(DRAFT) - Taxonomy
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
TAXONOMY
NAME - SANDALWOOD, LANAI
OTHER COMMON NAMES - SANDALWOOD, LANAI; SANDALWOOD and LANA'I; 'ILIAHI;LA'AU'ALA
ELEMENT CODE -
CATEGORY - Angiosperm
PHYLUM AND SUBPHYLUM - MAGNOLIOPHYTA,
CLASS AND SUBCLASS - MAGNOLIOPSIDA,
ORDER AND SUBORDER - SANTALALES,
FAMILY AND SUBFAMILY - SANTALACEAE,
GENUS AND SUBGENUS - SANTALUM,
SPECIES AND SSP - FREYCINETIANUM, VAR.
SCIENTIFIC NAME - SANTALUM FREYCINETIANUM VAR.
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Lanai Sandalwood
Santalum freycinetianum var. lanaiense (Wagner) Rock
KINGDOM: Plant GROUP: Angiosperm
DIVISION: Magnoliophyta CLASS: Magnoliopsida
ORDER: Santalales FAMILY: Santalaceae
The Lana'i sandalwood is a small, gnarled tree with leaves that
vary from nearly round to 2.5 times as long as broad. The leaves are
dark green on the upper surface and brighter below, with red veins and
a short, pointed tip. The tree bears small clusters of bright red
flower, about 16 mm long. The fruit is ovoid, about 15 mm long,
fleshy, deep purple in color, with a single, hard pit (01,02,04,05).
Joseph F. Rock discovered this sandalwood on the island of Lana'i
in 1910, and formally described it in 1913. Historically, it has been
collected and/or reported from several widely spaced localities on the
island. Sandalwood trade prior to this plant's discovery by the
scientific community could have already reduced the number of trees by
an unknown amount. Thirty-nine individuals of the variety are now
known and are widely spaced over its range. These can be divided into
two populations, one near Kanepu'u and the other near the summit of
the island. Both populations occur on private lands owned by Castle
Taxonomy - 1 (DRAFT) - Taxonomy
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
and Cooke, Inc. (02).
The taxon was formally described by Joseph F. Rock in 1913 as a
variety of the endemic Hawaiian species, Santalum freycinetianum Gaud.
Rock 10061, collected in July of 1910 "on the highest ridge of
Lana'i", was designated the holotype and is in the herbarium of the
Bernice Pauahi Bishop Museum (01,04). Three years later Rock raised
the variety to the specific level, S. lanaiense (Rock) Rock (05).
Most botanist subsequently have followed this later treatment of the
taxon (06,07,09). Recent publications have recognized the taxon at
the varietal level (08,10,11). In 1980, Stemmermann (08) described a
new variety of sandalwood, S. freycinetianum var. auwahiense, from
Maui (Holotype: Auwahi Forest Reserve, 4000 ft. elev. L. Stemmermann
& J. Kjargaard 2149 BISH). This taxon is closely allied with both
S. haleakalae and S. freycinetianum and is synonymized under S.
freycinetianum var. lanaiense by Wagner, et al. (11), but this
taxonomic judgement has not yet been formally published. Variety
auwahiense was based on very minor differences in leaf shape and
inflorescence position. As these features do not consistently
separate populations from Maui and Lana'i, it was included under
variety lanaiense (11).
A line drawing is included in Stemmermann's paper (08) (as var.
auwahiense). Photographs are included in Carr's unpublished status
report for the taxon (01), and in Rock's monograph of the Hawaiian
sandalwoods (05). Distribution maps are included in the Carr status
report (01).
Additional common names or spelling variations of the common name
include: Lanai sandalwood, 'iliahi, and la'au'ala.
Taxonomy - 2 (DRAFT) - Status
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
STATUS
Coded Status
E: Federal Endangered
Commercial
Ornamental
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The Lana'i sandalwood (Santelum freycinetianum var. lanaiense)
has been designated as Endangered pursuant to the Endangered Species
Act of 1973 (50 CFR 17.12; P.L. 93-205, 87 Stat. 884; 16 U.S.C.
1531-1540), as amended. The Lana'i sandalwood has this status
wherever found including the State of Hawai'i.
This species is protected by the Lacey Act (P.L. 97-79, as
amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to possess
any wild plant (including roots, seeds, and other parts) within U.S.
territorial or special maritime jurisdiction (as defined in 18 U.S.C.
