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Elevational Relationships of Introduced
Woody Species on the Islands of Lanai
and Hawaii
Jack D. Brotherson
..:
~ "
(Carlquist 1980). Much of the angiosperm flora evolved in
place, making the Hawaiian archipelago an unparalleled
example of insular evolution (Wagner and Funk 1995).
The islands of the Hawaiian archipelago, like most islands
in the Pacific, have experienced major perturbations oftheir
biota, resulting in loss of species and ecosystem stability.
The conversion of the natural ecosystems in Hawaii, often
into actively manipulated cultural landscapes, began with
Hawaii's colonization by the Polynesians about 400 A.D.
(Kirch 1982; Cuddihy and Stone 1990). Captain James Cook
came to the islands in '1778. In the 200 years following that
first European visit to Hawaii, human intruders have inundated the islands' fragile ecosystems with some 5,000 introduced plant species, and 3,000 insect and animal species. In
time, these aliens have displaced hundreds of native species.
For example, Dolan" (1990) states that, ''much of native
Hawaii-its lush rain forests and noisy, colorful birds that
inhabit them-is gone forever, and its remnants are hurling
toward some oblivion in the worst extinction crisis in the
nation." The change in plant communities has been caused
by several major factors: (1) the introduction oflarge herbivorous ungulates; (2) the introduction of exotic plants;
(3) the burning of the vegetation in the lowland areas of the
islands and the concurrent expansion of agriculture; and
(4) commercial logging oflowland forests for firewood, sandalwood (Santalum sp.), and Koa (Acacia koa) (Tomich 1969;
Kirch 1982; Cuddihy and Stone 1990).
The alteration of the native vegetation in the Hawaiian
Islands has progressed upslope from sea level since settlement by the Polynesians, but has been much accelerated
since the coming of the Europeans in 1778. Estimates are
that 10 percent of the native Hawaiian flora has gone extinct
with another 40-50 percent being threatened with extinction
(Cuddihy and Stone 1990). Particularly impacted are the dry
and mesic ecosystems below 3,000 ft (914 m) where invasion
by alien grasses have resulted in an increase in the frequency, size, and intensity of fire (Muller-Dombois and
others 1981).
Because of the enormous human impact on the native
Hawaiian ecosystems and their corresponding floristic elements, it becomes evident and important to establish baseline
data. Stone and Stone (1989) suggest that available time and
resources be focused on field studies and analyses on native
Hawaiian plants and their associated ecosystems, to provide
data on the biology, evolution and ecosystem relationships of
important native taxa. Such data will contribute to developing conservation and management strategies of these highly
impacted Hawaiian ecosystems. Management of native areas in Hawaii is possible, although expensive, and an added
benefit of basic field studies is that knowledge gained will
Abstract-The extreme isolation of the Hawaiian island chain has
produced a highly unusual native island flora. Scientists estimate
the native flora to consist of about 1,000 species of flowering plants,
89 percent of which are endemic. The native plant communities
have been heavily impacted by the introduction of alien animal and
plant species thought to be in excess of 5,000 taxa. Especially
impacted have been the land areas" below 1,500 ft. elevation where
the native plant communities have been almost totally decimated .
These introductions are partitioning the available niches in ways
uncharacteristic to their native habitat. An assessment of the
patterns of colonization and distribution of several introduced
woody species onto the islands of Lanai and Hawaii shows the
invading species to have adapted to moisture regimes and distributed themselves in almost perfect bell-shaped curves along the
entire elevational gradient. The areas of colonization stretch across
the lower slopes of three shield volcanos from sea level to 3,000 ft.
(914 m) elevation.
