Ecology Final Notes

Lecture 1: Ecology & evolution: the origin and dissemination of species

 Central variable: distribution (dispersion – where, density- how many)
organisms-surroundings.
 Genotype: all genetic characteristics of an individual that together
determine the characteristics of the individual.
 Phenotype: the actual expressed characteristics of the individual
 Phenotypic plasticity: the extent to which variation in phenotypes of an
individual is expressed depending on the environment.
 Co-evolution: mutual selection of organisms in interaction. Defence
(glucosinolates), detoxification and resistance.
 Little exchange and strong selection leads to ecotypes
 Ecotypes (similar to subspecies) populations of a species with a different
phenotype adapted to local conditions.
 Speciation: long-term isolation can lead to speciation: no hybrids with
fertile offspring occur between the species.
 Effects of complex patterns of allopatric speciation:
Subspecies: populations of a species that differ in characteristics, but
produce fertile offspring when hybridized
Ecotypes: (similar to subspecies) populations of a species with a different
phenotype adapted to local conditions
Ring species: complex of subspecies which can interbreed with adjacent
populations, but for which ‘’end’’ populations are too distantly related to
interbreed.
 Allopatric speciation: species arise in isolation (for example on an island).
Isolation and difference in ecological selection regimes
 Sympatric speciation: species arise when they live side-by-side
 Phytochoria. Classification according to the occurrence of orders of higher
plants (taxonomic characteristics)
 Zoogeographic regions do not completely overlap with phytochoria: due to
later terrestrial colonization; animals did not dissipate until the plates
(continents) were further apart.
 Parallel evolution. Starting point: common ancestor, but geographically
separated. Organs: analogue (the same form and function) and homologue
(developed from the same ancestral organ). Example: placental mammals
and marsupials.
 Divergent evolution. Starting point: common ancestor. Organs:
homologous organs (developed from the same ancestral organ). Example:
development of forelimbs in mammals.
 Convergent evolution. Starting point: very different ancestors. Organs:
analogue (the same form and function). Example: wings in animals.
 DNA: code of life. Source of variation. ‘Means of transport’ for inheritance.
 Cause of variation: mutations
 3 major forces in evolution:
Mutation (random, slow) adds alleles to the population
Selection (environment, slow) subtracts alleles from the population
Genetic drift (random, fast) subtracts alleles from the population =
decreases genetic diversity.
  Polymerase Chain Reaction (PCR): multiple copying of small pieces of DNA;
makes analysis of the sequence possible
 Molecular research. Genetic divergence within a population: markers with
a high mutation rate. Colonisation of environment over time (e.g. 10.000
years): markers with low mutation rate.
From Course unit 1:
 Proximate explanation: in answering the question ‘’how’’, the
environmental factors that are active at the same time are addressed and
in this way explain a reaction of the organism. These factors are a cause
of the reaction of the organism. In a proximate explanation, a cause-effect
(causal-analytical) explanation is given.
 Ultimate explanation: the ‘for what purpose’ question concerns the
function. It addresses factors, that through evolutionary processes in the
past, have been the cause of certain traits that the organism has acquired.
The answer explains these traits with evolutionary reasoning.

