport or introduce

alien plants through seeds on their footwear and they can cause

erosion, trample plants, and start fires (Corn et al. 1979). Brighamia

insignis, Hibiscus clayi, and Peucedanum sandwicense have populations

next to trails and are considered to be immediately threatened by

recreational use of the areas in which they occur (Clarke and Cuddihy

1980; Takeuchi 1982; T. Flynn, pers. comm., 1991).

C. Disease or Predation

Browsing damage by goats has been verified for the following taxa:

Brighamia insignis; Exocarpos luteolus; Peucedanum sandwicense; and

Schiedea spergulina var. spergulina (HHP 1991y5; Takeuchi 1982; T.

Flynn, J. Lau, and S. Perlman, pers. comms., 1991; L. Mehrhoff, pers.

comm., 1993). The remaining taxa are not known to be unpalatable to

goats, deer, or cattle and, therefore, predation is a probable threat

where those animals have been reported, potentially affecting 15

additional taxa. These taxa include Delissea rhytidosperma, Diellia

pallida, Hedyotis cookiana, Hibiscus clayi, Lipochaeta fauriei,

Lipochaeta micrantha, Lipochaeta waimeaensis, Melicope haupuensis,

Melicope knudsenii, Melicope pallida, Munroidendron racemosum,

Nothocestrum peltatum, Phyllostegia waimeae, Pteralyxia kauaiensis, and

Solanum sandwicense. The lack of seedlings of many of the taxa and the

occurrence of individuals of several taxa only on inaccessible cliffs

appear to indicate the effect that browsing mammals, especially goats,

have had in restricting the distribution of these plants (HHP 1990b,

Takeuchi 1982).

Of the four species of rodents that have been introduced to the

Hawaiian Islands, the species with the greatest impact on the native

flora and fauna is probably the black or roof rat (Rattus rattus),

which now occurs on all the main Hawaiian Islands around human

habitations, in cultivated fields, and in dry to wet forests. Black

rats, and to a lesser extent house mouse (Mus musculus), Polynesian rat

(Rattus exulans), and Norway rat (R. norvegicus) eat the fruits of some

native plants, especially those with large, fleshy fruits. Many native

Hawaiian plants produce their fruit over an extended period of time and

this produces a prolonged food supply that supports rodent populations.

Black rats strip bark from some native plants (Cuddihy and Stone 1990,

Tomich 1986). Rats threaten Delissea rhytidosperma by damaging the

fruits and stems (Bruegmann 1990). Rats eat fruits of Exocarpos

luteolus, threatening this species as well. It is probable that rats

damage the fruit and stems of Cyanea asarifolia, Munroidendron

racemosum, and Pteralyxia kauaiensis, which have fleshy fruits and

populations in areas where rats occur (Lamoureux 1982; T. Flynn and D.

Herbst, pers. comms., 1991).

Black twig borer (Xylosandrus compactus) is a small beetle about

1.6 mm (0.06 in) in length that burrows into branches, introduces a

pathogenic fungus as food for its larvae, and lays its eggs. Twigs,

branches, and even the entire plant can be killed from such an

infestation. In the Hawaiian Islands, black twig borer has many hosts,

disperses easily, and is probably present at most elevations up to

2,500 ft (670 m). It is known to attack species of Melicope and is a

potential threat to Melicope haupuensis, M. knudsenii, M. pallida, and

M. quadrangularis, all of which grow in areas where the insect is

believed to be present (Davis 1970; Hara and Beardsley 1979; Hill 1987;

Medeiros et al. 1986; Samuelson 1981; S. Montgomery, pers. comm.,

1991).

D. The Inadequacy of Existing Regulatory Mechanisms

Hawaii's Endangered Species Act states, ``Any species of aquatic

life, wildlife, or land plant that has been determined to be an

endangered species pursuant to the [Federal] Endangered Species Act

shall be deemed to be an endangered species under the provisions of

this chapter * * *'' (HRS, sect. 195D-4(a)). Federal listing would

automatically invoke listing under Hawaii State law, which prohibits

taking of endangered plants in the State and encourages conservation by

State agencies (HRS, sect. 195D-4).

None of the 24 taxa in this final rule are listed by the State.

