© 2006 The Authors
© 2006 British
Blackwell Publishing Ltd
E. V. J. TANNER and P. J. BELLINGHAM*
Are more diverse ecosystems more or less resistant to disturbance? Does diversity
year study of four Jamaican forests, which differ in soil fertility and diversity, and which
were hit by Hurricane Gilbert in 1988; the decades were: pre-Gilbert (1974 –84), Gilbert
(1984 –94), and post-Gilbert (1994 –2004).
Diversity (Shannon index) was always higher in the three forests (Col H
2.91 and Slope H
signiﬁcantly lower in the Mor forest (H
N : P ratio 44). Diversity increased during the Gilbert decade in two of the more diverse
forests (Mull and Slope), it did not increase in the least diverse, Mor forest. The overall
increase in diversity during the Gilbert decade was due to the recruitment of eight,
mostly light-demanding, species and the increased abundance of uncommon species.
We used turnover rates (the average of mortality and recruitment of stems) as a
damage. Turnover increased during the Gilbert decade in all forests, but increased more
in the three more diverse forests (Mull 1.5% year
Slope 1.3–2.6; Col 1.5–3.2); than in the least diverse Mor forest (1.2–1.9).
Stem diameter growth rates pre-Gilbert were very low in all forests and were lowest
during the Gilbert decade and remained, in the post-Gilbert decade, double those of the
pre-Gilbert decade (Mor 0.6 mm year
increased growth more than larger stems. The stems recruited during the Gilbert and
post-Gilbert decades grew faster than those present in 1974.
Thus, in montane forest in Jamaica the least diverse forest was most resistant to
over 30 years including a hurricane, the hurricane increased diversity.
: diversity, hurricanes, mortality, recruitment, resistance
Natural disturbances affect all ecosystems, killing all
processes. With global loss of biodiversity a pressing
issue, determination of relationships between ecosystem
diversity and response to disturbance is of particular
interest: this has been the subject of studies over many
years (MacArthur 1955; Hooper
. 2005). However
response to disturbance remain elusive. While one recent
review concluded that ‘communities with greater diversity
tend to be stable’, with stable meaning ‘less oscillatory
and less susceptible to invasion by exotic species’
. 2001), others have concluded the opposite
Responses to disturbance may be anywhere in the
space deﬁned by the following two axes: (i) resistance
(‘staying essentially unchanged’, Grimm & Wissel 1997);
and (ii) resilience (‘returning to a reference state’, Grimm
Correspondence: E. V. J. Tanner (e-mail email@example.com).
Journal of Ecology
& Wissel 1997). Disturbances may be of many kinds,
for example wind, drought, ﬂooding, ﬁre, freezing and
herbivores, and attempts to generalize about how
ecosystems respond to disturbances, across different
disturbances and different ecosystems, are likely to
have many exceptions. If for no other reason, because
different disturbances cause different communities, for
example in an area of hundreds of hectares, with access
to the same species pool, an ecosystem disturbed by wind
is likely to be very different to an ecosystem disturbed by
ﬂooding. In addition, some recent experimental studies
of the relationship between diversity and ecosystem
properties have studied the ﬁrst disturbance to affect
the ecosystem (Tilman 1996). It is likely that natural
ecosystems have recurrent disturbances and that
susceptible species will have become locally extinct, or
have small population sizes, by the time a study of a
particular natural disturbance event is made. Thus,
patterns in nature will probably be very different from
patterns discovered in relatively short-term experiments.
Finally, it has been shown that it is not diversity itself
that deﬁnes ecosystem responses to disturbance but the
properties of the species in the ecosystem (Sankaran &
There are very few studies of how natural systems that
differ in diversity are affected by natural disturbances,
and even fewer of forests despite the likelihood that
major disturbances determine the species composition
of large areas of forest (Oliver 1980). Tropical rain forests
are among the earth’s most diverse ecosystems, and many
are subject to major disturbances such as hurricanes,
which have major effects on their growth and composition
(Everham & Brokaw 1996). Research into the effects of
hurricanes shows that while some of the initial effects
are destructive, some of the longer term, decadal, effects
may be positive, for example increased growth of surviving
trees has been reported following hurricanes (Merrens
& Peart 1992; Bellingham
. 1995; Scatena
The effects of disturbance on diversity in tropical rain
forests have been less well studied. In Kolombangara,
plots with higher diversity suffered more canopy damage
and had higher recruitment and turnover of the 12
commonest species (only 12 species were recorded after
1964) following two cyclones in the 6 years after the
start of the study in 1964 (Burslem & Whitmore 1999).
