Connell Hypothesis

Compensatory Recruitment, Growth, and Mortality as Factors Maintaining Rain Forest Tree
Diversity
Author(s): Joseph H. Connell, J. G. Tracey and Leonard J. Webb
Source: Ecological Monographs, Vol. 54, No. 2 (Jun., 1984), pp. 141-164
Published by: Wiley
Stable URL: http://www.jstor.org/stable/1942659
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Ecological Monographs, 54(2), 1984, pp. 141-164
? 1984 by the Ecological Society of America
COMPENSATORY RECRUITMENT, GROWTH, AND MORTALITY
AS FACTORS MAINTAINING RAIN FOREST TREE DIVERSITY'
JOSEPH H. CONNELL
Department of Biological Sciences, University of California,
Santa Barbara, California 93106 USA
J. G. TRACEY2 AND LEONARD J. WEBB3
Rainforest Ecology Unit, CSIRO Laboratories, Division of Plant Industry,
Indooroopilly, Queensland, 4068 Australia
Abstract. One general hypothesis to explain how forest tree diversity is maintained is that rarer
species are favored over commoner species in their reproduction, growth, and/or mortality. Mecha-
nisms acting in this way would continually compensate for the tendency of some species to increase
at the expense of others, and would reduce the chance of local extinction of rare species. Two hypotheses
concerning such compensatory mechanisms were tested in subtropical and tropical evergreen rain
forests in Queensland, Australia.
Hypothesis 1: on the scale of 1 -2 ha, commoner species have lower rates of recruitment and growth
and higher rates of mortality than do rarer ones. This hypothesis was tested using abundances either
of adults or of members of the same size-class, and was rejected for growth and mortality and for
recruitment of over-story species, but not rejected for recruitment in subcanopy and understory species
in either forest.
Hypothesis 2: the close proximity of other individuals is more deleterious (i.e., causes slower growth
or higher mortality) if they are the same species than if they are different species. This hypothesis was
accepted for growth or survival of nearest neighbors in several of the seedling size-classes in both
forests. In contrast, increased densities of the same species in quadrats had no more deleterious effect
on growth and survival than did increased densities of different species. At the scale of proximity to
adults, hypothesis 2 predicts that young trees have higher mortality nearer conspecific adults than
farther away. In both forests, 90% of the species tested showed no such pattern of mortality of seedlings
or saplings, nor was the strength or direction of the deviation from equal mortality correlated with
the abundance of adults of that species. Field experiments gave the same results.
In summary, tests of both hypotheses showed that some compensatory trends occurred and that
these were very similar in the two forests. The mechanisms producing these compensatory trends may
be attacks by natural enemies (grazers, pathogenic fungi, etc.), interference, or, less likely, competition
for resources.
Key words: community structure; compensatory mechanisms; density dependence; diversity; fre-
quency-dependence; growth; mortality; population dynamics; rain forest; seedling recruitment; sub-
tropical; tropical.
INTRODUCTION
are reduced, with the reverse for species that become
rarer, this will continually compensate for the tenden-

A general problem of ecology is how species diversity
cies of some species to increase at the expense of others.

is maintained in natural communities. Since 1963 we
For this reason such general frequency-dependent pro-

have investigated this question using trees in tropical
cesses have been called "compensatory mechanisms"

and subtropical rain forests in Queensland, Australia.
(for general reviews see Connell [1978, 1979]). If such

In this paper we analyze our results to test one general
mechanisms were operating at a site, rare species would

hypothesis concerning the mechanisms by which the
be protected from local extinction, and common species

diversity of tree species could be maintained. The gen-
could not continually increase at the expense of rarer

eral hypothesis is that at any one site, rarer species are
ones. Therefore several to many species could be main-

favored over commoner ones. If, as a species becomes
tained on a site indefinitely, without immigration.

commoner, its rates of recruitment, growth, or survival
This hypothesis assumes that commonness or rarity
is not a property of a species; a species is only tem-
' Manuscript received 12 July 1982; revised 11 February
porarily rare or common on a site. Given time, rare

1983; accepted 9 May 1983; final version received 23 June
1983.
species will tend to increase and common ones to de-
2 Present address: Division of Forest Research, CSIRO,
crease. Over a long period the different species will
Atherton, Queensland, 4833 Australia.
fluctuate within upper and lower bounds that are de-
3Present address: School of Australian Environmental
fined stochastically, i.e., a population will exceed them
Studies, Griffith University, Brisbane, Queensland, 41 1 1 Aus-
tralia. only with some low probability (Chesson 1978).
142 JOSEPH H. CONNELL ET AL. Ecological Monographs
Vol. 54, No. 2
This assumption contrasts with another commonly

pothesis concerning compensation. Hypothesis 2: the
held one that each species has an equilibrium popu-

close proximity of other individuals is more deleterious
lation size toward which it returns if perturbed away.

(i.e., causes slower growth or higher mortality) if they
Under this latter assumption, common species remain

are conspecific than if they are not. In situations such
common, fluctuating around a high equilibrium pop-

as the forests studied here, where rare species are more
ulation size, while rare species do the same around a

intermingled with other species than are common
lower population level. In this framework, compen-

species, this hypothesis will produce compensation. We
satory mechanisms are density-dependent changes in

tested this hypothesis at two scales: between individ-
reproduction, mortality, etc., as the population is either

uals of about the same size, and between young indi-
above, or below, its equilibrium. By contrast, in our

viduals and adult trees.
framework compensation acts in response to the species'

Hypothesis 2 can be tested using data from only
local abundance relative to that of other species, and

common species. If it holds within a species or a size-
hence is frequency dependent rather than strictly den-

class, then a mechanism has been demonstrated which
sity dependent. Either assumption will tend to prevent

would tend to be unfavorable for common species but
local extinction of rare species. The hypothesis to be

not for rare ones. This is not as direct a demonstration
tested in the present paper makes no assumptions about

of compensation as the first hypothesis, but given the
equilibrium levels; it simply suggests that when species

sampling difficulties posed by rare species, this ap-
are relatively rare they will tend to increase, and when

proach seems a good alternative for the smaller spatial
relatively common, to decrease.

scales.
When studying long-lived organisms such as trees,
and particularly tropical species that cannot be reliably
AREA AND METHODS
aged, it is very difficult to document directly increases
The study plots
or decreases in adult populations. Therefore we have
looked for mechanisms that would produce such Area 1: Davies Creek, North Queensland. -This site
changes. We have done this by comparing differences is located at 17'05'S, 145034'E, at -850 m elevation
in recruitment, growth, and mortality with differences on Davies Creek in the Lamb Range, map reference
in abundance, either among species or among samples Bartle Frere CB 535, 155 on the R631 Series 1:10 000
within species. Ideally, a direct comparison should be

topographic survey. The annual rainfall at Mt. Haig,
made between common and rare species. However, the at about the same elevation 10 km south, was 230 cm,
problem with rare species is that it is difficult to obtain

mostly falling in the cloudy, warmer months. (Com-
samples large enough for statistical analyses. We ap-

monwealth Scientific and Industrial Research Orga-
proach this problem in two ways. First, for each study

nization 1982; Fig. 1.) The mean annual air temperature
plot we combined all individuals of a species over the

at Mt. Haig was 18.90C, with a mean annual range of
whole plot, thereby increasing the sample size of the 13.40; at the nearest meteorological station, Atherton,
rarer species. Using these data we calculated recruit-

at 808 m, the same statistics are 19.90 and 9.40 (Com-
ment for each species or growth and mortality for each monwealth of Australia 1971, Commonwealth Scien-
size-class of each species. Then we tested the following

tific and Industrial Research Organization 1982). The
hypothesis of compensation, which we will refer to as

site lies on a slope facing west to southwest and extends
hypothesis 1: on the scale of the whole plot, commoner from a small creek upwards to a ridge 50 m higher.
species have lower rates of recruitment and growth,

Slope varies from nearly level on the ridge to 350 at
and higher mortality, than do rarer ones. These com-

some places on the slopes below. The parent rock is a
parisons are done among species; in the regression of,

deeply weathered gray biotite granite with muscovite
for example, recruitment rate vs. adult abundance, each

and some tourmaline, from which is derived a deep
point represents a single species.

acid brown earth of intermediate nutrient status. The
At finer spatial scales (nearest neighbors and quad-

soils near the creek are deep, while those on the ridge
rats), the sample sizes for rare species are too small, so
are shallow. The mapped plot, 1.68 ha in area, is ap-
the only data one has to work with come from the
proximately square with irregular borders; a contour
commoner species. Our approach to overcoming this

map is shown in Fig. 2. One portion of the plot near
problem was to look for processes that would be del-

the ridge is a 0.4-ha site chosen by Mr. E. Volck of the
eterious for commoner species but not for rarer ones.

Queensland Forestry Department as one of a series of
For example, as an analysis below will show, trees of

"increment plots" to measure growth rates; it is site
rarer species tend to be intermingled with those of other

number 207/4 in State Forest 557 (Dinden). In 1951
species, whereas trees of commoner species more often

all the trees >6 m in height were marked, and their
have neighbors of the same species. Also, those seed-

girths (circumferences) at breast height (gbh) have been
lings arising from seeds carried away from the vicinity

measured at intervals since then. In 1963 we mapped
of the parent tree are more likely to occur near other

the location of these trees and also extended the map
adults of their own species in common than in rare

along the ridge and into the gully below.
species. Given these facts, we formulated the next hy- This plot has its closest affinities with two types of
June 1984 RAIN FOREST TREE DIVERSITY 143
Cn 30- two separate plots of about equal area 0.6 km apart.
Both are in the form of long strips, either 20 or 40 m

< 20 AREA 1
3E20 /+
wide, lying parallel to the contours (Fig. 3). The section
at the lower elevation (880 m) changes direction slight-

10a AREA 2
ly to follow the contour, while that at the higher ele-
60 - I
vation (920 m) is straight, since the ridge at that point
Z 0D
0
is almost level.
E 50AREA 1 This plot is located in a typical evergreen complex
40-
notophyll vineforest on basalt; in a floristic classifi-
30 cation of Australian rain forests (Webb and Tracey
1981 b), it is grouped with the warm wet seasonal sub-
Z 20 1 \
tropical forests having Argyrodendron trifoliolatum and
I10 AREA -
Dysoxylum fraseranum as characteristic canopy trees.
0
However, the altitude of the plot and the presence, as
IJ F 'M AM J JA SON
common trees, of Argyrodendron actinophyllum and
Orites excelsa suggest it is on the transition to cool wet

FIG. 1. Average monthly rainfall and number of days
with rain at stations near Area 1 (Mount Haig, 4-6 yr of subtropical forests. The floristic analysis links these
records) and Area 2 (O'Reilly's Guest House, 18 yr of records).

two groups at the next highest division in the hierarchy.
(Mean ? 2 S.E.)
Similar forests to those found on the Area 2 plot are
found at Binna Burra, Springbrook, Mount Mistake,
evergreen rain forest in North Queensland. The forest

and Mount Tamborine in South Queensland and Rose-
on and near the ridges has structural characteristics of

wood creek on Lower Bellengen River and Mount Wil-
simple notophyll evergreen vineforests while that in

son in Koreela State Forest in New South Wales. The
the gully resembles the complex mesophyll type. A

ecological relationships of these forests are discussed
cluster analysis of 146 sites in North Queensland using

further in Webb and Tracey (1981 a) and McDonald
presence and absence data for 740 species of trees has

and Whiteman (1979). In summary, these analyses in-
been made and will be published elsewhere (Williams

dicate that the two study plots are floristically similar
and Tracey 1984). In this analysis, the Davies Creek

to other Queensland rain forests at equivalent eleva-
plot was split into two sites, data from the ridge portion
tions and physical and climatic characteristics.
being analyzed separately from that from the portion