Twelve taxa have populations located on privately owned land. Two taxa,

Melicope quadrangularis and Schiedea spergulina var. leiopoda, are

found exclusively on private land. Peucedanum sandwicense is found on

City and County of Honolulu land and federally managed land, as well as

State land. At least one population of each taxon except Schiedea

spergulina var. leiopoda and Melicope quadrangularis occur on State

land. Eleven of the taxa are located in State parks, NARs, or the

seabird sanctuary, which have rules and regulations for the protection

of resources (DLNR 1981b; HRS, sects. 183D-4, 184-5, 195- 5, and 195-

8). One or more populations of each of the 24 tax a except Schiedea

spergulina var. leiopoda is located on land classified within

conservation districts and owned by the City and County of Honolulu,

the State of Hawaii, or private companies or individuals. Regardless of

the owner, lands within these districts are regarded as necessary for

the protection of endemic biological resources and for the maintenance,

enhancement, or conservation of natural resources. Activities permitted

in conservation districts are chosen by considering how best to make

multiple use of the land (HRS, sect. 205-2). Some uses, such as

maintaining animals for hunting, are based on policy decisions, while

others, such as preservation of endangered species, are mandated by

both Federal and State laws.

Requests for amendments to district boundaries or variances within

existing classifications can be made by government agencies and private

landowners (HRS, sect. 205-4). Before decisions about these requests

are made, the impact of the proposed reclassification on ``preservation

or maintenance of important natural systems or habitat'' (HRS, sects.

205-4, 205-17), as well as the maintenance of natural resources, is

required to be taken into account (HRS, sects. 205-2, 205-4). Before

any land use change proposed to occur on county or State lands and

funded, in part or in whole, by county or State funds, or would occur

within land classified as conservation district, an environmental

assessment is required to determine whether or not the environment

would be significantly affected (HRS, chapt. 343). If it is found that

an action would have a significant effect, preparation of a full

Environmental Impact Statement is required. Hawaii environmental policy

and, thus, approval of land use, is required by law to safeguard ``* *

* the State's unique natural environmental characteristics * * *''

(HRS, sect. 344-3(1)) and includes guidelines to ``Protect endangered

species of individual plants and animals * * *'' (HRS, sect. 344-

4(3)(A)). Federal listing, because it automatically invokes State

listing, would trigger operation of these other State regulations

protecting the plants.

State laws relating to the conservation of biological resources

allow for the acquisition of land as well as the development and

implementation of programs concerning the conservation of biological

resources (HRS, sect. 195D-5(a)). The State also may enter into

agreements with Federal agencies to administer and manage any area

required for the conservation, management, enhancement, or protection

of endangered species (HRS, sect. 195D-5(c)). If listing were to occur,

funds for these activities could be made available under section 6 of

the Federal Endangered Species Act (State Cooperative Agreements). The

DLNR is mandated to initiate changes in conservation district

boundaries to include ``the habitat of rare native species of flora and

fauna within the conservation district'' (HRS, sect. 195D-5.1).

Ten of the taxa are threatened by plants considered by the State of

Hawaii to be noxious weeds. The State has provisions for eradication

and control of noxious weeds on State and private land in conservation

districts and other areas (HRS, chapt. 152; DOA 1981, 1991). State and

Federal agencies have programs to locate, eradicate, and deter

marijuana cultivation, which is a potential threat to six taxa (HHP

1990c). Federal listing of these 24 plant taxa would reinforce and

supplement the protection available under the Hawaii State Endangered

Species Act and other laws. The Federal Endangered Species Act would

offer additional protection to these 24 taxa because, if they were to

be listed as endangered or threatened, it would be a violation of the

Federal Endangered Species Act for any person to remove, cut, dig up,

damage, or destroy any such plant in an area not under Federal

jurisdiction in knowing violation of State law or regulation or in the

course of any violation of a State criminal trespass law.