7); or to import, export, transport, sell, receive, acquire, or
purchase in interstate or foreign commerce any wild plant (including
roots, seeds, and other parts) taken, possessed, transported, or sold
in violation of any State law or regulation. It is also unlawful to
import, export, transport, sell, receive, acquire, or purchase any
wild plant (including roots, seeds, and other parts) taken or
possessed in violation of any U.S. law, treaty, or regulation or in
violation of Indian tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
STATE: Hawai'i
DESIGNATED STATUS: Endangered
ADMINISTRATIVE AGENCY: State Department of Land and Nat. Res.
STATE STATUTE: Hawai'i Revised Statutes, Chapter 195D.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
None.
Status - 1 (DRAFT) - Status
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
ECONOMIC STATUSES:
None currently. However, from about 1790 to 1840, the sale of
sandalwood to China was the first profitable export trade of Hawai'i.
The wood is valued for its fragrance and beauty and was used in making
incense and indecorative woodworking.
75/07/01:40 FR 27823/27924 - Notice of review
76/06/16:41 FR 24523/24572 - Proposed rule
78/04/26:43 FR 17909/17916 - Summary of comments on proposed rule
79/12/10:44 FR 70796/70797 - Withdrawal of proposed rule
80/12/15:45 FR 82480/82569 - Notice of review update
85/03/06:50 FR 09092/09095 - Proposed rule - Endangered
85/09/27:50 FR 39526/39527 - Notice of Review
86/01/24:51 FR 03182/03185 - Final rule - Endangered
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
LAND USE -
Evergreen Forest Land
Mixed Rangeland
COMMENTS ON HABITAT ASSOCIATIONS -
The Lana'i sandalwood occurs on Lana'i Island which is a
shield volcano built by eruptions at the summit and along three rift
zones. The summit later collapsed and partially filled to form the
area known today as the Palawai Basin. Beyond the basin the land
rises to form Lana'ihale, the highest point on the island (1,027 m or
3,370 ft). Water has carved Lana'ihale into narrow ridges and
steep-sided gullies, while wind has been a major force in shaping
local lowland topographies in historic times since the removal or
degradation of native vegetation. Apparently all the lavas of Lana'i
are tholeiitic basalts, ranging from olivine-free theoleiites through
olivine tholeiites to olivine-rich oceanites. Historically, the
Lana'i sandalwood ranged from the lower, gentle slopes of the island,
where it was a component of the dryland forest, to the wet summit of
Lana'ihale (01,12).
The soil type at the Kanepu'u site basically is a Kanepu'u silty
clay (Oxic Palenstalfs soil group of the order Alfisols) with the
surface layer varying from intact to partly removed by wind erosion.
There are 100 to 125 cm (40 to 50 in) of subsoil over bedrock. The
summit areas are mostly classified as Rough Mountainous Land with the
solid mantle very thin (2.5 - 25.5 cm or 1 - 10 in) over saprolite in
most locations. The soil that is present is of the Typic or Histic
Placaquepts soil groups of the order Inceptisols. Kahanui silty clay
occurs on the narrow ridgetop of Lana'ihale. It is of the Petroferric
Acrohumox soil group of the order Oxisols because of its many
ironstone fragments in its A horizon and the presence of a
discontinuous ironstone sheet in its B horizon. There is a cloud
cover over Lana'ihale during most of the afternoons and nights, and
the soil receives much of its moisture in the form of fog drip,
therefore the soil under the trees is usually poorly drained (13).
Lana'i has a subtropical climate. It is very dry and windy at
Kanepu'u, suffering drought conditions eight months of the year. Less
severe in the summit areas which experience more precipitation and
lower temperatures. The precipitation ranges from about 53 cm (21 in)
per year at Kanepu'u to about 94 cm (37 in) per year at the Lana'ihale
population. However, the effective precipitation amounts to 152 cm
(60 in) to 203 cm (80 in) annually at Lana'ihlae due to cloud cover
and fog drip (01,03,13,14).
The vegetation of the Kanepu'u site has been compared to that of
ridge vegetation on other islands. It is a dryland forest with
vegetation that is low in stature as is also that of Lana'ihale.
Windward branches tend to be stunted. The Kanepu'u forest is composed
of woody plants from two to eight m (6.5 - 26 ft) in height with
interlocking crowns and sparse ground cover. Although it does not fit
well, it is closest to the sclerophyllous forest, shrubland, and
grassland regional vegetation type according to the Kuchler system.
The dominant canopy species at Kanepu'u are Nestegis sandwicensis and
Planchonella spathulata, while Metrosideros polymorpha is the dominant
tree on Lana'ihale (01,03). The vegetation of the Lana'ihale site is
a wet forest vegetation most closely fitting the 'ohi'a lehua forest
type of the Kuchler system.