Lecture 2: Environmental conditions, resources, biomes

 Biomes: ‘’geographic regions that contain communities composed of
organisms with similar adaptations’’.
 Convergent evolution: species descending from unrelated ancestors, but
with similar adaptations.
 Biomes: current vegetation is not equal to potential vegetation. Biomes
are related to potential vegetation. Current vegetation can differ (due to
land use).
 Primary succession: no previous vegetation
 Secondary succession: some vegetation left after disturbance
 Aquatic biomes categorized by salinity, flow, depth
 Freshwater biomes:
Lotic systems (streams, rivers): flowing fresh water, shallow
Ponds & lake: non-flowing fresh water, shallow or deep
Swamps: non-flowing, very shallow, emergent trees
Marshes: non-flowing, very shallow, non-woody plants
Bogs: non-flowing, very shallow, acidic (rainfed) water
 Saltwater biomes:
Intertidal: very shallow, daily flooding
Mangroves: very shallow, woody plants
Salt marsh: very shallow, regular floods, non-woody plants
Coral reefs: shallow, warm
Open ocean: deep, different zones
 Environmental conditions: ‘’abiotic environmental factors that vary over
time and in space, to which different organisms respond differently’’. An
environmental condition is variable, is not consumable. Examples:
temperature, pH, salinity.
 Resources: ‘’abiotic and biotic matter that varies over time and in space
and that can be consumed by organisms’’. A resource is variable, is also
consumable. Examples: Radiation (PAR=photosynthetically active
radiation), CO2, nutrients, water.
 Soil pH. Acidification leads to cation exchange. This leads to plant species
replacements (and sometimes species loss).
 Plant resources: light (or PAR), CO2, water, nutrients.
 Closed vegetation absorbs almost all the radiation. Little light reaches
lower levels.
 Leaf adaptations to light vs. shade: leaf morphology.
Sun-grown leaf: thick layer of photosynthetically active cells, higher
carbon investment
Shade-grown leaf: thin layer of photosynthetically active cells, lower
carbon investment
 Without light, sun-grown leaf loses more carbon than shade-grown leaf
Sun-grown leaf needs more light to have a neutral carbon balance
Maximum photosynthetic rate is higher for sun-grown leaf.
 CAM plants: cacti, crassulaceae
 Nutrients (e.g. N, P,K)
 .
 Environmental conditions: role of temperature. Signalling or stimulus
effect (e.g. germination after cold period, vernalisation) or extreme factor
(frost). Influence on metabolic rate and physiology.
 Ectothermic (cold-blooded) organisms versus endothermic (warm-blooded)
animals.
 Role of temperature on ectothermic animals. Great influence on metabolic
rate, growth, development. Lifecycle determined by physiological time Vt,
time needed to finish development. (=product of physiological rate
(depending on temperature) and time).
 Calculation of physiological rate (V) for ectothermic organisms
(=development of the organism). V=c(T-a) – c=proportionality constant,
a=threshold value.
 Role of temperature on endothermic animals (birds, mammals). Regulation
by internal thermostat. Requires much additional energy. Energy req.
M=aW0.67
 Bergmann’s rule: in cold regions you find larger species or subspecies
(applies to related, endothermic animals)  due to reduced surface area/
volume ration, this decreases heat loss and hence energy demand.
 Light and water pressure. Adaptation to dark: luminescent fin ray to lure
prey, pheromones, dark skin. Water pressure: bones of cartilage (instead
of calcium carbonate)
 Resources for animals: food, nest cavities, territories. Food differs between
herbivores (grazers and browsers), predators, parasites, decomposers.
 Mutualism in herbivores – micro organisms have enzymes (such as
cellulase) that make digestion of cellulose and hemicellulose possible.
Various adaptations in ruminants (grazers, browsers, intermediaries).
Foregut digestion
 Defences: mechanical (spines, thorns, armour), chemical defence, escape,
camouflage, aposematism and mimicry (Bates)
 Competition occurs when different individuals (from the same, or different
species) need the same, limited resource.
 Exploitative competition: individuals consume resource up to the point
that other individuals cannot persist.
  Interference competition: individuals interact in such a way that they do
not allow the other to get excess to a resource.
 Different biomes, characterized by different plant growth forms and animal
communities, exist on Earth
 Climatic conditions (on land), salinity, depth and water flow (in the water)
determine the distributions of biomes.