E. Other Natural or Manmade Factors Affecting Its Continued Existence

One or more of the almost 50 taxa of introduced plants threaten 21

of the 24 plant taxa in this rule and potentially threaten the other

three. The original native flora of Hawaii consisted of about 1,000

taxa, 89 percent of which were endemic. Of the total native and

naturalized Hawaiian flora of 1,817 taxa, 47 percent were introduced

from other parts of the world and nearly 100 taxa have become pests

(Smith 1985, Wagner et al. 1990). Naturalized, introduced taxa degrade

the Hawaiian landscape and compete with native plants for space, light,

water, and nutrients (Cuddihy and Stone, 1990; D. Lorence, pers. comm.,

1991). Some of these taxa were brought to Hawaii by various groups of

people, including the Polynesian immigrants, for food or cultural

reasons. Plantation owners, alarmed at the reduction of water resources

for their crops caused by the destruction of native forest cover by

grazing feral animals, supported the introduction of alien tree taxa

for reforestation. Ranchers intentionally introduced pasture grasses

and other taxa for agriculture and sometimes inadvertently introduced

weed seeds as well. Other plants were brought to Hawaii for their

potential horticultural value (Cuddihy and Stone 1990, Scott et al.

1986, Wenkam 1969).

A small tree, Acacia confusa (Formosa koa), was introduced to

Hawaii for reforestation purposes and is naturalized in dry to mesic,

disturbed habitats on most of the Hawaiian Islands (Smith 1985, Wagner

et al. 1990). Acacia mearnsii (black wattle) was introduced as a

cultivated plant and has naturalized on five islands in pastures and

dry to mesic forests (Wagner et al. 1990). It threatens Exocarpos

luteolus (T. Flynn, pers. comm., 1991). Two subshrubs in the genus

Ageratina have naturalized in the Hawaiian Islands and are classified

as noxious weeds by the State (Hawaii, Department of Agriculture (DOA)

1981). Ageratina adenophora (Maui pamakani), naturalized in dry areas

to wet forests on four islands, is also classified as a noxious weed by

the Federal government (7 CFR 360). It threatens Peucedanum sandwicense

(HHP 1991u16, Wagner et al. 1990). Ageratina riparia (Hamakua pamakani)

is naturalized in disturbed, dry to mesic areas and wet forest on four

islands and is a threat to Lysimachia filifolia, as well as to

Peucedanum sandwicense (HHP 1991u16; HPCC 1990g2, 1990j1, 1990j3;

Wagner et al. 1990). Ageratum conyzoides (maile hohono), an herb that

is a common weed in many areas of the main Hawaiian Islands, threatens

Brighamia insignis in some areas (HHP 1991a1; Wagner et al. 1990).

Although it is the official State tree of Hawaii, Aleurites

moluccana (kukui) is not a native Hawaiian plant. It was originally

native to Malesia. It was brought to Hawaii by the Polynesian

immigrants and is now a component of mesic valley ecosystems on all of

the main islands except Kahoolawe (Wagner et al. 1990). One or more

populations of Hibiscus clayi, Lipochaeta fauriei, Munroidendron

racemosum, and Pteralyxia kauaiensis grow in areas with kukui, which

may compete with these native species for space. Hibiscus clayi and

Lipochaeta fauriei do not grow under a dense canopy, so kukui could

prevent them from regenerating in an area. Munroidendron racemosum and

Pteralyxia kauaiensis, overstory trees in native forests, are displaced

when kukui dominates (HHP 1991h2, 1991s1, 1991s4, 1991s5, 1991s8,

1991s10, 1991s15, 1991w1, 1991w4, 1991w5; HPCC 1990d3; Lamoureux 1982;

T. Flynn, J. Lau, and S. Perlman, pers. comms., 1991). Araucaria

columnaris (columnar araucaria), planted in Hawaii for reforestation

and timber production and now found on all the main islands, is

reseeding and threatens Hibiscus clayi (Little and Skolman 1989; Neal

1965; D. Bates, pers. comm., 1991). Bidens pilosa (Spanish needle), an

annual herb naturalized on all the main Hawaiian Islands, is a threat

to Peucedanum sandwicense (Ganders and Nagata 1990, HHP 1991u15).

Classified as a noxious weed by the State of Hawaii, Clidemia hirta

(Koster's curse) is an aggressive shrub found in mesic to wet forests

on at least five islands in Hawaii (Almeda 1990, DOA 1981). It is a

threat to Melicope pallida and Peucedanum sandwicense. It is a

potential threat to Cyrtandra limahuliensis (HHP 1990c; T. Flynn and S.

