Testing the intermediate disturbance hypothesis: Diversity pattern

differs between natives and invasives?

Kwo Mun Wong
Abstract
The Intermediate Disturbance Hypothesis (IDH) posits that species diversity in an ecosystem is
maximized at intermediate levels of disturbance intensity or frequency. This study explores the
applicability of IDH to both native and invasive species across five distinct habitats including
fragmented habitats, riparian zone, forest and hemlock forest with varying levels of disturbance.
The results indicate that, in general, the diversity patterns of both native and invasive species
follow a bell-shaped curve consistent with IDH, with peaks in diversity at medium levels of
disturbance. However, some variations exist, and it is important to interpret these results with
caution. Furthermore, the study identifies differences in diversity patterns between native and
invasive species. Invasive species tend to thrive in habitats with higher disturbance levels, while
native species peak in diversity at intermediate disturbance levels. This suggests that native and
invasive species may respond differently to disturbance, potentially due to variations in life
history traits and characteristics. The study also highlights the influence of habitat type and
disturbance type on the diversity of native and invasive species. This research provides insights
into the application of IDH to native and invasive species in different habitats and disturbance
levels. While the study generally supports the IDH concept, further research is needed to better
quantify disturbance levels and explore additional factors that contribute to the complex
relationship between disturbance and species diversity and its relationship between native and
invasive species.
Introduction
The intermediate disturbance hypothesis (IDH) states that the highest levels of species diversity
are found at intermediate levels of disturbance intensity or frequency within an ecosystem
(Connell, 1978). Disturbance takes on various forms, natural events like fire, storm and
anthropogenic events like pollution and habitat destruction. Some papers define disturbance as
any event that increases resource availability which includes biotic interactions like grazing
(Carford et al. 2012, Connell, 1978). Moreover, disturbance could be separated into frequency
and intensity, duration, timing, spatial variability (Carford et al. 2012, Shea et al. 2004). For
Simplicity, this study will focus on disturbance frequency. According to Connell (1978), low
disturbance favors late successional species which outcompetes other inferior species, reducing
the species diversity. At high disturbance, it eliminates late successional species as they require
more time to grow, and only fast reproducing species survive. Thus, Connell (1978) conclude
that a moderate level of disturbance maximizes species diversity. However, studies have found
mixed results on IDH. A study found evidence of IDH across shrubs, grass, trees with varying
anthropogenic disturbance (Mayor et al., 2012). While Bonger et al. (2009) found that IRH only
explains species richness in dry tropical forests but not in wet and moist tropical rainforests,
indicating disturbance is less important in maintaining species diversity. They also noted that

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density-dependence might slow down the decrease of species richness under low disturbance,
making IDH harder to detect (Bonger et al. 2009).
The invasion of species poses a significant threat to the stability of natural ecosystems, resulting
in annual economic losses amounting to billions of dollars worldwide (Mack et al. 2000). Recent
studies suggested natives fits IDH but invasive species does not and benefits from disturbance
due to its rapid life cycle and high reproductive output (Catford et al. 2012, Jauni et al. 2015).
Luong et al. (2023) reported natives and invasive bunchgrass exhibit different response to
disturbance. In a meta-analysis, Jauni et al. (2015) found that diversity and abundance of
invasive plants are significantly higher at disturbed sites depending on the type of disturbance.
(Vujnovic et al., 2002) found evidence of IDH in grasslands in Alberta and show that invasive
species increase significantly with magnitude of disturbance. However, not all research findings
have indicated a positive correlation between disturbance and plant invasions. Hester and Hobbs
(1992) found that fire does not increase invasion and even increase abundance of natives. While
(Stohlgren et al., 1999) found no significant difference between natives and invasives in response
to grazing. Other studies even found negative relationships between disturbance and invasive
grass species (Cox and Allen 2008).
Limited studies have explored whether native species are favored under low disturbance. On the
other hand, The positive relationship between disturbance and invasion could be explained by the
difference between life history traits in natives and invasives. High reproductive output, early
and quick germination, traits that are common in invasive plants allow them outperform natives
under disturbance (Orbán et al., 2021). Moreover, Invasive plants in general have higher
phenotypic plasticity than natives and can better adapt to disturbance (Davidson et al. 2011). In a
Broader perspective, disturbance increases resource availability and opens up invasive windows
and favors invasive species with high resource use efficiency (Davis et al. 2000). Other studies
have reported disturbance altered resilience of native communities and favors invasion (Manea et
al. 2016). This is supported by Collinge et al. (2011), which shows that invasive species benefit
by altered community structure. Scale of study might explain the insignificant relationship found
between disturbance and invasion. It is reported that disturbance induced invasions are stronger
at small scales (Juani et al. 2015).
This study aims to investigate (1) whether the intermediate disturbance hypothesis is supported
in our sites for both native and invasive species. (2) whether the diversity patterns of native and
non-native species differ and what makes species more or less able to thrive at different levels of
disturbance?
I hypothesize that both native and invasive species would follow IDH. I also hypothesize that at
higher disturbance, there will be more invasive species than natives.

