Journal of Vegetation Science 6: 211-222, 1995
© IAVS; Opulus Press Uppsala. Printed in Sweden
- Primary succession of Hawaiian montane rain forest -
211
Primary succession of Hawaiian montane rain forest
on a chronosequence of eight lava flows
Kitayama, Kanehiro1,3*, Mueller-Dombois, Dieter1 & Vitousek, Peter M.2
1Department of Botany, University of Hawaii at Manoa, 3190 Maile Way #101, Honolulu, Hawaii 96822, USA;
2Department
of Biological Sciences, Stanford University, Stanford, CA 94305, USA;
author; 3present address: Forestry & Forest Products Research Institute, P.O. Box 16,
Tsukuba Norin Kenkyu Danchi, Ibaraki 305, Japan; Tel. +81 298 73 3211; Fax +81 298 74 3720
*Corresponding
Abstract. The primary-successional sere of a Hawaiian
montane rain forest was inferred from an age sequence of eight
closely located ‘a‘ Ç flows (clinker type lava); 8, 50, 140, ca.
300, ca. 400, ca. 1400, ca. 3000 and ca. 9000 yr, on a windward
slope of Mauna Loa, Hawaii. All study sites (0.2 ha each) were
at 1120 - 1250 m a.s.l. with 4000 mm mean annual rainfall.
The 400-yr, 1400-yr, and 9000-yr flows had younger volcanic
ash deposits, while the others were pure lava. Comparisons of
tree size and foliar nutrients suggested that ash increased the
availability of nitrogen, and subsequently standing biomass.
An Unweighted Pair Group Cluster Analysis on the samples
(flows) using quantitative vascular species composition revealed that clusters were correlated with age regardless of the
substrate types (pure lava vs. ash), and an indirect ordination
on the samples suggested that the sequence of sample scores
along axis 1 was perfectly correlated with the age sequence.
Although ash deposits increased biomass, they did not affect
the sequence of the successional sere. Both pubescent and
glabrous varieties of Metrosideros polymorpha (Myrtaceae)
dominated upper canopy layers on all flows ≥ 50 yr and ≤ 1400
yr, but the pubescent variety was replaced by the glabrous on
the flows ≥ 3000 yr. Lower layers were dominated initially by
a matted fern, Dicranopteris linearis, up to 300 yr, and subsequently by tree ferns, Cibotium spp., to 9000 yr. The cover of
Cibotium declined slightly after 3000 yr, while other native
herb and shrub species increased. A ‘climax’ stage in the
conventional sense was apparently not reached on the observed age gradient, because the sere changed continuously in
biomass and species; this divergent successional phenomenon
may be unique to Hawaii where the flora is naturally impoverished and disharmonic due to its geographic isolation in contrast to more diverse and harmonic floras in continents.
Keywords: Cluster analysis; Metrosideros polymorpha; Nutrient availability; Oceanic island; Ordination; Species diversity; Volcano.
Nomenclature: Wagner et al. (1990) for flowering plants; the
1987 version of the checklist by Wagner & Wagner (unpubl.)
for pteridophytes.
Introduction
The Hawaiian Islands are volcanically derived and
spatially arranged in a chronological sequence from
youngest Hawaii to oldest Kauai. The island of Hawaii
shows various geological stages ranging from active to
dormant to extinct volcanoes. Mauna Loa (4169 m) is
the second youngest volcano on this island; its slopes
are very gentle (e.g. rising 4 km over 86 km distance on
a NE slope) and consist of numerous lava flows. Ages of
historic lava flows are known precisely, and recent
geological surveys using 14C dating on buried charcoal
have produced a detailed map of prehistoric surface lava
flows on Mauna Loa (Anon. 1986; Lockwood et al.
1988).
The NE slopes of Mauna Loa and other high mountains in the Hawaiian Islands receive persistent, moist
trade winds, 65 - 80 % of the time during a normal year
(Blumenstock 1961). The orographic uplift of trade
winds on Mauna Loa results in a concentric pattern of
isohyets with maximum rainfall at ca. 700 m a.s.l.
(Giambelluca et al. 1986). Air temperature changes predictably (lapse rate 0.55 °C/100 m) below the trade wind
inversion, which frequently is as low as 1800 - 1900 m
a.s.l. (Blumenstock 1961; Giambelluca & Nullet 1991).
This climatic pattern together with lava flows of known
ages forms a laboratory-type matrix of environments on
Mauna Loa (Vitousek et al. 1992).
Patterns of succession can be investigated through
direct monitoring using permanent plots (Tsuyuzaki
1991; del Moral & Wood 1993), or can be inferred from
a sequence of closely located sites which differ only in
age or time after disturbances but are controlled for
other sources of variation (Jenny 1941, 1980; Drury &
Nisbet 1973; Mueller-Dombois & Ellenberg 1974). Primary succession in Hawaii up to a few hundred years
was studied with the latter approach by Atkinson (1970),
Eggler (1971), and Uhe (1988). However, dates of prehistoric flows were not available to those authors, and
212
Kitayama, K. et al.
climatic factors were not well controlled in their studies
(Drake & Mueller-Dombois 1993). Doty (1967) and
Smathers & Mueller-Dombois (1974) monitored the
beginning of succession for several years on new lava
flows. Populations of the major woody species in Hawaiian dry forests were compared on two flows of
different ages in each of climatically controlled sites
(Stemmermann & Ihsle 1993).
