Module 7: Ecosystems As Complex Systems
Module 7: Ecosystems As Complex Systems
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 analyzed, 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.
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.
Learning Outcomes
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
systems 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 enable 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).
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.
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).
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).
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 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.
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.
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.
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.
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).
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.
Business Ecosystem
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).
Guide/Study Questions
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.
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.
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.
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.
a. 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.
b. 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.
c. 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.
Guide/Study Questions:
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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