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Materials and methods
Habitat description
Plot one is an isolated piece of land that is made of sediments like soil and rocks in the middle of
a stream located in Morningside Park in Scarborough, Ontario (43.780790, -79.182290). This
habitat represents the highest disturbance site. Disturbance might include flooding, erosion wind
and anthropogenic events (Pattison et al. 2016). Moreover, plot one is a fragmented habitat, and
it is found that rate of invasion is generally higher in these habitats (Vilà & Ibáñez 2011).
Plot 2 is located more uphill than plot 1 (43.780682, -79.182017). This site represents medium
high disturbance. Plot 2 is in the riparian zone, which is defined as the biotic communities along
streams and lakes and is known as the transitional zone between the stream and uplands (Naiman
& Décamps, 1997). Common disturbances in the riparian zone include windthrow, insect
disturbance, treefall caused by snow loading, bank erosion, floods and debris flows (Moore and
Richardson 2012). It is also reported that trees in the riparian zone have shallower rooting
depths, making it more vulnerable to disturbance (Moore and Richardson 2012).
Plot 3 is more upwards compared to plot 2 on the same latitudinal gradient (43.780570, -
79.181866). This site represents medium disturbance. It could be categorized as the riparian zone
since the spatial extent of the zone is hard to precisely delineate (Moore and Richardson 2012). It
is likely that this area is also influenced by the elevated water table and contributes organic
matter to the stream and could be included in the riparian zone (Moore and Richardson 2012).
Common disturbance includes erosion, wind, snow loading and anthropogenic disturbance. Since
the area is distant from the stream, it is less affected by floods and debris flow and represents
medium disturbance. Also, it is important to note that this area is located next to a road, which
could increase wind penetration and solar radiation, affecting its species composition and
microclimate (Herunter et al. 2003).
Plot 4 is a well-established forest on a slope located closer to the University of Toronto
Scarborough (43.781428, -79.185205). This site represents medium low disturbance since it is
populated with hardwood trees that are late-successional species. Disturbances include erosion,
surface runoff, snow loading and wind and potentially anthropogenic sources. Landslides might
also be a source of disturbance.
Plot 5 is a hemlock forest dominated by late successional species that is adjacent to the trail
leading to University of Toronto Scarborough (43.781965, -79.187370. Hemlock is considered
the most dominant species in most sites but requires minimum disturbance to be established
(Woods, 2000). In other words, hemlock forest is a good indicator of an area with low
disturbance. Common disturbances in hardwood forests include wind, drought, insects, lighting
strikes, fires and storms(Frelich & Lorimer, 1991; Woods, 2004). It is also reported that
hardwood forests have a rotation period of 1200 years for a catastrophic disturbance (Frelich &
Lorimer, 1991).

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Data collection and analyzation
Five habitats with varying levels of disturbance (from high to low) are assessed. Multiple groups
of students collected species on 18th September within a 4*4 quadrate, this includes leaves from
trees that intersect with the quadrate and plants on the ground. Samples are stored in separate
labeled bags and were identified in the lab on 2 nd October. Name of plants and its native or
invasive status are identified using Plants of Southern Ontario (Dickinson & Royer, 2014). Data
were inputted into excel and descriptive data analysis is used to spot any patterns.
Results
Both native and invasive species diversity approximately followed a hump-shaped curve (figure
1). The number of invasive species and native species peaks at plot two and plot three
respectively (figure 1). Only plot one has more invasive species than native (figure 1). There are
slightly more native species in plot five than plot four (figure 1). If we integrate both species
diversity, there is a rising trend in the total species diversity from plot one to three with plot three
having the maximum number of species (figure 1). Total species diversity dropped dramatically
in plots four and five in which plot four have less species than plot two and slightly less species
than plot five (figure 1).
Discussion
Is the intermediate disturbance hypothesis supported by both native and invasive species?
The overall pattern combining both native and invasive species approximately fits the IDH
except Plot four having one less species than plot five although this difference is likely not
significant (figure 1). Native and invasives, if analyzed separately also approximately follow a
hump-shape curve with invasives peaking at plot 2 and natives peaking at plot 3 (figure 1). The
IDH is supported by large amounts of studies in different areas, different types of habitats, under
different types of disturbance, with 46% conformance in a meta-analysis (Vujnovic et al. 2002).
Mayor et al. (2012) showed evidence of IDH in boreal forest which is the type of forest in
Canada and in our site. Although my data fits the IDH, it does not necessarily mean IDH could
explain the whole variation of species diversity. Different frequency and intensities of
disturbance interact, forming the diversity-disturbance relationship, the same IDH could be
reproduced with different intensity and frequency (Miller et al. 2011). Moreover, life history
traits and any other tradeoffs like colonization-competition, reproduction-survival are likely to
affect species’ response to disturbance (Catford et al 2012). The most common explanation of
IDH is the reduced competition at intermediate disturbance, allowing more species to co-exist.
Roxburgh et al. (2004) showed that there are other mechanisms, for example storage effect and
relative nonlinearity could also produce the same IDH disturbance diversity relationship
(Roxburgh et al. 2004). The storage effect is defined as species “storing” gains in a favorable
period so that during disturbance it suffers less impact and relative nonlinearity is based on
different species reacting differently to fluctuations of resources, allowing co-existence
(Roxburgh et al. 2004). To add to the complexity, it is found that the same magnitude of
disturbance could result in different patterns of species diversity (Svensson et al. 2009). Since I