The Hawaiian archipelago is one of the most isolated
island chains in the world. It is 2,390 miles (3,854 km) from
California, 2,300 miles (3,710 km) from Alaska, 2,400 miles
(3,871 km) from the Marquesas Islands, and 3,850 miles
(6,210 km) from Japan. The island system comprises a chain
of about 132 islands, reefs and shoals of volcanic origin in the
mid-Pacific ocean (Annstrong 1993). The islands are thought
to have formed successively over a fixed ''hot spot" beneath
the northwestward moving Pacific tectonic plate (Carson
and Clague 1995). Because of the extreme isolation of the
islands, they support one of the most unusual and remarkable oceanic island biota's in the world (Carlquist 1980'
Carlquist 1995). For example, Wagner and Funk (1995)
state that endemism in this biological array ranges from 50
percent for the mosses, 89 percent for angiosperms, and 99
percent for insects. The colonization of these islands by plant
species is hypothesized to have occurred in a variety of ways
(Carlquist 1980) and from multiple sources (Funk and Wagner
1995). It is thought that the approximately 1,000 angiosperm
species originated from about 235 original introductions,
these coming in at the rate of about one per 100,000 years
I?: Barrow, Jerry R.; McArthur, E. Durant; Sosebee, Ronald E.; Tausch,
Robm J., comps. 1996. Proceedings: shrubland ecosystem dynamics in a
changing environment; 1995 May 23-25; Las Cruces, NM. Gen. Tech. Rep.
INT-GTR-33~. Ogden, UT: U.S. Department of Agriculture, Forest Service,
Intermountam Research Station.
Jack D. Brotherson is Professor of Botany and Range Science, Department
of Botany and Range Science, 435 WIDB Brigham Young University Provo
UT 84602.
'
,
,
36
contribute to our ability to manage highly' invaded island
ecosystems, and will ultimately prove useful for conservation of both island and continental systems (Loope and
others 1988). Also, those ecosystems that have been damaged or degraded by alien introductions and past land use
may have the "capacity for significant recovery" if the agents
of disturbance are understood and controlled (J okobi and
Scott 1985).
To establish baseline data on conditions existent in the
vegetative communities between sea level and upland areas
(3,000 ft/914 m elevation), the colonization and distribution
patterns of several introduced woody species were studied.
The purpose was to determine alien tree and shrub species
distribution and to correlate those patterns with elevation
and precipitation patterns.
Methods
centered on the study site, using abundance classes as follows:
1 =rare; cover small, less than 1 percent; 2 =scarce; cover small,
less than 5 percent; 3 =infrequent; cover between 5-25 percent;
4 = frequent; cover between 25-50 percent; 5 = prevalent; cover
between 50-75 percent; 6 = abundant; cover between 75-100
percent. Average cover was computed for each species encountered by site and then compared with the abundance
estimates of the same site as a check of that species importance at that site. Elevation was determined at each site
using a Thommen TX mechanicallbarometer altimeter. Precipitation was estimated for each site from island maps with
isohyetal estimation lines.
Results
-----------------------------------
Fourteen species of woody plants with origins from 11
different geographical areas of the world (table 1) were
encountered on the study sites located along the three
transects. All have origins in tropical or sub-tropical climates and with one exception, were introduced to Hawaii
before 1900. Areas colonized by these species have been
altered to such an extent that only one indigenous taxon and
no native endemic taxa were encountered at the 65 study
sites. Numbers of species encountered at each study site
varied from 2 to 9, averaging 5.7 per site.
All species were not present on each of the three transects
(figs. 1-3), but when a single species occurred on more than
one transect, its relative importance varied from transect to
transect. For example, Opuntia ficus-indica was found only
on the Saddle Road, and Psidium guajava and Schinus
terebenthifolia showed greater importance on Hawaii than
on Lanai. Conversely, Leucaena leucophala showed greater
importance on Lanai. The individual species showed changing patterns of importance with respect to the elevational
gradient. For example, Schinus terebenthifolia was important at the higher elevations, Leucaena leucophala and
Lantana camara were important at mid-elevations, and
Prosopis pallida and Waltheria indica were important at or
near sea level. The individual plant species placement along
the elevational gradient in relationship to other species was
highly predictable. For example, Prosopis pallida was always found along the sea coast and below Lantana camara
along the gradient which was always below Schinus
terebenthifolia. These patterns would suggest colonization,
and subsequent success by invading species are random
with respect to location, but controlled by evolutionary
history as to the species interaction with the environment.
Species distribution patterns show that while some are
narrowly restricted in distribution to a single elevational
level (figs. 1-3), the majority peak in importance at specific
points along the gradient and then decline as elevation
either increases or decreases. Where the elevational gradient and the corresponding moisture gradient change gradually, as they do on all three of the transects, the species sort
along the gradient where they form nearly perfect ''bellshaped" curves relative to their patterns of distribution. The
elevational distribution ofthe individual plant species growing along the three gradients was generally similar in
position with respect to all other species. However, an
individual species relative position with respect to elevation
varied some from transect to transect. For example, Indigofera
----------------------------------
Three transects (65 study sites) were placed below 4,200 it
(1,280 m) on the lower slopes of three volcanos (Lanai,
Mauna Kea, Mauna Loa) on the islands of Lanai and Hawaii.
