Objectives

LSM3254 Ecology of Aquatic Environments

To learn about selected ecological processes and concepts at various scales in freshwater environments Scope Populations
Growth models Life history strategies

Population, community, ecosystem ecology in freshwaters

Communities
Aquatic communities Species interactions Community structure and dynamics

Ecosystems
Ecosystem concept Primary production Secondary production Chemical cycles

Darren Yeo Dept of Biological Sciences

References: - Dodson, S. 2005.Introduction to Limnology. McGraw-Hill Chapters 6-10 - Molles, M. C. Jr., 2008. Ecology: concepts and applications, 4rd Ed. McGraw-Hill

Models of population dynamics

Predict changes in population size and structure. Applications in:
Fisheries management Biological control Invasive species spread

Previous conventional wisdom: mainly bottom-up regulation of birth rate by food availability
E.g., Falling algal/Daphnia densities Prediction: falling Daphnia birth rate expected But insteadhigh Daphnia birth rate (based on population model estimates)

Two useful models of population growth for aquatic organisms:

Exponential growth model
Near-optimal conditions; rapid reproduction and development, and population growth Goal: production of the most offspring as quickly as possible Egg bank e.g., Daphnia in arctic pondssubitaneous or diapausing eggs? All subitaneous (more females more propagules) long growing season

Egg bank (resting stages stored in aquatic habitat) model

All diapausing (only Suboptimal conditions (time and resource propagules) short growing constraints); rapid population growth not possible Goal: production of propagules (diapausing stage) season to withstand harsh environmental conditions Combination of both

Paradigm shift - new hypothesis: Daphnia death rate from predation > birthrate, i.e. Top-down regulation of aquatic species population dynamics by predation

Life history strategies

Underlying aims:
Maximise genotype contribution to the next generation Benefit is to the population/species, not individual

Life history strategies

Seasonal timing
Timing of life cycle stage to coincide with specific environmental conditions, i.e. adaptation for specific seasonal pattern

Life history adaptations. Characters that:

Increase reproductive success Can be inherited (i.e. genotypically derived vs environmentally derived) Are affected by environmental conditions (especially aquatic species)
Less food fewer, smaller offspring Higher/lower temperature suboptimal population growth

Size vs Number of offspring

Tradeoff between size and no. of offspring Influenced by
Predation Need to complete life cycle inland before being flushed to sea

Other factors:
Biological constraints (interacting species) Tradeoffs for limited resource (e.g., food or time)

Age-specific allocation of energy

Division of labour: juvenile growthadult reproduction Tradeoff between growth and reproduction

Reproductive speed vs Resting egg production

Life history strategies

Reproductive strategies and age-selective predation
Tradeoff between reproduction and survival

Aquatic communities
Major aquatic communities
Generally distinct Based on habitat/microhabitat Facilitate study and Shallow, understanding of aquatic shoreline ecosystems waters

Open, deeper waters

Life span
Reproductive output in few, old, large individuals vs many, young, small individuals Tradeoff between life span and locomotion
Branchiopod (Daphnia pulex) short lifespan
Fast reproduction, large reproductive effort Little escape ability

Lake communities
Pelagic Littoral Benthic Periphyton (aufwuchs)

Copepod (Diaptomus ashlandi) long lifespan

Slow reproduction, small reproductive effort Excellent escape ability

Stream communities
Channel Riffle and hyporheic Pool Periphyton (aufwuchs) Riparian floodplain

Aquatic communities
Food webs
Typical of community ecology
Benthic feeding fish

Osprey

Aquatic communities
Models of community assembly
Piscivorous fish

Reflect species interactions among members of a community

Competition Predation

Null or neutral assembly

Based on adaptation of species (independently of other species) to physico-chemical conditions of the environment

Crab

Planktivorous fish

Niche assembly

Snail Large zooplankton

Descriptive models
Components (boxes) Groups of organisms that have the same predator(s) and prey Interactions (arrows) Indicate diet food ! consumer Different from trophic webs, which reflect energy flow

Small zooplankton

Based on interactions between species adapted to particular roles in the environment Major drivers (species interactions): Competition Predation

Macrophytes

Periphyton

Phytoplankton

Species interactions: competition

Competition
Limited resources. Inter- or Intraspecific Exploitative vs interference competition
Independent (indirect) competition for resource vs direct prevention of competitor from utilising resource

Species interactions: competition

Avoiding/reducing competition maintains high diversity/community complexity
Environmental specialisation
Species with different environmental requirements (e.g., temperature, pH, DO, salinity)

Competitive exclusion of one species by a competitor

Important process in niche assembly model Should result in reduced no. of species, yet many complex communities with higher nos. of species

Habitat partitioning
Species with similar general environmental requirements live in different parts of habitat

Selective feeding/resource partitioning

Species exploit different aspects of same general food resource
e.g., zooplankton feeding strategies: individual capture vs bolus capture