From Course unit 2:



Lecture 3: Populations
 Population: a group of individuals of a single species
 Unitary organisms: organisms with a highly determinate form and
predictable development (e.g. birds, fish, humans)
 Modular organisms: organisms that are build up of modules that can
change shape/function in response to the environment (e.g. plants, corals)
 Clonal organisms: modular organisms in which the structural units
(modules) also survive independently of other units.
 Genet (genetic individual): the product of the zygote, which consists of
many modules and ultimately forms an individual
 Ramet: the independent unit of a clonal organism (e.g. a plant shoot with
its own roots).
 How to count individuals:
Census: total population, not statistical
Sample: part of the population, estimation, statistical
Tagging and recapturing: indirect, statistical
Marking and recapturing: indirect, statistical
N=Mn/m
 Reproduction. Semelparous: an organism has offspring only once.
Iteroparous: an organism has offspring several times during its lifespan.
 Counting iteroparous organisms.
Cohort: all individuals born in the same period (usually a year) (also called
a generation)
Static life table: counting the number of individuals of a certain age group
to gain insight into the population structure.

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 
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 Dispersal: unfocused, dispersal of individuals or a part of the population.
Density-dependent: dispersal due to an increase in local population density.
Age-dependent: dispersal depending on the age of the organism; for example; young animals
often colonise new areas.
 Migration: focused and often massive
Repeated migration: multiple return ticket; Dailu migration similar to tides
or daily commute between sleeping and foraging sites. But also between
habitats, as with migratory birds.
Migration with return: one return ticket only; birth in habitat A, reaching
adulthood in habitat B and migrating back to habitat A for reproduction
(salmon, eel) Eel migration is age-dependent
One-time migration: one-way only, migration in a single direction.
 Intraspecific competition and population growth.
Exponential growth: dN/dt=r*N
When resources are unlimited: maximum natality and minimum mortality;
growth rate r=intrinsic growth rate (rmax)
 Exponential growth: unrestrained growth when resources are unlimited
Logistic growth: feedback on population growth through intraspecific
competition for unlimiting resources
K: carrying capacity of an area (max. population size limited by resources)

r-selected species: exhibit exponential population growth, individuals have many small offspring.
Environmental conditions cause collapse of the population. r-selected species are often found in
unstable environments.

K-selected species: the population grows towards the carrying capacity and will fluctuate around the
carrying capacity. Individuals have few but relatively large offspring and invest more in their young.
K-selected species are found in stable environments.
 Interference competition: populations are limited because individuals physically exclude each
other from resources.
 Exploitation competition: individuals compete by using resources, but do not exclude each
other physically. Parasitic wasps: larvae of parasitic wasps that depend on a limited resource,
such as the caterpillar host, compete for food. Larvae usually stay smaller when density is
high.
 .
 Grimes plant strategies: r/K classification extended for the plant realm
 Stress-tolerant species (S): slow-growing and long-living species that dominate in high-stress,
low-competition environments
 Ruderal species (R): similar to r-selected species; fast-growing species, massive reproduction
and short-lived. They dominate in low stress, high disturbance environments.
 Competition dominant species (C-competitors): similar to K-selected species; large, fast-
growing plants that rapidly absorb available resources. They dominate in low stress, low
disruption environments.
 .
 Predation
True predators: usually kill their prey immediately after attacking, consume multiple prey
animals over their lifetime
Grazers: attack multiple/many prey organisms in their lifetime, consume only part of their
prey, usually do not kill their prey
Parasites: consume only part of their prey (in this case called a host), usually do not kill their
prey, only attack a single prey in their lifetime and are usually heavily dependent on their
host.
 Cyclical patterns: predators inhibit prey population growth and are dependent on prey
populations for their own population growth.
 Grazers (predation): cause changes in vegetation structure. Inactive buds become active when
other growing points are consumed. In this way plants compensate for damage caused by
herbivores.
 Plant parasitism can lead to changes in competitiveness and thus vegetation composition.
 Prey populations grow in the absence of a predator: dN/dt=rN
 Predators inhibit the growth of that prey population depending on the efficiency of the
predator (a): dN/dt=rN-aPN
 Predator populations decrease due to absence of prey, depending on mortality rate (q):
dP/dt=-qP
 However, predator populations grow due to the presence of the prey and that growth is
determined by the consumption rate (aPN) and the efficiency of converting food into
offspring (f): dP/dt=faPN-Qp
 What causes the prey population to stop growing? dN/dt=0=rN-aPN. The predator population
(P) is therefore r/a
 What causes the predator population to stop growing? dP/dt=0=faPN-qP. The prey population
(N) is therefore q/fa
 In short: the prey and predator populations interact cyclically!
 Many predators result in the prey population decreasing, a small prey population results in the
shrinkage of the predator population, few predators result in an increased population of hares
 Predation: optimal foraging. If E=energy uptake and h=processing time, then
optimisation=maximisation E/h
 Therefore if s is small: predator is a specialist (lion), if s is large: predator is a generalist
(insect-eating songbirds search for camouflaged prey)