The only studies of primary succession over an
extended period with a controlled climatic condition in
Hawaii were conducted on a NE slope of Mauna Loa by
Drake & Mueller-Dombois (1993) and Aplet & Vitousek
(1994). The former focused on the population development of four major canopy species and one genus of tree
fern on six closely located lava flows of different ages at
the same altitude (1200 m). The latter analyzed species
composition and biomass in a matrix of five ages, from
5 to 3400 yr, at six altitudes, from 914 m to 2438 m. The
present study was conducted to include the whole community on the same chronosequence as Drake & MuellerDombois (1993), with two additional flows.
Study area
This study was conducted on an age sequence of
eight lava flows which were located close to each other
on a gentle NE slope of Mauna Loa, the island of Hawaii
(Fig. 1). The ages of the flows were 8, 50, 140, ca. 300,
ca. 400, ca. 1400, ca. 3000 and ca. 9000 yr at the time of
the study in 1992. The 8-yr and 9000-yr flows were in
addition to those used by Drake & Mueller-Dombois
(1993). The flows ≤ 140 yr old are historic, while those
≥ 300 yr were dated from a 14C analysis of buried charcoal (Anon. 1986; Lockwood et al. 1988; Lockwood
pers. comm.). All sites, except for the 9000-yr flow at
1120 m, are located at 1200 to 1250 m a.s.l and all sites
receive a mean annual rainfall of 4000 mm (Giambelluca
et al. 1986). Mean daily air temperature at 1130 m a.s.l
on the NE slope (Jan. 92 to May 93) reveals a maximum
value of 18.0 °C in Sept. and a minimum of 12.7 °C in
Feb.; the diurnal difference between the mean daily
maximum and minimum air temperatures at this station
is 11.6 °C (Juvik & Nullet in press).
The sites are underlain by chemically and physically
uniform ‘A‘Ç lava flows (Macdonald et al. 1983; Drake
& Mueller-Dombois 1993). ‘A‘Ç lava is composed of
rough-textured porous clinker-like rubble, and is thus
extremely pervious to water. The 400-yr, 1400-yr and
9000-yr flows are located close to the youngest
neighboring volcano, Kilauea, and have thin layers of
volcanic ash and fine ejecta (10-15 cm thick). The age
of ash is unknown, but obviously younger than the
underlying ‘a‘Ç lava. Volcanic ash weathers relatively
fast, releasing greater mineral nutrients in a shorter time
than lava flows (Balakrishnan & Mueller-Dombois 1983;
Vitousek et al. 1983; Drake & Mueller-Dombois 1993).
Therefore, it was anticipated that lava with ash would
display a community development pattern different from
that on pure lava.The vegetation at all sites was intact
and not affected by the widespread dieback (Jacobi
1993; Mueller-Dombois 1993). The only potential disturbance factor in the area was feral pigs. Pigs were,
however, limited to small patches on the forest floors of
the 1400-yr, 3000-yr, and 9000-yr flows because rugged
lava restricted their activities and effects of pigs were
judged to be inconsequential. All flows were narrow
bands < 0.5 km width at the study sites, surrounded by
intact vegetation with the same flora. Therefore, the
supply of disseminules to newly created flows was not
limited substantially (Drake & Mueller-Dombois 1993).
Methods
Vegetation
Fig. 1. Map showing the eight lava flows studied on the NE
slope of Mauna Loa, Hawaii.
Five plots (each 20 m × 20 m quadrat, total 0.2 ha)
were placed more-or-less contiguously from a randomly
selected point on each flow. Except on the 8-yr and
9000-yr flows, the plots were located on the transects of
Drake & Mueller-Dombois (1993). In each of the plots,
the vegetation was stratified into structural layers and all
vascular species present in each layer were identified
and inventoried with Braun-Blanquet’s cover-abundance
scale (the relevé method, Mueller-Dombois & Ellenberg
1974). In the same plot, five tallest canopy trees of
- Primary succession of Hawaiian montane rain forest Metrosideros polymorpha, the dominant species in the
Hawaiian rain forests (Mueller-Dombois 1987), were
measured for height and diameter at breast height (DBH).
The enumeration was limited to five trees per plot to
ensure that all trees measured belonged to uppermost
canopies in all plots.
Vegetation data were combined by flow; the coverabundance scales of enumerated species were converted
into midpoint values (Mueller-Dombois & Ellenberg
1974) and means were computed by flow. This procedure resulted in a data table of species and samples
(flows). Subsequently, reciprocal similarities among the
eight flows were computed based on the data table using
the percent similarity index, PS (Kovach 1990). The 8yr flow was omitted because its vegetation was extremely sparse and its inclusion would distort relationships of the other flows.
An Unweighted Pair Group Cluster Analysis was
applied to the resulting similarity matrix. An indirect
ordination of the flows was performed on the same data
table using Detrended Correspondence Analysis (program DECORANA, Hill 1979). The 8-yr flow was
omitted from the ordination for the same reason. Species diversity was computed for each flow based on
Simpson’s index:
D = 1/Σ pi2
(1)
where pi = relativized cover-abundance value of the i-th
species. The cluster analysis and the computation of
species diversity were performed with MVSP Plus 2.0
(Kovach 1990).
213
micro Kjeldahl procedure with concentrated sulfuric
acid, and the ammonium produced was measured by the
indophenol blue method on an autoanalyzer. Phosphorus was extracted in the modified Truog solution acidified to pH 2.04 (Ayers & Hagihara 1952). Soil pH was
determined on 1:1 fresh soil to de-ionized water. Finally, a subsample of each composite was oven-dried at
105 °C for 48 h to determine water content. Soil mass
and chemical data of the 140-yr and 3000-yr flows were
used from Vitousek et al. (1992). The 8-yr and 50-yr
flows were not analyzed because they did not have a
measurable amount of soils.