Habitat Associations - 1 Specific environmental requirements, if any, of this taxon are
unknown.
Habitat Associations - 2 (DRAFT) - Food Habits
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
FOOD HABITS
TROPHIC LEVEL -
AUTOTROPH
Food Habits - 1 (DRAFT) - Environment Associations
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Coastal Features: Vegetated offshore islands
G
G
G
Environment Associations - 1 (DRAFT) - Life History
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
LIFE HISTORY
HABIT:
The Lana'i sandalwood is a tree (05).
LIFE CYCLE:
Perennial (16).
TYPE OF REPRODUCTION:
Sexual (16). Other varieties of this species are known to
reproduce through root suckers, however, this has not been observed in
the Lana'i variety (16).
REPRODUCTIVE PHENOLOGY:
Little is known about the reproductive phenology of the Lana'i
sandalwood. Leafing is not seasonal because this tree is an
evergreen. Fruiting probably depends upon seasonal rains (16).
SEX OR SPORE STATUS:
Monoclinous (01).
POLLINATION, SPORE AND SEED DISSEMINATION:
Pollen and seed dissemination is unknown (01,16).
SEED BIOLOGY:
Unknown. Some trees have been observed to produce abundant
seeds. One lot of more than 100 seeds was germinated in a greenhouse
with intermittent misting; after about one year's time, over 50
percent of the seeds germinated (01).
POPULATION BIOLOGY:
The taxon is presently represented mostly by widely spaced
individuals showing virtually no signs of reproduction. The fruit is
usually destroyed by rats before falling from the tree. Only a single
sapling is known and it occurs at the Kanepu'u site. It is difficult
to define populations of this taxon since a total of only 39 mostly
widely scattered individuals are known. Perhaps the best known
"population" is the one at Kanepu'u where five individuals are known;
locations of four of these are plotted on Spence and Montogomery's map
(03). They are widely spaced in a triangular patch of about 12 ha
(30 ac) of dry forest. About 9.5 km (6 mi) away is the nearest
"population" of four individuals on Kaiholena Ridge. Six individuals
are known from a pie-shaped section of ridges extending windward from
the summit region between Maunalei and Hauola gulches, an area of
about 525 ha (1300 ac); the remaining 24 known individuals are in an
extensive pie-shaped wedge of ridges extending windward from the
summit area between Hauola and 'Awehe gulches (about 810 ha or
2000 ac) (01).
ECOLOGICAL/EDAPHIC FACTORS:
Specific habitat and soil requirements, if any, of this taxon are
unknown. The soil type at the Kanepu'u site basically is a Kanepu'u
silty clay (Oxic Palenstalfs soil group of the order Alfisols) with
Life History - 1 (DRAFT) - Life History
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
the surface layer varying from intact to partly removed by wind
erosion. There are 100 to 125 cm (40 to 50 in) of subsoil over
bedrock. The summit areas are mostly classified as Rough Mountainous
Land with the soil mantle very thin (2.5 - 25.5 cm or 1 - 10 in) over
saprolite in most locations. The soil that is present is of the Typic
or Histic Placaquepts soil groups of the order Inceptisols. Kahanui
silty clay occurs on the narrow ridgetop of Lana'ihale. It is of the
Petroferric Acrohumox soil group of the order Oxisols because of its
many ironstone fragments in its A horizon and the presence of a
discontinuous ironstone sheet in its B horizon. There is a cloud
cover over Lana'ihale during most of the afternoons and nights, and
the soil receives much of its moisture in the form of fog drip,
therefore the soil under the trees is usually poorly drained (13).
Lana'i Island is a shield volcano built by eruptions at the
summit and along three rift zones. The summit later collapsed and
partially filled to form the area known today as the Palawai Basin.
Beyond the basin, the land rises to form Lana'ihale, the highest point
on the island (1,027 m or 3,370 ft). Water has carved Lana'ihale into
narrow ridges and steep-sided gullies, while wind has been a major
force in shaping local lowland topographies in historic times since
the removal or degradation of native vegetation. Apparently all the
lavas of Lana'i are tholeiitic basalts, ranging from olivine-free
theoleiites through olivine tholeiites to olivine-rich oceanites.
Historically, the Lana'i sandalwood ranged from the lower, gentle
slopes of the island, where it was a component of the dryland forest,
to the wet summit of Lana'ihale (01,12).
