Module 7

Ecosystems as Complex Systems

Jay T. Dalet, PhD.

Introduction

To deduce how a particular living system works you must understand that you are dealing with a
conglomeration that is greater than the sum of many heterogenous things taken together. However, the
approach by which all complex systems are analysed in terms of their components has not revealed the
fundamental laws that govern this entire system as a whole (Cohen & Harel, 2007). So far we have the
technology to acquire fragments of evidence. But to piece the information altogether, it would require
machine learning. In this module you will be introduced to the concept of complexity applied to biological
ecosystems which are formed by the non-linear interaction of a community of living organisms with their
physical environment. You will also be given the opportunity to be familiar with how complexity exist in
other ecosystems dubbed as “analogues” of biological ecosystems.

Ecological complexity is an emerging multidisciplinary field of study that makes use of tools and concepts
developed from the essential disciplines of complex systems science (CSS) namely physics, mathematics
and computer science (Complex Environmental Systems Lab, 2014). It provides special emphasis on the
relationships between pattern and process in natural systems. According to Complex Environmental
Systems Lab based in Canada that ecological complexity is characterized by local interactions between
individual ecosystem components, feedbacks between processes occurring at different scales,
amplification of minor variations in initial conditions, and the emergence of patterns in the absence of a
global controller. Ecological theory has provided the fundamental assumptions for figuring out possible
solutions to major social, economic and environmental problems that comes into view indistinctly.

Our existence is greatly influenced by social organization, economic situation and natural environment
that have direct or indirect relations with the provisions that we derive from ecosystem services such as
food, energy source, human population, and natural resources. Analyzing the connection between
changes in both rural and urban environments with the phenomenon observed at a single or different
points in space and time is highly relevant in addressing the feedbacks brought about by the perturbations
across domains or microcosms (composed of different elements) that comprise complex systems.

On the concept of social social-economic-natural complex ecosystem Rusong (1984) stated “Complex
problems that we experience today such as climate change, emerging and re-emerging diseases, inflation
and wars can be regarded as neither social nor economic ones separately but as ones of the social
social-economic-natural complex ecosystem.” Economic and natural systems possess different
characteristics as indicated by their own structures, functions and developmental rules. However, their
path to maturity are influenced by the structures and functions of the others (Rusong, 1984). In this
perspective appreciating the role of keystone species, ecosystem engineers and ecosystem components
are essential in understanding their impact on how complex ecosystems share common properties with
each other. In this learning process you will better understand the interplay between different ecological
scenarios. Although, the in-depth use of computer applications cited in this module is not required of you
familiarizing oneself how these computer technologies work will help focus one’s lens on measures of
ecological complexity as useful ecological indicator. Diagrams, illustrations and graphs are provided in
this module to facilitate your learning process.

Learning Outcomes

After studying this module, the student should be able to:

1. discuss unique characteristics of an ecological complex system;

2. explain the significance of models of ecological complex ecosystem;

3. interpret relationships between various measures of complexity among graphical displays of data
and

4. discuss agreements between biological ecosystem and its analogues based on adaptation,
survival and extinction

7.1 Ecosystem as a Complex System

Ecosystems are very good examples of complex systems governed by laws of physics and chemistry.
Each component of an ecosystem is predisposed to be involved in many different interactions that
generate emergent properties -- patterns at higher levels emerge from localized interactions and selection
processes acting at lower levels (Levin, 1998 and Cohen & Harel, 2007). Such interactions between
components is non-linear. The outcomes are determined by external conditions (i.e. the environment) and
the extent by which ecosystem components organized themselves.

7.1.1 Complex System

Here you will learn different examples of complex systems. Just to perk you up, other than the ecological
systems, our brain and the global economy in itself are all examples of complex systems
(Peltoniemi & Vuori, 2004 and Brown, 1994).

Activity 1 (Total duration 53.98min)

You will watch video presentations (soft copies will be provided by your teacher prior to your first
meeting).

A. These videos will help you gather and refresh prior knowledge essential to this learning process.
1. Understanding the Complex Systems Around Us: Martin Schmidt at TEDxMcDonogh
Link: https://www.youtube.com/watch?v=nGFLegJEZTY
 Duration: 17:18
Central Idea: In a general sense what is a complex system? What are examples of complex
systems? And where can we find them? And why are they important?

2. Complex adaptive systems - Igor Nikolic – TEDxRotterdam

Link: https://www.youtube.com/watch?v=jS0zj_dYeBE
Duration: 16:06
Central Idea: How changes in the local environment may have impact on global conditions?

B. These videos will help you understand ecosystem as a complex system.

1. What is a Complex System? (Complexity Labs, 2017)
Link: https://www.youtube.com/watch?v=vp8v2Udd_PM
Duration: 10:23
Central Idea: What is the working definition of a complex system? What is emergence? How
emergence is related to self-organization? Why you cannot simply remove or isolate one
component or reduce the levels of organization to one?

2. Complexity Theory Overview (Complexity Labs, 2010)

Link: https://www.youtube.com/watch?v=i-ladOjo1QA
Duration: 10:51
Central Idea: What are the four areas that encompass the major perspectives in a complex
system?
Guide/Study Questions for Activity 1

1. What are the four primary parameters that make up a complex system?

2. Define phase transition? Why is its occurrence relevant to our understanding of a complex
system?

3. What is the significance of heightened connectivity as indicated by networks seen in complex

systems in the integrity of the biosphere? Are these networks good or bad indicators of a healthy
society?

a. How is resiliency achieved in this context?

Discussion (Guide to study questions)

According to Complexity Labs (2010) complexity can be defined as the primary product of four primary
parameters. It is initially defined by the number of elements at different hierarchy within a living system.
Another dimension to complexity is nonlinearity where non-additive interactions and feedback loops over
time can give us exponential relations between the input and output to systems and lead to phase
 transition a period of rapid change where non-linear systems may grow or decay at exponential rate.
Complex system are able to shift or flip into whole new regimes within very brief periods of time. Some
small change to the input value may trigger large systemic effect (sensitivity to initial conditions). Another
driver of complexity is connectivity that often appear as networks in a higher level that indicates the
degree of how things flow in the network. In this regard resiliency is achieved in the presence of
alternative species. Autonomy and adaptation enables self-organization and the process of evolution
that shapes complex systems on macro scale. Typical examples of complex systems include:
ecosystems, economies, transportation networks and neural systems (i.e. brain).