Eight to ten mature sun-leaves of Metrosideros polymorpha were collected from each of five to eight trees
on all flows, using a shotgun where sun-leaves could not
be obtained by hand. The youngest fully mature whorl
of leaves (behind an expanding whorl or a mature bud)
was sampled in each case. Fresh leaves were measured
for leaf area, and then oven-dried for three days at 70 °C
to determine leaf mass per area (LMA). Leaf samples
were ground and acid-digested using peroxide/persulfate
procedure for N and P; for foliar cations, they were dryashed at 500 °C for 4 h, dissolved in nitric acid, and
analyzed by atomic absorption spectrophotometry. At
some flows, pubescent and glabrous varieties of
Metrosideros coexisted. They were collected, analyzed,
and reported separately as Vitousek et al. (1992) described significant differences in foliar nutrients between the two varieties.
Results
Soils and foliar nutrients
Soil
Soil depth was measured at 20 points in 1 m-intervals along the center line of each plot. Soil samples were
collected from three of the five plots on each of the 300yr, 400-yr, 1400-yr, and 9000-yr flows in a systematic
manner; ten cores (37 mm diameter, 15 cm depth or to
the surface of lava if shallower) were collected in 2-m
intervals along the center line of each plot, and mixed
into a composite sample for chemical analyses. Additionally, to determine soil mass, five soil cores (5 cm
diameter) were collected to the surface of lava in 5-m
intervals along each of four 25 m transects where soils
were deeper. It should be noted that an unknown amount
of organic, and weathered mineral matters filters into
interstices of ‘a‘Ç rocks and these could not be sampled
by our method. In the laboratory, exchangeable Ca, Mg
and K were extracted in 1-N NaCl and analyzed by
atomic absorption spectrophotometry. Organic C was
determined by the modified Walkley-Black wet digestion method (Heanes 1984). Total N was digested by the
Mean soil depth increased linearly with increasing
age expressed on a log scale (r2 = 0.98, p < 0.01) on the
lava series without ash (Fig. 2). The ‘soils’ on these
flows are highly histic (see below) and the linear relationship indicates continued organic matter accumulation at an ever-decreasing rate at least up to 3000 yr.
Mean depth was 0.2 cm on the 50-yr flow, and 19.4 cm
on the 3000-yr flow. There were abundant live and dead
lichens on the 8-yr flow but no derived soils were found.
The soil depth on the 400-yr, 1400-yr, and 9000-yr
flows deviated above the regression line due to deposits
of younger volcanic ash.
Soil organic C concentrations were extremely high
on the 140-yr and 300-yr flows (47.8 % and 45.4 %,
respectively), suggesting that the soils were purely organic and that weathering of the underlying lava was
extremely slow (Table 1). The C concentration on the
3000-yr flow was reduced to 29.2 % probably due to
minerals incorporated from the underlying lava. The pH
214
Kitayama, K. et al.
Fig. 2. Soil depth on the ‘a‘Ç flows without ash ( ), and with
ash ( ). Mean values ± standard errors based on 100 points per
flow.
differed between the substrate types; it was lower on the
lava series than the ash series. Extractable cation concentrations (Ca, Mg and K) were high in the soils from
the two youngest lava flows analyzed (140-yr and 300yr), and abruptly decreased on the oldest lava without
ash (3000-yr) and in all soils derived from the ash series
(Table 1).
Fig. 3. Foliar nitrogen (N) and phosphorus (P) concentrations
of the glabrous-leaved variety of Metrosideros polymorpha
along the age sequence of the eight lava flows. The values are
means ± standard errors. Open symbols with solid lines denote
leaves from pure lava, and filled symbols with dashed lines
from lava with ash. The variety on the 8-yr flow is unknown.
Foliar nutrients
Mean LMA of Metrosideros (including glabrous
and pubescent varieties) from the lava series was initially low on the 8-yr flow, increased to 270 - 290 g/m2,
and then declined to 190 g/m2 on the 3000-yr flow
(Table 2). Similarly, LMA for the ash series declined
from ca. 250 g/m2 on the 400-yr and 1400-yr flows to
180 g/m2 on the 9000-yr flow.
Mean foliar N concentrations of the glabrous-leaved
variety of Metrosideros (i.e. the most common variety)
increased with substrate age on the lava series (Fig. 3).
Foliar N concentrations from the ash series were slightly
higher than those for the same ages on the lava series
(Fig. 3). Mean foliar P concentrations of the glabrous
variety did not show any age-related trends on the lava
Table 1. Soil mass above the surface of lava, and top soil (15 cm or less) concentrations (oven dry basis) of organic carbon (O-C),
total nitrogen (T-N), extractable phosphorus (P) and cations (Ca, Mg, and K), and pH (H2O) on the six lava flows studied for soil
analyses on Mauna Loa. Values are means with standard errors in parentheses.
Age
yr
pH
T-N
P
%
Pure lava
140
300
3000
*
*#
Lava+ash
400 #
1400
9000
*Cited
O-C
Mg
K
ppm
Mass
kg/m2
4.1 (0.0)
4.5 (0.1)
4.7 (0.0)
47.8
45.4 (1.2)
29.2
1.1
2.0 (0.0)
1.3
41 (3)
2 (0.2)
55 (3)
3770 (380)
3786 (228)
2200 (240)
700 (60)
544 (20)
120 (15)
430 (20)
589 (15)
180 (20)
4.4 (0.3)
3.6 (0.5)
22.2 (3.9)
5.3 (0.1)
5.1 (0.1)
5.3 (0.1)
14.0 (1.0)
14.1 (0.7)
13.0 (0.6)
1.0 (0.1)
1.1 (0.0)
1.1 (0.0)
16 (1.4)
14 (1.7)
17 (2.5)
1331 (52)
2157 (116)
1798 (87)
266 (8)
265 (9)
238 (18)
84 (14)
82 (1)
76 (3)
27.0 (2.0)
57.7 (5.4)
44.2 (3.6)
from and modified based on Vitousek et al. (1992).
collected to the surface of lava, depth ranging from 15 to 25 cm.
