Unit II: Ecosystem Science (Ch 22—24)

We’ll start with limnology (lakes) and aquatic ecology

• Readings are spread out in several chapters:
18-19, 70-71, 104-110, 486-487, 516-525
• Limnology has always provided excellent
examples of integrated approaches to
ecology (recall our discussion of the two
approaches to ecology)
• Lake structure (shape, depth, location) is
closely linked to its functioning Lake Baikal
• Water chemistry is closely linked to biology
• How do we classify lakes? What controls
lake productivity? The answers involves
nutrients, lake shape and depth, and climate
as well as the type of organisms found in the
lake (particularly phytoplankton,
zooplankton and fish).
From Chapter 1: the ecosystem approach to ecology describes
organisms and their activities in terms of common “currencies’
(energy, water, carbon, nutrients).

The systems approach to ecology (box and arrow, pools and fluxes
diagrams).
One of the analytical / quantitative skills you need to learn in this
class.

Energy,
Carbon,
Nitrogen,
Mercury,
etc
 An ecosystem example
Boxes are pools (amounts of stuff) while arrows are
fluxes (rates of transfer). Rates always have units of
stuff per unit time, while boxes have units of stuff.
The diagrams assume “steady state”. At steady state
inputs = outputs for all pools. A system may be open or
closed

Residence times = pools / flux = stuff / stuff x time-1

= 1 / time-1 = time
A global diagram
Turnover rates = flux / pools = time -1
 The Global Water Cycle (Fig 23.4)
Calculating turnover rates and residence times if you know pools (compartments)
and fluxes (transfers): pay attention to the units (calculations also explained on
P.477). The atmosphere contain 13 TT (1018 grams) of water vapor on average.
Inputs each year are (71+425) = 496 TT/yr (496 TT yr-1). Your book has the same
numbers in Fig 23.4. Note that 1 TT = 1000km3 of water.

Residence times have units of

time (for example yrs).

Pool / flux = TT/(TT/yr) = yrs

13TT / (496 TT/yr) = 0.026

yr

0.026 yr x (365 days/1 yr) =

9.5 days
The residence time of water
vapor in the atmosphere is
about 9.5 days. The turnover
rate is 1/(residence time) =
1/0.026yr = 38 times per year
 What’s the residence time of water in global
rivers?
• What units do we want for the answer? Time
• Do we divide the pool (2000 km3) by the flux (45,500 km3/yr) or
the flux by the pool?
• Pool / flux = (2000 km3) / (45,500 km3/yr) = 0.0439 1/1/yr
• =0.0439 yr or 16 days

• If we had divided the flux by the pool, the answer would have had
units of 1/yr or yr-1. That would have been the turnover rate.
• 1 / 0.0439 yr = 22.78 per yr

• Remember to use only one set of fluxes (either inputs or outputs,

but not both) in these calculations
A lesson on
pools and
fluxes, Science,
8/26/2006

pools
fluxes
The Global Water Cycle from Oti and Kanae (Science, 2006)

What’s the residence time of water in global rivers?

 The global water cycle - global pools and
residence times
• Although the atmosphere and surface water (lakes and rivers)
are not particularly big pools of water globally, they are very
dynamic (short residence times). Note that these numbers don’t
match your text exactly - they’re just guestimates! Pay attention to the
qualitative (relative) differences.
• Atmosphere = 13,000 km3 of water, residence time = 8.9 days
• Oceans = 1,350,000,000 103 km3 , rt = 3,060 yrs
• Polar ice
– Arctic = 2,400,000 km3 , rt = 4500 yrs
– Antarctica = 22,000,000 km3 , rt = 14,000 yrs
• Continents
– Lakes and rivers = 231,000 km3, rt = 6.6yrs
– Groundwater = 8,200,000 103 km3 , rt = 600yrs
 Limnology
Some definitions:
– Limnology:
• The scientific study of bodies of fresh water for their biological,
physical and chemical properties (includes lakes, wetlands and
streams).
– Lake:
• a (considerable) inland body of standing water.
– Wetland:
• Lands on which water covers the soil or is present either at or near
the surface of the soil or within the root zone, all year or for varying
periods of time during the year, including during the growing season
Distribution of Water on Earth
 The Caspian and Black Seas are usually not
considered as fresh water (land-locked seas)