7.1.2 Measures of Complexity in an Ecosystem

Knowing how complex an ecosystem under study will help you understand the importance of species
interactions and the relationships between patterns and processes in it. In the next activity you will learn
the various sources of complexity in an ecosystem. Also, you will be familiar with some computational
tools in biology used in measuring these sources of complexity.

Activity 2 (Total duration 30min)

Integration of data from various sources of complexity helps provide overall assessment of various
ecosystem services as rendered by an ecological complex system. Watch the accompanying video
entitled “Understanding Ecological Complexity” to familiarize you with fundamental sources of complexity
in an ecosystem and understand relationships between various measures of complexity among graphical
displays of data.

There are at least three dimensions in ecology where complexity can be measured (Loehle, 2004; Anand
et al, 2010 and Parrot, 2010).

Spatial

Complexity in ecosystem can be measured based on the manner species are organized in a given
geographical location (Parrot, 2010). Example of sources of spatial complexity includes species
distribution and vegetation patterns (Figures 4 to 6). For example in Figure 4, a bubble chart was used
to profile the population density (bubble diameter) and actual locations of four keystone species marked
by their respective coordinates (x, y) that were derived using geographical positioning system (GPS)
device.

Another example of reporting spatial distribution of species is shown in Figure 5. Here the presence of
three fruit bearing trees along the elevation of a mountain named “Mount Amihan” are plotted. Elevation is
defined as the distance of your subject above a reference point (i.e. sea level) in this example are fruit
bearing trees. During data gathering a researcher hikes up on a mountain (increasing elevation) and
records the presence of fruit bearing trees. Patterns resulting after data has been plotted as shown in
Figure 5 could possibly be attributed to some biophysical factors (i.e. levels of humidity and/or availability
of pollinators or agents of seed dispersals agents) that may have influence in the growth of vegetation in
those areas of elevation (Parrot, 2010 and Mexia et al, 2018).

Figure 4. Distribution of four keystone species in GRA in year 1984(A) and

1987(B). Population densities are indicated by the bubble diameter. Actual
locations in the forest are determined by the recorded X and Y coordinates.

Figure 5. Availability of three fruit bearing trees along the

elevation of Mount Amihan in year 2000 (A) and 2005 (B).
A heat map is a diagrammatic representation of data whose values are represented as colors. The heat
map shown in Figure 6 is an example of plant species richness profile in a given locality named “Sierra
Pacifica”. Vegetation patterns can provide a certain degree of better ecosystem services (i.e. food
security) in the indicated areas of a local community (Mexia et al, 2018).

A. 2D Maps Plot of Plant Species Richness in B. 2D Maps Plot of Plant Species

Richness in
Riparian Forest Sierra Pacifica (Year 1995) Riparian Forest Sierra Pacifica
(Year 2000)

Figure 6. Overall heatmap profile of plant species richness in different areas

(patches) in Sierra Pacifica in year 1995(A) and 2000(B).

Temporal

Its measure characterizes time series of different variables describing the dynamics of a system (Parrot,
2010). It can be derived from dynamics or changes in population, effects of changes in climate and
weather, extinction, invasion and successions and predator-prey cycles (Loehle, 2004). A very good
example is shown in Figure 7A and 7B. Take note of the periodically repeated phenomenon of increasing
and decreasing predator and prey density. You would notice that wolves’ and shrews’ initial state (density)
is restored (as sort of a memory) in the indicated duration (no. of years) as show in Figure 7A.
 Figure 7. Population dynamics of wolves, shrews and insects in a duration of
ten years.

Such dynamics reflect a stable supply and demand and entails resiliency in ecosystem. However,
perturbations affect ecosystem stability and complexity (number of interacting species and the
connectivity between species) resulting in unfavorable conditions. One very good real life example was in
the case of Yosemite National Park when they eliminated the wolves, the elks predominated which
affected the density of plant species essential for the life of other organisms (i.e. beavers). In Figure 7B
the decline in wolf population impacts on the increase of shrew population modifying insect population
whose participation in the food web is essential in pollination of fruit bearing trees. Another situation is
brought about by anthropogenic factors causing soil contamination with
 Figure 8. Biomass complexity decreased incrementally after 28 years as
exemplified by the death of calving industry due to soil contamination by heavy
metals and biphenyl derivatives.

unwanted toxic substances. Such changes may negatively impact on the communities serviced by the
ecosystem. Such conditions can be analyzed by plotting pertinent measures of data whose patterns can
be visualize in both the spatial and temporal dimensions (Spatio-Temporal). An example of this is shown
in Figure 8 indicating changes in the biomass complexity of an ecosystem named “Makabuhay Forest”.
Suppose you are analyzing the effect of heavy-metals and biphenyl derivatives to calving industry. You
would notice the gradual decrease of biomass from 1995 to 2017. Such a scenario would require
bioremediation processes that would entail number of years for the land to restore its integrity (Chaerun,
2014).

Structural

Refers to relationships within the ecosystem as exemplified by food web and species interaction networks
in Figure 9. Take note of the arrows indicating the connectivity of each organism situated in their
respective habitats and/or habitat patches.

Figure 9. Example of a food web structure.