Two were located on Hawaii and one on Lanai. Those on
Hawaii were located at South Point, the southern most point
ofthe island, and along the saddle road about 20 miles (32 km)
north of the town of Kailua-Kona. The Lanai transect was
located on the slopes of the old shield volcano about
5 miles (8 km) northwest of Lanai City. Transect placement
was in the more arid areas on the volcanos leeward sides
where the climate tends to be sunny and dry. Transects and
study sites were positioned on the shield volcano's lower
slopes where little or no erosion has taken place and where
the precipitation gradient mimics the elevation gradient,
increasing gradually as elevation increases. Transects began at sea level and extended upslope. Study site number
and placement varied depending on transect length and
steepness of the elevation and on associated precipitation
gradients.
The South Point transect contained 20 study sites placed
at 100 foot (30.5 m) elevational increments, was 11 miles (18
km) long, and ranged from 0 to 1,900 ft (0 to 579 m) in
elevation. The Saddle Road transect contained 22 study
sites placed at 200 foot (61 m) elevational increments, was 22
miles (35 km) long, and ranged from 0 to 4,200 it (0 to 1,280 m)
elevation. The Lanai transect contained 20 study sites placed
at 100 foot (30.5 m) elevation increments, was 7 miles (11 km)
long, and ranged from 0 to 1,900 it (0 to 579 m) in elevation.
The steepness ofthe elevational gradient along each transect
varied. Precipitation patterns also varied ranging from 10 to
50 inches (256 to 1,282 mm) along the elevational gradient
at South Point, and from 10 to 40 inches (256 to 1,026 mm)
on the Saddle Road and Lanai transects.
Data collection was accomplished using two 100 yard (91
m) transects at each study site, one placed perpendicular to
the elevation gradient and one placed parallel to the elevation gradient. These two transects intersected at a cross at
their 50 yard (46 m) midpoints. Each site was subsampled
with 20, one m 2 quadrats placed every 10 yards (9 m) along
the transects. Canopy coverage by species was estimated
according to Daubenmire (1959). Abundance of each species
was ocularly estimated at each study site along a 440 yard
(400 m) walking transect, set parallel to the slope and
37
Table 1-Woody species encountered along the three transects on the lower slopes of the shield
volcanos-Lanai, Mouna Kea, and Mouna Loa. Species' common names (in parenthesesHawaiian names underlined), origins, and dates of introduction (where known) are
included. All species are alien introductions having naturalized in Hawaii, except Sida fa/ax
which is indigenous.
Species
Origin
Introduction date1
Agave sisalana (AGSI)
(Sisal, Sisal hemp, Malina)
Yucatan, Mexico
1893
Casuarina equisetifolia (CAEQ)
(Common ironwood, faina)
Australia
1882
Chamaecrista nictitans (CHNI)
(Partridge tea, l..au.Ki)
Neotropics
1871
Indigofera suffruticosa (INSU)
(Indigo bush,1nik2,lniJsQ.a, KQLu)
Java
1836
Lantana camara (LACA)
(Lanatana, J...alsana, J...a:au.IsalaIsaI.a,
Lanakana, Mikinolia bibiY)
West Indies
1858
Leucaena leucocephala (LELE)
Neotropics
1837
Opuntia ficus-indica (OPFI)
(Prickly pear, Eanini,~)
Mexico
Before 1809
Prosopis pallida (PRPA)
(Algaroba, Mesquite, ~
Peru, Colombia, Equador
1828
Psidium guajava (PSGU)
(Common guava, ~
Neotropics
Early 1800's
Schinus terebenthifolia (SCTE)
(Christmas berry, ~, Nanjohilg)
Brazil
Before 1911
Sida falax (SIFA)
Pacific islands to China
?