Lake Baikal
contains
about 20%
of the
earth’s
freshwater

The Great Lakes contain about The lakes of the east African rift
20% of the earth’s freshwater valley contain about 20% of the
earth’s freshwater (e.g. Lake
Tanganyika and Lake Victoria)
Most of the remaining freshwater lakes are in recently glaciated
regions of N. America, Europe and northern Asia
 Nebraska surface water
• 10th in nation in stream miles
• 16th in nation in wetland area
• About 2000 natural Sandhills lakes
• Dozens of reservoirs
• About 800 sandpit lakes along major rivers
• Oxbow lakes along the Missouri River
• example- Desoto Bend National Wildlife Refuge
north of Omaha
• This doesn’t count Nebraska’s massive reserve of
Niobrara River
groundwater. Nebraska contains 65% of the
groundwater stored in the High Plains Aquifer (the
main body of the ground water in the Great Plains,
covering parts of 8 states). Maintaining the size and
quality of that aquifer is a major natural resources
issue in the Great Plains, but one we won’t have time
to get into.
an oxbow lake
 Lake Formation
1. Tectonics 1.
2. Glaciation
3. Volcanic
4.
4. River meandering

2.
3.
 Lake Classification
• Chemical Classification
– Salt water
– Freshwater

• Classification based on Mixing patterns

• Trophic Classification:
– Oligotrophic – low productivity, low nutrients
– Eutrophic – high productivity, high nutrients
 Salt Creek Watershed
to the Platte River

So why do we have lots of

reservoirs in Lancaster
County?
 The Sandhills are the largest sand dune complex in the western hemisphere.
Today, they are almost completely stabilized by grass. Because of the high water
table, many of the valleys between dunes are filled with thousands of wetlands,
lakes or wet meadows.

Region of
alkali lakes
 Lakes and wetlands are dynamic (especially in the Sandhills)
UNL’s Barta Brothers Ranch
Late June
2010

one
mile

Late June
2008
Water Balance - the balance water inputs and outputs.
• Water balance determines lake size, seasonal water fluctuations and
chemistry.
• Water inputs include precipitation, streams, surface runoff and
groundwater (both seepage and springs).
• Water outputs include streams, seepage to groundwater and evaporation.
• In the west, many lakes have closed basins (the only output is
evaporation).

Line where annual evaporation generally

exceeds precipitation
 Great Salt Lake
• A good example of a closed basin lake
is the Great Salt Lake. Because the lake
has no surface water outflow, inputs of
salts and solids have collected and been
concentrated by evaporation over
hundreds of thousands of years, leaving
high concentrations of dissolved salts
and solids. The Great Salt Lake is 6
times saltier than seawater.

• So, second in importance to the

water budget of a lake (the balance
of water in and out) are the inputs
and outputs of stuff in that water
(nutrients, salts, sediments, etc).
In the Great Plains, low precipitation, high evaporation and soils combine to
create closed basin lakes in some areas. In the Sandhills, most lakes are
connected to groundwater and are fresh, but if a lake is perched above the water
table and/or sealed underneath from ground water (perhaps by a layer of clay) it
can become hypersaline. The white dried out lake bed below is largely salt, the
lake behind it is also salty. The types of organisms found in such lakes are very
different from fresh water lakes only a mile or so away .

Evaporation basin in
Sandhills. Photo by T.
Barrow from the
Sandhills Lake
Classification Project
(Headed by John Holz,
SNR/UNL)
Lake Zonation
 littoral zone - shallow zone with rooted
Lake vegetation (pondweeds, cattails,etc).
Zonation
limnetic zone - essentially the open water

benthic zone - the bottom of the lake, usually

covered with sediments

photic zone - the portion of the lake where light

can penetrate. The depth of this zone depends on
water clarity.