Both sites have a dense closed evergreen canopy with
in and near the gully. The three classification methods

an average height of -30 m. Basal area is high, 61.5
used (Mulclas, Minimum Spanning Tree, and Net-

and 63.4 m2/ha for Areas 1 and 2, respectively. Except
work) all gave similar results, placing the ridge site with
the simple notophyll vineforests characteristic of gra-
nitic uplands and highlands, e.g., Tinaroo Range, Her-
Scale 0 10 20 30 AREA 1
berton Range, Mount Spec, Kirrama Range (Type 8 830 ~ NORTH QUEENSLAND
metres
on the Vegetation Map of the Humid Tropic Region
840 ~~850 860
of North Queensland [Tracey and Webb 1975]). The
gully site was classified with complex mesophyll vine-
forests (type lb, e.g., Maalan, Boonjie) and complex
notophyll vineforests (type 5a, e.g., Koolmoon creek
840~~~~~~~~~~6
and Cannabullen) on wet cloudy uplands and high-
820 1 ' 4-
lands. Our plot, therefore, is located at the overlap of
rain forest types representative of a wide area of North
Queensland uplands (for further discussion see Tracey
820\~~~~~~~~~~~~~~5
1982).
Area 2: O'Reilly's, Lamington National Park, South
Queensland.-This site is located at 28o14'S, 153010'E,
at 900 m elevation on the Lamington Plateau on well-
drained basaltic krasnozem soils. The rainfall at
O'Reilly's Guest House 1 km distant averages 191 cm/
yr, with most of it falling in the warm months, but
840
with less seasonal variation than at Area 1 (see Fig. 1).
820 830
The mean air temperature at the nearest meteorological
FIG. 2. Map of Area 1 showing contours. Large trees were

station, Mount Tamborine at 654 m, is 17.5?, with a
mapped on the whole plot; medium trees were mapped on
mean range of 8.40 (Commonwealth of Australia 1971).
the wider strips shown along the survey lines; saplings and
The site faces westerly, lying just below the crest of a
seedlings were mapped on the areas shown in shading along
ridge. The mapped plot, 1.94 ha in area, consists of certain survey lines.
Vol. 54. No. 2
AREA 2
cm gbh (medium trees) were mapped on parallel belt
SOUTH QUEENSLAND
transects 6.1 m wide centered on the survey lines; these
constituted 30.3% of the map. Individuals < 8 cm gbh
(saplings and seedlings) were mapped on narrower belt
transects centered on the same survey lines. The lo-
...............................:..:._ .cale: : 0 10 20 30
cations and widths of all transects are shown on Figs.
metres
2 and 3. All individuals of all woody species except
NI
lianes and climbing palms were identified, tagged, and
measured; for saplings and seedlings, the height was
measured rather than the girth. Tags were nailed to
medium and large trees, tied loosely around the base
of smaller individuals. As shown in Figs. 2 and 3, the
FIG. 3. Map of Area 2. The areas on which the different-
strips on which the medium- and smaller-sized trees
sized trees were mapped are shown, as in Fig. 2 (see text).
were mapped encompassed the whole extent of both
The two halves of the plot are separated by ,O.6 km; in this
plots, so they should represent a good sample of the

figure they are placed together to save space. The upper plot
was angled to follow the land contour.

microhabitats on the whole plot on which the large
trees were mapped.
Large and medium trees were mapped between April
and December 1963, together with a small sample of
in treefall gaps, direct sunlight reaches the ground only saplings and seedlings which were mapped on a narrow
in sunflecks (Bjorkman and Ludlow 1972). These sites strip 0.91 m wide along one side of all survey lines on
have been very little disturbed by humans. In Austra- Area 2 only. In August 1965, the sample of saplings
lia, aborigines never practiced agriculture, so these rain and seedlings was increased by expanding the width of
forests have never been cut for shifting cultivation as the strips on Area 2 to 3.6 m; this was also the first
have those in most other regions of the world. Com- time that this small size-class was mapped on Area 1.
mercial timber cutting began in the vicinity of Area 1 Censuses were repeated on precisely the same mapped
in the early 1950s. At that time the site was set up as areas in February 1967, and in August or September
a growth measurement plot and protected from all cut- of 1969, 1971, 1974, and 1978 and, as described below,
ting or treatment by the Queensland Forestry Depart- in 1980-1981. At each census, notes were made of all
ment. In the region of Area 2, a few trees may have individuals that had died or been damaged since the
been removed before the area became a national park previous census. In addition, except for the 1967 cen-
in 1915. However, no evidence of previous cutting has sus, all new seedlings that had appeared since the pre-
ever been found on the Area 2 mapped plot. An old vious census were mapped, tagged, measured, and
bridle track, used to reach the guest house before the identified. All treefalls and any light gaps produced by
road was built in the mid 1 930s, runs diagonally through them were also mapped over the whole plot in the
the upper part of the site; it has not been used since initial census of 1963 and at each subsequent census.
then and is overgrown but still detectable. In August and December 1980, the censuses for mor-
tality were repeated on areas 2 and 1, respectively, and
Field methods
in September 1981, for mapping of new seedlings on
Mapping and periodic censuses. -Within each of the both areas. In 1978 on area 1 and in 1980 on area 2,
plots the trees were mapped as follows. Several parallel the sizes of all surviving individuals were remeasured.
survey lines were laid out 20.1 m apart, with steel Identification of species in rain forest is often diffi-
stakes every 7.5 m along each line. Rectangular coor- cult, particularly of the young seedlings. J. G. Tracey
dinates were measured to each tree as follows. A tape

has learned to identify the smaller size-classes by doing
measure stretched along the line served as one coor- many years of extensive fieldwork and by germinating
dinate, and the distance at right angles from it to the many of their seeds in the laboratory. This work, plus
tree, the other. The latter was measured using an optical extensive collecting and consultation with taxonomists
range finder accurate to within -0.4 m at 10 m dis-

of the Queensland Herbarium, makes us confident in
tance, more accurate at shorter distances. One person the identification of all of our material on Area 2 and
stood beside the tree with a battery light attached to most of it on Area 1. On Area 1, - 3% of the seedlings
his belt. The light was used as the target for the range
have not been identified to species; these have usually
finder since it could easily be seen through the dense been identified to genus or family, and are not included
foliage. A right-angle prism and mirrors in the range in the present analyses.
finder enabled the observer on the survey line to orient Field experiments. -In addition to the periodic cen-
the line of sight at right angles to the line. For distances suses, field experiments were done at various places
<3 m from the line, a tape measure was used. either on the mapped plots or just outside them to test
All trees -32 cm gbh (hereafter referred to as large hypothesis 2. Seeds were either planted or placed on
trees) were mapped on the whole plot, those 8-31.9 the surface near and far from adults, and at high and
low density. We did the experiments using those seeds

the Introduction, we needed to calculate rates of re-
that happened to be falling at the times of our visits;

cruitment, growth, and mortality for each species sep-
therefore, only species with seeds large enough and

arately. Recruitment for each species was defined as
common enough to be collected easily from the forest

the number of new seedlings that became established
floor were used. In some instances, cages were placed

between 1965 and 1981 on the whole plot. Recruitment
over seeds or over existing seedlings to see whether

rate was defined either as this number per species, or
herbivores might be responsible for differences in

as a per capita rate calculated by dividing it by the
growth and mortality. The cages were made of either

number of adults of the species on the plot. Since cen-
large-meshed (1 cm diameter) wire screening to exclude

suses were repeated at intervals of 2-4 yr, the new
vertebrates and large insects, or fine-meshed nylon

seedlings that became established between censuses
stretched over large mesh to exclude small insects as

were the survivors of an unknown larger number that
well. Controls for the effects of the caging itself were

had germinated in the interval. If seedlings of a species
either roofs of large mesh with two open sides, or large

happened to germinate soon after a census, they would
meshed cages with the fine nylon covering only the

be exposed to the risk of mortality over a longer period
upper half. Since these cages were used in shade or in

to the next census than seedlings of another species
small treefall gaps in which the sunlight came only from

that happened to germinate just before the census. Thus
directly above, these control roofs probably had the

in this paper "recruitment" refers to the addition of
same shading effect as the complete cages. Both the

seedlings which have experienced an unknown amount
cages and the controls excluded most of the litterfall;

of early mortality between germination and the time
at each examination, we placed litter inside the cages.

they were first mapped. Since seedlings were mapped
Results of the experiments are given in Table 12.

on only part of each plot, this rate is not a measure of
To study possible competitive effects of neighboring

total recruitment per adult. However, it is accurate for
plants, root competition was experimentally reduced.

comparing different species, as used here. Adults were
Trenches -25 cm deep were dug around some plots,

defined as trees -32 cm gbh for species which normally
cutting the roots to prevent larger plants outside from

reach the canopy and ?8 cm gbh for species which
removing soil nutrients or water from the seedlings

normally exist as subcanopy and understory trees.
within the plots. Most of the roots were < 1 0 cm deep;

(Some of the latter flower at a very small size in heavy
our trenches therefore probably cut the majority of

shade.) We excluded from the latter category the few
surface roots, although we cannot rule out the possi-

light-demanding ephemeral species that do not persist
bility of some large deeper roots being left intact. We

in the understory of these forests. Species were placed
then lined the trench with plastic sheeting before re-

in these categories using the extensive experience of J.
turning the soil; this allowed us at intervals to cut any

G. Tracey and L. J. Webb in many Queensland rain
new roots growing back into the plot along the plastic.
forests. Recruitment rates for each species, as defined
above, were then regressed against the abundance of
Analytical methods adults, expressed either as the number or basal area of
that species over the whole plot. The former empha-
The diversity of species in each size-class was cal-
sizes the influence of the more numerous smaller trees,
culated using several different indices. The first is the
whereas the latter emphasizes the influence of the fewer
number of species, S, which gives all species equal
larger trees. All species having at least two adults or at
weight. The second is the Shannon-Wiener index, which
least 0.01 m2 of adult basal area, and at least 10 seed-
takes into account both the number and relative abun-
lings, were included; they are listed in the Appendix.

S
dance of species. Its equation is H' =- P I In pi, in

Growth and mortality rates for each species sepa-
rately were calculated as follows. Because these rates
vary a great deal with the size of individuals, we sep-

which pi = n/ ni, and ni refers to the number of
arated the trees into five size-classes: seedlings (<31
species i in that size-class on the study plot. We used
cm high), small saplings (31-77 cm high on Area 1,
exp(H') as suggested by Peet (1974). As a measure of
31-108 cm on Area 2), large saplings (from the upper
evenness of relative abundance we calculated E =
limit of small saplings to the height, ranging from 3.5
[exp(H')]/S (Hill 1973). Lastly we calculated an index
to 6 m, at which they reach 8 cm gbh), medium trees
of pattern diversity, D, as described by Pielou (1966).
(from 8 to 31.9 cm gbh), and large trees (32 cm gbh
We used each individual and its first- and second-near-
and greater). The limits of these classes were decided
est neighbors as a triplet to calculate D. Each such

as follows. New seedlings appearing in each of the later
triplet could be composed of 1, 2, or 3 species. The
censuses (and therefore never older than 2-4 yr) were
index represents a comparison between the observed

almost always <31 cm in height, so for the original
distribution of triplets in these three categories and the

mapping in 1965, this was taken as their upper limit.
expected distribution if the triplets were drawn at ran-

To obtain sufficient sample sizes, the division point
dom from the total population.
chosen between the two sapling classes separated them
For the purpose of testing the hypotheses given in
into approximately equal-sized groups; this point was
Vol. 54, No. 2
different in Areas 1 and 2. The medium- and large- bor is beyond the line. To calculate this probability we
tree classes were mapped on different-sized areas, so drew an imaginary circle around each central tree with
they needed to be analyzed separately. a radius equal to the distance to the nearest neighbor.
The seedlings were also separated into six classes, If the central tree was closer to the boundary than to
according to the years in which they were initially the neighbor this circle would intersect the boundary,
mapped, i.e., the censuses in which they first appeared: and we took the proportion of the area of the circle
1965, 1969, 1971, 1974, 1978, and 1981 (all these within the map as the probability that the measured
censuses were done in August or September). The 1965 neighbor was the true one. We then used only those
seedlings were simply the individuals <31 cm height central trees which had at least a 0.9 probability that
present at the initial census, whereas the last five classes the measured neighbor was the true one.
contain only those seedlings that had become estab- As a check on the accuracy of our results, we also
lished during the interval since the previous census. found the second nearest neighbor of each central tree,
For each class, growth and mortality rates were cal- and carried out the analysis on both first and second
culated from the date of original mapping to the final nearest neighbors. Since the second was usually also
census in 1980 or 1981. Although some individuals quite close to the central tree, it probably had only
grew into a larger size-class during this period, for these slightly less influence than the first. Therefore we felt
calculations they were included in their original size- that if the results were similar for both first and second
or year-class. nearest neighbors, we could rely on them as being more
The problem of a sufficient sample size at the smaller accurate than if only the first nearest neighbor had been
spatial scales is a particularly difficult one in these di- considered. These neighbor analyses were the only in-
verse forests. To ensure objectivity, only species sat- stances in which different species were combined be-
isfying the following criteria were analyzed. For all fore calculating a rate.
regressions there must have been at least five different We did a second analysis to test hypothesis 2, com-
points on the graph, and both variables must have had paring the growth or mortality rate of a species with
a range of at least five units. For rates of growth and either its abundance or the combined abundance of all
mortality, a sample size of at least two and five indi- other species in quadrats. As discussed above, common
viduals, respectively, was required. species tend to have conspecifics as neighbors, whereas
To test hypothesis 1 for growth or mortality, the rarer species more often have different species as neigh-
following regression for each size- or year-class was bors. Hypothesis 2 predicts that local crowding by
calculated. The growth or mortality rate of a species members of the same species will have a greater del-
was plotted against the abundance of individuals either eterious effect on growth and survival than crowding
of the same size-class or of adults of that same species by members of other species. Therefore, if such effects
over the whole plot. Thus each point in these regres- occur commonly among species, they should produce
sions refers to a single species so that the regression a compensatory trend, favoring rarer over commoner
compares all species with a sufficient sample size within species. The quadrats consisted of contiguous rectan-
that size- or age-class. gular sections of the belt transects, 1.8 x 12.2 m; they
Hypothesis 2 states that the close proximity of other had to be this large to obtain a sufficient sample size,
individuals is more deleterious (causes slower growth given the low densities of most species. Each regression
or higher mortality) if they are the same than if they analysis refers to a single species, with the dependent
are different species. To test this hypothesis we did two variable the growth or mortality rate of the given species,
sorts of analyses. In the first we compared the growth and with two independent variables, the abundances
or mortality rates of individuals whose nearest neigh- of the same species and of all different species. Multiple
bors were either the same or different species. Since regression analysis was used to evaluate the effect of
we were interested in the influence of a neighbor, we each independent variable after statistically removing
did not consider an individual to be a neighbor unless the effect of the other one.
it was about equal in size to, or larger than, the one Our criteria for minimum sample size for this anal-
being considered. For this reason we recorded as a ysis were at least five quadrats, each having, for growth
neighbor the nearest individual that was at least 0.9 rate, at least two individuals of the size-class being
times the size of the central individual under consid- considered, and for mortality, five individuals. Mor-
eration. This assumes that much smaller neighbors ex- tality of the seedling classes was calculated only over
ert little influence on a plant, which we regard as a the first census interval after mapping, since the den-
reasonable assumption. Also, if the central tree was sities of seedlings changed rapidly due to high early
closer to the boundary of the map than to its nearest mortality; mortality of sapling and larger classes was
neighbor within the boundary, there was a degree of calculated over the period 1965-1980. The abundances
uncertainty, since there might have been an even closer were calculated as either the numbers or the sum of
unknown neighbor just beyond the boundary. If, for squared sizes either for the same size-class as the de-
example, the central tree is directly on the boundary pendent variable or for all individuals larger than that
size-class, averaged over the period for which mortality