 Predation
 Functional response Type 1: predation (P) increases with increasing density of prey (N). The
predator has no processing time but will become satiated. For example: a water flea that
 swallows prey completely becomes satiated when the intestinal tract is full of blood.

 Functional response Type 2: predation (P) increases with increasing density of prey (N). The
predator does have processing time and will become satiated. For example: most large
predators catch prey that are too large to swallow at once. The prey must first be torn into
pieces to be able to consume it. They have processing time, but these predators also become
satiated.

 Functional response Type 3: predation (P) increases with increasing density of prey (N). The
predator does have processing time, will become satiated, but will opt for another prey
(switching) if the density of the initial prey becomes too low. For example: a blackbird
searches for earthworms, but switches to other prey if the density of the earthworms becomes
low.

 Metapopulations. Local populations have unstable predator-prey relationships.

Metapopulations have a cyclic relationship between predator and prey because the prey can
escape the predator in individual populations.
Local populations have larger fluctuations than the metapopulation.
Larger patches often harbour a population for longer periods of time.
Small patches are more likely to have a temporary population (smaller target for colonisation
and greater chance of extinction).
  Most important aspects:
Counting by marking and recapturing
Birth, mortality and population growth
Semelparity versus iteroparity
r- and K-selected species
Dispersal versus migration
Intraspecific competition inhibits the growth of a population
Predators and optimal foraging
Metapopulations and the stability of predator-prey relationships.

Lecture 4: Interspecific competition and species richness

 Early succession: higher species richness, heterogeneity, number of
niches.
 Later succession: higher competition, dominance; lower species richness.
 Infraspecific competition: individuals of the same species compete for or
exclude each other from the use of resources, resulting in reduced growth,
survival or fitness.
 Interspecific competition: individuals of different species compete for or
exclude each other from the use of resources, resulting in reduced growth,
survival or fitness.
 Interspecific competition is one of the most fundamental processes in
ecology and partly determines the dispersion and success of a species, as
well as the evolution of a species. The result is that interspecific
competition is decisive for the species composition of an ecological
community.
 Experimental evidence shows that the effects of competition are powerful.
For example, when measuring competitive exclusion.
 Important observations in the field. Competing species live together on a
larger spatial scale, but have clear differences in distribution on a small
scale. Species are excluded from areas they could populate if interspecific
competition was not a factor.
 In a stable environment, two species can only live together sustainably
(coexist) when they occupy different niches. If there is no niche
differentiation, one competing species will drive the other species out.
 Interspecific competition: Lotka-Volterra model.


 Fundamental niche (without interspecific competition): the combination of

conditions and resources that make the occurrence, development and
reproduction of a species possible.
 Realised niche: the conditions and resources that make the occurrence,
development and reproduction of a species possible in the presence of
other species that limit survival (interspecific competition).
 When interspecific competition is lost, the realised niche expands towards
a fundamental niche.
 Niche differentiation (change in fundamental niche)
 Evolutionary consequences:
Character displacement: a morphological adaptation due to competition to
reduce interspecific competition.
Mongoose competition: in the presence of large competing species, the
upper jaw of H. janvanicus becomes smaller to exploit an open niche of
small prey (niche differentiation).
 Character displacement in Darwin’s finches.
Niche differentiation: adaptations of species to a new environment and/or
resource. Therefore: character displacement is the result of competition
(current or in the past). Niche differentiation can be the result of
competition, but not necessarily.