Analyzing connectivity of habitat patches is highly relevant in understanding the movement of elements
(i.e. genes, individuals, populations, and species) on different time scales (Minor & Urban, 2008). (1)
Juvenile dispersal and recolonization of empty habitat patches as influenced by connectivity are usually
observed in short time periods (Clergeau & Burel 1997 and Minor & Urban, 2008); (2) migration and
continuing metapopulations are recorded at intermediate time scales (Hanski & Gilpin 1991; Ferreras,
2001 and Minor & Urban, 2008); (3) species range expansion in response to natural forces such as
climate change are observed in largest time scales (Opdam & Wascher 2004 and Minor & Urban, 2008).
Why do you think habitat connectivity is important? In the event of global warming for example organisms
will follow the range of coldness or hotness of the environment they are used to. Commonly they migrate
toward the poles (north or south) or high up in the mountains (Montgomery, 2017). Similarly, organisms
will respond to land conversion, heavy industrialization, logging, major urbanization etc. Unpredictable
changes in the ecosystem is likely to occur particularly when habitat is rare, fragmented, or the
distribution is wide which is crucial in preparing a reserve design (Flather & Bevers, 2002; King & With,
2002 and Minor & Urban, 2008). Fragmented habitats limit the gene pool for a particular species since the
population is too isolated, more chances of inbreeding and disease are likely possible (Crain, 2015).

Based on a widely recognized Graph Theory there are four kinds of connectivity networks or matrices
namely: planar, random, scale-free and small-world that can be constructed depending on the food
web structure, community composition, networks of competition and facilitation of the ecosystem under
study (Minor & Urban, 2008).

Figure 10. Four kinds of networks: (a) planar, (b) random, (c) scale free,
and (d) small world. Black dots are nodes; lines are edges (Minor and
Urban, 2008).

In Figure 10, you can see four different patterns possessed by each kind of network (a- d). Common to
all of these networks have black dots called nodes and black lines called edges. The nodes represent an
individual organism situated in a habitat patch while the edges represent the relations between two
nodes. Hence, a graph or connectivity network is composed of a set of nodes and edges. Nodes are the
elements in the network while edges connect nodes (Minor & Urban, 2008).

This time look at Figure 11A and 11B for you to see how the concept of Graph Theory is applied. The
positions marked by the dots correspond to those of the components of the food web structure shown in
Figure 11B. These dots are connected by edges to come up with nodes organized in a connectivity
matrix.
 Figure 11. Connectivity matrix of 12 nodes (Panel A). Each
node is situated in their corresponding habitat patches (Minor
and Urban, 2008). Information deciphered in this matrix is
applicable in the analysis of the food web structure shown in
Panel B.

From a conservation standpoint, according to Minor & Urban (2008) that among the four kinds of
networks (Figure 10) the more complex ideal habitat network may resemble a scale-free network with
several large hubs connected to multiple smaller patches. In Figure 11A, node 12 is designated as a hub
because it is connected to many other nodes (7, 8, 9, 10 and 11). Since the number of its connections is
higher compared to those of other hubs, node 12 is regarded as a high-degree node.

Notice that nodes 1–5 (Figure 11A) are highly clustered since they are highly bundled up with their
connections, whereas nodes 7-12 on the right side of the graph are highly compartmentalized
characterized by the presence of low degree nodes (7, 8, 9, 10 and 11) connected to hub node 12. What
is the significance of this observation?

Highly clustered nodes facilitate: (1) spread of disturbances which are temporary changes that could
possibly result from natural causes (i.e. weather and wildfires), the activities of humans or from disease
and invasion of exotic species and (2) dispersal (i.e. of seeds) that aids recovery from an environmental
disturbance. Also, nodes 1-5 may be more resilient to patch removal (disaster or calamity) due to many
redundant pathways. High compartmentalization in nodes 7-12 slows movement through a network and
may isolate the potentially cascading effects of disturbance. Scale-free networks are highly resistant to
random disturbances but vulnerable to deliberate attacks on the hubs (Barabasi & Bonabeau, 2003 and
Minor & Urban, 2008). Looking at a scale-free habitat network, if a habitat or a patch is chanced upon by
certain sickness or unwanted species there is low probability for this sickness to spread far or quickly due
to few patch connections. However, if the targeted entry was into a hub, the disease would rapidly spread
in the entire network. In the same manner network connectivity would be less affected if almost all of the
smaller patches were taken away. But such connectivity would break apart in a brief period of time if hubs
were taken away by a certain circumstance that could either be man-made or natural disaster (Minor and
Urban, 2008).

Hence, to maintain the complexity of an ecosystem it entails conservation and monitoring efforts that
would be best spent on hub patches. Taking into consideration the habitat connectivity matrix constructed
for a riparian forest (Figure 12). This is an example of a conceptual model of a complex ecosystem.
Applying what you learned from Graph Theory, take note of the link phrases found along the lines or
edges (i.e. prey on, feed on, habitat for, promote, etc.) that connect the nodes represented by circles
labeled with the names of organisms and/or habitats. The edges actually represent the flow of energy
from one node (element or component) to another. Also you would notice how clustered and
compartmentalized the nodes in the left and right side, respectively. At this point you would see how
crucial the role of these organisms in this complex system. They are not only the keystones but also
ecosystem engineers acting on their own to maintain the integrity of this given ecosystem. See a potential
scenario that may have taken place if an environmental disturbance is introduced in this forest. Take note
of the structural changes that could be brought about by the indicated perturbations (Figure 13).

The rampant use of pesticides and the occurrence of heavy industrialization in GRA resulted in the
destruction of high-degree nodes as indicated by low species count (Figure 13). As a consequence this
led to the promotion of contaminated waterways and rivers. The introduction of invasive species in the
river brought the loss of beneficial species. The reduced complexity of this ecosystem impacted on the
overall economy and health of local communities as indicated by loss of both main and alternative native
species; high food prices and malnutrition; diarrhea and infectious diseases and decline in provisioning
services.
Figure 12. Connectivity matrix of 20 nodes is shown in Panel A. Each node is presumed to
be situated in their corresponding habitat patches. The matrix shown in Panel B will serve as
a guide in the analysis of the food web structure shown in Panel A.

Figure 13. Connectivity matrix of 14 nodes that resulted from pesticide and heavy-metal
contamination of habitat patches in GRA. The introduction of invasive species is indicated by
red circle on the right.
 Figure 14. Different sources of complexity analyzed for links between changes in the
ecosystem and those of socio-economic dimensions.

In the long run if the aforementioned anthropogenic disturbances persist such changes may result in
hybrid systems retaining some original characteristics as well as novel elements, whereas larger changes
will result in novel systems, which comprise different species, interactions and functions (Hobbs, Higgs
and Harris, 2009).