Fig 5.20 is a bit misleading: both the light

blue and the dark blue are part of the
limnetic zone. Only the area next to the
bottom is the benthic zone.
 Light and the Photic Zone

Photic
Zone

Secchi disk
Lake Structure – Vertical Zones

wind
Epilimnion - warm water, well mixed
with the atmosphere (O2 and CO2),
contains the photic zone.

Thermocline or Metalimnion- the zone

of rapid transition between warm upper
waters and cold deep waters. The mixing
of water driven by surface waves and
winds stops here.

Hypolimnion - deep cold water. In

eutrophic lakes, this layer may go anoxic
during the summer. Often rich in
nutrients because of proximity to the
sediments
Temperature and Density
 Seasonal stratification of temperate lakes (Fig 4.12)
• In the winter, water <4oc is at the top of the lake near the ice.

• In the spring, the lake warms from the top until the entire lake is approximately 4 oc. At
that time, all of the water in the lake has the same density and is easily mixed by wind and
wave disturbance.
• In the summer, the upper
water warms and a density
gradient develops that isolates
the shallow (epilimnion) and
deep (hypolimnion) waters
making the lake ‘stratified’.

• In the fall, the epilimnion

cools until it matches the
hypolimnion and the lake
‘turns over’ again

• Lakes that behave like this

Epilimnion are referred to as ‘Dimictic’
Thermocline
hypolimnion
Another text’s figure
showing stratification
and lake turnover
Not all lakes have
seasonal stratification
and turnover.

Arctic lakes (A) may

be uniformly cold;
tropical lakes (D) may
be uniformly warm.
(Fig. 24.17)
 Types of lake stratification
• Dimictic - turnovers twice a year, the typical temperate lake we just described
• Warm monomictic lakes - ice free conditions cause mixing throughout the winter,
but stratification occurs in the summer.
• Polymictic lakes - lakes too shallow to develop a thermocline in the temperate
zones (many shallow Sandhills lakes) or uniformly warm lakes in the tropics
• Meromictic lakes - permanently stratified lakes, the epilimnion never mixes with
the hypolimnion. The deep waters become permanently anoxic and chemically
different. For example they contain large quantities of dissolved methane and
hydrogen sulfide. In this case, a “chemocline” separates the upper and lower strata.
• Meromixis can happen if:

–the lake is very deep and protected from the wind

–Natural or human inputs of salts create a deep layer of
salty water that is denser and restricts mixing with fresh
open water.
–Deep lakes in tropical environments have stable
stratified water temps. Lake Tanganyika (see P.19) is
permanently stratified: only the top 200m mix, the
bottom 1200m do not.

Crawford Lake, Ontario

 Lake Nyos, Cameroon, Africa
The lake does not normally turnover, BUT on August 21, 1986 it
did for the first time in decades (or centuries). 1700 people died
within hours from the toxic cloud. In this case, volcanic activity
contributed to the high concentration of gases in the hypolimnion.
http://www.pbs.org/wnet/savageplanet/01volcano/01/indexmid.html
 Lake Classification
• Chemical Classification
– Salt water
– Freshwater

• Classification based on Mixing patterns:

– Dimictic,
– Monomictic
– Polymictic
– Meromictic

• Trophic Classification:
– Oligotrophic – low productivity, low nutrients
– Eutrophic – high productivity, high nutrients
 Lakes

 
• Lake communities
– Typical foodweb

                                 

                         

                                  

                          
 Lakes
• Lake communities
– Typical foodweb Littoral zone
– Primary producers Macrophytes
Water Lily

Coontail

Chara

Pondweed
 Lakes
• Lake communities
– Typical foodweb Limnetic zone
– Primary producers Phytoplankton

Diatoms Volvox Anabaena

 Lakes
• Lake communities
– Typical foodweb
– Primary producers Zooplankton
– Consumers

Daphnia Copepod Bosmina

 Lakes Chaoborus

• Lake communities
– Typical foodweb
– Primary producers
– Consumers
– Secondary
Walleye
consumers/predators

Lake Trout

Perch
 Lake Function and Productivity
• Productivity is primarily influenced by the
– balance of nutrient inputs and outputs (“nutrient loading”)
– lake shape and mixing pattern
– trophic dynamics (the types and numbers of different trophic levels)
– climate. Unlike terrestrial vegetation, where climate has a large direct impact on
productivity, the impact of climate on lake productivity is mainly its indirect effect
on mixing patterns. Cold northern lakes can be very productive and warm southern
lakes can be relatively unproductive under typical conditions.