line, there is a 50% chance that the actual nearest neigh-
20 -
scale of proximity to adult trees as follows: we defined
the immediate vicinity of the adult as a concentric
H-
r 1 5 _-8
circle of radius 1.5 times the radius of its crown. An
annulus outside this circle, of width equal to its radius,
1 0 0? 00 * i
was taken as being under lesser influence of the adult
Z 00 ~ ~~~~~ 00 0 0 0
than the inner circle. Small trees beyond this outer ring
were expected to be even less influenced by the adult.
Q I IA) 0 I 0
The tree's crown radius was calculated from a regres-
sion of crown diameter on the gbh, for a subsample of
0 40 80 120 160 200 240 280 320

trees measured in the field; this relationship is shown
GIRTH BREAST HEIGHT (cm)

in Fig. 4. Since rectangular coordinates had been re-
corded for all the trees, the computer simply compared

FIG. 4. The relation of crown diameter to girth of trees
on Area 2: 0 Argyrodendron actinophyllum, C Orites excelsa, the distance between each adult tree and each seedling
o several other species. The regression equation for large trees
or sapling to either the radius of the adult's inner circle
is y = 7.28 + 0.33 x; that for medium trees is y = 1.05 +
or twice that radius, to see whether the young tree was
0.84 x.
in the inner circle or outer ring, respectively. This pro-
cess was repeated with all the adult trees of a species.
If a juvenile tree occurred in the inner circle of one
or growth was calculated. However, because a single

adult and in the outer ring of another, it was assigned
species was almost always much less abundant than all

to the inner ring; if neither, it was assigned to the "far"
other species combined in a quadrat, this meant that

category. Thus each juvenile was categorized as to its
we were usually comparing regressions in which the

proximity to the nearest adult of its own species. Only
range of the two independent variables overlapped lit-

those adults alive at the start of a census interval were
tle if at all. In the usual case the number of individuals

used for seedlings that became established in that in-
of the same species ranged from 1 to 10, while those

terval. The mortality rates of the groups in the inner
of all other species ranged from 5 to 100 or more.

circle vs. the outer ring, or of these two groups com-
We also tested the second hypothesis at the larger

bined vs. individuals farther away, were then com-
TABLE 1. Sample sizes and diversity indices for different size-classes at the date of mapping and of the survivors in 1980.
See Analytical Methods for definitions of size-classes. The seedlings of 1965 were those present at the initial mapping in
1965 that were <31 cm high; each other seedling class represents only those newly established in the interval since the
previous census. The 1980 data refer to survivors of those shown on the same line on map date. The sample size used to
calculate pattern diversity is smaller than the N shown here because some individuals were closer to the border of the map
than to the first- or second-nearest neighbor (see Table 13). See text for definitions of indices.
On map date Survivors of each size-class in 1980
No. of No. of
individu- No. of Diver- Even- Pattern individu- No. of Diver- Even- Pattern
Year als species sity ness diversity als species sity ness diversity
Size-class mapped (N) (S) exp(H') (E) (D) (N) (S) exp(H') (E) (D)
Area 1, North Queensland
Large trees 1963 1371 120 59.1 .49 .98 1224 116 56.3 .48 .98
Medium trees 1963 928 123 56.8 .46 .98 819 120 55.1 .46 .99
Large saplings 1965 1149 130 61.6 .47 .97 979 121 60.3 .50 .97
Small saplings 1965 1149 109 45.6 .42 .94 806 99 44.3 .45 .96
Seedlings 1965 1221 101 34.8 .34 .85 516 84 40.0 .48 .93
Seedlings 1969 1823 99 16.8 .17 .85 447 73 25.3 .35 .96
Seedlings 1971 2036 103 26.3 .26 .85 275 70 38.1 .54 .93
Seedlings 1974 1189 93 39.2 .42 .92 173 62 43.0 .69 .96
Seedlings 1978 4373 99 8.7 .09 .78 2458 85 4.6 .05 .99
Seedlings 1981 6016 97 8.1 .08 .51
Area 2, South Queensland
Large trees 1963 1295 73 27.7 .38 .97 1140 69 26.3 .38 .97
Small trees 1963 1238 76 27.1 .36 .98 1117 75 26.6 .35 .99
Large saplings 1965 1087 69 30.0 .43 .96 962 67 29.4 .44 .98
Small saplings 1965 1255 63 26.3 .42 .93 970 60 24.5 .41 .94
Seedlings 1965 744 48 21.8 .45 .85 311 40 21.5 .54 .96
Seedlings 1969 491 47 25.3 .54 .91 217 40 26.3 .66 .96
Seedlings 1971 3477 52 7.5 .14 .72 268 38 17.6 .46 .96
Seedlings 1974 330 33 18.5 .56 .83 90 23 16.3 .71 .97
Seedlings 1978 920 41 5.1 .12 .97 251 34 9.7 .28 1.05
Seedlings 1981 2612 33 4.7 .14 .68
Vol. 54, No. 2
AREA 1
RESULTS
(a) AREA 1 SUBCANOPY
CANOPY SPECIES PLUS UNDERSTORY
Species diversity
,, 1000 1 00
z @..
Seedlings had lower diversity than did larger indi-
viduals, for all indices on both sites (Table 1), due

wJ 100 * 10 3.
0 *< mainly to the fact that in the short interval (2-4 yr)
0 rr~~~~~~~~~~
between censuses, not all species in the larger classes
10 s* 1-
recruited seedlings. In the 16 yr between 1965 and 1981
m . * .00
about half of the species of large trees contributed al-

D r r=-0.25 r=-0.76
Z I , , I 0.1 I I I_ l l l l l _ _ , l
most all the seedlings. By 1980, after a considerable
1 10 100 1000 1 10 100 1000
amount of mortality had occurred, the number of

NUMBER OF ADULTS
species and diversity of seedlings had declined, but
evenness and pattern diversity had risen almost to that
AREA 1
(b) AREA 1 SUBCANOPY of the larger classes. We will consider the causes of the
increases in evenness and pattern diversity of seedlings
E 1000 * 10,000 later in the paper.
U)w U. 0
OK~~~~~0 *X
_00
Intermingling of common vs. rare species
I 100 *: 1000 *
0<~~0
If individuals are distributed randomly, commoner
U)mc
species will tend to have conspecific neighbors more
0 D 10 _ 100 _
often than will rarer species. Alternatively, rare species
might be more highly clumped, and thus, despite their
U_ L _ r =-0.27 r =-0.83
DO 1 l , l l 10o, I,, 1 I I I _ I I ,,,

rarity, be as likely to have conspecifics as neighbors as
Z 0.01 0.1 1 10 0.01 0.1 1 10
common species. To check this we compared the pro-

BASAL AREA OF ADULTS (m2)
portions of all neighbors that were the same species.
The null hypothesis was that these proportions for
FIG. 5. Seedling recruitment of species of various adult
common vs. rare species are equal. Since neighbors
abundances on Area 1. a) Seedlings per adult in relation to
were chosen only from trees at least 0.9 times the size
the number of adults of the same species b) Seedlings per
square metre of adult basal area, in relation to total basal area

of the central tree, we used as the measure of abundance
of adults of the same species. Recruitment is the total number
of new seedlings of a species appearing at all censuses between
1965 and 1981. All species were included which had at least
AREA 2
two adults, a total basal area of at least 0.01 m2 and at least
(a) AREA 2 SUBCANOPY

10 seedlings.Where the slope is significantly different from
zero, the regression line is included.
Ca) 100 100
Z*
D 10 . 10
Cf) C
0 Cr1 t < 0
pared, using the G test; all species with sufficient sample
size for this test were included.
The analyses described so far have all correlated suc- D r -0.24 r -0.70
0.1 10I | , , 0, 0 0.1 I , - I,_l_ l_ ,l _ , , j

cess in recruitment, growth, and survival with abun-
1 10 100 1000 1 10 100 1000
dance or proximity to other individuals of the same
NUMBER OF ADULTS
species. A difficulty with this approach is that each
AREA 2
comparison consists of several different observations
(b) AREA 2 SUBCANOPY
(points in a regression, etc.), each of which is in a

5000 CANOPY SPECIES 5000 PLUS UNDERSTORY
different location within the study plot, often using
51000 50000
trees of different ages, in quadrats within which local
) 1000 1000
(Dinr 00
densities vary a great deal, etc. Such variability can be
Z <~~~~~~~
reduced by doing field experiments in which proximity
<U) 100 * 100 \
to neighbors is held constant, all individuals are of the Uj <al10 _ 10 _
cf) Co
wH-
same age, and replicates are set up in the same locality.
A disadvantage of such experiments is that few species

0 10 10 1 0
Lj <
are available, i.e., only those having large seeds or fruits
0 r -0.1 1 *r -0.79
which happened to be falling at the time of our visits
Z i 1 i 1 1 I_ I_ _ _ I_ _ I_ _ _ I_ I I_ _
0.1 1 10 100 0.01 0.1 1 10

to the site. In contrast, the statistical analyses cover a
BASAL AREA OF ADULTS (m2)
broader range of species and ages, using every species
that had a sufficient sample size. Thus, each method
FIG. 6. Seedling recruitment of species of various adult
complements the other.

abundances on Area 2. (See Fig. 5 for format and details.)
TABLE 2. Seedling recruitment rate vs. abundance of adults. In canopy species, adults were defined as trees -32 cm gbh;
for subcanopy and understory species, they were trees -8 cm gbh. Both variables were transformed to logarithms. All
species were included that had at least 10 seedlings and either two adults or an adult basal area (b.a.) of 0.01 M2.
Dependent variable (recruitment rate)
Absolute no. of seedlings No. of seedlings per adult
Independent variable mapped 1965-1981 or per m2 adult b.a.
Independent variable
(adult abundance) Tree stratum No. spp. r No. spp. r
No. of adults Canopy 31 .42* 31 -.25
Subcanopy 13 .21 13 -.80**
Understory 13 .11 13 -.83**
Subcan + Underst 26 .04 26 -.76**
Adult basal area (M2) Canopy 33 .37* 33 -.27
Subcanopy 12 .63* 12 -.85**
Understory 8 .22 8 -.87**
Subcan + Underst 20 .27 20 -.83**
No. of adults Canopy 14 .75** 13 .24
Subcanopy 7 .35 7 -.83*
Understory 6 .84* 6 -.49
Subcan + Underst 13 .59* 13 -.70**
Adult basal area (M2) Canopy 14 .65* 13 -.11
Subcanopy 7 .54 7 -.86*
Understory 5 .35 5 -.57
Subcan + Underst 12 .36 12 -.79**
* .05 > P > .01, ** .01 > P > .001.
all trees of the same size-class or larger that occurred