Montgomery, pers. comms., 1991). Cordyline fruticosa (ti) is a shrub

that was brought to Hawaii by the Polynesian immigrants. Its original

range is unknown, but in Hawaii it is now naturalized on all the main

islands except Kahoolawe in hala forest and mesic valleys and forests,

sometimes forming dense stands (Wagner et al. 1990; J. Lau, pers.

comm., 1991). One or more populations of the following taxa compete for

space with ti: Delissea rhytidosperma; Hibiscus clayi; Lipochaeta

micrantha var. exigua; Lysimachia filifolia; Munroidendron racemosum;

and Pteralyxia kauaiensis (HHP 1991d2, 1991h1, 1991h2, 1991j1, 1991s1,

1991w7; HPCC 1990c, 1990e, 1990g2; J. Lau, T. Flynn, and S. Perlman,

pers. comms., 1991). Corynocarpus laevigatus (karakanut), a tree

introduced to Hawaii for reforestation, is now found on four islands

and is a threat to Exocarpos luteolus (Wagner et al. 1990; T. Flynn,

pers. comm., 1991).

Brought to Hawaii as a cultivated herbaceous plant, Erigeron

karvinskianus (daisy fleabane) is naturalized in wetter areas of four

islands (Wagner et al. 1990). Invasion by daisy fleabane threatens

Exocarpos luteolus, Lipochaeta micrantha var. micrantha, Melicope

pallida, Nothocestrum peltatum, and Peucedanum sandwicense (HHP 1991k1;

HPCC 1990f, 1990i4, 1990j2; T. Flynn and K. Wood, pers. comms., 1991).

Furcraea foetida (Mauritius hemp), a large rosette plant naturalized on

most islands in Hawaii on rocky ledges, slopes, and in pastures,

threatens Schiedea spergulina var. leiopoda (Wagner et al. 1990; T.

Flynn, pers. comm., 1991). Grevillea banksii (kahili flower),

considered a noxious weed by the State of Hawaii, was introduced as a

cultivated tree and has naturalized in disturbed, dry to wet forests on

most of the main Hawaiian Islands (DOA 1981, Wagner et al. 1990). It

threatens Cyrtandra limahuliensis (T. Flynn, pers. comm., 1991).

Grevillea robusta (silk oak) was extensively planted in Hawaii for

timber and is now naturalized on most of the main islands (Smith 1985,

Wagner et al. 1990). Silk oak threatens Lipochaeta waimeaensis and

Peucedanum sandwicense (HPCC 1990j1; S. Perlman, pers. comm., 1991).

Three species of Hedychium (ginger), native to the Himalayas and

surrounding areas, were brought to Hawaii as ornamentals and are now

naturalized in mesic or wet forests. Two of these species threaten one

or more of the 24 taxa in this rule. Their rhizomes produce rapid,

vegetative growth, forming dense ground cover that excludes other

plants. The Wainiha population of Cyrtandra limahuliensis is threatened

by H. flavescens (yellow ginger) (T. Flynn and K. Wood, pers. comms.,

1991). Hedychium gardnerianum (kahili ginger) produces red seeds that

are distributed by alien fruit-eating birds; it threatens Solanum

sandwicense (Cuddihy and Stone 1990; HPCC 1990m; Nagata 1990; Smith

1985; T. Flynn and K. Wood, pers. comms., 1991). Kalanchoe pinnata (air

plant) is an herb that occurs on all the main islands except Niihau and

Kahoolawe, especially in dry to mesic areas (Wagner et al. 1990).

Populations of Brighamia insignis and Peucedanum sandwicense are

threatened by competition with air plant (HHP 1991u15, Takeuchi 1982).