 Species richness: number of species in an area

 Biodiversity index: an index that weighs both species richness and the
distribution of individuals across species (evenness)

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  Regional: isolation, size, energy/productivity, age (time)
Local. Environmental conditions: competitive exclusion niche
differentiation, predation, disturbance, Spatial and temporal factors: island
biogeography, metapopulation.
 High availability of nutrients can reduce species richness: species that can
use nutrients efficiently will dominate and outcompete other species.
 Local niche use. The quantity of potential resources and the way species
use these resources determines species richness.
 Effects of predation. Predators ensure greater species richness by
preventing competitive exclusion between prey species.
 Effects of parasites. Predators ensure greater plant species richness by
preventing competitive exclusion.
 Effects of disturbance.
Initial situation: niches overlap substantially with expected competitive
exclusion (equilibrium theory around K)
Disturbance: prevents competitive exclusion due to fluctuations in
environmental conditions (non-equilibrium theory) and ensures that strong
competitor species do not dominate.
Abiotic disturbance: wildfires, storms, floods
Biotic disturbance: herbivores, predation
 Effects of disturbance on vegetation:
After disturbance (humans and climate): pioneer vegetation and new
succession to climax vegetation
After intensive grazing: interspecific competition, strongly competitive
plants are suppressed, grazing-tolerant plants remain.
 Explanations of species richness: spatial factors (island theory)
Colonisation (distance effect): the greater the distance from an island to
the mainland, or the smaller the island, the smaller the chance of
colonisation by species.
Extinction (area effect): the smaller the island, the smaller the population
and the greater the risk of extinction.
Migration effect (I): as more species are present on an island, fewer
species will migrate there.
Extinction effect (E): as more species are present on an island, extinction
increases.
 Species richness equilibrates where the immigration and extinction curves
cross.
 Most important aspects:
Niche differentiation
Fundamental versus realised niche
Character displacement
Lotka-Volterra model for competition
Effect of disturbance and predation on species richness
Disturbance and succession in vegetation
Island theory

Lecture 5: Food webs, energy, nutrient fluxes and cycles

 Food webs are representations of how species feed on each other in a
community; graphically shown as networks.
 Trophic levels:
Consumer subsystem: Tertiary consumers, secondary consumers, primary
consumers
Autotrophic subsystem: primary producers
Many organisms are omnivores (eat organisms from multiple trophic levels
Guilds: species within a trophic level with similar food.
Detritivore subsystem: decomposers & detritivores
 A trophic cascade
 Bottom-up versus top-down control
 .
 Energy fluxes
 Primary productivity: the rate at which solar energy or chemical energy is
captured and converted into chemical bonds by photosynthesis or
chemosynthesis.
 Gross primary productivity (GPP): the rate at which energy is captured and
assimilated by producers in a given area. Unit: energy per area per year
 Net primary productivity (NPP): the rate at which energy is assimilated by
producers and converted into producer biomass in a given area. Unit:
energy per area per year
 NPP=GPP-respiration
 Standing crop (standing biomass): biomass of producers in an area. Unit:
biomass per area
 Measuring primary productivity: GPP=NPP+respiration
 Secondary production: the rate of biomass accumulation of consumers in a
given area.
 Consumption efficiency=consumed energy / NPP lower trophic level
 Assimilation efficiency=assimilated energy/ consumed energy
 Net production efficiency=Net production energy/ assimilated energy
 Ecological efficiency=net production energy/ net production energy lower
trophic level
 Biomass=net productivity* biomass residence time
 Biomass residence time: average time that biomass spends in a trophic
level.
 SB (standing biomass)=mean residence time (MRT) *NPP
 Very high soil N and P availability, relatively high precipitation  high NPP
 Energy fluxes drive trophic pyramids
 .
 The carbon cycle