2). In all comparisons of canopy species, the commoner
in the narrow strips in which the seedlings and saplings

ones had more seedlings than did the rarer ones. For
were mapped. The category of "commoner species"

subcanopy and understory species analyzed separately,
was determined by ranking species in order of abun-

six of the eight comparisons showed no differences
dance and cumulating their numbers in order of rank between common and rare species.
until at least half of the total number of individuals in

These rates refer to species with at least two adults
that size-class on the plot was reached. For every size-

and in which 10 or more seedlings became established
class except the 1974 seedlings, the group of commoner

in the 16-yr period since 1965. There were 60 such
species had a significantly higher percentage of neigh-

species on Area 1 and 26 on Area 2; they are listed in
bors of the same species (for different size-classes, it

the Appendix. However there were many other species
ranged from 8 to 46% of neighbors) than did the rarer

with two or more adults in which <10 seedlings be-
species (ranging from 1 to 10%), judged from chi-square

came established during this interval. There were 76
tests. The results for both first and second nearest

such species on Area 1 and 35 on Area 2. They form
neighbors were consistently the same. a significant portion of the tree biomass on the study
sites; on Area 1 they constituted 30.5% of the basal
Recruitment of common vs. rare species
area, on Area 2, 29.4%. They apparently require, for
on the whole plot
seedling establishment, circumstances that did not oc-
For subcanopy and understory species combined,

cur during the study period. Some clearly require more
recruitment rates per adult were lower for common

light than is received in the treefall gaps that occurred
than for rare species, as predicted by hypothesis 1 (Figs.

on the plots, or more-disturbed soil, or both, since their
5 and 6), suggesting a very strong tendency toward

seedlings were common along roadsides near the plots.
compensation on both study areas. When the two strata

In other species, seedlings were never found in any
were analyzed separately, subcanopy species showed

numbers on or off the plots.
this trend in both study areas, but understory species
Growth in relation to size of individuals
did so only on Area 1 (Table 2). Canopy species showed
no compensatory trend on either study area. This sug-

Because growth rates may vary with size of individ-
gests that common species should have more seedlings

ual, we looked for such a relationship in all trees within
than will rare species on the plot as a whole. Compar-

each of the size-classes. There is a significant relation-
ison of the total number of seedlings appearing between

ship between growth and size within some size-classes
1965 and 1981 for each species with the average adult
when all species are lumped, but it is weak, judging by
abundance during that period demonstrates this (Table

the low coefficients of determination (Table 3). Two
Vol. 54, No. 2
TABLE 3. Growth rate as a function of individual initial size. Within each size-class all individuals, regardless of species,
were analyzed together. Growth increment measured from the year that size-class was mapped to 1978 (area 1) or 1980
(area 2); number of measurements equal to the number of survivors in 1980 (Table 1). Dimensions of large and medium
trees are girths, of saplings and seedlings, heights. * .05 > P > .01, ** .01 > P > .001, *** .001 > P > .0001.
Area 1 Area 2
Growth incre- Mean Regression Growth incre- Mean Regression
ment (cm) int ssioncm) initial statistics
Y ear __ _ _ _ _ _ _ _ size size__ __ _ _ _ _ _ __ _ _ _ _ __ _ _ _ _ __ _ _ _ _ _
Size-class mapped Mean SD (cm) r2 Slope Mean SD (cm) r2 Slope
Large trees 1963 4.68 6.39 75.4 .114 .043*** 3.68 5.90 76.5 .043 .026***
Medium trees 1963 1.09 1.90 15.9 .023 .045*** 1.34 2.07 14.8 .002 .017
Large saplings 1965 7.13 53.34 180.7 .021 -.087*** 12.60 75.58 197.2 .013 -.128**
Small saplings 1965 15.24 32.70 52.4 .0005 -.052 26.66 46.31 68.0 .003 -.114
Seedlings 1965 11.76 17.46 22.6 .005 .221 23.97 24.38 21.3 .00001 .019
Seedlings 1969 7.04 12.90 16.1 .011 -.210* 27.41 40.87 16.8 .047 1.19**
Seedlings 1971 6.77 8.60 10.1 .009 -.145 13.40 20.88 11.8 .004 .159
Seedlings 1974 5.21 11.37 9.7 .030 -.359** 8.04 15.02 10.6 .025 .431
significant relationships were consistent in both Area

Growth in common vs. rare species
1 and 2. First, the taller individuals in the large-sapling

on the whole plot
class grew more slowly than the shorter ones. This
reflects the common observation that during the period

To test hypothesis 1 we compared the growth of
1965-1980, many of these taller saplings were seen to

seedlings of a species with the abundance of adults of
have died back, or to have broken and then resprouted.

the same species. We analyzed canopy and subcanopy
Second, in the largest size-class, larger trees grew faster

species separately to see whether the differences found
than smaller ones. This is the size-class in which trees

in recruitment were maintained during later growth.
range from a subcanopy to a canopy position, passing

Because of the different periods of growth of different
from limited to full light conditions, which presumably

seedling year-classes, each was analyzed separately.
allows them to grow faster. In general the correlation

These seedlings were the survivors of the ones used in
coefficients were very small in all but the largest size-

the recruitment analyses (Table 2, Figs. 5 and 6). Only
class on Area 1, so that we felt it would be reasonable

one of the regressions was significant (Table 4); clearly
to use the mean growth rate within a size-class in our

the abundance of adults has little effect on growth of
regression analyses.
their young seedlings on the scale of the whole plot.
TABLE 4. Correlations between the abundance of adults and either the growth or the percentage mortality of seedlings of
the same species.
Adult numbers Adult basal area
Year Canopy species Subcanopy species Canopy species Subcanopy species
seedlings No. of No. of No. of No. of
Area mapped species r species r species r species r
Seedling growth rate vs. abundance of adults of same species
1 1969 20 .25 15 -.18 20 .31 10 -.03
1 1971 16 .10 16 .002 16 .04 11 -.14
1 1974 15 .15 14 .09 15 .13 11 -.18
2 1969 12 -.27 11 .46 12 -.24 10 .73*
2 1971 9 .44 9 .63 9 .11 9 .26
2 1974 6 -.78 6 .30 6 -.52 6 .58
2 1978 7 -.27 7 .44 7 .47 6 -.21
Seedling mortality vs. abundance of adults of same species
1 1969 25 .30 18 -.09 25 .33 18 .13
1 1971 24 .06 20 .39 24 .24 20 .16
1 1974 19 .11 17 .03 19 .07 17 .01
2 1969 12 .47 11 .39 12 .32 10 .19
2 1971 12 .41 12 -.06 12 .37 12 .24
2 1974 8 .41 7 -.11 8 .15 7 .28
2 1978 9 .23 9 -.24 9 .25 8 -.24
*.05 > P > .02.
TABLE 5. Multiple regressions of growth rates of a species vs. abundance and average individual size, within the same size-
class over the whole plot of Areas 1 and 2. Growth rates of large and medium trees are in girth, and for saplings and
seedlings, in height. X2 is the same in both of the multiple regressions shown.
Slope of multiple regression
Regression A Regression B
Average Average sum Average
Growth rate per sp. (cm) Average no. individual of squared individual
Year No. of individuals size sizes size
Size-class mapped species Mean Range (XI) (X2) (XI) (X2)
Area 1
Large trees 1963 85 5.03 -2 to 18 -.005 .044** -.00001 .045**
Medium trees 1963 88 1.19 -1 to 5 -.002 .051 -.0001 .054
Large saplings 1965 91 11.80 -108 to 98 -.007 .029 -.0002 .044
Small saplings 1965 77 21.18 -26 to 160 -.008 .488 -.002 -.434
Seedlings 1965 57 12.60 -5 to 49 .001 .060
Seedlings 1969 46 7.53 -2 to 25 -.0002 -.108
Seedlings 1971 44 8.50 -1 to 37 -.002 .052
Seedlings 1974 44 7.71 -2 to 118 -.007 .317
Seedlings 1978 23 10.70 3 to 26 .0001 -.984
Area 2
Large trees 1963 51 4.01 0 to 17 -.004 .063** -.0001 .070**
Medium trees 1963 61 1.57 0 to 7 -.001 .080 -.00004 .084
Large saplings 1965 62 22.19 -55 to 137 -.011 -.274* -.0002 -.234
Small saplings 1965 56 34.44 -37 to 140 -.006 -.199 -.001 .168
Seedlings 1965 26 26.00 -2 to 71 -.006 .826
Seedlings 1969 34 31.09 -5 to 165 -.016 2.46
Seedlings 1971 26 13.69 -8 to 36 .0003 .055
Seedlings 1974 13 5.52 Oto 11 .005 .024
Seedlings 1978 25 5.55 0 to 44 -.001 .080
* P = .05; ** .001 > P > .0001 that slope is equal to zero; all other P > .05.
TABLE 6. Relation between mortality of a species and its mean abundance (number or sum of squared sizes) within the same
size or year-class over the whole plot. Mortality and mean abundance calculated from year each size class was mapped to
1980. Dimensions of large and medium trees are girths, of saplings and seedlings, heights. The values given in the range
of sums of squared sizes are m2 x 10.
Relationship to mortality
Numbers Sum of squared sizes
No. % mortality
of per species Range Range
Year spe- initial no. z (sizes
Size-class mapped cies Mean Range per sp. r2 Slope per sp.)2 r2 Slope
Area 1
Large trees 1963 60 12.6 0-67 5-116 .044 -.001 1.04-169.51 .045 -.0001
Medium trees 1963 50 13.1 0-45 5-81 .015 -.001 0.07-4.20 .023 -.0001
Large saplings 1965 53 16.2 0-50 5-105 .022 -.001 0.12-3.40 .073 -.000 *
Small saplings 1965 56 28.4 0-80 5-120 .001 -.0003 0.01-0.40 .0002 -.0001
Seedlings 1965 44 50.3 0-89 5-161 .029 .002
Seedlings 1969 51 68.1 18-100 5-765 .003 .0002
Seedlings 1971 52 79.7 33-100 5-554 .004 .0007
Seedlings 1974 44 82.3 40-100 5-145 .012 .002
Seedlings 1978 58 59.2 0-100 5-2343 .047 -.0004
Area 2
Large trees 1963 41 13.2 0-50 5-230 .028 -.0005 1.88-316.26 .036 -.0000
Medium trees 1963 41 11.4 0-54 5-241 .010 -.0003 0.10-9.25 .006 -.0000
Large saplings 1965 53 14.2 0-68 5-143 .003 -.0002 0.17-6.04 .003 -.0002
Small saplings 1965 41 28.7 0-91 5-140 .062 -.002 0.02-0.77 .079 -.0004
Seedlings 1965 23 49.5 0-86 5-52 .006 .002
Seedlings 1969 29 51.9 17-82 5-62 .007 .002
Seedlings 1971 31 79.4 47-100 6-1635 .049 .0009
Seedlings 1974 16 67.6 37-100 8-40 .017 .005
Seedlings 1978 21 51.6 0-91 5-615 .031 .003
* .05 > P > .01.
Vol. 54, No. 2
TABLE 7. Growth rate in relation to the species of first- and second-nearest neighbor. Growth rates (height in cm) apply
over the period from the year mapped to 1978 (area 1) or to 1980 (area 2); all measurements were made in August or
September. F and t statistics compare samples from same vs. different species. If F was significant, the t test applied was
that for unequal variances (Winer 1971).
Neighbor same species Neighbor different species
Mean Mean
Year Nearest growth growth
Size-class mapped neighbor (cm) SD n (cm) SD n F t df
Area 1
Large saplings 1965 First 12.3 32.2 38 5.4 49.6 654 2.37* 1.23 48
1965 Second 12.3 36.5 30 5.1 49.0 504 1.80 0.79 537
Small saplings 1965 First 11.1 28.8 50 14.0 30.1 610 1.10 0.66 658
1965 SecOnd 10.5 14.7 55 12.6 24.2 520 2.70* 0.92 88
Seedlings 1965 First 8.1 15.1 63 11.9 16.3 441 1.17 1.71 502
1965 Second 11.4 15.5 52 11.2 16.6 399 1.15 0.07 449
Seedlings 1969 First 5.9 10.0 89 6.9 13.6 371 1.82* 0.81 174
1969 Second 6.6 23.1 70 6.8 10.5 352 4.89* 0.05 75
Seedlings 1971 First 4.4 4.8 46 6.9 8.7 262 3.33* 2.84** 107
1971 Second 4.8 4.6 45 6.9 8.9 242 3.81* 2.35* 118
Seedlings 1974 First 1.8 3.9 23 7.0 21.4 208 30.6* 3.07** 191
1974 Second 2.1 3.1 13 6.2 20.2 200 41.3* 2.47* 113
Area 2
Large saplings 1965 First 4.0 83.5 46 9.5 74.1 546 1.27 0.48 590
1965 Second 21.3 78.6 36 8.9 74.2 443 1.13 0.96 477
Small saplings 1965 First 17.8 30.2 78 24.5 46.6 626 2.37* 1.71 128
1965 Second 18.9 34.4 80 23.8 47.9 531 1.93* 1.12 130
Seedlings 1965 First 17.0 15.1 27 25.6 26.0 225 2.96* 2.53* 47
1965 Second 27.3 21.7 28 25.0 26.2 207 1.45 0.45 233
Seedlings 1969 First 21.7 28.7 17 27.3 40.4 152 1.99 0.56 167
1969 Second 22.6 49.2 17 27.8 40.0 136 1.51 0.49 151
Seedlings 1971 First 16.7 25.1 39 13.0 19.9 155 1.58 0.86 192
1971 Second 11.9 16.9 42 13.8 20.2 135 1.43 0.56 175
Seedlings 1974 First 8.7 5.9 7 8.0 17.4 56 8.55* 0.22 23
1974 Second 3.7 6.6 5 9.0 17.0 56 6.63 0.70 59
Seedlings 1978 First 5.3 12.2 45 4.3 4.9 116 6.19* 0.50 50
1978 Second 4.7 4.0 33 4.7 8.7 116 4.76* 0.03 117
* .05 > P > .01.
** .01 > P > .001.
We then compared growth in each species with abun-