Lantana camara (lantana), brought to Hawaii as an ornamental plant,

is an aggressive, thicket-forming shrub that can now be found on all of

the main islands in mesic forests, dry shrublands, and other dry,

disturbed habitats (Wagner et al. 1990). One or more populations of

each of the following taxa are threatened by lantana: Brighamia

insignis; Delissea rhytidosperma; Diellia pallida; Hibiscus clayi;

Lipochaeta fauriei; both varieties of Lipochaeta micrantha; Melicope

haupuensis; Melicope knudsenii; Munroidendron racemosum; Nothocestrum

peltatum; Peucedanum sandwicense; Pteralyxia kauaiensis; and both

varieties of Schiedea spergulina (HHP 1991a1 to 1991a3, 1991e3, 1991i1,

1991j1, 1991k1, 1991o1, 1991p2 to 1991p4, 1991s1, 1991s5, 1991s11,

1991s15, 1991t7, 1991u1, 1991u3, 1991u5, 1991w4, 1991w7, 1991y5; HPCC

1990a, 1990d1, 1990d2, 1990e, 1990f, 1990k1, 1990k2; T. Flynn, R.

Hobdy, D. Lorence, and S. Perlman, pers. comms., 1991).

Leucaena leucocephala (koa haole), a shrub naturalized and

sometimes the dominant species in low elevation, dry, disturbed areas

on all of the main Hawaiian Islands, threatens the following plants:

Lipochaeta waimeaensis; Munroidendron racemosum; and Schiedea

spergulina var. leiopoda (Geesnick et al. 1990; HHP 1991s3; Lamoureux

1982; T. Flynn and S. Perlman, pers. comms., 1991). Lonicera japonica

(Japanese honeysuckle) is becoming naturalized in mesic to wet areas on

Kauai and Hawaii and threatens Solanum sandwicense Bruegmann 1990, HPCC

1990m, Wagner et al. 1990). Melastoma candidum, a small cultivated tree

that is now naturalized in mesic to wet areas of Kauai, threatens some

populations of Cyrtandra limahuliensis. Melia azedarach (Chinaberry), a

small tree widely cultivated and naturalized on most of the main

Hawaiian Islands, threatens Diellia pallida, Munroidendron racemosum,

and Schiedea spergulina var. spergulina (HHP 1991e3, 1991y5; HPCC

1990h; Wagner et al. 1990). The aggressive Myrica faya (firetree) has

become a dominant plant in many mesic to wet forests on five Hawaiian

Islands. Populations of Exocarpos luteolus, Munroidendron racemosum,

and Peucedanum sandwicense are threatened by firetree (HHP 1991u3; HPCC

1990h; S. Perlman, pers. comm., 1991). Opuntia ficus-indica (prickly

pear, panini) is a cactus found in dry, disturbed habitats on five

islands and poses a threat to Lipochaeta waimeaensis (Solomon 1990; S.

Perlman, pers. comm., 1991).

Passiflora edulis (passion fruit) is a woody vine that occurs on

five Hawaiian Islands in mesic forests and shrublands and threatens

Nothocestrum peltatum (Escobar 1990, HPCC 1990i3). Passiflora ligularis

(sweet granadilla) is a woody vine that now occurs in diverse mesic

forest and wet forest on four islands and threatens Delissea

rhytidosperma (Escobar 1990; S. Perlman, pers. comm., 1991). Passiflora

mollissima (banana poka), another woody vine, poses a serious problem

to mesic forests on Kauai and Hawaii by covering trees, reducing the

amount of light that reaches trees as well as understory. It causes

damage and death to trees by the weight of the vines. Animals,

especially feral pigs, eat the fruit and distribute the seeds (Cuddihy

and Stone 1990, Escobar 1990). Banana poka threatens Delissea

rhytidosperma, Nothocestrum peltatum, Peucedanum sandwicense,

Pteralyxia kauaiensis, and Solanum sandwicense (HHP 1991d1, 1991u5;

HPCC 1990i3, 1990m; D. Herbst, R. Hobdy, and J. Lau, pers. comms.,

1991). Pluchea carolinensis (sourbush), a shrub naturalized in dry,

coastal areas and mesic and wet forest on all of the main Hawaiian

Islands, threatens Lysimachia filifolia and Peucedanum sandwicense

(HPCC 1990g2; Wagner et al. 1990; R. Hobdy, pers. comm., 1991).