In Nicaragua a comparison of diversity between ﬁve
forests, four in one area hit by a hurricane and one in an
area not hit by the hurricane, showed higher diversity in
the hurricane damaged forests (Vandermeer
the high diversity when the plot was established was due
to a hurricane 15 years previously, though a hurricane hit
in 1998 caused no increase in diversity in that plot by 2000
(Weaver 2002). None of these studies have unambiguously
shown increased diversity caused by a hurricane (or
cyclone) for different reasons: Burslem & Whitmore (1999)
had no complete species records after their cyclones;
. (2000) had no pre-hurricane records;
records he found no increase in diversity.
Our study has both pre- and post-hurricane records
collected over a 30-year period from permanent plots
in four Jamaican montane rain forests. We address the
question of whether there is a relationship between tree
species diversity (on the one hand) and resistance and
resilience (on the other), using a natural gradient of
diversity/fertility and a natural disturbance, a hurricane.
In addition to the between-forest comparison of diversity
and resistance and resilience we report on the overall
effect of the hurricane on diversity and growth, by using
the combined data set from the four forests. Thus, our
null hypotheses are: (i) that there was no relationship
between diversity on the one hand and resistance and
resilience on the other; (ii) that diversity was unchanged
by hurricane damage; and (iii) that growth was unaffected
by hurricane damage.
We studied four Jamaican upper montane rain forests
within 300 m of each other (18
1600 m): Col forest (Gap forest of Tanner 1977, 0.09 ha
sampled); Wet Slope forest (0.1 ha); Mull Ridge forest
(0.1 ha) and Mor Ridge forest (0.06 ha); hereafter
abbreviated to Col, Slope, Mull and Mor (the ‘forests’
in the current paper are the ‘sites’ of Tanner 1977). The
forests were selected in 1974 (for a study of nutrient
cycling) as representative of forests in the western Blue
Mountains, the ﬁrst 10
subjectively positioned in ‘representative’ forest, the
other plots in each forest were contiguous. All stems
3 cm d.b.h. (at 1.3 m) in permanent 10
were tagged, painted with a ring at 1.3 m, identiﬁed, and
measured in 1974, 1984, 1989, 1991, 1994 and 2004. Multi-
stemmed trees were recorded. In total we had 2745 stems,
2171 individuals and 68 species; nomenclature follows
Adams (1972), except where other authorities are listed.
Soil (0 –10 cm depth) total C, N (CHN analyser), P (acid
digest) and Bray 1-extractable P were determined by
Brookside Laboratory Association Inc. (Knoxville,
Ohio, USA) from cores collected and air-dried in July
and August 2004, from three randomly chosen plots
Hurricane eyes pass over, or within 15 km of, the Blue
Mountains of Jamaica on average every 25 years. During
our study period (1974–2004) Hurricane Gilbert struck
in 1988; it was the most powerful hurricane recorded in
the Caribbean during the 20th century (Dodge
forests ‘immediately’ (Bellingham
. 1992) and 13%
crowns from 4% of stems and completely defoliated 19%
of stems, thus increasing light availability at the forest
ﬂoor for up to 33 months after the hurricane (Bellingham
. 1995, 1996). We report results over three decades
(1974 –84), Gilbert decade (1984 –94), and post-Gilbert
decade (1994 –2004).
For the tree stems (not the individuals) we report Fisher’s
= number of species in
= number of stems in the sample,
alpha) and the Shannon index, H
all species of (
= abundance of
= total number of stems in the sample). We
between (unreplicated) forests
following the worked example in Magurran (1988); in
our data cumulative H
two plots in each forest. We carried out rarefaction
procedures using the ECOSIM program (Gotelli &
Entsminger 2006), in which we used the number of stems
(1716) per species in 1984 (Appendix S1 in Supplementary
Material) to estimate the number of species for 1716 stems
from the data sets for 1994 and 2004. For each decade we
calculated mortality, recruitment and turnover. Mortality
as percentage is: 100(
= 1 – [1 – (
= number at the beginning of a period,
of survivors at the end of a period, and
= time in years.