 Anoxic respiration: common form of respiration in swamps or bogs;
produces methane, an important greenhouse gas.
 The soil stores around 70% of the terrestrial organic carbon.
 .
 The nitrogen and phosphorus cycle
 Nitrogen fixation: atmospheric nitrogen is converted to ammonium. Biotic
fixation is carried out by cyanobacteria and some bacteria associated with
plant roots. Abiotic fixation is carried out by lightning, combustion and
fertilizer production.
 Nitrification: nitrifying bacteria convert ammonium into nitrite and then
into nitrate.
 Assimilation: producers take up either ammonium or nitrate. Consumers
take up nitrogen by eating producers.
 Mineralization: decomposers in soil and water break down biological
nitrogen compounds into ammonium.
 Assimilation+waste: tertiary, secondary, primary consumers, primary
producers
 Decomposers & detritivores: assimilation +mineralization
 Denitrification: in a series of steps, denitrifying bacteria in oxygen-poor
soil and stagnant water convert nitrite into nitrate, nitrous oxide and
nitrogen gas.
 Atmospheric N decomposition. Human activities  increased N
decomposition in nature areas  increased plant productivity  loss in
plant biodiversity due to light competition.
 The phosphorus cycle. Excess phosphorus on land runs off the surface or
leaches out of the soil and into aquatic habitats. Phosphates can also
return to the ocean via wastewater.
 P eutrophication causing dead zones
 .
 Decomposition: consumption by large detritivores, maceration, direct
decomposition, leaching, breakdown of lignin and cellulose by fungi,
breakdown of numerous organic compounds by bacteria, soluble minerals
organic compounds
 Decomposers help recycling nutrients
 Decomposability different materials: from highest to lowest: lignin,
hemicellulose, cellulose, soluble sugars.
 Decomposition rates also influence soil layers
 From lower to higher more active soil fauna, more nutrient-rich:
 Mor (sandy soil): litter, decomposing litter, humus, as mull, but with a
higher inorganic content.
Moder: litter, decomposing litter, as mull, but with a higher inorganic
content
Mull (clay soil): litter, organic material mixed with mineral soil.
 In the absence of soil fauna: slow decomposition.
 Global warming can decrease C stocks in tundra. This can cause positive
feedback, i.e. global warming accelerating itself.
 .
 Organisms are part of foodwebs, and changes in the abundance of one
species can (indirectly) alter other components of foodwebs.
 Biomass, energy and nutrients are produced by primary producers (e.g.
plants), and transferred (often with losses to e.g. the atmosphere) to
higher trophic levels.
 As such, energy, biomass and nutrient cycles are inherently related to
each other.
 Decomposers recycle carbon and nutrients
 Human activities can alter foodwebs, nutrient and carbon cycles, often
with very complex outcomes.