#Samples
Ca
- Primary succession of Hawaiian montane rain forest -
215
Table 2. Foliar nutrient concentrations on a dry mass basis, and leaf mass per area (LMA) for sun leaves of glabrous (G) and
pubescent (P) leaved varieties of Metrosideros polymorpha on the eight lava flows on Mauna Loa. Mean values with standard errors
in parentheses. ‘SN’ indicates sample numbers and ‘Both’ unweighted means of pubescent and glabrous leaves.
Flow
N
P
(SN)
Ca
Mg
K
LMA
g/m2
%
Pure lava
8#
0.61 (0.06)
0.072 (0.006)
2.12 (0.30)
0.16 (0.03)
0.66 (0.02)
196
50
G(3)
P(5)
Both
0.62 (0.06)
0.56 (0.02)
0.59 (0.03)
0.078 (0.010)
0.056 (0.003)
0.064 (0.005)
0.86 (0.11)
0.96 (0.08)
0.92 (0.06)
0.08 (0.01)
0.08 (0.01)
0.08 (0.01)
0.62 (0.01)
0.38 (0.04)
0.48 (0.06)
238 (21)
317 (21)
280 (19)
140
G(2)
P(4)
Both
0.72 (0.02)
0.56 (0.04)
0.61 (0.04)
0.053 (0.001)
0.048 (0.003)
0.050 (0.002)
1.02 (0.20)
0.73 (0.08)
0.82 (0.09)
0.13 (0.01)
0.10 (0.01)
0.11 (0.01)
0.62 (0.06)
0.43 (0.02)
0.49 (0.04)
268 (5)
300 (12)
290 (15)
300
G(3)
P
Both
0.65 (0.01)
0.63 (0.02)
0.64 (0.02)
0.054 (0.003)
0.060 (0.008)
0.058 (0.005)
1.33 (0.16)
1.04 (0.08)
1.15 (0.09)
0.09 (0.00)
0.09 (0.01)
0.09 (0.01)
0.50 (0.06)
0.35 (0.04)
0.41 (0.04)
243 (8)
283 (14)
269 (12)
3000
G
1.00 (0.04)
0.078 (0.004)
0.78 (0.07)
0.13 (0.01)
0.53 (0.02)
191 (13)
Lava+ash
400
G(2)
P(3)
Both
1.11 (0.10)
0.84 (0.03)
0.95 (0.08)
0.072 (0.009)
0.061 (0.005)
0.065 (0.005)
0.80 (0.09)
0.56 (0.10)
0.68 (0.09)
0.21 (0.06)
0.11 (0.03)
0.16 (0.04)
0.65 (0.10)
0.54 (0.18)
0.59 (0.09)
226 (39)
258 (9)
247 (15)
1400
G(3)
p(3)
Both
0.96 (0.10)
0.76 (0.03)
0.86 (0.06)
0.065 (0.002)
0.047 (0.004)
0.056 (0.005)
0.72 (0.01)
0.51 (0.19)
0.61 (0.10)
0.16 (0.04)
0.14 (0.03)
0.15 (0.02)
0.55 (0.02)
0.41 (0.05)
0.48 (0.05)
237 (7)
279 (19)
255 (13)
9000
G(4)
P(1)*
Both
1.07 (0.03)
1.01
1.06 (0.02)
0.080 (0.003)
0.073
0.078 (0.003)
0.67 (0.06)
0.79
0.70 (0.05)
0.16 (0.03)
0.15
0.16 (0.02)
0.59 (0.04)
0.33
0.52 (0.07)
170 (7)
227
182 (12)
(5)
# Variety unknown.
* Found in the close vicinity of the study plot. No pubescent leaves actually found inside the plot.
series and differences in the concentrations between the
substrate types (lava vs. ash) were not significant.
Cation concentrations (Ca, Mg and K) were high in
the leaf samples from the 8-yr flow (Table 2). There
were no consistent differences in foliar cation concentrations between the substrate types of comparable ages.
Vegetation
Mean height and DBH of Metrosideros canopy trees
on the lava series (Fig. 4) increased linearly with increasing substrate age (on a log scale, r2 = 0.99, p < 0.01
for height and DBH). The 8-yr flow was at an early
colonization stage and no measurable trees were found.
Mean height and DBH-values on the ash series deviated
significantly above the regression lines on the 400-yr
and 1400-yr flows. However, the values conformed
closely to the regression on the 9000-yr flow. The relationship between DBH and height varied greatly among
flows (Fig. 4), suggesting that it is highly substrate ageand type-specific.
Quantitative vascular species composition on the
eight lava flows varied markedly with age (App. 1).
Metrosideros polymorpha dominated the upper layers
throughout the age sequence ≥ 50 yr. Dicranopteris
linearis dominated the lower layers on the flows ≥ 50 yr
and ≤ 140 yr, and was replaced by Cibotium glaucum
and Sadleria pallida on the older flows. The cover of
rhizomatous hemicryptophytes, Athyrium sandwichianum and Thelypteris sandwicensis, in herb layers steadily increased on the flows ≥ 300 yr.