Two shrubs or small trees, Psidium cattleianum (strawberry guava)

and Psidium guajava (common guava) were brought to Hawaii and have

become widely naturalized on all the main islands, forming dense stands

in disturbed areas. Strawberry guava, found in mesic and wet forests,

develops into stands in which few other plants grow, physically

displacing natural vegetation and greatly affecting Hawaiian plants,

many of which are narrowly endemic taxa. Pigs use strawberry guava for

food and, in turn, disperse the plant's seeds through the forests

(Smith 1985, Wagner et al. 1990). Strawberry guava is considered to be

the greatest weed problem in Hawaiian rain forests and is known to pose

a threat to Brighamia insignis, Cyrtandra limahuliensis, Hibiscus

clayi, Lipochaeta fauriei, and Lipochaeta micrantha var. exigua, and it

is a potential threat to Melicope quadrangularis (HHP 1991a1; HPCC

1990c, 1990e; Smith 1985; T. Flynn, pers. comm., 1991; D. Lorence et

al., in litt., 1991). Common guava invades disturbed sites, forming

dense thickets in dry as well as mesic and wet forests (Smith 1985,

Wagner et al. 1990). Common guava threatens Brighamia insignis,

Cyrtandra limahuliensis, Hibiscus clayi, Lipochaeta fauriei, Melicope

pallida, Munroidendron racemosum, Peucedanum sandwicense, and

Pteralyxia kauaiensis (Lamoureux 1982; HHP 1991a1, 1991a4, 1991s1,

1991s4, 1991s5, 1991u3, 1991u16; HPCC 1990d1, 1990h; T. Flynn, R.

Hobdy, and J. Lau, pers. comms., 1991).

Rubus argutus (prickly Florida blackberry), an aggressive alien

species in disturbed mesic to wet forests and subalpine grasslands on

four islands, is considered a noxious weed by the State of Hawaii (DOA

1981, Smith 1985, Wagner et al. 1990). Prickly Florida blackberry

threatens Exocarpos luteolus, Melicope pallida, Melicope

quadrangularis, Nothocestrum peltatum, and Solanum sandwicense (HHP

1991z18, 1991z25; HPCC 1990i3, 1990i4, 1990m; T. Flynn, D. Herbst, R.

Hobdy, J. Lau, S. Perlman, and K. Wood, pers. comms., 1991). Schefflera

actinophylla (octopus tree), brought to Hawaii as a cultivated tree, is

shade tolerant and becomes established in undisturbed forests (Lowrey

1990, Smith 1985). It is now naturalized on at least four islands and

is a threat to Lysimachia filifolia as well as a potential threat to

Peucedanum sandwicense (HHP 1990c, HPCC 1990g2).

After escaping from cultivation, Schinus terebinthifolius

(Christmas berry) became naturalized on most of the main Hawaiian

Islands (Wagner et al. 1990). It threatens Hibiscus clayi and is a

potential threat to Peucedanum sandwicense (HHP 1990c, 1991h1; HPCC

1990j1, 1990j3; T. Flynn, pers. comm., 1991). Four species of the genus

Stachytarpheta have naturalized in the Hawaiian Islands, usually in

disturbed areas (Wagner et al. 1990). These alien herbs or subshrubs

threaten Brighamia insignis and Peucedanum sandwicense (HHP 1991a1,

HPCC 1990j1). Syzygium cumini (Java plum), a tree naturalized in

disturbed mesic forests on most of the main Hawaiian Islands, threatens

Brighamia insignis, Hibiscus clayi, Melicope quadrangularis, and

Peucedanum sandwicense (HHP 1991a1, 1991a2, 1991h1, 1991h2, 1991u1,

1991u3; HPCC 1990a; Wagner et al. 1990; K. Wood, pers. comm., 1991).

Triumfetta semitriloba (Sacramento bur) is a subshrub now found on four

Hawaiian Islands and considered to be a noxious weed by the State of

Hawaii (DOA 1981, Wagner et al. 1990). Populations of Munroidendron

racemosum and Schiedea spergulina var. spergulina are threatened by

Sacramento bur (HHP 1991y5, HPCC 1990h). Toona ciliata (Australian red

cedar), a tree now naturalized on four Hawaiian Islands, is quickly

spreading in forests of the Waianae Mountains on Oahu and threatens

Melicope pallida (Wagner et al. 1990; S. Montgomery, pers. comm.,

1991).