Growth rate in relation to nearest neighbors
dance and average size within the same size-class.
Before we compared the effects of species of neigh-
Growth was never correlated with abundance (Table
bor, we tested to see whether distance to neighbor had
5). In the largest size-class, species with trees of larger
any effect on growth. There was no effect of distance
average size grew faster on both study areas. In the
to either first or second nearest neighbor on growth in
case of large saplings on Area 2, species of larger av-
44 of the 48 comparisons on both study areas. When
erage size grew more slowly than those of smaller size.
there was an effect of distance it was not consistent in
direction, so we felt that it could be ignored in the
Mortality of common vs. rare species
analysis.
on the whole plot
To test hypothesis 2 we compared the growth rate
We found no relationship between the abundance of

of individuals with conspecific neighbors with that of
adults of a species and the mortality of new seedlings

individuals having different species as neighbors (Table
of the same species in either study area (Table 4). Also,

7). On Area 2 there was no effect of species of neighbor
when mortality was compared to abundance within the

on growth rate for 11 of the 12 comparisons. This single
same size-class (Table 6) there were no significant re-

significant finding indicates that any compensatory
lationships on either Area 1 or Area 2, and for the

trends in growth are probably weak on Area 2. On
comparisons against the index of biomass (sum of

Area 1 there was no effect for 8 of the 12 comparisons.
squared sizes), only that for large saplings on Area 1

However in the four comparisons of the two most re-
was significant. Among this many comparisons one

cent seedling classes, growth was significantly faster
should achieve significance (P < .05) by chance. Thus

when either the first or second neighbor was a different
no compensatory trend was shown in mortality over

species, a compensatory effect. In these latter two classes,
the whole plot.

the period in which growth was measured included
TABLE 8. Mortality rates of seedlings and saplings in relation to the species of, and distance to, their first- and second-nearest
neighbors. Mortality rates apply to the period between the map date and 1978 (area 1) or 1980 (area 2). Near/far categories
of distance were separated by omitting pairs with intermediate distances as follows. First nearest neighbors in Area 1: 1965
and 1969, 17-26 cm; 1971, 8-11 cm; 1974, 8-17 cm. Second nearest neighbors in Area 1; 1965, 29-38 cm; 1969, 23-38
cm; 1971, 23-32 cm; 1974, 20-29 cm. First nearest neighbors in Area 2: 1965, 9-26; 1969, 16-34; 1971, 3-12; 1974, 12-
25; 1978, 3-9. Second nearest neighbors in Area 2: 1965, 26-43; 1969, 30-46; 1971, 10-27; 1974, 27-37; 1978, 4-27. For
sapling classes the sample sizes were too small to permit this separation. Rates compared with G statistic. Within distance,
* .05 > P > .01; ** .01 > P > .001. Between distance classes, single underline signifies .05 > P > .01, double underline
.01 > P > .001.
Near nearest neighbor Far nearest neighbor
N % mortality N % mortality
Differ- Differ- Differ- Differ-
Year Nearest Same ent Same ent Same ent Same ent
Size-class mapped neighbor species species species species species species species species
Area 1
Large saplings 1965 First 46 788 17 13
1965 Second 42 627 17 15
Small saplings 1965 First 79 825 37 * 25
1965 Second 91 748 33 ** 26
Seedlings 1965 First 83 230 72 ** 42 68 369 60 ** 48
1965 Second 79 286 75 ** 50 71 368 70 ** 47
Seedlings 1969 First 215 342 79 ** 58 148 440 82 ** 71
1969 Second 174 272 84 ** 61 120 427 80 ** 69
Seedlings 1971 First 212 307 89 ** 82 157 907 89 ** 81
1971 Second 232 570 93 ** 82 78 578 78 82
Seedlings 1974 First 53 187 87 74 43 574 86 81
1974 Second 40 248 85 75 34 534 88 79
Area 2
Large saplings 1965 First 68 740 22 13
1965 Second 52 604 17 14
Small saplings 1965 First 116 894 25 24
1965 Second 109 772 23 25
Seedlings 1965 First 44 41 91 * 61 39 274 64 * 62
1965 Second 40 50 93 * 68 33 305 67 * 60
Seedlings 1969 First 22 65 73 62 22 165 65 55
1969 Second 15 52 73 64 17 204 53 56
Seedlings 1971 First 617 101 98 ** 93 485 1139 96 ** 89
1971 Second 626 108 99 ** 94 379 971 95 ** 90
Seedlings 1974 First 25 38 84 68 28 149 82 74
1974 Second 19 39 74 74 20 169 85 72
Seedlings 1978 First 216 19 88 ** 84 137 477 69 ** 68
1978 Second 150 5 89 ** 80 143 162 72 ** 69
between near and far neighbors of seedlings, mortality

only the first years after establishment; for previous
was significantly higher when conspecific neighbors were

year classes, because all remeasurements were made in
closer. In contrast, of the 18 comparisons when neigh-

1978, growth included older individuals. Thus it is
bors were different species, only 2 were significant, and

reasonable to infer that on Area 1 compensatory effects
on growth may exist very soon after seedling estab- these indicated higher mortality when neighbors were
lishment, but not thereafter. more distant. Thus distance seems to have a marked
effect on mortality mainly when the neighbors are the
same species. The result from different-species pairs
Mortality in relation to nearest neighbors
may have been due to chance, since it happened in
We first analyzed the effect of distance to neighbor only 1 of the 9 year-classes compared.
on mortality of both seedlings and saplings. To provide Mortality was significantly higher when the nearest
a stronger contrast between near and far neighbors we neighbor was the same species in 13 of 20 comparisons
omitted from the analysis the seedlings with neighbors

on Area 1 and in 12 of 24 comparisons on Area 2
at intermediate distances. The distances of the ap- (Table 8). In no case was mortality significantly higher
proximately one-third that were omitted are given in when neighbors were different species. In fact, even
the title of Table 8. Because of small sample sizes, the among the nonsignificant comparisons, only 1 of 19
two sapling classes were not separated into distance had higher mortality when neighbors were different
classes as the seedlings were. In 6 of the 18 comparisons species. Since commoner species have conspecifics as
Vol. 54, No. 2
TABLE 9. Mortality in regard to whether the first or second nearest neighbor is the same species, genus, or family. The
underlining connects classes within which the mortalities did not differ significantly (G test, P > .05). Percent mortality
calculated from the year mapped to 1978 (area 1) or 1980 (area 2). Dotted lines connect numbers not significantly different.
First-nearest neighbor is: Second-nearest neighbor is:
Same Same Same
genus, Same genus, family,
differ- family, differ- differ-
Year Same ent differ- Different Same ent ent Different
Size-class mapped species species genus family species species genus family
Area 1, North Queensland*
Large 1965 N 47 35 749 17 21 600
saplings % mort. 17 14 13 17 9 15
Small 1965 N 84 78 779 91 51 692
saplings % mort. 37 22 25 33 33 34
Seedlings 1965 N 215 31 52 788 187 25 51 715
% mort. 69 52 38 48 72 32 47 48
Seedlings 1969 N 517 35 67 988 482 41 49 917
% mort. 81 57 49 66 84 71 57 64
Seedlings 1971 N 466 23 94 1184 378 16 99 1155
% mort. 89 61 85 80 88 63 84 80
Seedlings 1974 N 126 19 36 884 90 17 38 845
% mort. 80 84 67 79 84 71 66 79
Area 2, South Queensland*
Large 1965 N 68 70 670 52 58 546
saplings % mort. 22 7 14 17 12 14
Small 1965 N 116 134 758 109 129 641
saplings % mort. 25 17 25 23 24 25
Seedlings 1965 N 136 80 440 113 70 436
% mort. 78 63 53 74 57 56
Seedlings 1969 N 65 31 344 48 26 330
% mort. 68 58 57 60 42 60
Seedlings 1971 N 1634 55 1513 1493 62 1504
% mort. 97 89 89 97 88 90
Seedlings 1974 N 72 12 222 59 14 223
% mort. 82 67 72 84 71 71
Seedlings 1978 N 457 15 389 389 30 399
% mort. 80 47 66 82 70 65
* In those size-classes with only 3 columns of data per neighbor, the middle column refers to instances in which the neighbor
was a different species in the same genus or family.
neighbors more often than do rarer species, these re- nificance of this species specificity will be discussed
sults clearly are in the direction of compensation. later.
To see whether these effects extended beyond the
Growth and mortality in relation to variations
species level, we calculated mortality of those having
in local abundance of the same
nearest neighbors of the same species, of the same ge-
or different species
nus but of different species, of the same family but of
different genus, and of a different family. The effect of Hypothesis 2 was also tested by comparing the effects
nearest neighbor was quite species specific (Table 9). of increasing density in quadrats on Area 1, as de-
For the comparisons in which the mortality was sig- scribed in the Methods section above. Few species were
nificantly higher if the neighbor was conspecific, the common enough at this scale to be analyzable using
mortality of individuals with congeneric or confamilial the sample size criteria described earlier. Of the 20
neighbors was not significantly different from the mor- regressions of growth on density for seedling size-classes,
tality of individuals whose neighbors were from dif- only 2 were significant, and both were in the direction
ferent families in all but one case. In cases in which of compensation. However this evidence is quite weak,
mortality was not significantly higher with conspecific since the probabilities were close to .05, and with 20
neighbors, adding individuals with congeneric or con- tests, one such would be expected to occur by chance.
familial neighbors would not change the outcome, since For the sapling size-classes, 4 out of 24 regressions were
the latter two often had lower mortality than did in- significant. However, the direction of 1 of the 4 went
dividuals with neighbors of a different family. The sig- in the opposite direction from compensation. Taken
TABLE 10. Mortality rates of seedlings and saplings in relation to their proximity to adults of the same species. The inner
circle was an area within 1.5 x the crown radius of the adult; the outer ring was an annulus concentric to and of width
equal to the radius of the inner circle. All species with sufficient sample size to apply a G test are included.
No. of species in which the mortality closer to adult,
as compared to farther away, was
Significantly lower No significant Significantly higher
nearer adult difference nearer adult
Inner circle vs. outer ring
Area 1: North Queensland 0 22 1**, 5*
Area 2: South Queensland 1* 22 1**, 1*
Inner circle + outer ring vs. farther from adult
Area 1: North Queensland 1**, 2* 54 3**, 4*
Area 2: South Queensland I**, 3* 39 2**, 3*
* .05 > P > .01.
** P < .01.
as a whole, the evidence for compensation in growth

Mortality in relation to the proximity to adults
at the scale of these large quadrats is very weak.
With respect to mortality, 19 of the 22 species tested

We tested hypothesis 2 at the scale of individual
showed no significant relationship to density. In one

adult trees, calculating the mortality of seedlings and
case (the 1971 year-class of Beilschmiedia sp. aff. ob-

saplings in the inner circle and outer ring around each
tusijfolia), mortality decreased with increased abun-

conspecific adult as well as in the area beyond the outer
dance of existing trees of the same species, but showed

ring (Table 10). The results were identical for Area 1
no effect of variation in abundance of different species.

and Area 2; mortality was greater in the inner ring than
The two significantly negative relationships were for

in the outer circle in only 1 of 23 comparisons (using
the 1978 year-class of Planchonella sp. nov. with either

a significance level of.0 1). Comparing the "near" zone,
density or sum of squared sizes of existing trees. With

defined as the combined inner and outer rings, with
22 comparisons, three significant results (P < .05) do

the area beyond ("far" zone), 3 of 58 comparisons in
not constitute strong evidence. In summary, there is

Area 1 and 2 of 42 in Area 2 showed greater mortality
little evidence of compensatory mortality at this spatial

nearer the adult. The reverse, lower mortality near the
scale for the period of study.