The number of vascular species per 0.2 ha rapidly
increased from 10 on the 8-yr to 64 on the 300-yr flow,
and then slightly declined and reached a steady state
216
Kitayama, K. et al.
Fig. 4. Relationships between flow age and values for mean (±
standard deviation) height (circles) and DBH (triangles) of
Metrosideros polymorpha canopy trees. Open symbols are for
pure lava, and solid symbols for lava with ash. For Metrosideros
top-canopy trees on pure lava without ash:
H = 8.80 log(age) – 11.81; r2 = 0.99, p < 0.01;
DBH = 15.88 log(age) – 23.36; r2 = 0.99, p < 0.01.
(Table 3). The greatest richness of canopy tree species
(7 taxa) was found on the 300-yr flow and all flows with
ash (App. 1). Overall, the greatest number of species
were provided from pteridophytes and their number
rapidly increased from 2 on the 8-yr to 32 on the 300-yr,
and then declined (App. 1). Alien species were very low
in number and cover throughout the age sequence.
The highest species diversity (0.81) based on
Simpson’s index except for the colonizing stage (the 8yr flow) was found on the 300-yr flow (Table 3); here,
the total number of species also attained a maximum 64. Interestingly, the 8-yr flow had a higher diversity
index - 0.87; this is merely a reflection of high species
evenness due to an extremely sparse vegetation cover.
There was no distinct linear trend in the diversity index
along the age sequence.
An Unweighted Pair Group Cluster Analysis applied to the data set in App. 1 demonstrated that sample
(flow) clusters were correlated with age regardless of
Fig. 5. Results of the Unweighted Pair Group Cluster Analysis applied to the flows studied. The 8-yr flow was omitted
from the analysis.
the substrate types (Fig. 5). Community similarities
between substrates close in age are generally high (> ca.
50%). There are four clusters at 60 % similarity level: 1)
50-yr; 2) 140-yr; 3) 300-yr, 400-yr + ash, and 1400-yr +
ash; and 4) 3000-yr, and 9000-yr + ash.
The Detrended Correspondence Analysis showed
that the highest species variation among the flows was
found only along axis 1 (eigenvalue = 0.65) and that
three other axes explained little variation (all eigenvalues
≤ 0.1, Table 4). The sequence of sample scores along
axis 1 from high to low was perfectly correlated with the
age sequence (Spearman’s rank correlation r2 = 1). In
contrast, soil depth was not correlated with the sequence
of sample scores across the eight flows (Spearman’s
rank correlation r2 = 0.51, p > 0.05).
Fig. 6 depicts the species abundance curves for dominant taxa selected in each of the tree (tree-fern included)
and pteridophyte groups. Species changed unimodally
in cover along the age sequence of the eight lava flows
(Fig. 6). The modes of the species were staggered along
the age sequence and the width of the curves varied
greatly among the species, from the order of hundreds of
years to thousands of years. Noteworthy is the successional displacement between the two conspecific varieties of Metrosideros polymorpha. The pubescent var.
Table 3. Diversity of vascular species (taxa) on the eight lava flows based on Simpson’s index. Diversity values on the 8-yr flow
(*) should be downgraded; see the text for explanation.
Flow age
8
50
140
300
400
1400
3000
9000
Diversity
Evenness
0.87*
0.87*
0.73
0.52
0.66
0.42
0.81
0.45
0.74
0.41
0.77
0.43
0.74
0.42
0.80
0.45
10
25
36
64
63
62
60
60
No. taxa/0.2ha
- Primary succession of Hawaiian montane rain forest -
Fig. 6. Species abundance curves along the chronosequence
of the eight lava flows. Species are plotted against mean midpoint cover values (see text), therefore the curves do not
strictly reflect absolute species values. a. Eight top-ranked
dominant tree and tree-fern species: MPG = Metrosideros
polymorpha var. glaberrima; MPP = M. polymorpha var.
polymorpha; CG = Cibotium glaucum; CC = Cibotium
chamissoi; CT = Cheirodendron trigynum; IA = Ilex anomala;
CO = Coprosma ochracea; ST = Styphelia tameiameiae.
b. Seven top-ranked dominant herbaceous species: SC = Sadleria cyatheoides; DL = Dicranopteris linearis; LC = Lycopodium cernuum; SP = Sadleria pallida; TS = Thelypteris sandwicensis; AS = Athyrium sandwichianum; MA = Machaerina
angustifolia.
Table 4. Eigenvalues and sample scores of the lava flows on
the four ordination axes extracted by Detrended Correspondence Analysis. The youngest flow was omitted from the
analysis.
Flow age
50
140
300
400
1400
3000
9000
Eigenvalue
Axis 1
Axis 2
Axis 3
Axis 4
294
267
52
30
28
22
0
65
97
132
142
118
53
0
21
16
4
86
67
7
0
1
35
10
91
68
22
0
0.65
0.10
0.02
0.00
217
polymorpha was displaced by the glabrous var.
glaberrima at a later stage; their curves widely overlapped, but the modes were clearly separated.
Fig. 7 shows the cumulative cover values of selected
dominant species, and four successional phases by combinations of these species. The dominant species selected include Cibotium glaucum, Metrosideros polymorpha var. glaberrima, M. polymorpha var. polymorpha, Dicranopteris linearis, Sadleria pallida; the
rest of the species are lumped into one category. Phase 1
(< 140 yr) was characterized by the dominance of
Dicranopteris linearis, and by the absence or low cover
values of Cibotium glaucum; phase 2 (140 - 300 yr) by
the abundance of Dicranopteris linearis and Cibotium
glaucum; phase 3 (300 - 1400 yr) by the dominance of
Cibotium glaucum and the coexistence of the two varieties of Metrosideros polymorpha; and phase 4 (> 1400
yr) by the displacement of the pubescent by the glabrous
variety of Metrosideros polymorpha, and by a slight
decline of Cibotium glaucum.