Lecture 6: The influence of humans: pollution and nature conservation

 The values of biodiversity:
Intrinsic values
Instrumental values (Cultural, Supporting, Regulating, Provisioning
services)
 Cultural services (aesthetic, spiritual or recreational values)
 Supporting services (benefits of biodiversity that allow ecosystems to
function)
 Regulating services (Benefits of biodiversity that make Earth’s
environment more liveable for humans). Climate regulation, flood control,
water purification
 Provisioning services (Benefits of biodiversity that provide products for
human use). Food, timber, medication, fibres (clothing)
 .
 Biodiversity in declining
 Estimation of current extinction rates: based on e.g. species-area
relationships. These can also be used to project future species losses.
 .
 Causes of biodiversity decline: habitat loss, overharvesting, climate
change, introduced species, pollution.
 Current extinction rates. The more habitat you lose, the more negative the
consequences for biodiversity.
 Habitat fragmentation
 Overharvesting
 Effects of climate change on animals and plants:
Changes in phenology (e.g. growing season, flowering time, reproductive
behaviour)
Changes in behaviour (e.g. habitat selection, diurnal cycles)
 Changes in performance (survival, growth, reproductive success)
Changes in distribution
 Responses to elevated CO2: prairies and savannahs: high abundance of C4
grasses. C4 photosynthetic pathway: efficient with CO2  C3 plants profit
more from elevated CO2 levels  shrub encroachment
 Biological feedbacks (also called vegetation-climate feedback): increases
in photosynthetic rates (CO2 uptake); increases in decomposition and
respiration rates (CO2 release)  these changes can either buffer or
accelerate climate change.
 Introduced species. Examples: spread of ornamental (garden) plants,
escaped pets/ farm animals, hunting game, pest control species,
accidents.
 Pollution. Examples: eutrophication, acidification, pesticides, heavy
metals, light, sound.
 .
 Biodiversity conservation
 Targets: maximise species richness, protect rare species, protect
‘charismatic’ species, species with high ‘cultural’ values
 Metapopulations: a collection of sub-populations that live in isolated
patches that are linked by dispersal.
 Sub-populations are often unstable (and sometimes locally extinct)
Colonization depends on the degree of isolation
Extinction risk is larger for smaller populations in small patches.
 Addressing pollution: policies (fertilizers, pesticides), grazing/mowing (to
remove nutrients); topsoil removal (to remove nutrients)
 There are various reasons why humans value biodiversity
 Biodiversity is declining rapidly due to various causes, including habitat
loss, overharvesting, climate change, invasive species and pollution.
 Nature conservation policies and management activities aim to reverse
these causes, or compensate for them, thereby restoring biodiversity.

Tutorial 1: Demography
 r = lnR = lnR0 / 1 = lnR0

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 
 mx= the average number of female calves born per surviving female
buffalo per year in age class x

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 N 1=N 0∗R

where R=e r
t
N t =N 0∗R

 R0= The mean number of female descendants that a mean female in

the population produces over her entire life

Tutorial 2: Intraspecific competition and harvesting models

dN
 Growth of population: =r∗N
dt
r∗t t
 N t =N 0∗e =N 0∗R
dB
 Biomass growth= =r∗B
dt
r∗t
 Bt =B 0∗e where B is the biomass of the population
 ln Bt =ln B0 +r∗t
 Carrying capacity: the maximum biomass that a population can reach
under certain circumstances.
 Intraspecific competition: individuals in the population start competing for
the limiting resource
 dB B
 The logistic growth equation: =r∗B∗(1− )
dt B∞
dB
∗1
 The intrinsic growth rate: dt
=r
B
 MSY=Maximum Sustainable Yield=dB/dt
 If we assume that a certain fraction of the biomass can always be
harvested with a certain fishing effort (fixed effort), this means that a
population decrease will automatically result in a smaller harvest.
 The advantage of a logistic transformation: the slope yields the intrinsic
growth rate

Tutorial 3: Interspecific competition

 RYT (Relative yield total)
 Interspecific competition – competition between species
 Replacement design – to measure the competition between 2 plant
species
 For each species the logistic growth model is characterised by the intrinsic
growth rate (r) and the carrying capacity (K).
 Competition coefficient reflects the relationship between interspecific and
intraspecific competitiveness.
 The competition coefficient α 12 indicates how, for individuals of species 1,
the interspecific competitiveness of one individual of species 2 relates to
the intraspecific competitiveness of species 1. In other words, the first
index (in this case species 1) indicates on which species an effect is
exerted, while the second index indicates by which species (in this case
species 2) the interspecific competition is exerted).
 The Lotka-Volterra model for interspecific competition:
dN1 ( K −N 1−α 12 N 2 )
=r 1 N 1 1
dt K1
dN2 (K 2−N 2−α 2 1 N 1 )
=r 2 N 2
dt K2
 The zero isocline of species 1 is the line with combinations of population
sizes of species 1 and species 2 at which the population growth of species
1 is exactly zero.
 Only one of the 2 populations prevails (competitive exclusion), or a stable
equilibrium develops in which both species coexist.
 Both species can continue to exist side-by-side in a steady state. This is
called a coexistence.
 Niche differentiation, where both species partially use different resources,
or retrieve the resources from a different location.
 