adult, occurred once in each of Area 1 and Area 2. In
The field experiments in which seeds were placed on

summary, compensatory mortality was occurring in
the ground in dense vs. sparse treatments constitute

4/4% of the cases when inner vs. outer zones were
an experimental test of hypothesis 2. The differences

compared, and in 5/5% of the cases when these two
in survival between the two treatments were insigifi-

zones were combined and compared to areas more
cant for Cryptocarya corrugata and Eugenia brachy-

distant from the adult. These frequencies are so low
andra seeds brought to Area 1 from Area 2. Almost

that they could well have occurred by chance.
all the plants (N = 900 for Cryptocarya in the dense

A lower mortality rate nearer adults than farther
treatment, 100 for its sparse treatment, and for both

away is the reverse of compensation; however, this
Eugenia treatments) disappeared over the observation

could still produce a compensatory change overall if it
period of 9.5 mo.

tended to happen among the rarer species, while the
TABLE 11. The pattern of mortality of seedlings (or saplings) near vs. far from adults of the same species in relation to the
abundance of adults. Each Spearman rank correlation compares the adult abundance (basal area) of each species with the
values of the G statistic calculated for that species by comparing the mortality of seedlings or saplings in the two zones
around adults. A significant negative r would signify that more abundant species had higher mortality nearer adults. All
species that had at least six individuals in each zone were included. * .05 > P > .01. Dash indicates sample size too small
for analysis.
Inner zone vs. Outer ring Inner + outer vs. elsewhere
Small Large Small Large
Seedlings saplings saplings Seedlings saplings saplings
No. species in correlation 18 7 - 32 19 12
Correlation coefficient r -0.11 -0.04 - -0.09 0.30 0.37
No. species in correlation 11 7 10 17 8 20
Correlation coefficient r 0.03 0.21 0.21 0.56* -0.10 0.21
Vol. 54, No. 2
TABLE 12. Field experiments on survival of seeds and seedlings placed under adults of the same vs. different species. Length
of life not different between treatments for any cohort (Mann-Whitney U test, P > .05).
A. Survival from seed to end of experiment
No. Seeds per Length of % survival at end, for each replicate
repli- repli- experiment Under adult Under adult of
Species Treatments cates cate (mo) of same sp. different sp.
Planchonella Trenched 2 98 36 34, 30 86, 73
sp. nov. (Area 1) Untrenched 2 98 36 49, 45 74, 57
Cryptocarya Caged 3 25 22 0, 0, 0 0, 0, 0
corrugata Roofed 3 25 22 0, 0, 0 0, 0, 0
(Area 1) Control 3 50 22 1, 0, 0 0, 0, 0
Insecticide 3 50 22 0, 0, 0 1, 0, 0
Eugenia Caged 4 25 24 8, 0, 20, 16 0, 12, 4, 8
brachyandra Roofed 4 25 24 8, 0, 0, 4 0, 0, 0, 0
(Area 2) Control 4 50 24 0, 4, 0, 0 0, 0, 2, 6
Insecticide 4 50 24 0, 2, 0, 0 0, 4, 0, 0
B. Average length of life calculated for the year following germination
No. of seedlings at start Average length of life (mo)
Not under Not under
Period when cohort Under adult adult of Under adult adult of
Species germinated of same sp. same sp. of same sp. same sp.
Eugenia Dec 1969-Feb 1970 37 38 4.7 4.9
brachyandra Apr-May 1970 25 22 5.4 4.0
(Area 2) Jul-Sep 1970 27 24 5.7 3.9
reverse happened among the commoner species. To adults, the reverse of compensation. We conclude that
investigate this possibility we used the results of the there was no evidence for compensatory mortality of
comparison of mortality between inner and outer rings, young trees due directly to the influence of adults.
and between these two combined and elsewhere, com- In field experiments, seedlings of one species (Plan-
paring ranks based on the G statistics with the ranks chonella sp. nov.) had lower survival closer to their
of basal area for each species as shown in Table 1 1. Of adults; survival of Cryptocarya corrugata and Eugenia
the 11 comparisons, only 1 showed a significant cor- brachyandra seeds was equally poor near and far from
relation, and this was positive, indicating that the com- their adults (Table 12). Once Eugenia brachyandra
mon species tended to have lower mortality closer to seedlings had become established, they survived as well
SEEDLINGS SAPLINGS TREES
Sm. Lg. Med. Lg.
a) Change dC b
1. - Tde : | ! i 0 | AE
Inc
t 1981 1978 1974 1971 1969 1965 1965 1965 1963 1963
0.6
1981 1978 1974 1971 1969 1965 1965 1965 1963 1963
YEAR MAPPED
FIG. 7. Pattern diversity for all size-classes at the time of original mapping and in 1980 after some mortality had occurred.
* significant difference (z test, P < .05) between times within a year- or size-class. Letters indicate significant differences at
that date between a small size-class and larger one(s) as follows: a, b: comparison to next larger class; c: seedlings vs. small
and large saplings; d: seedlings vs. large saplings only; e: seedlings vs. small saplings only.
TABLE 13. Pattern diversity (D of Pielou [1966]) of the different size- or year-classes at the time of first mapping and in
1980 after some mortality. N is the number of triplets (tree and first- and second-nearest neighbors) in each analysis.
Significance tests were performed on the differences in D between each seedling class and the two sapling classes and between
the sapling classes and medium trees. Tests were also done for the same size-class between dates. Significant differences
between size-classes shown as: (a) large saplings compared to medium trees, (b) small saplings compared to large saplings,
(c) seedlings compared to small and large saplings, (d) seedlings compared to large saplings, (e) seedlings compared to small
saplings; significant differences between times within size-classes shown as *. All values of D except those underlined were
significantly different (smaller) than 1.0, the random expectation. All comparisons used the z test, P < .05.
Area 1 Area 2
Year Map date 1980 Map date 1980
Size-class mapped N D SE N D SE N D SE N D SE
Large trees 1963 1161 .979 .004 1009 .976 .004 1192 .969 .006 1047 .966 .007
Medium trees 1963 550 .984 .006 457 .988 .006 481 .984 .006 457 .988 .006
Large saplings 1965 798 .970 .006 482 .972 .007 652 .962a .008 * 583 .984 .008
Small saplings 1965 911 .940b .007 506 .961 .009 862 .927b .008 672 .941b .009
Seedlings 1965 1010 .852c .010 * 379 .926c .013 594 .851c .014 * 237 .958 .014
Seedlings 1969 1521 .851c .012 * 301 .963 .017 405 .912d .012 * 165 .956 .013
Seedlings 1971 1784 .850c .009 * 218 .927d .017 3107 .723c .011 * 191 .956 .020
Seedlings 1974 1023 .922d .007 * 108 .957 .015 381 .831c .022 * 72 .973 .025
Seedlings 1978 3987 .776c .011 * 2050 .992 .223 816 .975 .029 216 1.050e .034
Seedlings 1981 5825 .51Ic .009 2448 .676c .013
near as far from adults. Of the experimental treatments,

the 10 seedling classes on both areas. However in the
caging increased survival of seedlings in Eugenia, but

eight instances among the older classes, there was only
trenching or insecticide application apparently had no

one significant increase (and no significant decreases)
effects on it or on other species. Thus the field exper-

in pattern diversity. Thus most of the change in the
imental results were similar to the statistical analysis,

degree of intermingling of species directly observed
showing little evidence of compensation.

occurred between the seedling and sapling stages, prob-
ably because of the high mortality rates of the seedlings.
Pattern diversity
Since there are significant differences between small
We found earlier that seedlings with nearest neigh-

and large saplings, the process of increasing intermin-
bors of the same species tended to have higher mor-

gling probably continues into these older age-classes.
tality rates than those with different species as neigh-
bors (Table 8). Since most of the data come from the
DISCUSSION
commoner species, in time this process should cause
Maintenance of diversity
the different species to become more intermingled. Pie-
lou (1966) devised a method to estimate the degree of

If rarer species tend to be favored over commoner
intermingling of species, which she called the pattern

ones, some degree of species diversity may be main-
diversity of a community. In all but one size- or year-

tained. We have searched for evidence that variations
class (1978 seedlings on Area 2, Fig. 7 and Table 13),

in the recruitment of seedlings and in the growth and
the pattern diversity values at the time of original map-

mortality of seedlings and saplings was such as to favor
ping were significantly < 1.0, indicating that individ-

rarer species. At some spatial scales this compensatory
uals of all sizes tend to have the same species as neigh-

trend seems to be the case, in others not. Over the
bors more often than expected by chance.

whole plot, in both study areas, seedling recruitment
Does pattern diversity increase with age as one would

of subcanopy and understory species was lower for
expect from the results of our analysis of mortality?

commoner than for rarer species at both sites. For the
Pattern diversity of seedlings when first mapped after

species that reach the canopy, recruitment rate per adult
initial establishment was lower than that of small or

was not related to adult abundance. This meant that
large saplings, or both, in 1 1 of the 12 year-classes on

there were many more seedlings of the commoner can-
the two study areas (Table 13, Fig. 7). The small sap-
opy species than of rarer ones becoming established in
lings also had lower pattern diversity than large sap-

these forests. Growth and mortality rates of seedlings
lings in both areas, and the large saplings had lower

did not vary with abundance of either adults or smaller
pattern diversity than did medium trees in Area 2.

size-classes at this scale. Field experiments in which
Thus as size increases, the trees become more inter-

the density of seeds was varied showed no consistent
mingled; the trend predicted from the mortality rates trend in mortality.
of seedling nearest neighbors extends up to medium

At the scale of the size of crowns of adult trees,
trees.
mortality was higher close to adults in a few species,
Pattern diversity also increased significantly between

but in other species it was the reverse. In the majority
the time of initial mapping and 1980 for all but 1 of

of species there was no difference, nor were these dif-
Vol. 54, No. 2
ferences in adult effects related to adult abundance.

diversity were increased, then the chance of a dead
Field experiments gave similar results; compensation

adult tree being replaced by a different species would
seems to be occurring in some species at this scale, the

be increased. Our results support this expectation, since
reverse in others, with the majority showing neither.

pattern diversity was indeed greater in the saplings than
At the smallest scale, between nearest neighbors,

in the seedlings.
growth very soon after seedling establishment seemed
Mechanisms of compensation
to be compensatory, being slower if the neighbor was
the same species than if it was different. This trend

The compensatory trends between nearest neighbors
apparently does not continue after the age of -6 yr.

could be a result of intraspecific competition. Young
Mortality was compensatory for the majority of size-

individuals of commoner species tend to have con-
classes tested (seedlings and small saplings), and was

specifics as nearest neighbors more often than do rare
often greater when the neighbor was closer. This com-

species, and at the very short distances between them,
pensatory mortality was species specific, there being

competition might be expected. If such competition or
no indication of higher mortality if neighbors were of

interference is more deleterious between conspecifics
the same genus or family.

than between different species, this would produce a
The general result is that all the compensatory trends

compensatory effect. Competition for light is intense
we have discovered apply to the youngest individuals:

in these dense forests, but it seems unlikely that shading
initial recruitment per adult and early growth and mor-

could have stronger effects against conspecifics than
tality. Aside from initial recruitment, compensation

against other species. Intraspecific competition or in-
seems to be strongest at the smallest scales, that of first

terference has been shown to be more deleterious than
or second nearest neighbors, which for these young

interspecific for some herbaceous species in laboratory
trees involves differences of only a few tens of centi-

experiments (e.g., Newman and Rovira 1975, Harper
meters.
1977, Parrish and Bazzaz 1982) and field experiments
These results are not a reflection of the fact that

(Rahman 1976, Kroh and Stephenson 1980). For woody
sample sizes are larger for younger individuals; growth

species, the present trenching experiments with Plan-
and mortality of these same young individuals were

chonella (Table 12, see also Connell 1971) also indi-
also analyzed at larger spatial scales without revealing

cated that intraspecific interactions were more dele-
compensatory trends. Also, growth rates are faster for

terious than interspecific ones. In contrast, Fonteyn
larger trees, so that any results of growth analyses should

and Mahall (1981) found that interspecific interference
be as accurate or more so for the larger size-classes

was equal to or stronger than intraspecific in field ex-
than for smaller ones. Only in the case of mortality

periments with woody desert bushes. Although many
rates is it possible that we missed finding compensatory

other trenching experiments have been done in forests
trends. In the larger size-classes, mortality is so low

(reviewed by Connell 1975), to our knowledge only
than 17 yr is probably too short a time to record sig-

one other study of woody plants has been designed
nificant differences.
expressly to compare the strengths of intraspecific vs.
How could these compensatory trends in the re-

interspecific interactions. Webb et al. (1967) found del-
cruitment, growth, and mortality of young seedlings

eterious effects of adults on seedlings of a rain forest
contribute to the maintenance of diversity of adult trees? tree, Grevillea robusta, in plantations in Queensland.
First, compensatory trends in any size-class should help