The cumulative cover-abundance of the five dominant species (outlined by shaded areas in Fig. 7) increased rapidly from 8 yr to 300 yr, and then attained a
near-constant value. Overall cumulative values (indicated by the uppermost line), however, still increased
after 300 yr due to the steady increase of other nondominant species.
Discussion
Influences of volcanic ash
The mean height and DBH of Metrosideros polymorpha canopy trees on the 400-yr and 1400-yr ashincorporated flows were substantially greater than those
extrapolated from the regression lines for the pure lava
series (Fig. 4). Ash deposits apparently caused canopy
Metrosideros trees to increase in mean height and DBH.
Because the montane rain forest studied was strongly
dominated by Metrosideros polymorpha with only a
few subordinate canopy species, ash deposits resulted in
elevating the entire standing biomass, as Drake &
Mueller-Dombois (1993) also observed for basal area.
Metrosideros foliar N concentrations were initially
low, but increased steadily with age on the lava series,
and the concentrations were greater in leaves from the
ash than from the lava series (Fig. 3). This was consistent with the findings by Vitousek et al. (1983), who
demonstrated that net N turnover was greater in soils
derived from ash than from the lava (pÇhoehoe, solid
smooth-surfaced type) of comparable ages. Ash deposits in our sites led to an improved N availability, which
was responsible for increasing biomass. Ash deposits
218
Kitayama, K. et al.
Fig. 7. Four phases, recognized by
combinations of dominant species,
in the primary successional sere of
the Hawaiian montane rain forest.
The y-axis indicates cumulative
cover values of constituent species.
CG = Cibotium glaucum; MPG =
Metrosideros polymorpha var. glaberrima; DL = Dicranopteris linearis; MPP = M. polymorpha var. polymorpha; SP, Sadleria pallida;
OTHER, all other species combined.
may also increase the availabilities of other soil nutrients but the enhanced growth by greater N can dilute
concentrations of other nutrients in the leaves (Tanner et
al. 1992).
In contrast to its effects on biomass, volcanic ash
does not seem to have changed the sequence of species
displacements in the sere. Axis 1 in the ordination
perfectly defined the age sequence (Table 4), and flows
of similar ages formed cohesive clusters regardless of
ash deposits (Fig. 5). Therefore, the age sequence of the
flows (Figs. 6 and 7) can represent the chronosequence
of rain forest development on a single flow. We speculate that thin ash deposits may not completely devastate
forest canopies, leaving at least dominant trees to recover from the disturbance event. This phenomenon
was observed in ash deposits from the 1959 eruption of
Kilauea Iki (Smathers & Mueller-Dombois 1974).
Inferred primary-successional sere on an ‘a‘Ç lava flow
After the formation of an ‘a‘Ç lava flow, the lichen
Stereocaulon vulcani (Bory) Ach. densely colonizes
exposed porous materials on the flow (Atkinson 1970;
Eggler 1971; Mueller-Dombois 1987). Several native
species establish individuals within at least 8 yr, including Metrosideros polymorpha, Vaccinium calycinum,
Hedyotis centranthoides, Dubautia scabra, Coprosma
ernodeoides, Machaerina angustifolia, Nephrolepis
cordifolia, and Polypodium pellucidum. The establishment of readily dispersible Metrosideros polymorpha
seedlings is constrained only by the availability of ‘safe
sites’ for germination (Drake 1992). Such safe sites are
not limited on ‘a‘Ç lava flows because there are numerous pockets with fine mineral particles. The development of the Hawaiian montane rain forest after the
establishment of Metrosideros seedlings proceeds as
follows from phase 1 to phase 4 (Fig. 7).
Phase 1 (≤ at least 140 yr): A widely open-canopy
Metrosideros scrub with a dense ground cover of
Dicranopteris linearis. The mean upper canopy height is
ca. 6.5 m in 140 yr. The canopy consists of both varieties
of Metrosideros with a greater cover of the pubescent
var. polymorpha than the glabrous var. glaberrima.
Phase 2 (> at least 140 yr, and ≤ at least 300 yr): An
open-to somewhat closed-canopy Metrosideros lowstature forest. The mean upper canopy height becomes
ca. 10 m in 300 yr. The canopy consists of both varieties
of Metrosideros with a greater cover of the pubescent
than the glabrous variety. An accompanying canopy
tree species, Cheirodendron trigynum, starts increasing
in cover. Cibotium glaucum prevails in the shrub layer.
Dicranopteris linearis still persists. Species diversity
becomes maximal in this phase because early colonizers
and late-seral species coexist.
Phase 3 (from as early as 300 yr up to < 3000 yr): A
somewhat closed-canopy Metrosideros forest. Mean
upper canopy height is expected to be ca. 15 m on ‘a‘Ç
lava without ash at 1000 yr (extrapolated from the regression in Fig. 4). Both varieties of Metrosideros still
coexist, but the glabrous variety increases until it completely displaces the pubescent one. Cibotium glaucum
forms a closed, dense shrub layer.
Phase 4 (≥ at least 3000 yr): A closed-canopy
Metrosideros forest. Extrapolated from Fig. 4, the mean
upper canopy height is ca. 20 m on the ‘a‘Ç lava without
ash in 4000 yr. The pubescent variety of Metrosideros
has been displaced by the glabrous one. Cibotium
glaucum starts to decline in cover, while other native
species increase.