20-30 spp. chydorid water fleas found in littoral zone of temperate lakes
Different body sizes associated with ability to exploit a habitat in different ways

Species interactions: predation

Predation
Killing individuals of one species (prey) by another species (predator) Intense summer predation ! significant effects on freshwater community assembly in temperate zones
No population growth or decline in prey populations Changes in species composition

Many species of shorebirds ( waders ) feeding in the shore/littoral zone habitat of aquatic habitats
Different shapes/sizes (overall, bills, legs) and behaviours help partition the habitat exploiting various microhabitats and different aspects of common food resource (benthic invertebrates)

Three different spp. of freshwater crabs in different parts of the same stream habitat Three different spp. of freshwater prawns in the same stream with different feeding strategies

Species interactions: predation

Size-selective predation strategies
Invertebrate predation
Larger zooplankton selectively feed on smaller zooplankton

Species interactions: predation

Avoiding/defending against predation based on selective nature of predators - maintains high diversity/community complexity
Behavioural defences
Migration to refuges, e.g., DVM Inconspicuous movement Swarming/schooling

Morphological defences
Thick coverings/shells, spines - response to chemical cues from predators (kairomones) Cyclomorphosis: seasonal morphological forms between generations

Vertebrate predation
Major factor affecting zooplankton species composition in lakes
Fish selectively feed on larger zooplankton Lakes with more fish predation ! more smaller zooplankton ! more larger phytoplankton

Life history defences

Timing and/or duration of life history stages Adult body size Number and size of offspring

Behaviour, morphology, life history are interconnected

Limited energy budget - therefore tradeoffs expected
E.g., longer spines but longer development time; faster development but smaller size

Species interactions: predation

Predation vs Competition: which is more important in shaping a community?
Studies on zooplankton
In general, competition (food limitation) rather than predation affects zooplankton abundance However, intense predation in summer can affect zooplankton abundance and species composition

Effects of predation on community mediated through intermediate trophic level species

How do predatory fishes decrease algal densities?

Species interactions: trophic cascades

Traditional water chemistry perspective: aquatic species interactions thought to have little effect at community scale
E.g., Managing lake algal blooms caused by nutrient enrichment Top-down chemical poisoning approach Bottom-up control of physical, chemical factors

Alternative approach: biomanipulation based on a trophic cascade hypothesis

Top-down approach using predator-prey interactions to control algal blooms ! improve water quality Increase large herbivorous zooplankton that prey on nuisance phytoplankton
Achieved indirectly by increasing (stocking) piscivore abundance (directly increasing zooplankton/decreasing planktivore abundance not feasible)

Absence of predatory fishes increase algal densities Predator-prey interactions exploited for biomanipulation

Increase in piscivore population

TROPHIC CASCADE HYPOTHESIS

Community structure
Community structure is described in various ways Species diversity characterised by:
Richness number of species
In a lake, depends on:
Taxonomic group richness inversely related to size scale of organisms Lake size richness directly related to lake size Lake production highest species richness in mesotrophic lakes

Decrease in planktivorous minnow population

Increase in herbivorous large zooplankton population

Decrease in nuisance phytoplankton (biomass)

Evenness relative abundance of species Richness and evenness

Decrease in rate of primary production

Diversity indices (e.g., Shannon Index) Rank-abundance curves

Community structure
Species composition (community composition): a list of species in the community
Turnover - change over time Global associations
Common evolutionary/biogeographical history

Community dynamics
Community dynamics:
Changes in structure of communities over time (various time scales) Important driver: disturbance (see LSM2251) Stability: persistence of community despite disturbance (see LSM2251): Resistance Resilience

Local associations
Local adaptation (spp. with similar environmental requirements) Symbiotic relationships

Short term changes:

DVM Stream drift
Daily downstream drift of macroinvertebrates Highest in summer, at night, and when fish predators present Compensated by:
Upstream movement of benthic insects Upstream egg deposition by flying adults

Size-spectrum: relative abundance or biomass of organisms or particles across a range of sizes

Seasonal/Annual succession

Community structure
Long term change processes:
Invasions invasive species impact community structure Immigration and Extinction
Equilibrium Model of Island Biogeography (lake=island)species richness based on dynamic balance between immigration and extinction rates

Ecosystem concept
Ecosystem:
Organisms and their physical environment as an integrated system Ecosystem models: describe flow of energy (trophic dynamics) or chemicals (chemical/nutrient cycles) between components (trophic levels or reservoirs) Movement affected by physical, chemical and biological processes

Trophic dynamics models

Energy flows from energy sources through to successively higher trophic levels (sinks), with loss of some energy at each level Trophic levels: Position of component in an ecosystem based on common source of energy and number of energy transfers from primary producers to that level
Primary producers: autotrophs (energy transfer from 1 source) Primary consumers: herbivores (energy transfer from 1 producers) Secondary consumers: primary carnivores (energy transfer from 1 consumers) Tertiary consumers: top carnivores (energy transfer from 2 consumers)

