Theoretical Plant Population Dynamics and Practical

Management Strategies for the Restoration of Degraded

Ecosystems
M. J. Crawley and J. R. Mauremootoo

Abstract
Management of alien pest populations is reviewed in the light of modern population dynamics theory.
The evident lack of any useful life history traits for predicting either the probability of invasion, or the
controllability of established aliens, has meant that past attempts at eradication or reduction have tended
to be piecemeal efforts based on trial and error. With multiple infestations, where several invasive alien
species require simultaneous management, the prospect of simultaneous control is often greatly
diminished, because population expansion of the least effectively controlled species can compensate for
(and undo) all of the benefits contained by reduced abundance of the most amenable pests. The
management implication of contrasting models of community dynamics are discussed (competitive
lotteries, seed limited systems, herbivore limited systems, spatially structured systems, Lotka-Volterra
dynamics, etc.)

Introduction
In this paper we take a theoretical ecologist’s look at what lessons we can learn from practical attempts
to restore ecosystems. The aim is to find out how this might help us to generalise and hence to carry out
future restorations more efficiently, more quickly and on a larger scale.

This paper covers 4 separate themes which are as follows:

1. What makes a good invader? Are there traits of species that allow us to separate good invaders from
less good ones?
2. Population dynamics: The process of invasion. Is there anything about the dynamics of populations
that we can learn and that might help us restore degraded ecosystems?
3. What makes a community invasible? What do multi-species, multi-trophic dynamics tell us about
the invasibility of a community?
4. Techniques and monitoring. How can we construct protocols to tackle questions like those raised in
the above sections?

What makes a good invader?

In the early days of ecological thinking about biological invasions this was the question that everybody
focused on. There was a lot of information and people believed that there would be patterns in this
information which would allow us to separate the traits of good and bad invaders. In the 60s and 70s
there was a lot of thought given to the notion of what would constitute the ‘ideal weed’, the word ‘ideal’
being used in an ironic sense. There was some extremely influential work by Baker (1965). ‘Baker’s
list’ was a list of the attributes of the ideal weed.
When scientists looked at the list critically they discovered that it did not give reliable
predictions (Noble 1989). Not a single trait in Baker’s list stood up to critical analysis. The reason for
this was that the list paid no heed to the basic ecological notion of trade-offs (Stearns 1989). For
example increased reproductive rate almost inevitably has a consequence in terms of reduced
subsequent survival. Baker’s list was devoid of ecological context. It was as if you could think about the
ecology of a species without thinking about the ecosystem within which it was placed. Information such
as the successional stage of the ecosystem, the competitors of the species of interest and its herbivores
were not addressed.
If some theoretical ecology had been applied from the beginning then people would have
realised that you cannot have traits that separate good and bad invaders. The most important single fact
in population dynamics theory is summarised in a piece of jargon known as the invasion criterion which
states that ‘all species must be able to increase when rare’ (Crawley 1997). If this wasn’t true then
every time there was an environmental calamity the species population would be knocked down in
abundance. If it was unable to increase in number the population would be further reduced after the next
calamity and so on until it inevitably became extinct. This means that the change in numbers over time
(dN/dt in the terminology of population ecology) must be greater than zero when N is small. You can
take the integral of dN/dt as a constant rN (r is known as the intrinsic rate of increase). We can ignore
density dependence because we at the beginning of an invasion and population density is low. This
means that the invasion criterion is simply an r max of greater than zero, which is to say that  (ermax , the
net multiplication rate) has to be greater than 1. Given that this has to be true and that all species have
to be able to increase when rare, this means that all species possess the traits for invasion.
In spite of this assertion there are some patterns in conservation weeds that are extremely simple
but all-pervasive. First, they all form thickets within which recruitment both of themselves and of other
species is precluded. Secondly, they are dispersed by birds. These are patterns but they are not very
useful to the conservation manager after the invasion has already occurred.