The intraspecific deleterious effects were much stronger
prevent commoner species from increasing at the ex-

in Grevillea plantations than were interspecific effects
pense of rarer ones, thus reducing the rate of local

in plantations of Araucaria cunninghami. The delete-
extinction of rare species. Second, any process that

rious effect may have been produced by competition
increases the number of species or the evenness of

(which the authors judged unlikely), interference from
relative abundance of seedlings or saplings present in

toxic exudates from the roots, or microorganisms (see
a gap created by a fallen adult should help maintain

below). Clearly, intraspecific competition or interfer-
species diversity, since it is likely that the existing young

ence remains a possible compensatory mechanism.
trees in light gaps constitute the group from which the

Indirect deleterious influences, via natural enemies
succeeding adult will come.

such as microorganisms, fungi, herbivores, etc., are a
Seedlings of shade-tolerant species tend to become

third possibility. Florence and Crocker (1962), Flor-
established densely near the parent trees, since these

ence (1965), and Webb et al. (1967) suggested that
species often have large seeds or fruits that fall near

microorganisms associated with roots may have del-
the parent. Since the mortality of conspecific seedlings

eteriously affected tree seedlings, although direct in-
was no greater nearer adults (Table 10), one would

terference by toxins from the roots themselves was not
expect such adults to have many conspecific seedlings ruled out. For natural enemies to produce the observed
nearby, so that the adult would probably be replaced

compensatory trends in growth and mortality between
by another of the same species. However if the original nearest neighbors they probably need to be relatively
pattern of clumps of conspecific seedlings were changed

species specific in their attacks. This does not mean
to one of intermingled species of saplings by the higher that specialist enemies cause all the mortality; many
mortality of conspecific neighbors, i.e., if the pattern

rain forest seedlings undoubtedly are killed by non-
specific agents, both physical and biotic in origin. How-

examples of switching in populations in natural cir-
ever conspecific neighbors must in addition suffer some

cumstances as yet exist, to our knowledge. In summary,
species-specific mortality. We suggest that the most

natural enemies, either specialists concentrating their
likely agents producing this additional mortality are

attacks in denser prey patches or generalists switching
either herbivorous animals or pathogenic fungi. Fox

to commoner prey, could have produced the pattern
and Morrow (-1981) have shown that herbivorous in-

of lower growth or higher mortality rates observed be-
sects with generalized diets over their entire geographic

tween conspecific nearest neighbors.
range often have quite specialized diets in local com-

In regard to the compensatory trend toward reduced
munities. They found that the proportion of herbivo-

recruitment in subcanopy species with commoner
rous insects feeding on only one species of plant at a

adults, the mechanisms may have operated through
local site varied from 18 to 50%. Some pathogenic root-

either reduced flowering or fruiting on the adult tree,
infecting fungi are very specialized, though others are

or mortality of seeds before or after dispersal, or early
not (Wheeler 1968, Garrett 1970, Brian 1976). Thus

mortality during (and soon after) seedling establish-
some insects and fungi (and perhaps other natural ene-

ment. Although we have no direct information on events
mies) are sufficiently specialized to produce the addi-

that could influence flower and fruit production, stud-
tional mortality observed between conspecific seedling

ies by Lowman (1982) indicate that herbivorous insects
nearest neighbors.
could have had a compensatory effect on adult trees.
Even if natural enemies are specialized, this in itself

In rain forests in northern New South Wales, 250 km
is not sufficient to explain the lower growth and higher

south of Area 2, she found that leaf-eating insects con-
mortality rates of conspecific nearest neighbors. Such

sumed a higher proportion of the canopy leaves of
a pattern requires that the attacks be proportionately

commoner than of rarer species. (Many of the species
greater where the same species of seedlings occur in

in these forests also occur in or near Area 2 of the
close proximity, for example in single-species clumps.

present study.) For example, canopy leaves of adult
Although we have no direct evidence of this in the

Doryphora sassafras were consumed at a higher rate
present study, a considerable body of work indicates

in a warm-temperate forest, where it was a common
that natural enemies may act in this way. First, natural

species, than in either a cool-temperate or a subtropical
enemies are usually highly aggregated in their attacks

forest, where it was a rarer species. Shade leaves and
on individual prey. For example, female moths (Cac-

leaves located nearer to ground level suffered more
toblastis cactorum) lay their eggs in an aggregated pat-

grazing, so subcanopy species might be expected to lose
tern on cactus (Opuntia inermis) in Australia (Monro

proportionately more leaves than canopy species. Thus
1967), and similar patterns have been shown with par-

if greater leaf consumption by herbivores leads to lower
asites (Anderson 1979). Second, some natural enemies

production of flowers and/or seeds, commoner sub-
concentrate their attacks in places with higher densities

canopy species would have lower production than rarer
of prey; they are either attracted there (e.g., insect para- ones.
sitoids attracted to concentrations of the food of their

Mortality of seeds is very great. In one experiment
host [Rotheray 1979]), lay more eggs there (e.g., insect

with fruits of Halfordia scleroxyla on Area 1, done at
predators [Clark 1963]), or remain longer in such

the same time and with the same design and sample
patches (e.g., insect predators [Banks 1957], spiders

sizes as for Cryptocarya corrugata, not a single one of
[Turnbull 1964]). Tahvanainen and Root (1972) found

the 900 seeds germinated. The mortality may have
that herbivorous beetles were more abundant on col-

been due to attack by insects earlier on the tree or by
lards grown in monoculture than on those grown ad-

various animals after they fell. For example, casso-
jacent to diverse natural vegetation. The monocultures

waries, which eat huge numbers of seeds, have been
were colonized more rapidly and had greater feeding

observed on this plot, and other seed-eating animals
damage.
are common. However as discussed above, the mor-
Of additional interest is the situation in which at-

tality before and soon after germination in most of the
tacks by nonspecialized natural enemies are dispro-

field experiments done by us and others did not have
portionately greater against whichever of two prey

a compensatory pattern. In summary, several possible
species is commoner. Thus, the predator switches its

mechanisms could have produced the compensatory
attack to the commoner prey. Such behavior has been effects found.
demonstrated in the laboratory for the following pred-
Compensatory processes in forests in general
ators or parasitoids: marine snails, freshwater fish,
aquatic insects, protozoans, and insect parasitoids

Most of the previous work on compensatory or fre-
(Murdoch 1969, Lawton et al. 1974, Murdoch and

quency-dependent processes in forest trees has con-
Oaten 1975, Cornell and Pimentel 1978). The only

sisted of describing the distribution of the understory
published examples of switching by herbivores con-

in relation to the canopy. Much of the evidence in the
cerned wood pigeons feeding in the field on peas and

early literature tended to be anecdotal rather than
beans (Murton 1971, in Murdoch and Oaten 1975) and

quantitative. Aubreville (1938) in his "mosaic theory
European quail in the laboratory fed artificial bait of

of regeneration" (Richards 1952), suggested that species
two colors (Manly et al. 1972). No well-documented

seldom regenerate beneath themselves. Jones (1945)
Vol. 54, No. 2
states that: "there is a very strong belief amongst con-

petition for water. Thus both hypotheses were accepted
tinental foresters in the tendency of one species to re- in this temperate zone study.
generate beneath the canopy of another species."

In other instances only one of the hypotheses was
Janzen (1970) and Connell (1971) proposed two hy-

studied. I tested the distance hypothesis with Plancho-
potheses that formalized this general idea. They sug-

nella sp. nov. on Area 1 (Connell 1971). Seedlings
gested that mortality of seeds or seedlings would be

planted at the same density had higher mortality in the
greater either nearer adult conspecifics (the "distance"

inner than in the outer annulus around adults, as de-
hypothesis) or at higher density ofjuvenile conspecifics

fined in the present paper. Similarly, Vandermeer ( 1977)
(the "density" hypothesis). Since the distribution of

found that Welfia georgii palm seedlings had higher
seedfall usually results in higher densities nearer adults,

mortality under adults (from being crushed by the shed
when the advantages of seed dispersal are being con-

fronds) than elsewhere. In both cases the distance hy-
sidered, the two hypotheses are sometimes combined pothesis was accepted.
into an "escape hypothesis" (Howe and Smallwood

The density hypothesis has been tested in several
1982). When all species with a sufficient sample size

instances. On forest edges in Guatemala, Vandermeer
were analyzed in the two present study sites, the dis-

(1974) found greater infestation of seeds by insect lar-
tance hypothesis was rejected for most species (Tables

vae in trees of Calliandra grandiflora that were closer
10 and 11), but the density hypothesis was accepted

to conspecific neighbors. In dry forests in Costa Rica,
for the majority of seedling year-classes (Tables 7, 8,

Janzen (1975) found no difference in mortality of seeds
and 9).
of Acaciafarnesiana from dense, almost monospecific
Other tests of the hypotheses have dealt with single

stands vs. those in which the trees were sparse and
species, either observationally or with field experi-

intermingled with other species. In the same region
ments. In some instances, both hypotheses have been

Silander (1978) found that seed production of Cassia
tested separately on the same species. Let us consider

biflora bushes was greater when they were closer to-
first the field experiments on the present study sites.

gether, but that mortality of seeds did not vary with
Cryptocarva corrugata on Area 1 showed no effects of

density of bushes. In a study of seeds of Juglans nigra
either distance or density on seed mortality (Connell

buried by squirrels in Kansas, Stepanian and Smith
1971, 1979). Field experiments with Eugenia brachy-

(1978) found that seeds at higher density were eaten
andra on Area 2 showed no effects of distance (Connell

more readily. Culver and Beattie (1 980) found no effect
1979). The density hypothesis was tested on this species

on seedling emergence of planting seeds of two species
by bringing seeds to Area 1 where no adults occur; no

of Viola at different densities at sites in southern En-
effects of density were found (Connell 1971). Wilson

gland. In summary, the density hypothesis was rejected
and Janzen (1972) found no difference in the mortality

in two of the three tropical studies and one of the two
of seeds of the palm Scheelea rostrata placed at high temperate studies.
density near vs. far from adults, but seeds placed far
Since the distribution of seedfall usually produces
from adults at low density had much lower mortality

higher densities nearer adults, the results of many ob-
than those at high density. Thus for this species in a

servational and experimental studies do not separate
dry tropical forest in Costa Rica, Wilson and Janzen

the effects of distance and density. For example, in an
(1972) rejected the distance hypothesis but accepted

evergreen forest in Puerto Rico, Janzen (1972a) found
the density hypothesis. D. A. Clark and D. B. Clark

similar percentages of nonviable seeds of Euterpe glo-
(personal communication) found the same result by

bosa beneath the crown, where the density was higher,
following the survival of juveniles of the rain forest

than farther away at lower density. Janzen (1 97 1) found
tree Dipteryx panamensis in wet tropical rain forest in

that juveniles of the vine Dioclea megacarpa in a dry
Costa Rica. When the effect of distance to adult was

Costa Rican forest were more heavily grazed closer to
analyzed separately from that of juvenile density, sur-

adults where they were also at higher density. In the
vival over 2 yr was found to be a function of density,

same forest, Janzen (1975) found similar mortality of
not distance. In these subtropical and tropical forest

seeds of Spondias mombin beneath the tree at high
studies in which both hypotheses were tested, the dis-

density and at the edge of the seed shadow at lower
tance hypothesis was rejected in every case, whereas

density. In the same region, seeds of Sterculia apetala
the density hypothesis was accepted for two of the four

were placed along lines radiating from an adult tree;
species.
the density of fallen seeds also decreases with distance
Platt (1976) tested both hypotheses with field ex-

(Janzen 1972b). Bugs that destroy the seeds appeared
periments on the perennial herb Mirabilis hirsuta, in

more quickly on the closer, denser seeds, but within 4
an Iowa prairie. Fruits placed at different distances

h of placement the number of bugs was similar at all
from adults were eaten by rodents at a higher rate closer

distances and densities. Also in the same forest type,
to the adults. Fruits placed at different densities were

Janzen et al. (1976) found that seeds of Andira inermis
removed by ants more quickly from higher density

were killed by weevils at high rates on the ground be-
patches. Seedlings planted at different densities suffered

neath adults or beneath trees that served as roosts for
greater mortality at higher density, probably from com-

bats that had carried them away; both were also sites
of high seed density. Mortality was lower on seeds

assumed to be equally abundant, the number of species
scattered far from adults at low density.

that could coexist in the mathematical model was not
Howe and Primack (1975), working in a wet Costa

high. If relative abundances were unequal, as in all
Rican forest, found a lower rate of survival between

natural forests, many more species could coexist. In
the seed and seedling stages in Casearia nitida nearer

any case, the distance hypothesis applies to only a few
adults, where seed density was also higher, than farther

species in the forests studied here, although the density
away at lower seed density. Clark and Clark (1981)

hypothesis applies to juveniles. Either mechanism could
found seedlings of Bursera graveolens only far from

produce the trend toward reduced clumping with age
adults in a Galapagos Island site, whereas seeds were

found by Hubbell (1979).
much denser closer to adults. The same pattern was

Other quantitative studies on this subject have dealt
found for seeds and seedlings of Juniperus monosper-

mainly with pairs of species that regenerate beneath
ma in mountain woodlands in New Mexico (Salo-

each other. The first quantitative study of "alternation
monson 1978). H. Howe and E. Schupp (personalcomr-

of species" was that of Fox (1977). In four of the five
munication) placed seeds of Virola surinamensis on

temperate forests he studied that had two codominant
the ground in concentric annuli at different distances

species, the understory of each species occurred more
from 17 fruiting adults in a rain forest in Panama. Since

frequently than chance expectation beneath the canopy
the same number of seeds was placed on each annulus,

of the other species. Whittaker and Levin (1977) cite
those placed closer were also at higher density. The

a similar analysis by F. E. Smith from a forest of beech
percentage killed by insects was greater closer to the

and sugar maple. Woods (1979) and Woods and Whit-
adults and at higher density. Of the nine studies in

taker (1981) made a detailed analysis of a beech-maple
which the density and distance hypotheses were com-

forest that showed strong evidence of such "reciprocal
bined, it was rejected for three species and accepted in replacement."
the other six.