= 1 – (1 –
= number of recruits during a period and
average of mortality and recruitment. All comparisons
of mortality and recruitment were across equal decade
intervals so problems in assessing rates across different
census intervals were avoided (Kohyama & Takada 1998).
Mortality and recruitment of stems were compared
between forests using contingency tables. We used regres-
sion analysis to compare the number of tree stems
per species in 1984 with those in 2004 and compared the
ﬁtted line with a null hypothesis relationship of 1 : 1. We
compared the gains and losses of species overall (all
forests combined) using a binomial test, with the null
hypothesis that the number of species recruited should
equal the number lost. We calculated Euclidian distances
to show departure from initial biomass composition as
. (1982), biomass calculated using equations
were compared between forests within a decade by Mann–
-tests and between decades by paired
before and after the hurricane were related to trunk size,
we calculated, for the 977 trunks alive in 1974 and 1991,
the mean absolute growth in trunk diameter from 1974
to 1984 and divided it by 10, and the mean absolute
growth in trunk diameter from 1989 to 1991 and divided
it by 2. We plotted the difference between these two absolute
rates against trunk diameter in 1989 and ﬁtted a linear
regression; we compared the intercept and slope of the
ﬁtted line against a null hypothesis of zero for both.
Diversity was always highest in the three forests on more
fertile soils (Fig. 1) with lower C : N and C : P ratios
(Table 1); diversity was signiﬁcantly lower in the Mor
with very infertile soil (comparisons of H
for 1984: Mor
< 0.001; Mor vs. Col,
< 0.001; Mor vs. Slope,
< 0.001; all other
the hurricane in two of the three forests on more fertile
soils (Mull and Slope); in contrast, the least diverse Mor
forest showed no change in diversity throughout the
study (Fig. 1). Summed over the four forests there was
an increase in diversity in response to the hurricane
(Fig. 2, Appendix S1) due to the net recruitment of species
not previously recorded in the plots (but present, though
rare, in the surrounding forest). The increase in diversity
was a result of the increase in the number of stems because
rarefaction analyses, when the number of stems was kept
constant, gave no statistically signiﬁcant increase in species
post-Gilbert. Between 1984 and 1994, eight species were
recruited and none were lost (exact binomial test,
(occasional and rare = lower 40% of species in terms of
rank abundance), except for the invasive alien tree
, which by 2004 was, in terms of individuals,
The changes in populations of trees resulted in a drift
᭹ = Col; ᭢ = Slope; ᭺ = Mull; ᭞ = Mor.
away from the initial biomass composition, but less so
in the least diverse Mor forest (Fig. 3).
Mortality rates did not differ between the forests in the
pre-Gilbert decade (Fig. 4, Appendix S2); they increased
in all four forests during the Gilbert decade but increased
less in the Mor and Slope than in the Mull and Col
(Fig. 4). Recruitment rates in the pre-Gilbert decade
were lower in the Mor than the Mull and Col forests
(Fig. 4, Appendix S2) and, like mortality, increased
during the Gilbert decade but much less in the Mor
(Fig. 4, Appendix S2). Thus, the three forests on more
fertile soils were the most dynamic; that is, they had
higher turnover rates (mean of mortality and recruit-
ment) than the Mor, in the Gilbert decade (Fig. 4,
Across the four forests, 38% of the stems present in
1974 had died by 2004, and 47% of the stems present in
2004 were recruits since 1974; 32% of the basal area
present in 1974 was dead by 2004, but only 3% of the
basal area in 2004 was recruited since 1974.