Several hundred species of grasses have been introduced to the

Hawaiian Islands, many for animal forage. Approximately 100 grass

species have become naturalized. Melinis minutiflora (molasses grass),

a perennial grass brought to Hawaii for cattle fodder, is now

naturalized in dry to mesic, disturbed areas on most of the main

Hawaiian Islands. Formed mats smother out other plants and fuel more

intense fires than would normally affect an area (Cuddihy and Stone

1990, O'Connor 1990, Smith 1985). Plants threatened by molasses grass

are Brighamia insignis, Lipochaeta fauriei, and populations of

Peucedanum sandwicense (HHP 1991a1, 1991a3, 1991u3; HPCC 1990a; R.

Hobdy and S. Perlman, pers. comms., 1991). Oplismenus hirtellus

(basketgrass) is a perennial grass that is naturalized in shaded mesic

valleys and forests and sometimes in wet forests on most of the main

Hawaiian Islands (O'Connor 1990). Diellia pallida, Hibiscus clayi, and

Lipochaeta fauriei are threatened by basketgrass (HHP 1991h1; HPCC

1990c, 1990d3; W.H. Wagner, pers comm., 1991). The perennial grass,

Paspalum conjugatum (Hilo grass), naturalized in moist to wet,

disturbed areas on most Hawaiian Islands, produces a dense ground

cover, even on poor soil, and threatens Cyrtandra limahuliensis and

Hibiscus clayi (Cuddihy and Stone 1990, O'Connor 1990, Smith 1985; T.

Flynn and R. Hobdy, pers. comms., 1991).

Pennisetum clandestinum (Kikuyu grass), an aggressive, perennial

grass introduced to Hawaii as a pasture grass, withstands trampling and

grazing and has naturalized on four Hawaiian Islands in dry to mesic

forest. It produces thick mats that choke out other plants and prevent

their seedlings from establishing and has been declared a noxious weed

by the U.S. Department of Agriculture (7 CFR 360) (DOA 1991, Medeiros

et al. 1986, O'Connor 1990, Smith 1985). Kikuyu grass threatens

Melicope knudsenii (R. Hobdy, pers. comm., 1991). Rhynchelytrum repens

(Natal redtop) is an annual or perennial grass that is naturalized in

disturbed, usually dry areas on all the main Hawaiian Islands and

threatens Lipochaeta waimeaensis (O'Connor 1990; S. Perlman, pers.

comm., 1991). Setaria gracilis (yellow foxtail), a perennial grass

naturalized in wet to dry, disturbed habitat on most of the main

Hawaiian Islands, threatens Brighamia insignis, Melicope haupuensis,

and Peucedanum sandwicense (HHP 1991a1, 1991a3, 1991o1, 1991u3;

O'Connor 1990). A perennial grass naturalized in disturbed areas on

most of the main Hawaiian Islands, Sporobolus africanus (smutgrass)

threatens Brighamia insignis and Peucedanum sandwicense (HHP 1991a1,

1991a3, 1991u15; O'Connor 1990).

Because Hawaiian plants were subjected to fire during their

evolution only in areas of volcanic activity and from occasional

lightning strikes, they are not adapted to recurring fire regimes and

are unable to recover well following a fire. Alien plants are often

better adapted to fire than native plant taxa and some fire-adapted

grasses have become widespread in Hawaii. The presence of such taxa in

Hawaiian ecosystems greatly increases the intensity, extent, and

frequency of fire. Fire-adapted alien taxa can reestablish in a burned

area, resulting in a reduction in the amount of native vegetation after

each fire. Fire is a serious, immediate threat along the Na Pali coast,

especially during drier months. Fires are caused by people pursuing

recreational activities and prevailing winds spread fires to inland

areas. Fire could destroy dormant seeds as well as plants, even on

steep cliffs (Clarke and Cuddihy 1980, Corn et al. 1979, Cuddihy and

Stone 1990). Fire is a threat to Brighamia insignis, Exocarpos

luteolus, Melicope pallida, Munroidendron racemosum, Nothocestrum

peltatum, Peucedanum sandwicense, Pteralyxia kauaiensis, and Solanum

sandwicense.

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