Population dynamics: The process of invasion

It is very important to distinguish the 2 phases of an invasion. If we do not we get confused
because the rules that apply to one phase do not apply to the other. The first phase we call establishment
and spread and the second part we term equilibrium dynamics (Williamson 1989).
Establishment and spread
In this early part of an invasion the alien species has arrived in Mauritius, has been judged by the
environment and has passed the invasion criterion (dN/dt > 0) and its population is now increasing. The
key question is what determines its rate of spread through space? The answer is related to dispersal,
effective fecundity and so on. We have not looked at any of these sorts of problems during this
workshop for the simple reason that the problem conservation weeds in Mauritius have been here for a
long time.
Equilibrium dynamics
This phase is of far more relevance to the current situation in Mauritius. Whether or not these
weeds are precisely in equilibrium, we can still term the dynamics of the most important conservation
weeds in Mauritius equilibrium dynamics. The range has already been filled and so dispersal is
essentially irrelevant. The key questions now are questions like what determines interspecific
interactions? How important are herbivores? How important is the identity of competing plants? What
determines interspecific interactions with natives and other alien species? The key management
questions related to equilibrium dynamics are, once we set our target level of equilibrium abundance,
how exactly do we reach this and keep it? These are questions of population dynamics. These questions
can be related to control of alien species and measures to enhance the recruitment of native species.
Recruitment
Intuition and most theoretical models say the seed input per unit area at time t is directly
proportional to recruitment at time t+1 (Fig. 1.). It is critical to be clear about what we mean by
recruitment. This is the number of plants that will become reproductive adults, not seedlings as the vast
majority of these will die (Crawley 1997). If you assume that recruitment is directly proportional to
seed production (ignoring for the sake of simplicity such factors as seedbanks and immigration) then the
introduction of a seed predator as a biological control agent against your target weed is a step in the
right direction.
However, the real situation might not be so simple. The most obvious reason for this is microsite
limitation. This means that only a certain number of seeds could possibly become mature plants no
matter how benign the environment was (Grubb 1977). That would mean a change in our graph from a
linear to an asymptotic form. So once you get beyond a threshold density of seed, adding more seed will
not make any difference. This is often the situation for weeds so generally biological control involving
seed eating animals will not be useful (Huffaker and Messenger 1976). In our hypothetical example a
seed predator that reduces seed numbers by a factor of 2 will have no effect on recruitment.
Predator satiation is the other reason why recruitment may not necessarily be directly
proportional to seed production (Janzen 1970, Connell 1971). Imagine that you have the situation that
rats are eating the seeds of a plant that you are trying to encourage. If you add a small number of seeds
the rats will eat them and there will be no recruitment. So there is a threshold level of seeds you must
add before you get any recruitment.
The combined effect of these forces results in three zones where the relationship between
recruitment and seed production is highly non-linear (Crawley 1997). In Zone III you are essentially
feeding granivores rather than producing mature plants and in Zone I the plant is so numerous that it is
microsite limited so adding or taking away seeds makes no difference. Only in Zone II do you get an
apparently logical relationship between recruitment and seed production.
If you manage either for or against a particular species it would be useful to know what zone you
are in. This information can only be obtained experimentally. Simple observation will not suffice. The
sensible experiment to do is to randomly choose quadrats in which you add and do not add seeds. If you
find you get no recruitment in the areas in which you added seed you still do not know what zone you
are in. You may be in zone I but the asymptote may actually be zero, i.e. there are no suitable microsites
under present conditions. To determine what zone you are you must also have a seed predator
treatment. Some would say you should also look at fungal pathogens which can appreciably affect seed
mortality (Augspurger 1984). Sticking with predators only, the minimum experiment you must perform
is plus/minus seeds and plus/minus seed predators (rats and other granivores). If the study plant is
predator-limited you will get recruitment inside your exclusion cage but if it is microsite-limited you
will not. This is particularly germane to the problems in the forest because it is likely that nobody really
understands the recruitment dynamics of any of the trees in the forest in Mauritius. Such ignorance is
not restricted to Mauritius. It is also the case in Britain which has a comparatively well-studied flora.
a) b)
RA