Africa
1895
Mexico
Before 1871
Pantropical
Before 1779
(KQa~, ~,
J..ill.IsQg)
KY.awa
~,~UHnl,~)
(llimA)
Solanum linnaeanum (SOLI)
(Apple of Sodom, E.QJlQlQ. ~)
Verbesina encelioides(VEEN)
(Golden crown-beard)
Waltheria indica (WAIN)
O.lb.a/Qa, :AIa:ala 12M lQa, Hala'uhaloa,
tiIa/Qa, Kanakalga)
'Mostly from Wagner and others 1990.
suffruticosa peaked in importance at 400 ft (121 m) elevation
on the Saddle Road transect and at 900 ft (274m) on the
South Point transect. Similarly, Lantana camara peaked in
importance at around 750 ft (229 m) on South Point but was
most important at 1,500 ft (457 m) on Lanai.
When data from the three transects were combined (fig. 4),
the ecological relationships of the different species with
respect to each other and to elevation became much more
difficult to delineate. As shown in figure 4, the species
distribution curves that were developed from the combined
data intersect the elevation gradient across a much broader
segment of the gradient. For example, Sida faZax intersects
the elevation gradient from 200 to 700 ft (61 to 213 m) on
South Point gradient, while on the basis of the combined
data the species intersects the elevational gradient from 100
to 3,100 ft (33 to 945 m), a 6 times greater range of distribution elevationally. When all species are considered in this
analysis, the mean intersect of a species on the combined
transect gradient is significantly greater (p < 0.01), on
average of2.4 times greater, than the mean species range of
intersect on the individual transects.
Discussion and Conclusions ---The terrestrial flora of the Hawaiian Islands along with
aspects of its ecology and conservation has been well studied
(Carlquist 1970, 1974, 1980; Kay 1972; St. John 1973; Wallace
1973; Williamson 1981; Muller-Dumbois and others 1981;
Stone and Stone 1989; Stone and Scott 1985; Whiteaker 1983;
Cuddihy and Stone 1990; Wagner and others 1990; Stone and
others 1992; Wagner and Funk 1995). The island flora, as has
been stated previously, is highly endemic, and several ideas
have been proposed concerning its origin. Historically (precolonization by man), the majority of the species appear to
have had their origins in southeast Asia. The Carolina and
Marshall Islands of Micronesia appear to have acted as
bridges for the migration of plants from Asia to Hawaii. Only
about 20 percent of the native vascular plants appear related
to plant groups of North and South America (Carlquist 1970,
1980). The Hawaiian flora is most endemic in the more
advanced elements (i.e. 89 percent of the angiosperm species
are endemic) and less so in the ferns (65 percent), liverworts
(75 percent), mosses (65 percent), and lichens (38 percent).
38
·.....................................................······.·················_·•••I~.
OPFI
SCTE
r:::::::::::::::::::::
[::::::::::::::::::::::
................................
...................................................................................................
PSGU
........................................................................................
.........................................................................................
LELE
::::::::::::::::::C:::::::9.~~~:::::::::E::::::::::::::
F::::::::::::::::::::
~
·····::::::::::::::::::::::·····:::::::::::II--_
o
~
o
c
o
f::::::::::::·...·
SOLI
......................................................................................................."............... .
L_EL_E
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. . . . .1. . .•• •••• ••••• ............................... .
o
, ·...... ······ .... ·.. ·· .. ·· ....·..·.... ····· .. ····......1.············....................................
CD
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o
o
0)
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::::I::::,~~~::~::I·:·::·ili·:::·:.:::··:··:::::·::
.
[
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:
.
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.
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.
:
.
:
[
:
.
:
:
.
:
.
. . . . . . . . . . . . . . . . . .=. . . . . . . .,. . . . . . . . . . . . . . . . . . . . . ..
Sc
SIFA
~
~
o
c
................................
o
o ' ....................................
~................
.........
1:::::::
CD
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..... .............................................
.;;;;
,;,:; ................................. ..
CD
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::::::::::::::::::::::::::r··········iNSU··········l:::::::::::::::::::::::::::::::
ftI
C
.... ~ •••••••••••••••••••• 0 •••••• e . . . . . . . . . . . . a.a . . . . . . . . . . . . . . . . .
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:.
::::::::::::::::::::::::T..........~~~~·.....·· ..·..[:::::::::::::::
l..--.:..CH.-;,;,N...:..:..I_-.Jr
100
.......,.··1.···················.. ····· .. ·· .. ········· .. ·· .. ·· ....................................
....
1,000
3,000
Figure 2-The distribution of woody species along
the elevation/moisture gradient along the Saddle
Road transect on the island of Hawaii. See table 1 for
species symbols .