• High productivity requires a productive phytoplankton community, which means ample

light, nutrients and CO2.

• High productivity at higher trophic levels (zooplankton and fish) requires ample food
and oxygen.

• Oligotrophic= low phytoplankton productivity, high oxygen availability

• Eutrophic = high phytoplankton productivity, high nutrient availability
 Lake morphology affects its productivity
*Shallow lakes tend to be warmer, but warm water holds less dissolved
oxygen then cold water - shallow lakes are more prone to anoxia.
*the nutrient rich sediments are closer to the photic zone
*a higher proportion of the lake is in the photic zone
 Nutrient budgets (rate of nutrient inputs)
*phosphorus (P) tends to limit productivity in most (but not all!)
freshwater lakes, while nitrogen (N) tends to limit productivity in most
temperate terrestrial ecosystems. (why? N is inherently leakier than P in
terrestrial ecosystems).
*a classic experiment in Ontario by David Schindler, together with other
observations and experiments, settled the argument over N vs P limitation
in temperate lakes. (see fig 24.24, Page 520)

N
addition
P had little
addition effect
made lake
eutrophic
Top down vs bottom up control of lake function
* We’ll cover this more later, but want to introduce it in this limnology section.
So far we’ve only emphasized nutrients and lake productivity.
* Each trophic level affects and is affected by the trophic levels above and below
it.
* zooplankton grazing of phytoplankton can greatly reduce the abundance and
types of algae and diatoms. The number of zooplankton is, in turn, reduced by
predation from small fish.

*the abundance of small fish

Big fish depends both on the amount of food
(zooplankton) available and on the
number and efficiency of bigger,
Small fish predatory fish.

* Thus, the type and abundance of

zooplankton fish can control lake ecology from
the top, while the nutrient loading
can control lake ecology from the
phytoplankton bottom.
Mid-summer in a stratified lake. Oxygen availability is
decreasing in the hypolimnion as heterotrophs continue to grow
on organic matter coming down from above, consuming oxygen.
Meanwhile, nutrients being regenerated in the sediments and
hypolimnion are unable to reach the photic zone, where the
phytoplankton are. If the lake is eutrophic, the hypolimnion goes
anoxic (runs out of oxygen). (P.520 - study this carefully)

Chemistry of hypolimnion
summer in a Wisconsin Lake
 Lakes
• Nutrients, Oxygen and
Fish Kills

Low nutrient
content
 Lakes
• Nutrients, Oxygen and
Fish Kills

Low nutrient
content

High nutrient
content
 Lakes
• Nutrients, Oxygen and
Fish Kills

Low nutrient
content

High nutrient
content
Summer
fish kills
 Lakes
• Nutrients, Oxygen and
Fish Kills

Low nutrient
content

Winter
fish kills

High nutrient
content
Summer
fish kills
During eutrophication from pollution, the added nutrients increase productivity
in the epilimnion, and much of the extra organic matter produced by the
phytoplankton drifts down into deep water. Here it accelerates the food chain
initially (until oxygen runs out). Fishkills may occur if much of the lake (the
hypolimnion) goes anoxic. This is especially true following pollution events
when high-oxygen requiring fish such as lake trout run out of oxygen. Carp are
much harder to kill (they can get by on much less oxygen)!

Oxygen will be
replenished in deep water
when this lake turns over
in the fall, but unless the
productivity is decreased,
it will probably go anoxic
again next summer.

Fishkill in a eutrophic Colorado lake

The hypoxic “dead zone” in the Gulf of Mexico (P.522)