RB RA RB
Plant recruitment

0 B A 0 B A

c) d)
Zone III Zone II Zone I

0 0

Seed density
Fig. 1. Plant recruitment curves. A.) Linear relationship between seed input at time t and recruitment at time t+1. If
a seed predator reduces seed production from A to B, plant recruitment will be reduced proportionately (from R A
to RB). b.) An asymptotic relationship between plant recruitment and seed production when recruitment is
microsite-limited. The proportion of seed predation in diagram a will have no impact on plant recruitment
(RB=RA). c.) A threshold of recruitment beyond a point when seed predators are satiated. d.) A recruitment curve
showing the effects of both microsite limitation and predator satiation. This curve has three zones: Zone I -
microsite limited recruitment, Zone II - seed limited recruitment, Zone III - predator-limited recruitment (adapted
from Crawley 1997).

Dispersion - Complexities in real metapopulations

To add further complexity to the situation we have to address the fact that when you are thinking
about population dynamics on a landscape scale we ought to be thinking about metapopulations (see
Taylor 1990 for a review). We are now talking about patches in which the population dynamics are
different and the patches are not independent, being linked by dispersal. So local extinction is not a
disaster because we can get recolonisation. Therefore, the above discussion on populations needs to be
set in the context of metapopulations, the study of many patches when all sorts of different factors come
into play e.g. genetics, different patch qualities, catastrophes, the existence of corridors and different
patch shapes. So if we parameterise our models using data from a single patch we must not necessarily
assume that the predictions we make from that model will apply to other patches, even if these patches
are in close proximity to each other.

What makes a community invasible?

All communities are invasible. Both theory and experience support this. There is always a way
of designing a superior competitor to replace the current community dominant. The SCOPE programme
which examined biological invasions in the 1980s (Drake et al. 1989) suggested that some communities
are more invasible than others but the evidence is difficult to interpret. The data are hugely biased.
These findings are uncontrolled for introduction rate and species traits. If you analyse the data you
discover that that disturbance rate is extremely strongly positively correlated with both the richness and
abundance of alien species. Proximity to human settlement is also an extremely strong explanatory
variable. Most importantly, the introduction rate itself is a function of distance from human settlement.
For example you get more species introduced in towns which escape from gardens than you do on
mountain tops, simply because of isolation. Also the species that people introduce from other places are
obviously species that can live with people. If you introduce these species on to mountain tops it should
be no surprise that they do not do very well. The correct experiment to carry out to determine how
invasible communities such as those on mountain tops are would be to introduce species to mountain
tops from other mountain tops in other continents. Needless it is not recommended that we carry out
this experiment!
Lottery models
There are many methods by which we can use ecological theory to address community
dynamics. One method, illustrated here, is the lottery model. Lottery models are often used to assess
the influence of particular life history traits on the ability of the organism (usually a plant but
sometimes sessile animals such as coral) to occupy a given unit of space, a cell, in a landscape (Sale
1979). The key point is that you don’t model a number of individuals as we did in our population
dynamics model. Instead you model the proportion of space that is occupied by species of particular
types. So now X is not the number of individuals but the proportion of space occupied by populations
of a particular type of species, that is a fraction between 0 and 1. For example X for invasive aliens
such as Psidium might be 95% or .95 and the space occupied by natives would be 1-X which would be
5% or .05. What this model says is that the only way that new plants can be recruited is by the freeing
up of space by the death of plants that are already there. So space occupancy changes by two terms; A
negative term that says that space declines for species X at rate D, the death rate (which could be
density dependent or density independent), and a positive term given by adding all the deaths of all the
species in the community. This produces space which can be occupied by all the different species in the
community. In the example below we highlight a model for two plant species or two types of plant
species:

dX(t) = -DX(t) + D qX(t)F

dt qX(t)F+(1-X(t))F

where X(t) = fraction of cells occupied by plant 1

D = death rate (assumed to be the same for both species)
F = per capita fecundity (assumed to be the same for both species)
q = relative competitive ability of the 2 plant species

The proportion of that new space that is occupied by the species you are interested in is
determined by a more or less complicated function that has to do with the relative seed proportions or
the relative growth rates of the plants involved. In terms of weed control and threatened ecosystems
models like this are often going to tell you that after the control of a weed the space is going to be
occupied by something else and the chances are that if you control one weed species that this something
else is going to be another weed rather than a native. We have seen some nice examples from the USA
where this hasn’t happened (Randall and Mauremootoo, this volume). Unfortunately though there are
plenty of examples from other parts of the world where theory and experience tells us that you just get
replacement of one weed species by another.
The theoretical parts of this process that are interesting are linked to questions such as whether
herbivores are by and large good or bad in the process of recovery? And does herbivory by and large
help or hinder the regeneration of natives versus aliens? The areas covered above illustrate the kind of
framework within which you could make predictions which could then be tested by experiments.