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only used descriptive data analysis in this study, results should be interpreted with caution and
more studies are needed to support IDH.
Do the diversity patterns of native and non-native species differ and what makes species
more or less able to thrive at different levels of disturbance?
Our results show that diversity patterns differ between natives and invasives (figure 1). The
number of invasives is higher in all plots except plot one (figure 1). Although both natives and
invasives follow the IDH, invasives species diversity peaks at medium low disturbance, while
native species diversity peaks at medium disturbance (figure 1). Difference in response between
natives and invasives are expected and documented in other studies (Mayor et al. 2012, Jauni et
al. 2015). However, our results do not fully agree with these studies. Mayor et al. (2012) showed
that in boreal forests, only native species fits the IDH pattern and invasive species richness
increases linearly with disturbance. Jauni et al. (2015) reported contradicting results, they found
that disturbance favors invasive species but has no significant effect on natives. In contrast, my
result showed that both native and invasive species respond to disturbance and approximately
followed the IDH. In support of our results, Catford et al. (2012) mentioned that natives and
invasives are expected to respond to disturbance as they are constrained by similar tradeoffs.
The difference in life history traits and characteristics could probably explain the difference
observed between native and invasive species (Juani et al. 2015). Invasive plants have higher
plasticity than native species (Davidson et al. 2011), making it more adapted to disturbance.
Moreover, invasive plants often have a higher growth rate, resource-use efficiency, and higher
fecundity than native plants (Funk and Vitousek 2007, Pyšekand Richardson 2007, van Kleunen
et al. 2010). These traits could potentially affect species’ response to disturbance. Moreover, the
majority of invasive species are herbaceous pioneer or early successional plants which explains
why it outnumbers natives in the highest disturbance habitat. Davis et al. (2000) linked invasion
with the fluctuating of resources available due to disturbance. This potentially explains why
native species diversity is higher than invasives in most plots. At medium to high disturbance,
there are less resource fluctuations thus less “invasion windows”. (Trammell et al. 2020)
suggests that forests can resist plant invasions which might explain the low number of invasive
species in plot four and five.
Difference in Habitat and disturbance type might partly explain the difference between natives
and invasives. Plot one is an fragmented habitat, it is found that the rate of invasion is generally
higher in these habitats, fragments of small size also have more invasive species than natives
(Vilà & Ibáñez, 2011). The high disturbance frequency and intensity in plot one is also linked with
higher vulnerability to invasion due to resource availability (Catford et al. 2012). Plot 2 belongs
to the riparian zone and has been reported for being highly vulnerable to invasion (Meyer et al.
2021). Anthropogenic disturbance like modifications of the upland could alter the habitats
properties, potentially supporting more invasive species (Meyer et al. 2021). For plots four and
five, It is found that forests can resist plant invasions which might explain the low number of
invasive species (Trammell et al. 2020). Natives and invasives might respond to disturbance type
differently. According to Juani et al. (2015), anthropogenic disturbance favors plant invasions

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with nature disturbance only increases invasives diversity or abundance. More research is needed
to confirm whether natives and invasives respond to the site-specific disturbance differently.
Improvements and future directions
The scale of disturbance is not clearly defined in this study, thus intermediate disturbance is also
not clearly defined. The difference between low to intermediate and intermediate to high
disturbance is not the same. This is problematic as it does not capture the whole disturbance
variation in the site, making validation of IDH inaccurate. Future studies should quantify
disturbance for a more robust conclusion. Future studies could investigate additional factors
contributing to the IDH relationship as it is more than merely reducing competition.

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Figures

30 native species
invasive species
25
Number of Species per plot

20

15

10

0
1 2 3 4 5

Plot types

Figure 1. shows the number of invasive and native species in each location with varying types of
disturbance. All species are counted within a 4*4 quadrant in each habitat type and disturbance is
ranked from 1-5 with plot 1 having the highest disturbance and closest to the river and plot 5
being a hemlock forest that has the lowest disturbance.

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