At this point you are now much equipped to understand why ecosystem is in itself is a complex system
and is regarded as “a whole more than the sum of its parts”. Hence, ecosystem is a network of many
components whose aggregate behavior is both due to, and gives rise to, multiple-scale structural and
dynamical patterns which are not inferable from a system description that spans only a narrow window of
resolution (Complex Environmental Systems Lab, 2014 and Parrott & Kok, 2000). Nevertheless, data are
derived from different sources of complexity and analyzed for links between ecosystem change and those
of socio-economic dimensions (Figure 14).

You may explore some computational tools as shown in the video “Understanding Ecological Complexity”
to gain familiarity on how different measures of ecological complexity are studied.

Guide/Study Questions for Activity 2

1. Based on the type of vegetation ecosystems can be recognized and divided into three major
groups namely: terrestrial, water and artificial ecosystems. Give examples for each of these.
 2. Compare in general the temporal, spatial and structural signatures or profiles of simple,
disordered with that of a complex system.

3. What is the significance of understanding measures of complexity in the study of biological

ecosystems?

4. There are computational tools that are commonly used to analyze measures of complexity. Name
a few and describe how they are utilized.

Discussion (Guide to study questions)

Learning ecosystem as ecological complex system entails understanding the relationships between types
of ecosystems through the ensemble of different sources of complexity like vegetation patterns, species
distribution, population dynamics, weather and climate change, food web structure, soil formation, gene
expression and individual tree crown. The following are examples of each type of ecosystems:

1. Terrestrial Ecosystems: (a) deserts, (b) savannahs, (c) steppe, (d) temperate forest, (e) tropical
forest, (d) boreal forest (taiga), (e) tundra and (d) Mediterranean vegetation.

2. Water Ecosystems: (a) freshwater ecosystems (lakes and ponds, rivers and torrents, marshes
and swamps) and (b) marine ecosystems (reef, oceans, continental plateaus, nutrient upstream-
flowing areas and estuaries).

3. Artificial ecosystems: (a) urban-industrial ecosystems (metropolises), (b) rural ecosystems (small
towns) and (c) agro-ecosystems (farmlands).

Below is a summary of signatures/ profiles (Table 1) you should expect when the interaction of species is
indicated as simple, complex or disordered in an ecological system at different dimensions (Parrot, 2010).
The graphical or visual patterns that corresponds to each of these profiles are found in Figure 15 (Parrot,
2010).

Table 1. Summary of three different species interaction profiles in temporal, spatial and structural
dimensions.
Signatures/ profiles

Temporal Spatial Structural

Simple sinusoidal ordered regular network

Complex uniformly distributed diffusion over a coupled scale-free network

regular noise mapped lattice

Disordered uniformly distributed random spatial distribution random network

random noise

(L. Parrot, 2010)

 (L. Parrot, 2010)

Figure 15. Graphical representation of temporal, spatial and structural

signatures of simple ordered and disordered systems versus a complex
system.

To gain further knowledge and understanding about the possible sources of complexity you will read three
scientific review articles: (1) “Measuring Ecological Complexity” written by Lael Parrot. Focus only on
sections I (Introduction), II (Measuring complexity), and Figures 1 to 5. (2) Challenges of ecological
complexity by Craig Loehle and (3) Ecological systems as complex systems: challenges for an emerging
science by Madhur Anand et al (2010).

Research findings validated that common characteristic patterns of temporal, spatial, spatio-temporal and
structural signatures are shared across all types of complex systems (Parott, 2010). The advent of
computational tools in biology has helped better understand measures of complexity, thereby provides
both qualitative and quantitative assessments of ecosystems under study. There are two types of
measures of complexity that have been reported recently (Atmanspacher, 2007 and Parott, 2010): Type 1
measures increase with increasing disorder in the system in a linear manner, and Type 2 measures are
those of a convex function that assign their highest scores to systems whose regularity lies at the
intermediate level (Figure 16). The endpoint is to differentiate between simple systems versus those that
are complex regardless of what type of measure is used (Grassberger, 1988 and Parrot, 2010).
 Figure 16. Complexity versus regularity (Parrot, 2010).

As indicated by Dr. Parrot in her review article, recent research has thus focused on developing methods
to characterize the temporal, spatial or structural signatures of complex systems, classifying the system
along a gradient of order to disorder. Figure 16 is a plot of regularity versus complexity. It was argued that
both highly ordered and highly disordered (random) systems are simple systems and mostly complex
systems are found in the intermediate zone of regularity (Parrot, 2010). Properties of a complex system
are said to be found “at the edge of chaos” (Langton, 1992 and Parrot, 2010) in the middle of two
extremes of order (uniform spatial pattern or temporal equilibrium) and disorder (random spatial
distribution or white noise), demonstrating equal distribution between underlying regularity and complete
unpredictability (chaos) as shown in Figure 17 (Parrot 2010).
 Figure 17. Ecological complexity as an ecological indicator (Parrot, 2010).

According to Dr. Lael Parrot’s (2010) view on ecological complexity as an ecological indicator, an
ecosystem tends towards greater complexity via the process of self-organization, which draws the system
away from the two extremes of order and disorder to a state of maximal complexity. She further explained
that this state of maximal complexity is a site-specific attractor which is constrained by prevailing physical
and environmental conditions -- natural disturbance events may cause an ecosystems state to tend
towards greater disorder, whereas human intervention in the form of energy input can move the system
state towards greater order than might be attainable naturally. Descriptive data in Figure 17 (Parrot,
2017) purports that in an agricultural monoculture, such as a cornfield planted in rows, serves as an
analogy for an ordered ecosystem, whereas a recently opened gap in a forest, in which seeds have
randomly fallen and just begun to germinate, may be an example of a disordered ecosystem. An
undisturbed, ancient tropical rainforest (central image in Figure 17) is probably the best example of one of
the Earth’s most complex ecosystems.
 Figure 18. Spatial analysis in macroecology (SAM) Figure 19. Ecopath with Ecosim

Temporal, spatial and structural data would provide information that would help orient your point of view
on how to preserve or relieve an ecosystem of pressures depending on the case at hand. Below are
some outputs of open-source computer applications that are useful for measuring possible sources of
ecological complexities. Examples of such are computer program outputs from Spatial Analysis in
Macroecology (SAM v4.0) which aims to provide an integrated computational platform intended for spatial
analyses, identification, characterization of ecological and evolutionary forces that influence patterns in
body size, geographic range size, abundance, diversity and related macroevolutionary trends. The
software can be freely obtained from http://www.ecoevol.ufg.br/sam. The description of the software was
published in Global Ecology & Biogeography (Rangel, Diniz-Filho & Bini, 2006) and in Ecography
(Rangel, Diniz-Filho & Bini, 2010).