Tutorial 4: Predation
 The functional response of the predator is the relationship between
predation rate and prey density.
dN
 The prey equation: =rN −aPN
dt
r is the intrinsic growth rate of the prey population; N is the prey density
and P is the predator density; a is the captured fraction of the prey
population pre predator per unit of time (attack coefficient).

dP
 The predator equation: =faPN−qP
dt
f is the number of predators that can be produced by consumption of one
unit of prey (conversion efficiency); q is the mortality of the predator.
 aPN – absolute mortality rate of the prey due to predation = attack rate of
the predator
 [pred/prey] x [(1/(pred) x (1/time)] x [pred] x [prey] = [pred/time]
faPN – qP = dP/dt


 Type II: Predation rate depends on search efficiency and processing time:
finding more prey, needing more processing time, less time left to search.
  Type III:Like type II, but with increasing search efficiency or reduction of
processing time at higher N. Due to (1) limited number of hiding places for
the prey, or (2) learning behaviour of the predator with sufficient prey
supply, (3) passivity of thepredator at low N; (4) loss of interest from the
predator because it can switch to another prey that requires a different
search behaviour.


 .
 Many predators can consume more than one species of prey
(polyphagous). The characteristics of prey species can differ:
a) The time a predator needs to find prey (search time)
b) The time a predator needs to catch and eat prey (handling time)
c) Prey size (in terms of energy per unit of prey)
d) Prey quality (in terms of specific nutrients per unit of prey weight).
 Preference for prey is only expected when it provides a fitness benefit. A
fitness benefit is the result of a larger number of offspring per unit of time,
i.e. a larger basal reproduction rate (R 0) when the predator population can
find its preferred prey.
Tutorial 5: Processes in the ecosystem (Substance flows and cycles)

dP
 =I t−U t=P t+1 −P t
dt
 Pt +1=Pt + I t −U t
 U t =k∗Pt
 It=inflow rate; Ut=outflow rate; Pt=size of the pool
 When the two lines intersect, the pool has an equilibrium point (P*) at
which the influx is equal to the outflow, which causes the pool to remain
equal in size.
 The equilibrium is stable when I > U at Pt < P*, and I < U at Pt > P*
The equilibrium is unstable when I < U at Pt < P* and I > U at Pt > P*
The equilibrium is neutral when the lines of I and U coincide. (Equilibrium
interval)
 If the consequences of a process influence the rate of the process, this is
called feedback. Positive feedback is when the process is accelerated by
its consequences and negative feedback is when the process is
decelerated by its consequences.
 Positive feedback has a destabilising effect on a system, while negative
feedback has a stabilizing effect.
 The MRT (Mean residence time) is the time in which the amount flowing
through the pool is equal to the size of the pool.
 The opposite of the MRT is the turnover rate (TO). It is the fraction of the
pool that is replaced per unit of time.
Pt Ut
 MRT = and ¿=
Ut Pt
 TO: kg · kg–1year–1
MRT: year
 Resistance is the capacity of the system to withstand external influences
and recover from an external disruption.
 Resilience is the capacity of the system to restore the equilibrium after
disruptive external influences.
(I t −U t )
 Resilience= =¿
(P ¿−Pt )
 Consumption=NPP*CE
 Mortality=NPP*(1-CE)
 Assimilation=Consumption*AE
 Faeces=Consumption-Assimilation
 CE=Dc/(Sv+Fh)