Stem diameter growth was very slow in all forests,
but especially so in Mor (Mor 0.3 mm year
Slope 0.5, Col 0.6 from 1974 to 1984, Fig. 5); diameter
growth increased during the Gilbert decade and remained
Table 1 Soil (0–10 cm) nutrient concentrations, mean values from one core from each of three randomly chosen plots per forest
plot) in 1984 and 2004; solid line is best-ﬁt line ( y = 0.80 (SE
0.055), x + 0.32 (SE 0.064)). The slope of the line is signiﬁcantly
less than 1.00 (P < 0.05), and the intercept is signiﬁcantly
greater than 1.00 (P < 0.01), the expectation if numbers of stems
per species were the same in 1984 and 2004 (dashed line); thus
species were recruited, rare and uncommon species increased
and common species were unchanged.
composition using data for biomass per species in each forest,
calculated as in Leps et al. (1982):
᭹ = Col; ᭢ = Slope; ᭺ =
᭞ = Mor; arrow denotes impact of Hurricane Gilbert in
mortality of stems, in each of four forests in three decades;
mean per forest and standard error of the mean to show the
variation between plots within a forest. The variation between
forests (compared by contingency tables) was not signiﬁcant
(P > 0.05) for mortality or recruitment from 1974 to 84, but was
signiﬁcant for both mortality and recruitment in both 1984 –
94 and 1994–2004.
= Col; ᮀ = Slope; = Mull; = Mor.
Journal of Ecology,
higher in the post-Gilbert decade 1994 –2004 (Mor
0.6 mm year
, Mull 0.6, Slope 0.8, Col 1.1, Fig. 5). The
Gilbert growth was signiﬁcantly lower in Mor compared
with the other three forests, but in the last decade stem
growth in the Mor was not signiﬁcantly lower than in
the others (Appendix S3). Furthermore, in those stems
present from 1974 to 1991, growth in the Gilbert and
post-Gilbert decades increased more in smaller than larger
stems (Fig. 6, Appendix S4). The stems recruited during
the Gilbert and post-Gilbert decades grew faster both
absolutely (compare Fig. 5a with Fig. 5b) and relative
to trunk size than those present pre-Gilbert.
Mor forest, the least diverse throughout the study, is on
the crest of the main ridge of the Blue Mountains and
was as exposed to the hurricane as any of the four forests.
Although we studied in detail only one Mor forest, which
limits the statistical tests we can carry out, it is very similar
in species composition and physiognomy to other Mor
forests within 5 km. Despite the fact that 10.1% of
surviving stems in the Mor forest were severely damaged
(deﬁned as trees tipped up to > 40
° from their pre-
hurricane position and /or crown death) by the hurricane
(vs. 9.5% in the Mull forest, 7.0% in the Col forest, and
5.8% in the Slope forest), the Mor forest had lower
turnover than the other three forests. The low turnover
was probably a result of at least ﬁve factors. First, the
species in the Mor are less susceptible to being killed by
the hurricane. Evidence for this comes from the Mull,
where for the 277 individuals of species that also grow
in the Mor mortality was 16% (1984 –94); in contrast,
for the 245 individuals of species which grew in the
Mull but not the Mor, mortality was 35% (
P < 0.0001) (though there was no relationship between
mortality per species and wood density, just as there
was no correlation between wood density and damage
due to the hurricane, Bellingham et al. 1995). Secondly,
multiple-stemmed trees, which were more prevalent in
the Mor forest than the others (in 1974 38% of individuals
in Mor forest had multiple stems vs. 9–17% in the other
= 93, P < 0.0001), have a lower chance
of dying than single stemmed trees (of 1126 individuals
with single stems 21% died between 1984 and 1994;
in contrast, of 227 trees with multiple stems only 8%
= 14.4 P < 0.0001). Thirdly, there was a high
of the Mor (Bellingham et al. 1994), which reduced indi-
vidual mortality. Fourthly, the Mor forest has shorter
trees; this probably resulted in less wind-caused mortality.
Fifthly, low mortality in the Mor forest was probably
partly a result of the weak, c. 50-cm deep, mor humus
that allowed the trees to ﬂex in the wind, thus reducing
their canopy damage (although there was substantial
crown damage to one common tree in this forest, Cyrilla
racemiﬂora, Bellingham et al. 1995). The other half of
‘turnover’ is recruitment, this was also lower in the Mor,
due to the very infertile, acidic (pH c. 3.0) mor humus,
which we hypothesize limits the number of species that
can grow in the forest (Grubb & Tanner 1976). Evidence
supporting this hypothesis comes from an experimental
clearing of 10
× 10 m in the Mor forest; 10 years after
its creation the zone from which mor humus had been
removed, exposing the subsoil, had seedlings of 14 tree
Fig. 5 The absolute diameter growth rate of stems (mean
decade; (b) stems alive throughout, i.e. in 1974, 1984, 1994 and
= Col ; ᮀ = Slope; = Mull; = Mor.
growth 1989–91 and mean annual absolute growth 1974–84
), by stem size classes, for those stems alive in both
species, and 13 seedlings m
; in contrast, undisturbed
(Sugden et al. 1985). We conclude that characteristics
of the high resistance and low resilience to hurricane
damage, not low diversity per se.