Shown in Figure 20 is bird species richness profile in South America. Variations in birds’ species count in
the continent are emphasized by the red dashed lines for regions A, B and C.
 Figure 20. Bird species richness profile in South America.

A B

Figure 21. Bird species richness correlated with normalized difference vegetation index (NDVI) and
annual evapo-transpiration (AET) in South America.

Figure 21 A and B indicate that bird species richness is negatively and positively correlated with
normalized difference vegetation index (NDVI) and annual evapo-transpiration (AET), respectively.
Perhaps species richness of plants are reduced due to very high productivity resulting in an increase of
nutrients. But light becomes more and more of a problem for competing plants. Since there is increased
production of woody species in highly productive ecosystems (Oindo & Skidmore, 2002 and Prins & Olff
1998), bird species richness too increases with woody vegetation (Pomeroy & Dranzoa 1997). In the case
of AET, water plays a vital role in the life of an ecosystem. Evapotranspiration provides better capability
for a land to sustain life depending on the maximum amount of water regained per unit of water
consumed. Species distribution can be correlated to the rate of AET to examine how AET affects richness
of species in a particular geographical region.

Figure 22. Temporal data on biomass production by phytoplankton (blue) and zooplankton
(green) (EwE output using Anchovy Bay sample data).

Another program is Ecosim and Ecospace available in Ecopath with Ecosim (EwE) ecological modeling
software. Ecosim and Ecospace are applications with dynamic simulation capability at the ecosystem
level and a temporal-spatial dynamic computational capability, respectively. Ecospace is primarily
designed for exploring impact and placement of protected areas. It helps to allocate biomass across a
grid map while accounting for predation risk, feeding rate and species distribution. EwE is available at
http://ecopath.org/. Both SAM v4.0 and EwE will be provided by your teacher each with a sample data file.
Figure 22 shows temporal data on biomass production by phytoplankton (blue) and zooplankton (green)
(in (EwE Anchovy Bay sample data). Prey-predator population dynamics is clearly demonstrated here that
follows the principle behind the Lotka-Voltera model the simplest mathematical model of predator-prey
interactions. The model was developed independently by Lotka (1925) and Volterra (1926): It has two
variables (P, H) and several parameters: H = density of prey, P = density of predators, r = intrinsic rate of
prey population increase, a = predation rate coefficient, b = reproduction rate of predators per 1 prey
eaten and m = predator mortality rate (Sharov, 1996). The dynamics of predator and prey interaction can
be plotted as an estimate of predator and prey densities per unit time (Sharov, 1996). Analyzing the
predator-prey density profiles in time series at different locations provides an assessment of the temporal
complexity of an ecosystem under study. Take note of the uniformly distributed regular noise (as
mentioned in Table 1) generated by EwE from the temporal biomass data of both groups. Such temporal
signatures characterize that of a complex system.

Figure 23. Ecosim output of food web flow diagram in a marine ecosystem.

Figure 24. Energy flow diagram in a marine ecosystem.

Hence, in Figure 23 is an output of Ecosim showing the food web flow diagram of a given example of a
marine ecosystem. The actual flow of energy in this diagram is depicted in Figure 24. This emphasizes
the role of phytoplankton (with zooplankton as its predator) as primary producers and the decreasing
amount of energy towards the top of the pyramid.
Activity 3 (Total duration 10min)

This time you will apply what you learned about complexity to analyze a complex ecosystem. In this
activity you will be given a case scenario and you will observe how an ecosystem transition from a healthy
to an unhealthy one (modified from Rapport, 1994). You will do this by looking into two conceptual models
of a healthy and unhealthy native riparian forest that represents an ecological complex system found in
geographical region A (GRA), as shown in Figures 14 and 15, respectively. The story about the
ecosystem depicted in Figures 14 and 15 is provided in the case scenario given below. You will learn that
perturbations in an ecosystem (manmade or natural) may trigger changes that could be for better or
worse case scenarios in the biosphere. Read the case scenario given below:

Case Scenario for GRA

Based on the ecological survey that was conducted by your team ten years ago
Geographic Region-A (GRA) has been recorded to produce many resources valuable
to its local economy. It was estimated that about 3.1 million people depend directly on
this flood-plain for vegetables and raw materials such as cotton fabrics and leaves for
tobacco (Fig. 14).

At present water profiling indicates the presence of unwanted substances and or

factors. Flora and fauna survey showed preponderance of non-native and invasive
species. Forests that were formerly seen along river banks are no longer visible.
Evident are commercial plantations of Gossypium sp. and Nicotiana sp. Low grass
production potential affected the calving dairy herd (i.e. Paspalum notatum and P.
conjugatum) as well as biofuel supply (i.e. switch grass -Panicum virgatum) (Fig. 15).

Assessed relative richness of species in one habitat versus another indicate decreased
biodiversity (i.e. loss of riparian forest). Survey of the highlands indicates the
prominence of three shrews which apparently are mainly responsible for the dispersal
of Rafflesia sp., a local parasitic plant claimed to be devoid of chloroplast genome.
Both human and animal wastes are disposed of directly to trenches that are normally
used to supply dry land with water. Reduced food (low calorie availability, high food
prices and malnutrition) and water security (diarrhea and infectious diseases) were
observed in the local communities (Fig 15).
Figure 14. In panel A is a conceptual model of a healthy native forest riparian vegetation
representing the components derived from an ecological complex system scenario in geographical
region A (GRA) -- indicated by the components enclosed in different shapes in panel B. The case
scenario situates you as part of a team that conducted an ecological survey ten years ago in GRA
that was recorded to produce many resources valuable to its local economy.