Primary production
Primary production: fixation of carbon by photosynthesising autotrophs
Gross primary production (GPP): Total amount of energy fixed by autotrophs in an ecosystem Net primary production (NPP): Amount of energy remaining in ecosystem after autotrophs have met their own metabolic requirements (i.e. respiration) NPP = GPP respiration (autotroph)
= Amount of energy available to the rest of the ecosystem

Primary production
Freshwater photoautotrophs: photosynthetic autotrophs
Cyanobacteria Protists (euglenoids, diatoms, dinoflagellates, green algae, red algae) Macrophytes Photosynthetic pigments: Chlorophyll a and b Accessory pigments enhance light absorption; protection from photoinhibition
Carotenoids: xanthophylls (yellow), carotenes (orange) Phycobilins (red)

Chemoautototrophs: chemosynthetic autotrophs

Sulphur-oxidising bacteria Using H2S instead of H2O as electron donor to reduce CO2 to carbohydrate in absence of light

Secondary production
Secondary production: oxidation energy produced by autotrophs resulting in increased biomass of heterotrophs i.e. growth/production of consumers Ecological efficiency: ratio of energy entering one trophic level to energy entering the previous level. E.g.
Ecological efficiency of 1 producer = b/a Ecological efficiency of 1 consumer = c/b Ecological efficiency of 2 consumer = d/c
3 consumer d 2 consumer c 1 consumer b 1 producer a sunlight

Secondary production
Energy efficiency generally about 10%
i.e. only 10% of the energy entering one trophic level flows into the next level 90% energy lost by:
Incomplete assimilation Respiration Heat production

Energy losses accumulate across successive trophic levels ! pyramidshaped distribution of energy - limiting number of trophic levels
planktivores (2 consumers) piscivores (3 consumers) zooplankton (1 consumers) phytoplankton

In lakes
Positive correlation between fish production (2 production) and primary production/phosphorus input May lead to change in dominant fish species Extremely high primary production (hypereutrophication) ! reduced oxygen ! reduced fish production

Chemical cycles
Chemical cycles: ecosystem models that describe the cycling of energy chemicals between components (reservoirs) Reservoirs living and non-living components Chemicals cycle through ecosystems and between reservoirs (cf. energy flow in trophic dynamic models)
use, transformation between various forms, movement, re-use (cf. energy flow)

Phosphorus cycle in water

Important component of ATP, DNA, RNA, phospholipids. Most limiting nutrient for primary producers in freshwater (Next most limiting - Nitrogen)
Most P not directly available to producers Available P not abundant

Forms, examples:
Total phosphorus (TP) Soluble reactive phosphorus (SRP)
Readily available for uptake by producers Including orthophosphate (PO43-)
Buffering system (similar to carbonate) Particulate inorganic phosphorus (PIP) - insouluble orthophosphate attached to particles

Chemicals exist in various forms (oxidised/reduced; soluble/ insoluble; available/unavailable, etc.) Movement of chemicals between reservoirs caused by living (biological) and non-living (physical, chemical) processes

Phosphorus cycle in water: cultural eutrophication

Eutrophication: nutrient enrichment of water body. Accompanied by: Increased 1 production Abundant phytoplankton (algal blooms) Poor light penetration ! turbid water due to phytoplankton Reduced photic zone ! upper epilimnion Oxygen depleted (anoxic) hypolimnion Cultural eutrophication: large scale nutrient enrichment caused by human activities. E.g., agriculture, farming, sewage disposal, detergents

Strong positive relationship between algal abundance and TP

Reversing cultural eutrophication:

Chemical reduction of algal blooms, e.g., copper sulphate Chemical removal of excess phosphate, e.g., alum (hydrated aluminum sulfate) Reduction of phosphate input Point sources (e.g, piped effluent) Non-point sources (e.g., runoff) Biomanipulation

But not so simple to reverse

Alternate stable state driven by: Sediments storing phosphate Biological interactions

Nutrient cycling in streams

Downstream transport in streams - little nutrient cycling in one place ! nutrient spiraling instead Spiraling Length (S) is the length of a stream required for a nutrient atom to complete one cycle
e.g. phytoplankton ! zooplankton ! planktivore ! piscivore ! decomposer

Nutrient cycling in streams

Spiraling Length: S = VT
S = Spiraling Length (one cycle) V = Average velocity of a nutrient atom T = Average time to complete a cycle

Nutrient retentiveness tendency for ecosystem to retain nutrients Short Spiraling Length = high nutrient retentiveness.
Nutrient cycled rapidly, numerous times, before being washed out of stream ecosystem

Related to rate of nutrient cycling and velocity of downstream nutrient movement

Long Spiraling Length = low nutrient retentiveness.

Nutrient cycled slowly, few times, before being washed out of stream ecosystem