Halpern et al. (1990) and del Moral & Wood (1993)
- Primary succession of Hawaiian montane rain forest observed that chance factors had influenced the primary
succession on Mount St. Helens, USA, in many ways
(e.g. timing of eruption in relation to growing seasons,
availability of microsites, and distance to seed supplies).
By contrast, we did not find such evidence that chance
factors play significant roles in Hawaii and the inferred
sere may be generalized for the study area as far as the
environmental setting is the same.
The basal area of Metrosideros trees (≥ 5 cm basal
diameter) increases linearly with age (log) from phase 1
to an early stage of phase 4 according to Drake &
Mueller-Dombois (1993). They found a higher basal
area value on the 400-yr flow than that expected from
the regression and this is probably due to the favorable
effects of ash deposits.
There appear to be displacements of several
Metrosideros generations in the forest canopy during
the 9000-yr sere. Mueller-Dombois (1986, 1987) suggested that generation displacements proceed through
canopy dieback, a consequence of the cohort formation
and subsequent synchronous cohort senescence; and
that the next cohort’s recruitment in this shade-intolerant species follows the dieback. This pattern is more
pronounced on pÇhoehoe lava and poorly drained ash
than ‘a‘Ç (Jacobi et al. 1983), and we did not observe
strong evidence for the importance of dieback in these
sites. As Drake & Mueller-Dombois (1993) pointed out,
canopy dynamics on ‘a‘Ç lava seems to be driven initially through self-thinning, and gaps by wind thrown
trees. Small-scale gap formation by senescent trees probably becomes a driving force at later stages. In all cases,
conspecific varieties of Metrosideros polymorpha are
the only dominant canopy trees throughout the sere.
Seres in which a pioneer species persists as a dominant
at later stages despite relatively mild climates seem to be
unique to Hawaii and other isolated oceanic islands
(Itow & Mueller-Dombois 1988; Mueller-Dombois
1992; Drake & Mueller-Dombois 1993), probably because their floras are depauperate and disharmonic due
to isolation (Carlquist 1974).
Linear relationships of mean height and DBH with
age (log) ≤ 300 yr (Fig. 4) demonstrate growth patterns
for a single Metrosideros generation. Continued increases in mean height and DBH (Fig. 4) of this species
on the later flows involve several generations. There is
evidence that primary production at early stages of
primary succession in Hawaii is limited by N availability (Vitousek et al. 1987; L.R. Walker & Vitousek 1991;
Vitousek et al. in press), and total and available N
gradually increases on older flows (Vitousek et al. 1992).
The continued increase of mean height and DBH of
Metrosideros polymorpha after the first generation (Fig.
4) may be primarily due to increasing N availability
during ecosystem development; P may also be involved
219
in the development (T.W. Walker & Syers 1976) although we did not find direct evidence.
The mean DBH of Metrosideros polymorpha on the
9000-yr flow obviously declined compared with that of
the 1400-yr flow, in spite of the presence of ash (Fig. 4).
The cover of Cibotium glaucum also declined. The
reasons for the decline are not known, but perhaps the
availability of some nutrients decreases as both waterlogging and element leaching increase.
According to the above scenario, the Hawaiian
montane rain forest never reaches its ‘climax’ in the
conventional sense (Mueller-Dombois 1992). ‘Seral
communities’ of the Hawaiian montane rain forest become completely dominated by Metrosideros polymorpha var. glaberrima in a few thousand years, while
still continuing to change in biomass and species composition.
Acknowledgements. We thank Shinji Kitayama for his excellent assistance in fieldwork throughout the study. This study
would not have been possible without his help. Drs. Donald
Drake and Guillermo Goldstein commented on the manuscript. This study was supported by US National Science
Foundation grants BSR8918526 to DMD and BSR8918382 to
PMV.
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- Primary succession of Hawaiian montane rain forest -
221
App. 1. Species composition of the eight ‘a‘Ç lava flows studied. Figures indicate species dominance values by mean midpoints of
Braun-Blanquet’s cover-abundance scales.