.................................................~~~~............E::::::::::::::::::::::::::::
500
2,000
Elevation (ft)
'--_W_A_IN_----JI:::::::::::::::::::::::::::::::::::::::::·
o
1000
2,000
Elevation (ft)
Figure 1-The distribution of woody species along
the elevation/moisture gradient along the South
Point transect on the island of Hawaii. See table 1
for species symbols.
39
4,000
............................................................................................................
::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::[:::::~:::":I::::::::::::::::
sCTEI:::::::::::::::::::::::::::::::::::::::::::::::
::·.....:······::::::::::::r::::::§:::::::r:::::::::::
:::::::::::::::::::::::::::::f...·.....~~~~ ...····....
......
,
~-
,
~-
F
..-
.........................................................
.........................................................
.-.......... 1.·············································........... .
::::::::::::::::::::::::11-_....;..C_HN_I_--J~.
_1I._I_ ....
,.01• • • _ _ 11• • ····•·•···•········•·····•················· •...........•..
_...._.;_
_ .......................................................................................
,
-... .•.......................................................................................
-........-
PRPA
I:::::::::::::~::::::::::::::::::::::::·
..................................................................................
50
1,000
2,000
Elevation (ft)
Figure 3-The distribution of woody species
along the elevation/moisture gradient along the
Lanai transect on the island of Lanai. See table 1
for species symbols.
Figure 4-The distribution of woody species along the
elevation/moisture gradient. Data were combined from
the South Point, Saddle Road and Lanai transects on
the islands of Hawaii and Lanai, respectively.
Verbensina encelioides omitted from this figure. See
table 1 for species symbols.
40
100
1,000
2,000
Elevation (ft)
3,000
The coming of Polynesian and European settlers heavily
impacted native plant communities on all islands below
3,000 ft (914 m) elevation. These impacts paved the way for
species introduced into the islands to succeed and in many
cases to become serious pests (Stone and others 1992). The
aftermath of man's disturbance of natural ecosystems leads
to decreases in the diversity (species richness) ofthat system
(Carlquist 1980). Numbers of alien taxa encountered in this
study were low, ranging from 2 to 9 and averaging 5. 7 species
per site. These figures differ from work done by Anderson
and others (1992) in the Kipahulu valley in Haleakala
National Park on the island ofMaui who found that numbers
of alien species per site averaged 16.4 and varied from 10 to
25 for similar elevations. The communities they studied
showed both a decrease in numbers of alien species per site
as well as a decrease in vegetative cover provided by alien
introduced species as elevation increased. In contrast, this
study showed no pattern in either number of species per site
or vegetative cover provided by alien species with respect to
changing elevation.
Species introduced from widely divergent areas of the
world into the same environment, such as along the transect
at South Point on the island of Hawaii, come to the islands
preadapted to distinct elevation and/or climatic regimes
since they would have a genetic fitness for the environment
of their origin. Under such conditions they would be expected to partition the environment along encountered gradients and into unique geographical niches in their new
habitat. This creates a variation in dominant species extending across the island landscape that would match the variation of the island environment itself. Further, where the
introduced species encounter environments similar in character to the ones they emigrated from, they should be able to
establish viable populations where their greatest success
would be at the optimal conditions of their range. Also,
selection would impose severe limitation of the populations
at the periphery of their genetic tolerance, thus restricting
further expansion of range. However, if an introduced species carries with it unexpressed genetic capacity, and if the
unexpressed genetic capacity were to be adaptive in the new
habitat, species could become a severe weedy problem.
On Hawaii and Lanai the elevation and moisture gradients change gradually because of the lack of major erosion
effects on the shield volcanos. This gradual change in the two
gradients gives opportunity for species introduced to these
environments to spread and sort to positions along the
gradients where they are best adapted. Thus, the 14 woody
species encountered in this study exhibited distribution
patterns along gradients (figs. 1-3) and formed almost perfect bell-shaped curves. Studying plant species relationships with respect to these smooth gradients allows for quick
insight into the ecological relationships between the species
as well as the environments they occupy.