Mor forests are atypical in composition, structure and
response to Hurricane Gilbert (our Mor was resistant
not resilient) among Jamaican montane rain forests.
Why are resistant forests like the Mor forest not more
widespread? We think it is because in the steep Blue
Mountains there are very limited areas in which deep
acidic litter can accumulate and the process of feedback
begin and be maintained; even on ridge crests such
forests are conﬁned to knolls. However, in places where
topography does not limit litter accumulation and where
hurricane disturbance is frequent, the ‘resistant’ syndrome
we found in Mor forest also appears to be uncommon (e.g.
lowland Nicaragua, Vandermeer et al. 2000). Nutrient-
limited forests in Hawaii were found to be more resistant
to hurricane damage and less nutrient-limited forests
more resilient (Herbert et al. 1999). We propose that at a
landscape scale, nutrient supplies have to be limiting (as
shown by low leaf nutrient concentrations in montane
rain forests), which promotes the colonization by trees
typical of low nutrient systems (e.g. the Ericales), and
within such areas some sites will be especially nutrient
poor and by feedback mechanisms generate mor soils
and forests with their distinct species composition, which
are resistant to hurricane damage.
Forests on more fertile soils than Mor forests were
characteristically resilient to hurricane damage. In these
forests, diversity increased after hurricane disturbance
as a result of an increase in abundance of uncommon
species and recruitment of species not previously recorded
in the plots. These two phenomena are the same; it is a
matter of the scale at which the effect is measured. If the
plots had included all the forest in the area, the effect
would have been only an increase in abundance of the
rare and uncommon species. The promotion of the
uncommon species was a result of two patterns. First,
the canopy trees present in 1974 were mostly of more
light-demanding species that largely lacked seedlings in
the understorey. Of the 10 most common trees, two had
no seedlings and ﬁve had few seedlings before Hurricane
Gilbert (Healey 1990). Secondly, the damage to the
canopy trees allowed more light to reach the lower
layers of the forest, promoting germination of seeds of
light-demanding species and the growth of the seedlings
and saplings of the advance regeneration into the recorded
minimum size class (3 cm d.b.h.). Most of the species
that increased in abundance during the Gilbert decade
were light-demanding species (for example Hedyosmum
shade-tolerant tree also increased (Eugenia virgultosa).
Furthermore, we judge that most of the nine species that
were recruited after the hurricane were light demanding,
although we have little detailed information about their
ecology (almost by deﬁnition because they were mostly
rare): two are deﬁnitely light demanding (Brunellia
comocladifolia and Miconia dodecandra), four are likely
to be light demanding (Cestrum hirtum, Critonia parvi-
and two we cannot judge. However, one common
recruit, Pittosporum undulatum, which is an invasive
alien, has very shade-tolerant seedlings (Bellingham
demanding species, but at least two shade-tolerant
species also increased in abundance.
We think it likely that many hurricane-affected forests
will show an overall increase in diversity after hurricanes.
Indeed, any forest where the canopy trees have a lower
diversity than the advance regeneration, a pattern true
for most tropical forests on a scale of hundreds of square
metres, is likely to show an increase in diversity after
major wind storms, which severely damage the canopy
and increase light and thus growth of the more species-
rich advance regeneration. Such a response follows general
predictive models of short-term responses to disturbance
(Connell 1978; Huston 1979) and agrees with predictions
from a modelling study of higher diversity in hurricane-
damaged forest in Puerto Rico (Doyle 1981).
The theoretical predictions that hurricane disturbance
will increase diversity ﬁnd some support in the literature.