In other studies the evidence shows no such alter-
In summary, of the 26 instances in which hypothesis

nation of species. In a series of stands sampled in the
2 was tested for single species as either the distance or

hemlock-hardwood forest of the Great Lakes region,
density hypothesis or the two combined, it was ac-

Woods and Whittaker (1981) found that beech and
cepted in 14 cases and rejected in 12. When the distance

maple showed reciprocal replacement in some stands
hypothesis was tested separately it was rejected in 4 of

but not in others. Hemlock showed no reciprocal re-
7 tests, whereas the density hypothesis was rejected in

placement with any species, a result contrary to that
5 of 10 tests. These studies of single species vary in

from one of Fox's (1977) forests. Forcier (1975) found
experimental design and rigor and include a great range

a serial replacement of birch, maple, and beech, with
of habitats, regions, and life forms. In contrast, the no reciprocal alternation.
present study indicates the degree to which the hy-

If a forest has many species instead of two codom-
potheses apply to a less heterogeneous sample, namely

inants, it is very nearly impossible to perform this sort
all species that were abundant enough to be analyzed

of analysis because the number of possible pairs be-
on two similar rain forest sites. Both the set of studies

comes very large. In diverse forests the most fruitful
of single species and the present analysis demonstrate

analysis is to see whether intraspecies interactions are
that the hypotheses apply to some species but not oth-

more deleterious than interspecies ones, as we have
ers. The density hypothesis applies more strongly than

done in this paper. There is a wealth of anecdotal state-
the distance hypothesis in the two present study sites.

ments that species seldom regenerate beneath them-
In most populations, including trees in diverse rain

selves, but there are few quantitative studies. Beals
forests, species tend to be clumped rather than ran-

(1965) studied 20 remnant stands of Lebanon cedar
domly or uniformly distributed (Poore 1968, Whit-

and found that: "cedar seedlings thrive under hard-
more 1974, 1975, Hubbell 1979). In species to which

wood trees and shrubs but not under cedar trees, so
hypothesis 2 applies, the degree of clumping should

that some disturbance is apparently necessary for ce-
decline with age. Hubbell (1979) tested this prediction

dar-forest regeneration." Florence and Crocker (1962)
for trees in a tropical dry deciduous forest in Costa

and Florence (1965) found that in Eucalyptus pilularis
Rica. In about half of the 30 commonest tree species

in Australia and Sequoia sempervirens in California,
the adults were less clumped than the juveniles, a trend

growth of seedlings was poor in soil taken from beneath
predicted by the hypothesis. In the other half of the

adults. However if the microorganisms were killed by
species studied other mechanisms must have been op-

irradiation, seedling growth was greatly improved.
erating to maintain or increase clumping between ju-

In Queensland, attempts to establish pure planta-
venile and adult ages. In a mathematical model in

tions of six species of rain forest trees have not been
which the spatial pattern of canopy trees was deter-

successful, whereas two others (both Araucaria) were
mined solely by the rule that conspecific trees were

successful (Webb et al. 1967). The former six species
separated by trees of other species, more species could

seldom form pure stands in nature, whereas Araucaria
coexist the greater the spacing distance (Hubbell 1980).

species do. Experiments on one of the former species,
However even at great distances, when species were

Grevillea robusta, indicated that seedlings died if their
Vol. 54, No. 2
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APPENDIX
versity of Oxford, Oxford, England.
Continued.
1975. Tropical rain forests of the far east. Clarendon
Press, Oxford, England.
Basal No. of
Whittaker, R. H., and S. A. Levin. 1977. The role of mosaic
area adults
phenomena in natural communities. Theoretical Popula-
Species (m2/ha) on map
tion Biology 12:117-139.
Williams, W. J., and J. G. Tracey. 1984, in press. Network

Placospermum coriaceum 0.24 3
analysis of North Queensland tropical rainforests. Austra-

Eugenia cryptophlebia 0.19 3
lian Journal of Botany.

Gevuina bleasdalii 0.16 8
Wilson, D. E., and D. H. Janzen. 1972. Predation on Schee-
Area 1, subcanopy species (adults - 0.08 m gbh*)
lea palm seeds by bruchid beetles: seed density and distance
Brackenridgea australiana 0.83 125
from the parent palm. Ecology 53:954-959.
Apodytes brachystylis 0.59 51
Winer, B. J. 1971. Statistical principles in experimental de-
Synoum muelleri 0.48 5
sign. Second edition. McGraw-Hill, New York, New York,
Myristica meulleri 0.29 9
USA.
Cryptocarya sp. aff. rigida 0.23 25
Woods, K. D. 1979. Reciprocal replacement and the main-

Castanospora alphandi 0.21 5
tenance of codominance in a beech-maple forest. Oikos 33:

Alangium villosum 0.13 9
31-39.
Linociera axillaris 0.06 32
Woods, K. D., and R. H. Whittaker. 1981. Canopy-under-

Alectryon semicinereus 0.05 3
story interaction and the internal dynamics of mature hard-

Arytera lauteriana 0.03 4
wood and hemlock-hardwood forests. Pages 305-323 in D. Toechima lanceolatum 0.02 4
C. West, H. H. Shugart, and D. B. Botkin, editors. Forest Arytera divaricata 0.01 1
succession. Springer-Verlag, New York, New York, USA. Litsea sp. ("Mount Spec.") 0.01 4
Sarcopteryx martyana 0.01 3
Area 1, understory species (adults - 0.08 m gbh*)
Antirhea tenuiflora 0.43 51
Antirhea myrtoides 0.41 115
Polyosma alangiacea 0.35 18

APPENDIX
Rapanea achradifolia 0.15 20
Species reproducing during the study period. All species in

Polyosma rhytophloia 0.07 3
which, on the whole mapped plot, at least 10 seedlings became

Maba hemicycloides 0.05 9
established between 1965 and 1981, and which had at least

Steganthera australiana 0.05 28
two adults or at least 0.01 m2 basal area. No ephemeral (shade-

Mitrephora froggattii 0.01 5
intolerant, short-lived) species are included here.
Tapeinosperma pseudojambosa 0.01 3
Wendlandia inclusa 0.01 4
Basal No. of Bubbia semicarpoides <0.01 3
area adults Psychotria dallachyana <0.01 2
Species (m2/ha) on map

Caryospermum aborescens <0.01 4
Area 2, canopy species (adults - 0.32 m gbh*)

Area 1, canopy species (adults - 0.32 m gbh*)
Planchonella sp. nov. 4.82 112 Argyrodendron trifoliolatum 8.74 114
Flindersia pimenteliana 3.62 20 Argyrodendron actinophyllum 8.01 70
Ceratopetalum succirubrum 3.52 39 Doryphora sassafras 3.43 79
Eugenia sp. (Smithii complex) 3.10 8 Vitex lignum-vitae 3.35 29
Sterculia laurifolia 2.91 60 Orites excelsa 2.65 64
Flindersia bourjotiana 2.36 58 Litsea reticulata 2.30 11
Doryphora aromatica 2.19 30 Diospyros pentamera 1.97 44
Xanthophyllum octandrum 1.86 48 Eugenia crebrinervis 1.40 17
Cardwellia sublimis 1.35 15 Eugenia brachyandra 1.02 10
Garcinia sp. aff. hunsteinii 1.30 49 Synoum glandulosum 0.64 16
Elaeocarpus sericopetalus 1.06 14 Diploglottis australis 0.40 8
Opisthiolepis heterophylla 0.98 12 Elattostachys nervosa 0.38 18
Eugenia sp. ("u.r.p.") 0.97 16 Cinnamomum virens 0.22 9
Elaeocarpus domatia 0.90 7
Area 2, subcanopy species (adults - 0.08 m gbh*)
Citronella smythii 0.90 44
Baloghia lucida 4.52 274
Cryptocarya angulata 0.80 21
Sarcopteryx stipitata 0.69 38
Elaeocarpus largiflorens 0.73 6
Acronychia pubescens 0.37 82
Halfordia scleroxyla 0.72 18
Mischocarpus pyriformis 0.15 17
Beilschmiedia sp. aff obtusifolia 0.68 16
Guioa semiglauca 0.13 13
Carnarvonia sp. nov. ("Mount Haig") 0.65 31
Rhodomyrtus psidioides 0.09 4
Podocarpus amarus 0.62 1
Cupaniopsis serrata var. tomentosa 0.02 6

Darlingia darlingiana 0.42 11
Cryptocarya mackinnoniana 0.34 13

Area 2, understory species (adults - 0.08 m gbh*)
Eugenia sp. ("paper bark") 0.34 1

Randia benthamiana 0.47 101
Cryptocarya corrugata 0.32 6
Polyosma cunninghamii 0.08 9
Cryptocarya sp. aff. cinnamomifolia 0.32 10
Wilkiea huegeliana 0.08 114
Endiandra hypotephra 0.30 20

Decaspermum fructicosum 0.02 6
Galbulimima belgraveana 0.29 2
Psychotria simmondsiana 0.02 23
Eugenia cormiflora 0.25 11
Harpullia alata 0.01 5
Cryptocarya cinnamomifolia 0.24 7
* Girth at breast height.

Connell Hypothesis

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Compensatory Recruitment, Growth, and Mortality as Factors Maintaining Rain Forest Tree

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? 1984 by the Ecological Society of America

COMPENSATORY RECRUITMENT, GROWTH, AND MORTALITY

AS FACTORS MAINTAINING RAIN FOREST TREE DIVERSITY'

Department of Biological Sciences, University of California,

Santa Barbara, California 93106 USA

J. G. TRACEY2 AND LEONARD J. WEBB3

Rainforest Ecology Unit, CSIRO Laboratories, Division of Plant Industry,

Indooroopilly, Queensland, 4068 Australia

forests in Queensland, Australia.

are reduced, with the reverse for species that become

rarer, this will continually compensate for the tenden-

cies of some species to increase at the expense of others.

For this reason such general frequency-dependent pro-

cesses have been called "compensatory mechanisms"

(for general reviews see Connell [1978, 1979]). If such

mechanisms were operating at a site, rare species would

be protected from local extinction, and common species

could not continually increase at the expense of rarer

ones. Therefore several to many species could be main-

tained on a site indefinitely, without immigration.

This hypothesis assumes that commonness or rarity

is not a property of a species; a species is only tem-

' Manuscript received 12 July 1982; revised 11 February

porarily rare or common on a site. Given time, rare

2 Present address: Division of Forest Research, CSIRO,

crease. Over a long period the different species will

Atherton, Queensland, 4833 Australia.

fluctuate within upper and lower bounds that are de-

3Present address: School of Australian Environmental

fined stochastically, i.e., a population will exceed them

Studies, Griffith University, Brisbane, Queensland, 41 1 1 Aus-

tralia. only with some low probability (Chesson 1978).

Vol. 54, No. 2

This assumption contrasts with another commonly

held one that each species has an equilibrium popu-

lation size toward which it returns if perturbed away.

Under this latter assumption, common species remain

common, fluctuating around a high equilibrium pop-

ulation size, while rare species do the same around a

lower population level. In this framework, compen-

satory mechanisms are density-dependent changes in

reproduction, mortality, etc., as the population is either

above, or below, its equilibrium. By contrast, in our

framework compensation acts in response to the species'

local abundance relative to that of other species, and

hence is frequency dependent rather than strictly den-

sity dependent. Either assumption will tend to prevent

local extinction of rare species. The hypothesis to be

tested in the present paper makes no assumptions about

equilibrium levels; it simply suggests that when species

are relatively rare they will tend to increase, and when

relatively common, to decrease.

When studying long-lived organisms such as trees,

and particularly tropical species that cannot be reliably

AREA AND METHODS