Age of the lava flows (yr)
Tree species
Metrosideros polymorpha var. glaberrima
M. polymorpha var. polymorpha
Cheirodendron trigynum
Ilex anomala
Psychotria hawaiiensis
Myrsine lessertiana
M. sandwicensis
Xylosma hawaiiense
Shrub and tree-fern species
Cibotium glaucum
Coprosma ochracea
Broussaisia arguta
Styphelia tameiameiae
Cibotium chamissoi
Vaccinium calycinum (Hawaii)
Hedyotis terminalis
Coprosma pubens
Tetraplasandra oahuensis
Vaccinium reticulatum (typicum)
V. calycinum (coriaceous)
Clermontia montis-loa
Pelea clusiifolia
Hedyotis centranthoides
Pelea pseudoanisata
Rubus hawaiensis
Pipturus albidus
Nothocestrum longifolium
Perrottetia sandwicensis
Clermontia parviflora
Wikstroemia sandwicensis
Eurya sandwicensis
8
50
140
300
400
1400
3000
9000
0.01
5
2
14.5
21.5
0.01
7.5
24
2
0.6
0.01
0.1
0.1
9.5
33
2.5
1
0.1
0.6
0.1
14.5
42.5
5
4
1.5
1.1
0.1
42.5
72.5
5
1.5
7.5
4.5
2.5
0.6
0.1
0.01
72.5
19.5
1.1
7.5
4.5
1.1
0.1
87.5
4
2
0.1
0.1
0.1
2.5
0.6
3
82.5
0.5
1.5
87.5
72.5
0.1
0.1
0.6
0.6
0.1
2
10
0.1
0.1
0.1
0.01
0.1
0.01
0.1
0.1
0.01
0.1
0.01
0.1
0.1
0.1
0.1
0.6
0.1
0.1
0.01
2
0.1
2.5
Herbaceous species
Machaerina angustfolia
Peperomia hypoleuca
Uncinia uncinata
Astelia menziesiana
Nertera granadensis
Carex alligata
Isachne distichophylla
Deschampsia nubigena
Peperomia macraeana
0.6
0.1
0.1
0.6
0.5
2
1.5
0.01
0.1
0.01
0.01
0.1
0.1
0.1
0.1
0.1
0.1
0.1
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
Liana species
Freycinetia arborea
Alyxia oliviformis
Stenogyne calaminthoides
Smilax melastomifolia
Low shrub species
Cyrtandra platyphylla
Labordia hedyosmifolia
Vaccinium reticulatum (pahalae)
Cyrtandra lysiosepala
C. paludosa
Cyanea pilosa v. longipedunculata
C. degeneriana
Dubautia scabra ssp. scabra
Coprosma ernodeoides
1.1
0.1
1.5
1.1
0.1
0.1
1.5
0.1
0.01
0.01
0.1
0.1
0.1
0.01
0.6
0.1
0.1
0.01
0.1
0.1
0.01
2
0.6
0.6
0.1
0.6
0.1
1.5
1.5
0.6
0.6
0.1
0.1
0.1
0.1
0.1
0.1
0.1
0.1
0.01
0.1
0.1
0.1
0.1
1.1
0.6
0.1
0.1
1.5
1.5
0.6
2.5
1.5
0.1
0.1
0.01
0.1
0.01
0.1
0.1
0.1
0.1
0.01
4
7,5
0.01
0.01
0.01
0.1
0.1
0.1
0.1
0.6
0.01
0.01
0.1
0.01
2.5
0.1
0.1
0.1
0.1
0.01
0.1
2.5
0.01
1.1
222
Kitayama, K. et al.
App. 1. (Cont.)
Age of the lava flows (yr)
8
Pteridophytic species
Dicranopteris linearis
Sadleria pallida
Athyrium sandwichianum
Thelypteris sandwicensis
Lycopodium cernuum
Dryopteris wallichiana
Sadleria cyatheoides
Athyrium microphyllum
Diploterygium pinnatum
Sphaerocionium lanceolatum
Mecodium recurvum
Nephrolepis cordifolia
Adenophorus tamariscinus
Grammitis hookeri
Lycopodium venustulum
Coniogramme pilosa
Sadleria souleyetiana
Xiphopteris saffordii
Psilotum complanatum
Elaphoglossum hirtum
Polypodium pellucidum
Callistopteris baldwinii
Grammitis tenella
Elaphoglossum alatum
Asplenium lobulatum
Adenophorus hymenophylloides
Odontosoria chinensis
Marattia douglasii
Adenophorus tripinnatifidus
A. pinnatifidus
Thelypteris globulifera
Dryopteris glabra
Psilotum nudum
Microlepia strigosa
Elaphoglossum wawrae
Pleopeltis thunbergiana
Thelypteris keraudreniana
Sticherus owhyhensis
Asplenium contiguum
Ophioglossum pendulum
Elaphoglossum crassifolium
Vandenboschia davallioides
Pteris excelsa
Dryopteris fusco-atra
Adenophorus montanus
Huperzia serrata
Sadleria pall x soul
Thelypteris cyatheoides
Asplenium polyodon
Alien species
Arundina graminifolia
Andropogon virginicus
Pityrogramma austroamericana
Rubus rosifolius
Phaius tankarvilleae
Erechtites valerianifolia
Juncus planifolius
Hypericum mutilum
Total number of species
0.1
50
140
300
400
1400
3000
9000
28.5
0.1
77.5
1.1
3
19.5
0.01
7.5
7.5
4.5
15
9.5
15
12.5
4.5
0.6
33
19.5
4
12.5
2
0.01
6.5
42.5
0.01
1
1
5
3
0.6
0.5
0.1
0.1
0.1
0.1
1.1
2
1
0.1
0.01
0.1
0.1
0.1
0.1
0.01
0.1
1.5
0.1
0.1
0.1
0.1
0.1
0.6
0.01
0.1
0.1
0.01
0.6
0.01
0.1
0.1
0.01
0.01
0.01
0.1
0.1
0.1
0.01
0.01
0.1
0.01
0.01
0.1
0.01
2.5
4
0.6
0.1
0.1
0.1
0.6
0.1
2.5
2
1.5
0.1
0.6
0.1
0.1
0.01
0.1
0.1
0.01
0.01
0.01
0.5
0.1
0.1
0.1
0.1
0.01
0.01
0.1
0.1
0.01
0.1
0.01
0.01
0.1
0.1
0.01
0.01
0.01
0.01
0.1
0.01
0.01
0.01
0.01
0.1
0.1
0.01
0.1
0.01
0.1
0.01
0.1
0.1
0.1
0.1
0.1
0.1
0.01
0.01
0.1
0.1
0.1
0.01
0.1
0.01
0.01
0.01
0.01
0.1
0.1
0.1
0.01
0.01
0.1
0.01
0.01
0.01
0.01
0.1
0.1
0.1
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.1
0.01
0.01
0.01
0.01
0.01
0.1
0.6
0.6
0.1
0.01
0.1
0.01
0.01
0.01
0.01
10
25
36
64
63
62
60
60