Techniques and monitoring

The need for work of publishable quality
We should be aiming to carry out management experiments. That is using our management
manipulations in a way that makes them predictive and useful experiments. These management
experiments should be of publication quality. Not only that but they should be published! There is no
point in years of hard work disappearing into a report that is not readily accessible. Science belongs in
refereed journals where everyone can see it and everyone can benefit from the experience of the
investigators. To produce work of publishable quality you have got to replicate and randomise.
Replication improves reliability. If you do not randomise, all your results can reflect your personal
biases. Replication and randomisation are not luxury items, they are absolutely essential. If you do not
do these two things then your work is not of publication quality and if it is not of publication quality the
impact of what you do is reduced considerably.
Factorial designs
With this end in mind we should be thinking about doing factorial experiments (Mead 1988).
First of all factorial experiments are highly efficient in terms of the amount of information obtained for
the effort made in the field. A very important feature of factorial experiments is that they allow you to
assess the existence of interactions between processes. This is important because single factor
explanations in restoration ecology are rarely correct. The answer to a problem would seldom be
something like ‘herbivores matter’. Much more likely would be ‘herbivores matter under these
conditions but not under these conditions.’ This is what is meant by interaction.
A factorial experiment is an ideal but as ever there is a downside. The downside is that they can
rapidly become absurdly complicated. This can be illustrated by a simple example where we ask the
question what limits the recruitment of ‘bois makak’ Mimusops maxima in the Brise Fer forest? We can
come up with many possible influences. Here we illustrate seven which are defined as treatments. The
levels that any treatment can take can be infinite. In practice they must be limited. In this example we
chose 2 levels for each treatment. All treatments need to be replicated. The minimum amount of
replication is 2 so we chose this degree of replication to minimise the total number of experimental
plots needed (Table 1.).

Table 1. Number of plots needed in a hypothetical plant recruitment factorial experiment

Treatment Levels Cumulative number of
experimental units needed
+/- weeds 2 2
+/- seed addition 2 4
+/- pigs 2 8
+/- deer 2 16
+/- rats 2 32
+/- monkeys 2 64
+/- mulch 2 128
Replication (x 2) 256

These plots would have to big enough to measure recruitment realistically. Not only this but
each year is different. For example years may vary in levels of cyclonic activity. Places vary also, for
example in terms of slope, aspect, etc. This adds further complications. So how can such absurd
complexity be limited?
The experimenter must be selective and pragmatic. A decision must be made about what are the
most important factors to manipulate. This may involve assessing the effects of a combination of factors
rather than just one. In this case we would not be able to elucidate the exact role of each factor. Such
precision would be impossible anyway given the complex combination of alien species that we have in
Mauritius.
One should not forget the empty quadrats when assessing recruitment (Crawley 1990). That is
quadrats in which you do absolutely no manipulations but you do monitor. In other words you must
have complete factorial designs even if they are simplified. So many studies on plants are done in
quadrats that are put over the plant at the beginning of the study because the plants are there. So what
do you learn about plant recruitment? You learn that in many plant species recruitment almost never
happens because plant recruitment almost never does happens under a large parent plant (Whitmore
1990). It happens in a gap that doesn’t have parent plants in and if you do not study the empty quadrats
you will massively underestimate the rate of recruitment because you are looking for it in the wrong
place.
Last but not least, you must avoid pseudoreplication, that is you must know what a true replicate
is. There are many text books that spell out this issue in detail e.g. Krebs 1989 and Crawley 1989.
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