Hurricane-affected forests had higher diversity com-
pared with a less affected forest in lowland Nicaragua
(Vandermeer et al. 2000). In Kolombangara, plots that
were more damaged by cyclones had increased recruit-
ment and turnover of the 12 commonest species, but
effects on overall diversity were not recorded (Burslem
& Whitmore 1999). However, a 54-year study of diversity
(1946–2000) before and after hurricane damage in a
permanent plot in Puerto Rico did not show an increase
in diversity following two hurricanes, in 1989 and in 1998
(Weaver 2002). The lack of increase might be due to the
short interval, in which case later enumerations may
reveal an increase, or it could be due to the dominance of
two very light-demanding species (Cecropia schreberiana
and Psychotria berteriana) among the trees recruited after
the hurricane. If so, there may be areas in that forest
where increased light due to canopy damage was enough
to strongly promote the growth of pre-existing seedlings
but not so high that the seedling populations were
dominated by one or two strongly light-demanding
species. Short-term increases in diversity have also been
found after other kinds of disturbance in tropical rain
forests, such as logging, again due in part to recruitment
of light-demanding species (Molino & Sabatier 2001).
Recently it has been shown that in tropical rain forests
without large disturbances, populations of rare species,
in small quadrats, survive and grow better than those of
common species (Wills et al. 2006); however, note that
this study differs from the previous studies because
recruits were not included in the analysis by Wills et al.
Thus it seems that rare species fare better than common
species in a whole range of disturbance regimes.
Despite the signiﬁcant increase in rare and uncommon
species in Jamaica, and despite the fact that approxi-
mately half of the stems present in 2004 were recruits
since 1974 (and 32% of the basal area present in 1974 was
dead by 2004), there was a strong rank correlation in
species abundance between 1974 and 2004 (r
in Jamaica, across 30 years including a major hurricane,
is similar to the pattern across 54 years in Puerto Rico
(with hurricanes 45 and 52 years into the study, Weaver
2002), and similar to the pattern for the 12 most common
species across 30 years in Kolombangara (with four
cyclones, only two of which damaged the forests, 3–
6 years after the study started, Burslem et al. 2000).
Thus, even in hurricane-damaged forests the major
pattern is for species composition to change little over
Our results from Jamaica show more resistance to
hurricane damage in our least diverse forest. A similar
pattern was found across two forests in Puerto Rico, where
higher-altitude lower-diversity forest was more resistant
than higher-diversity lower-altitude forest (Walker et al.
1996). In Jamaica, resilience was also lower in the least
diverse forest when judged by recruitment of stems during
the Gilbert decade, though perhaps this is best seen as a
delayed resilience because recruitment increased in the
Mor in the post-Gilbert decade. Growth increase in the
surviving stems post-Gilbert, another form of resilience,
was also higher in Mor forest. Thus, our results do not
support the conclusion of Tilman et al. (2001) that
‘communities with greater diversity tend to be stable’.
Hoping for a generalization across very different
ecosystem types (forest to grasslands, e.g. this study vs.
Kahmen et al. 2005), and across different disturbances
(wind to drought), and across natural systems previously
exposed to disturbances as compared with experiments
where the ﬁrst disturbance is studied, is probably too
much, though it will probably be possible to generalize
for subsets of these categories. Our results from forest
in one area of the world show that low diversity was
associated with high resistance to disturbance by wind.
In summary, although many initial effects of hurricanes
on forests are destructive, because trees are destroyed,
crown areas reduced and emergent crowns destroyed
(Brokaw et al. 2004), this very destruction increases light
for smaller trees, thus increasing growth, and in this
study, diversity. Comparisons between forest differing
in diversity show that if there is an emerging pattern, it
is that least diverse forests are more resistant to severe
Earlier papers have acknowledged the collaborators and
funders of this long-term study. The 2004 enumeration,
funded by the Drummond Fund of Gonville & Caius
College, Cambridge, and by a Manaaki Tangata fellow-
ship from Landcare Research, New Zealand, was carried
out by the authors and Howard Beckford. N. Brokaw,
D. Burslem, D. Coomes, H. Grifﬁths, P. Grubb, D. Wardle
and an anonymous reviewer made valuable criticisms
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The following Supplementary material is available
online from www.Blackwell-Synergy.com
Appendix S1 The number of stems of each species in
1974, 1984, 1994 and 2004.
(a), recruitment (b) and turnover (c) between forests in
the three decades..
Appendix S3 Mean diameter growth rates per decade for
stems in the four forests alive in 1974 and 2004.
stem diameter growth 1989–91 and that from 1974 to 84,