Lana'i has a subtropical climate. It is very dry and windy at
Kanepu'u, suffering drought conditions eight months of the year. Less
severe are the summit areas which experience more precipitation and
lower temperatures. The precipitation ranges from about 53 cm (21 in)
per year at Kanepu'u to about 94 cm (37 in) at Lana'ihale. Due to
cloud cover and fog drip, the effective precipitation at Lana'ihale is
actually between 152 cm (60 in) to 203 cm (80 in) annually. The mean
annual soil temperature is approximately 21 degrees Celsius (70 F) and
16.6 degrees Celsius (62 F) for the Kanepu'u and Kahanui (Lana'ihale)
soil types, respectively (13).
TROPHIC STATUS:
Phototrophic and parasitic (15). Apparently hemiparasitic on the
roots of other plants. It is able to manufacture its own food by
photosynthesis, but obtains water and minerals from other plants
through its root haustoria (15).
CHARACTERISTIC DOMINANCE:
Component (01).
COMMUNITY ECOLOGY:
The vegetation at the Kanepu'u site is similar to the ridge
vegetation on other islands. Due to the dryness of the area, and
strong winds, the vegetation is low in stature and the windward
branches tend to be stunted. It is a dryland forest dominated by
Nestegis sandwicensis and Planchonella spathulata up to about 8 m
(26 ft) tall, with interlocking crowns. The ground cover is sparse
Life History - 2 (DRAFT) - Life History
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
and is comprised primarily of exotic annual herbs. Although it does
not fit well, it is closest to the sclerophyllous forest, shrubland,
and grassland regional vegetation type according to the Kuchler
system.
The vegetation of the Lana'ihale site is a wet forest vegetation
most closely fitting the 'ohi'a lehua forest type of the Kuchler
system. It is a wet forest, receiving 152 cm (60 in) to 203 cm
(80 in) of precipitation a year, about half of which is from cloud or
fog drip. The forest is of mixed native and exotic species with
Metrosideros polymorpha dominant (01). Community stability and
successional trends in the vegetation of this island have not been
studied.
SPECIES INTERRELATIONSHIPS:
Sandalwoods are hemiparasites; they are able to manufacture their
own food and other substances by photosynthetic processes, but must
obtain water and minerals at least partly from other plants through
root haustoria. They are not host specific (15).
OTHER LIFE HISTORY DESCRIPTORS:
None.
Life History - 3 (DRAFT) - Management Practices
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Suppressing wildfire
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Land Acquisition
Beneficial Maintaining Sperm/Seed Banks
Beneficial Stocking captive-reared wild-strain animals
Beneficial Controlling/Restricting Noncommercial Harvest
Beneficial Controlling/Removing Exotic Vertebrates
Beneficial Controlling/Removing Feral Animals
Adverse Commercial Exploitation
Existing Commercial Exploitation
Adverse Low Gene Pool
Existing Low Gene Pool
Adverse Predation
Existing Predation
Adverse Exotic/Feral/Introducted Species
Existing Exotic/Feral/Introducted Species
Adverse Erosion
Existing Erosion
Adverse Grazing
Existing Grazing
Adverse
Existing
Adverse Vegetation Composition Changes
Existing Vegetation Composition Changes
Adverse Fire
Existing Fire
Adverse Forest Alteration
Existing Forest Alteration
Adverse Harvesting
Existing Harvesting
COMMENTS ON MANAGEMENT PRACTICES -
The main factors which have contributed to the current status
of the Lana'i sandalwood are habitat alteration, grazing and browsing
by feral animals, rodent predation on fruit, and the potential threats
of taking and fire. The primary factor is the loss of habitat.
Natural vegetation has been eliminated over vast areas of Lana'i and
native dryland forests have been severely degraded. Agricultural
development has removed large tracts of native vegetation, first for
pasture and later for pineapple production. Cattle, sheep, and axis
deer, which have been introduced into the island have removed and
trampled vegetation, contributing to severe erosion of soil (01,02,
03).
Extensive removal of Hawaiian sandalwoods for trade occurred from
about 1790 to 1840. The wood is valued for its fragrance and beauty
and was used in making incense and in decorative woodworking.
Although the species is no longer common enough for profitable
commercial use, it may be threatened by individuals seeking its wood
Management Practices - 1 (DRAFT) - Management Practices
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
(02).
Other factors include rodent damage to fruit, and fire,
particularly in the dry Kanepu'u area. The low numbers of individuals
remaining has greatly reduced the gene pool for the taxon, and may
threaten its adaptive potential (02).
A resort is being planned for the Manele area of Lana'i.