Figure 15. In panel A is a conceptual model of unhealthy native forest riparian vegetation
representing the components derived from on ecological complex system scenario in geographical
region A (GRA) -- indicated by the components enclosed in different shapes in panel B --take note of
an invasive species of angler fish (I) and the occurrence of Rafflesia sp. a parasitic plant (P).
Unwanted toxic substance used in farming is indicated by the star in panel A. Death of some
keystone species is indicated by the white polygons – decreased species count hence, disrupted
ecosystem services. This reduces the complexity of GRA as represented in the model of locally
interacting components. This minor variations are amplified to loss of biodiversity, low productivity
resulting in diarrhea, emergence of infectious disease, malnutrition and high food prices.

The interaction between the components of an ecosystem occurs at three different levels. This interaction
can be viewed using a conceptual model (Figures 14A and 15A). Starting at the lowermost level are (1)
locally interacting heterogeneous components, followed in the middle by (2) emergent higher
entities at middle level and (3) global entity at the topmost level. You can also recognize the presence of
(4) feedbacks between processes occurring at different scales. A phenomena commonly observed in
such a system is amplification of minor variations (resultant change that accumulate through time) as
exemplified by changes brought about by manmade (anthropocentric) pressures, and emergence of
patterns in the absence of a global controller (due to the tendency of the components to self-
organize).

Concrete real life examples of locally interacting heterogeneous components (elements) are found
enclosed by shapes in Figure 14B. These shapes were used to represent them in Figure 14A. Observe
how the components are organized in Figure 14A and 14B. Also take note of the double headed arrows
that indicate a non-linear interaction between the given elements and the thick arrows that indicate a
bottom up direction. The feedback loops are represented by four bent arrows with short solid lines and
two long arrows with broken lines. Apply the same approach in observing the effect of changes in
ecological complexity shown in Figures 15 and 16.

Figure 16. The transition from species network A to C indicates decreasing species richness contributory
to decrease in ecological complexity of GRA, a condition brought about by prevalence in the use of toxic
pesticide in farming and introduction of exotic species in the river.

Deciphering the Model for GRA

The case can be easily analyzed by organizing and summarizing your observations in a table of three
rows with appropriate headings. Below is Table 1 composed of three rows with columns labeled as (A)
anthropogenic and natural perturbations; (B) changed ecosystem structure and function and (C) effects
on population and world economy. Each row was filled up with pertinent observations derived from the
narrative of the given case for GRA. The colored fonts will help you to cluster the observations per row.

Table 1. Summary of observations in GRA.

(A) Anthropogenic and natural perturbations
1. presence of unwanted substances and or factors
 2. preponderance of non-native and invasive species
3. commercial plantations of Gossypium sp. and Nicotiana sp.
4. prominence of three shrews
5. dispersal of Rafflesia sp.
6. human and animal wastes are disposed of directly to trenches

(B) Changed ecosystem structure and function

1. low grass production potential affected the calving dairy herd (i.e. Paspalum notatum
and P. conjugatum) as well as biofuel supply
2. decreased biodiversity
3. low productivity
4. resiliency negatively affected
5. aquatic foodweb structure has been badly affected

(C) Effects on population (plant, animal and human) and the world economy
1. reduced food (low calorie availability, high food prices and malnutrition) and water
security (diarrhea and infectious diseases)
2. decline in provisioning services
3. loss of both main and alternative native species

Events under each of these row headings (A) anthropogenic and natural perturbations; (B) changed
ecosystem structure and function and (C) effects on population and the world economy, are either
causative agents or consequences affecting the components in each of the levels of the model (1) locally
interacting heterogeneous components; (2) emergent higher entities and (3) global entity, respectively as
summarized in Box 1.

Box 1. Events corresponding to the levels found in the conceptual model of ecological complex system

Row C (effects on population and the world economy) → Global entity

Row B (changed ecosystem structure and function) → Emergent higher entities

Row A (anthropogenic and natural perturbations) → Locally interacting

heterogeneous groups

Now you know how to use the conceptual model of ecological complex system to analyze a given case scenario.
This time you will join your respective groups and construct a conceptual model of ecological complex
system for GRB. The case scenario that you will analyze is given below. A step by step guide on how to
construct your model is provided after the case narrative:

Case Scenario for GRB

In the meantime, another set of conditions has been profiled in Geographical Region-
B (GRB) about 800 miles from GRA. Based on the report made in 1980 GRB
is characterized by large forest covers, richness in mammals and abundance
 of marine and fresh water fishes. Currently, observed changes in land use and
land cover as evidenced by the presence of commercial resorts and private
agricultural lands, fish ponds and waterways has been reported. There was a
decline in provisioning services rendered by the regions ecosystem: (1) fish
stock, (2) food production, crop area, pasteur area, and area for biofuels and
(3) clean water resources. According to the most recent ecological
assessments that were conducted in the region excessive fishing, low soil
nutrient profile has contributed to low productivity. Resiliency has been
negatively affected due to loss of both main and alternative native species
brought about by excessive fishing and introduction of exotic species. The
aquatic food web structure has been badly affected as evidenced by a shift in
dominance from highly organized nearshore benthic-fish associations to the
relatively less organized offshore pelagic associations. These conditions led to
extinctions of native fish species, occurrences of algal blooms associated with
toxin producing dinoflagellates and predominance of exotic species. Hence, a
decline in the commercial value of GRB.

Step 1. Summarize your observations in a table of three rows similar to Table 1.

Step 2. Assign shapes for the organisms in GRB similar to Figure 14B and 15B.

Step 3.Construct the conceptual model for the first level. Use double headed arrows to indicate
the interactions.

Step 4. Construct the conceptual model for the second and third level. Use bottom up arrows to
emphasize the direction similar to Figures 14A and 15A.

Step 5. Indicate and label the feedback loops similar to Figure 14A and 15A.

Write your group output in a piece of paper. Use Figures 14A & 14B and 15A & 15B as your template.
Also include in your output two to three sentences stating the environmental significance of factors or
agents that triggered changes in the structure of species network resulting in transition from healthy to
unhealthy GRB ecosystem.