Plotting the distribution patterns of individual species
with respect to a single transect (figs. 1-3), in contrast to
combining data from several transects (fig. 4), yields more
precise information as to differences in the ecology of several
species growing together along the elevational and/or moisture gradient. Where data from several transects were
combined, differences in the elevationallmoisture requirements of the various species become much more difficult to
The first alien species (about 25 plant species plus the pig,
dog, jungle fowl, and rat) were introduced to the islands with
the coming of Polynesian immigrants roughly 400 A.D.
(Armstrong 1973). A second immigration of people into the
islands together with the plants they brought occurred
around 1200 A.D. (Kirch 1982). The islands were discovered
by Europeans in 1778, and since that time the introduction
of alien plant species into the island system has occurred at
an extremely elevated pace, making to day's flora a changing
mixture of exotic and endemic species (Diong 1982; Wagner
and others 1990). Strong attempts are presently being made
to control and eliminate the further introduction of alien
plant species, which are still freely being introduced to the
islands (Stone and Stone 1989).
The most important climatic factors governing plant distribution in Hawaii seem to be average annual rainfall at
elevations below about 5,000 ft (1,524 m), and temperature
and rainfall at higher elevations (Armstrong 1973). At
present, vegetation types are characterized and identified
by reference to their dominant species. When examining
vegetation maps of the Hawaiian Islands, it becomes apparent that the dominant or characteristic plant species in all
vegetation zones at lower elevations are species introduced
to Hawaii since 1778.
Studies done on the relationships of plant communities to
environmental gradients on the volcanos (Mouna Kea and
Mouna Loa) of the island of Hawaii placed the existent
vegetation into broad categories of zonation and emphasized
its correlation with moisture and temperature gradients
only in broad subunits (Muller-Dombois and others 1981).
Less emphasis has been placed on the relationship ofindividual species (especially introduced aliens) to these same
gradients. A single paper does exist which describes the
ecological distribution ofC 3 and C4 grass species in relation
to the altitudinal gradient on Mouna Loa volcano (Rundel
1980). Anderson and others (1992) discuss the distribution
and spread of alien plants as a group and their relationships
to elevation in the Kipahulu valley in Haleakala National
Park on Maui. The distribution and successful colonization
of alien plants, to island areas where they have invaded, is
closely related to the species adaptive potential, the climatic
factors and physical characteristics of the landscape as well
as to man's disturbance.
The characteristics of abiotic gradients and associated
vegetation in the Hawaiian Islands vary from sharp, unpredictable, highly dissected landforms (changing rapidly and
unpredictably across short distances) to smooth, predictable, non-dissected landforms (changing slowly and predictably across long distances). For example, gradients found on
highly eroded islands (Kauai) are highly disjunct and therefore so is the distribution patterns of associated plants.
Where smooth gradients have been disrupted by erosion of
the landscape, creating a wide variety of available habitats,
introduced species will invade and colonize available niches.
Under such conditions, species distribution becomes patchy,
making it difficult to perceive the habitat relationships
between the many alien species that have been freely introduced into the islands (Stone and Stone 1989). Conversely,
where gradients are smooth and predictable, invading alien
species distribution becomes continuous, with no sharp
boundary lines between species.
41
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Wagner, W. L.; Herbst, D. R.; Sohmer, S. H. 1990. Manual of the
Flowering Plants of Hawaii. 2 vol. Univ. of Hawaii Press, Honolulu. 1853 p.
Wagner, W. L.; Funk, V. A eds. 1995. Ha~aiian .Biogeog;ap~y:
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Williamson, M. 1981. Island Populations. Oxford Univ. Press, Oxford.
delineate. Although, combining data from several transects
does yield greater information about a single species distribution and ecology in the islands because of an increased
sample size, combining tends to obscure understanding of
interrelationships between multiple species because the
effects of microhabitat factors are masked when data are
lumped. For example, Sida falax peaked in importance at
about 450 ft (137 m) on the South Point transect and showed
a rather narrow range of distribution along that transect,
intersecting only 21.3 percent of the transects length. However, when the South Point, Saddle Road, and Lanai transect
data were combined, it became much more difficult to determine where along the elevation transect this species peaks.
Also, difficult to determine was how broad a tolerance range
Sida had relative to elevation since Sida in the combined
data intersected 80 percent ofthe measured gradient. It is not
unlikely that Sida grows across a wide elevational gradient,
but it is not readily apparent that its distribution encompasses 80 percent of the precipitation gradient as well.
References ____________________
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