Although there are no sandalwood trees known from this area, it may be
the first step in the development of the island. The potential
threats of resort development and recreation or urbanization
potentially could become important factors in the future
UNAPPROVED PLAN:
A recovery plan for this species has not yet been initiated. No
target date has been set for its inititation.
The prime objective of any management action would be to
perpetuate the natural populations of the Lana'i sandalwood from the
threats of trampling, grazing, and browsing by feral animals; from
fruit predation by rats; from the potential threats of taking and
fire; and from other detrimental factors, such as resort development
and recreation or urbanization, that may become important in the
future. This can be accomplished by:
(1) acquiring management rights for lands essential to the
continued existence of the tree;
(2) fence the areas and remove exotic and feral animals;
(3) control rodent predators;
(4) build a fire break around the Kanepu'u population, or
replace the existing vegetation with plants which are more
fire resistant or which provide less fuel;
(5) monitor the populations;
(6) propagate the taxon and replant within its historic range;
and
(7) store seeds in a seed bank.
A portion of the remnant dry-land forest, including most of the
sandalwood trees at Kanepu'u were fenced several years ago. Most of
the wild goats were hunted from the island, however, deer and other
feral animals still remain. No other recovery action is presently
ongoing.
Management Practices - 2 (DRAFT) - References
Species SANDALWOOD, LANAI
Species Id ESIS701044
Date 13 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 Carr, G.D. 1981. Status report on Santalum freycinetianum var.
lanaiense Rock. U.S. Fish and Wildl. Serv., Unpubl. rept. 34 pp.
02 U.S. Fish and Wildlife Service. 1986. Endangered and threatened
wildlife and plants; determination of endangered status for
Santalum freycinetianum var. lanaiense (Lanai sandalwood or
'iliahi). Fed. Reg. 51:3182-3185.
03 Spence, G.E. and S. Montgomery. 1976. Ecology of the dryland
forest at Kanepu'u, island of Lana'i. Hawaiian Bot. Soc.
Newletter 15:62-80.
04 Rock, J.F. 1913. The indigenous trees of the Hawaiian Islands.
Honolulu. Privately publ. 518 pp.
05 Rock, J.F. 1916. The sandalwoods of Hawaii. Board of Agri. and
For., Territory of HI Bot. Bull. No. 3
06 Skottsberg, C. 1927. Artemisia, Scaevola, Santalum, and Vaccinium
of Hawaii. B.P. Bishop Mus. Bull. 43 89 pp.
07 Degener, O. 1940. Fl. Hawaiiensis, fam. 100. Santalaceae;
Santalum Publ. priv., 2 pp.
08 Stemmermann, L. 1980. Observations on the genus Santalum
(Santalaceae) in Hawai'i. Pac. Sci. 34:41-54.
09 St. John, H. 1973. List and summary of the flowering plants in
the Hawaiian Islands. Pac. Trop. Bot. Gard. Memoir No. 1. 519 pp.
10 Stemmermann, L. 1980. Vegetative anatomy of the Hawaiian species
of Santalum (Santalaceae). Pac. Sci. 34:55-75.
11 Wagner, W.L., D.R. Herbst and S.H. Sohmer. (n.d.) Manual of the
flowering plants of Hawai'i. B.P. Bishop Mus. Accepted
12 Macdonald, G.A. and A.T. Abbott. 1970. Volcanoes in the sea.
Univ. Press of HI, Honolulu. 441 pp.
13 Foot, D.E., et al. 1072. Soil survey of the islands of Kauai,
Oahu, Maui, Molokai and Lanai, State of Hawaii. USDA.
14 Taliaferro, W.J. 1959. Rainfall of the Hawaiian Islands.
Honolulu: Hawaii Water Authority.
15 Heywood, V.H. 1978. Flowering plants of the world. Mayflower
Books, NY. 336 pp.
16 Herbst, D.R. 1986. Pers. knowledge. U.S. Fish and Wildl. Serv.,
Honolulu, HI.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 Carr, G.D. 1981. Status report on Santalum freycinetianum var.
lanaiense Rock. U.S. Fish and Wildl. Serv., Unpubl. rep. 34 pp.
02 U.S. Fish and Wildlife Service. 1986. Endangered and threatened
wildlife and plants; determination of endangered status for
Santalum freycinetianum var. lanaiense (Lanai sandalwood or
'iliahi). Fed. Reg. 51:3182-3185.
03 Spence, G.E. and S. Montgomery. 1976. Ecology of the dryland
forest at Kanepu'u, island of Lana'i. Hawaiian Bot. Soc.
Newletter 15:62-80.
References - 1