You can retrieve the answers online in UP Virtual Learning Environment (UVLE) (or any assigned UP
website). The link will be provided by your professor after your group activity.

Guide/Study Questions for Activity 3

1. Explain in your own words why ecosystem is a complex system?

2. How are feedback processes related to the local interactions between ecosystem components?
Why are there feedbacks?
 3. Describe the manner by which amplification of minor variations in initial conditions leads to
emergence of patterns in the absence of a global controller?

a. Give a possible consequence of a positive feedback and negative feedback mechanism.

b. What is homeostatic plateau? Why is it an important consideration in understanding

complex systems?

4. What do you think could be the reason why there is an increase in the consumption of Rafflesia
sp. seeds by tree shrews in GRA? Is it relevant to the rate and levels of productivity in plant
species found in that region?

5. Based on what you observed in GRB give one example of an adaptive mechanism demonstrated
by the freshwater ecosystem under study that led to more unfavorable conditions.

Discussion (Guide to study questions)

Ecosystem is a complex system because it has many parts that interacts with one another in a non-linear manner.
Interactions of components results in observable changes (phase transitions) that may be experienced
locally or globally. Hence, the term “ecological complexity” arises a measure that describes the state of an
ecosystem. Feedback processes reflect the impact of pressures (anthropogenic or natural) as indicated
by various measures of complexities on the components of the ecosystem. These may result in
adaptation, survival and or extinction of a given ecosystem. Reorganization of components into a more
adaptive structure (i.e. food web) may lead to survival of some components (old or new) resulting in a
newly emerging ecosystem (i.e. succession, benthic to pelagic organization, emerging or re-emerging
diseases etc.). Anthropogenic or natural pressures can modify the ecosystem structure (i.e. non-linear
interaction in the food web) may lead to conditions (emergence of patterns) that may be beneficial or
detrimental to the biosphere. Such conditions take place in the absence of a global controller.

In this regard, feedbacks indicate the consequences of emergent and global agents manifested in the local
heterogeneous interacting components of the ecosystem. Examples of positive feedback includes
extreme temperature conditions (climate change), disease outbreaks, eutrophication, pestilence and
death while negative feedback includes regulation of temperature conditions, homeostasis, and
maintaining the limits of carrying capacity of the environment (predators and prey interaction).
Homeostatic plateau is a relatively stable state of equilibrium that maintain balance between population
numbers and resources. It is characterized by the presence of a minimum and maximum state of the
ecosystem such that when exceeded may lead to unfavorable ramifications i.e. death and or extinction of
the ecosystem. Also, it is noteworthy to mention two examples of socio-economic consequences of the
poor ecosystem services rendered by the ecosystems from the two regions (GRA and GRB) you analyzed
in this activity. GRA: Reduced food for humans (low calorie availability, high food prices and malnutrition)
 and animals (i.e. calves dairy herd) and reduced water security (diarrhea and infectious diseases). GRB:
Decline in provisioning services rendered by the regions ecosystem: (1) fish stock, (2) food production,
crop area, pasteur area, and area for biofuels and (3) clean water resources; and decline in the
commercial value of GRB.

Introduced species may or may not alter the host habitats. Tree shrews like Crunomys fallax are common
inhabitants of subtropical and or tropical dry forest that is only found in the Philippines (Northern Luzon). It
is threatened by habitat loss. In this given case it was used as an example of one of the effects of
anthropogenic pressures that led them focus their diet on Rafflesia seeds which they brought to GRA -- a
lowland flood plain which could be a potential alternative habitat. Rafflesia is a parasitic plant that feeds
on the energy provided by a grapevine Tetrastigma sp. which however, can be indicative of tree shrews
habitat loss that has an impact on the rate and levels of productivity in highlands proximal to GRA. One
very good example of an adaptive mechanism demonstrated by the freshwater ecosystem that led to
more unfavorable conditions in GRB was the response to restructured aquatic food web due to the
presence of exotic species. This led to a shift in dominance from a highly organized nearshore benthic-
fish associations to the relatively less organized offshore pelagic associations (Rapport, 1998).

Ecological complexity emerges from interactions between organisms and their biotic and
abiotic environments. It describes the state of an ecosystem…
Anand, 2010

7.2 Concepts of Adaptation, Survival and Extinction in Other Disciplines

The term “ecosystem” is mostly noticed in the science of biology; nonetheless it frequently appears in
economy related literature, as well (Pilinkiene & Maciulis, 2014). These are the different analogies of
biological ecosystem. It could be inferred in certain respects that they exhibit ecosystem stages of
adaptation, survival and extinction. Examples of these are business, industrial, economy, digital business,
and social ecosystems (Pilinkiene & Maciulis, 2014 and Peltoniemi & Vuori, 2004). Certain causal factors
are possessed by these analogies that affect the performance of the entire complex system itself.

7.2.1 Different Analogies of Biological Ecosystem

The emerging paradigm of complexity as we understood it in the context of ecological complexity has
provided the lead to search for other applications in various fields including social and economic systems
(Peltoniemi and Vuori, 2004). Understanding the interoperability of the concept of “ecosystems as
complex systems” in other disciplines would provide an opportunity for us to learn their similarities with
biological ecosystems and appreciate their existence in a global continuum that interconnects everything
in our planet. As evolving systems, ecosystems are dynamic, constantly remaking themselves, reacting
to natural disturbances and to the competition among and between species (World Resources
Institute, 2000 & Peltoniemi and Vuori, 2004). In the two remaining short activities (Activity 4 and 5) put
your answers (per group) in a one-half sheet of paper.

Activity 4 (Total duration 30min)

Read the article entitled “Business ecosystem as the new approach to complex adaptive business
environments” by Mirva Peltoniemi and Elisa Vuori. Based on this article you identify which analogue of
biological ecosystem (industrial, economy, digital business, social and business ecosystems) is indicated
by the descriptions below (a - e).

a. A “digital environment” populated by “digital species”. These digital species can be software
components, applications, services, knowledge, business models, training modules,
conceptual frameworks, laws, etc. that can interact with each other, reproduce and evolve
according to laws of market selection.
b. All material is recycled infinitely and efficiently. Change of habits of both manufacturers and
consumers would help maintain standard of living without devastating the environment. Different
parties should co-operate by using each other’s waste material and waste energy flows as
resources.
c. Consists of firms and institutions such that when these components cease to function like a
community, they may break down. Organizations are always co-evolving and are fully
participating agents. This ecosystem is made up of all related businesses, consumers, and
suppliers, as well as economic, cultural, and legal institutions.
d. Extended system of mutually supportive organizations, communities of customers, suppliers, lead
producers, and other stakeholders, financing, trade associations, standard bodies, labor unions,
governmental and quasigovernmental institutions, and other interested parties. These
communities come together in a partially intentional, highly self-organizing, and even
somewhat accidental manner. Productivity defines its success. It is required to be robust such
that it can survive shocks from inside or outside of the ecosystem that threaten to destroy it. Also
it should have the ability to create niches and opportunities for new firms. The latter
requires a change in attitudes from protectionist to cooperative.
e. Development, and the social change flowing from it, is not shaped by society’s genes, but by its
accumulated technical knowledge. Technology, not people, holds the center stage.

You can retrieve the answers online through the UVLE link to be provided by your professor after the
activity.

Guide/Study Questions for Activity 4

1. How is virgin material and energy source conserved in industrial ecosystem?

2. In view of economic life who is the driver of economic development and social change?
 3. What are digital species? Why are they important to digital business ecosystem?

4. Why is social ecosystem consist of firms and institutions, and not of people?

5. Enumerate and describe the four stages in the life cycle of a business ecosystem.

Discussion (Guide to study questions)

In an industrial ecosystem different members or parties involved in the ecosystem cooperate by utilizing
each other’s waste material and waste energy flows as resources. There is a reduction in the use of virgin
material and virgin energy inputs, including waste and emission outputs of the system as a whole
because waste is used to substitute for the virgin material and energy sources. On the other hand, in view
of economic life, economic development, and the social change flowing from it, is shaped by its
accumulated technical knowledge – technology.

In digital business ecosystem, digital species plays a crucial role in shaping its ecosystem. Digital species
are the ones that populate the digital environment. Examples of digital species are software components,
applications, services, knowledge, business models, training modules, conceptual frameworks, laws, etc.
In digital business ecosystem, the environment enables species to behave like species in natural world.
Digital species are important because they are the key players that interact, express an independent
behavior and evolve or become extinct if the amount of individuals of a species is not sufficient. Simpler
species may form compositions, which allows more complex species to appear.

In social ecosystem it is the social consensus represented by the firms and institutions (i.e. businesses,
consumers, suppliers, as well economic, cultural and legal institutions), that influence the functioning of
the entire ecosystem itself such that when firms and institution cease to function like a community or
ecosystem, they may breakdown.

Among others based on existing literatures business ecosystem demonstrates clearly a life-cycle. Birth
stage of business ecosystem focus on more than just satisfying customers. During the expansion stage
the scale-up potential of the business concept is tested. In the leadership stage the business ecosystem
reaches stability and high profitability. At the final stage, self-renewal or death, is caused by the threat of
rising new ecosystems.

7.2.2 Focal Complexity Aspects in Business Ecosystem

Here you will learn that complexity concepts are also applied in business ecosystems. This provides a
better view on how a business community organizes itself; from self-organization how does new order
arise as an emergent property; how the concept of co-evolution (and adaptation) is relevant to business
ecosystem.
Activity 5 (Total duration 20min)

Based on the article of Peltoniemi and Vuori that you read in the previous activity indicate which of the
given real life scenarios below pertains to which complexity concept (self-organization, emergence, co-
evolution and adaptation). Provide clear explanations (1 – 2 sentences) for your answers.

a. The formation of a business ecosystem is a process, where participants are gathered

voluntarily and without external or internal leader. Goals are set in local interactions, where
companies negotiate and create new order. Companies are free to create the kind of structures
they prefer. This evolvement is continuing, new connections are created all the time and old ones
are dissolved.

b. A business ecosystem is always more than the sum of its parts. The result of interactions
between different units is something, which no one of those units could produce by oneself. This
is especially visible in R&D, where the result is consisted by the contribution of many factors.

c. The evolution of one company affecting the evolution of other companies this is exemplified by
the classical case of microprocessors and software. While microprocessor producers develop
more efficient processors, the software producers quickly make use of the new
opportunities and the software becomes heavier, which causes pressure to develop even
more efficient processors. Also strategic changes of one company affect strongly to possibilities of
other companies in its ecosystem. This is why managers should consider the broad impact of
their decisions over the whole ecosystem.

d. Governmental restrictions, taxes and tariffs are those constraints, which are set by the other party
generates structures of progressively higher performance among the business participants.

You can retrieve the answers online through the UVLE link to be provided by your professor after the
activity.

Guide/Study Questions for Activity 5

1. Is emergence independent of self-organization? Why?

2. What is the significance of negative and positive interactions in co-evolution?
3. Define fitness in the context of adaptation in business ecosystem.
Discussion
Complexity refers to “systems with many different parts which, by a rather mysterious process of self-
organization, become more ordered and more informed than systems which operate in approximate
thermodynamic equilibrium with their surroundings (Cowan, 1994; Peltoniemi & Vuori, 2004 and Santa Fe
Institute, 2018). On the other hand, complex systems contain many relatively independent parts
which are highly interconnected and interactive (Cowan, 1994 and Peltoniemi & Vuori, 2004). In which
case, emergence is not independent of self-organization. In fact emergent properties are the result of self-
organization. And together with self-organization emergence creates new order. In business ecosystem
we cannot do away with negative and positive interactions happening during co-evolution and adaptation,
nevertheless negative interactions as exemplified by predation and competition and positive interactions
as exemplified by mutualism and sharing indeed contributes in the generation of high or low performance
among business participants. Hence, in the context of adaptation in business ecosystem fitness is defined
as the measure of performance (overachievement or underachievement) in business.

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