Lecture from ENVT3363: Ecological Processes, third-year course at the University of Western Australia (UWA).
Lecture on long-term ecosystem development, patterns of plant diversity along soil chronosequences, and potential controls over plant diversity
Long-term ecosystem development and plant diversity
1. Long-term ecosystem development
and belowground controls over
terrestrial plant diversity
Etienne Laliberté
School of Plant Biology, UWA
ENVT3363 Ecological Processes
Sept 11, 2012
2. Organisms Climate Parent material Topography Time
Ecosystem
processes
Soils Soil abiotic Soil biotic
properties properties
Community
processes
Terrestrial plant
diversity
9. Maximum standing biomass
(‘climax’) does not persist
in the in the absence of
major disturbances:
• landslide
• glaciation
• volcanic eruption
Ecosystem decline or
retrogression
Wardle et al (2004) Science
12. Pedogenesis – Jurien Bay dunes
Ecosystem progression
Very young dune
(10’s—100’s years)
A
C
Very low N
High P
13. Pedogenesis – Jurien Bay dunes
Ecosystem progression
Very young dune Young dune
(10’s—100’s years) (~1000’s years)
A
A
C
C Highest N
Very low N High P
High P Peak fertility/productivity
14. Pedogenesis – Jurien Bay dunes
Ecosystem progression
Very young dune Young dune
(10’s—100’s years) (~1000’s years)
A
A
C
C Highest N
Very low N High P
High P Peak fertility/productivity
Ecosystem retrogression
Old dune
(~500,000 years)
A
low N
Ae low P
E
B1
B2
15. Pedogenesis – Jurien Bay dunes
Ecosystem progression
Very young dune Young dune
(10’s—100’s years) (~1000’s years)
A
A
C
C Highest N
Very low N High P
High P Peak fertility/productivity
Ecosystem retrogression
Old dune Very old dune
(~500,000 years) (>2,000,000 years)
A
low N O
Ae very low P A
E
B1 Ea
B2
E low N
extremely low P
‘terminal state’
16. Implications for Australia
Most
ecologists
work here Mt Michaud, Lesueur National Park
Productivity
Most of Australian
terrestrial ecosystems
are here
Soil age
18. Ancient soils, high plant diversity
Kwongan shrublands, SWA
Yasuní, Ecuador >70 species in 10x10-m plot
>1,100 tree species in 25-ha plot little dominance
weathered silty clay soils strongly leached sandy soils
Source: http://katerva.org
Valencia et al (2004) J Ecol Lamont et al (1977) Nature
19. Plant diversity along soil
chronosequences
Graham Zemunik
Laliberté et al (in preparation)
20. Nutrient availability and stoichiometry
Time
Pedogenic stage
Nutrient
availability and
stoichiometry
resource-ratio
model, productivity-
diversity (+/-)
Plant
diversity
21. ‘Humped-back’ model
• Low diversity at
high fertility
• Low diversity at
very low fertility
• Highest
diversity at
intermediate
fertility
Grime (1973) Nature
26. High diversity under strong P limitation
Strong P
N limitation Co-limitation Co-limitation P limitation limitation
Laliberté et al. (2012) J Ecol
27. Nutrient availability and stoichiometry
Time
Pedogenic stage
Nutrient • a role for productivity?
availability and • data inconsistent with resource-ratio model
stoichiometry
resource-ratio
model, productivity-
diversity (+/-)
Plant
diversity
28. Resource partitioning
Time
Pedogenic stage
Diversity
of N and Diversity of N and P forms
P forms
tend to increase in older soils
Plant
resource diversity
partitioning (+)
29. Nitrogen uptake and partitioning
Bever et al (2010) TREE
Hill et al (2011) Nature
Climate Change
30.
31. Phosphorus-acquisition strategies
P ‘miners’ = non-mycorrhizal/cluster roots
P ‘scavengers’ = AM fungi
Lambers et al (2008) Trends Ecol Evol
36. Soil spatial heterogeneity does not explain plant diversity
Smaller islands burn less often:
• last fire ~5000 years ago
• accumulate humus
• slower nutrient cycling
• lower productivity
• LOWER soil spatial heterogeneity
• HIGHER plant species richness
Arjeplog
island area
gradient,
Sweden
Gundale et al (2011) Ecography
37. Soil spatial heterogeneity
Time
Pedogenic stage
Soil spatial
Niche theory = classical
heterogeneity explanation, but does not
seem to actually be
important (at least in this
island system)
Plant
diversity
40. Mount St-Helens, USA
• volcanic eruption
1980
• high P, low N
• Lupinus lepidus = N2-
fixing legume
• Pathogens/herbivores
less abundant?
• Positive feedback =
high dominance?
Photo: John Bishop
42. Belowground heterotrophs
Time
Pedogenic stage
• Positive feedback may
Belowground
explain lower species heterotrophs
richness in young soils
• Negative feedback occurs
in old soils: a role for plant Plant
diversity
species coexistence?
• More data needed
43. Species pool hypothesis
Abiotic environmental
Time
conditions filtering (-)
Pedogenic stage Stage-
specific
species
pool size
Plant
diversity species pool
hypothesis (+)
46. Species pool hypothesis
Abiotic environmental
Time
conditions filtering (-)
Pedogenic stage Stage-
specific
species
pool size
Probably important in most systems
Plant
diversity species pool
hypothesis (+)
47. Multivariate controls over plant
diversity
Abiotic environmental
Organisms Climate Parent material Topography Time
conditions filtering (-)
time-area
Pedogenic stage Commonness hypothesis (+) Stage-
of habitat specific
species
pool size
Diversity Nutrient Soil spatial Belowground
of N and availability and heterogeneity heterotrophs
P forms stoichiometry
niche negative plant-
resource-ratio
theory (+) soil feedback (+)
model, productivity-
diversity (+/-)
Plant
resource diversity species pool
partitioning (+) hypothesis (+)
48. Conclusions
• Ecosystem ‘build-up’ followed by
ecosystem ‘decline
• Driven by loss of nutrients (e.g. P)
• Plant diversity often increases with soil
age
• Multivariate controls over plant
diversity:
– productivity
– resource partitioning (N and P forms)
– plant-soil feedback
– species pools
lecturer in plant biologysmall teaching loadi get to choose what i want to teachthe topic of thislecture is essentially the main theme of my current research
one of the key outcomes of this Unit is for you to gain a comprehensive understanding of ecosystems, including the links between its different aspectsthis lecture fits well with this outcome because it first starts with a description of ecosystem processes – how soils formand then explores the consequences this has on terrestrial plant diversity – community processesthe rationale is that you cannot understand how plant diversity changes during long-term ecosystem development if you do not have a basic understanding of soil and ecosystem development in the first place
How soils forms, and how plant communities respond to changes in soil conditions during soil formation, is one of the oldest and most-studied themes in plant ecologywhy? because it’s inherently interestingalso because it understanding how communities develop in time is the first step towards predictionHenry Cowles classic studies on vegetation succession along Lake Michigan dunesthis process is generally called ‘succession’
Equilibrium‘Self-perpetuating’
This view of a ‘climax’ or equilibrium also influenced ecosystem ecologyEugene Odum was one of the first and most influential ecosystem ecologistthis figure from his classic 1969 paper on ecosystem development in Science reinforces this view that ecosystems eventually reach a ‘climax’ or an ‘equilibrium’initially lots of resources or nutrients, therefore production rises rapidly (and so does respiration).but gross production rises more rapidly than respiration, therefore net production increases and biomass accumulateseventually resources get taken up so gross production declineshowever, as you accumulate biomass, you need to maintain that biomasstherefore net production starts to declinenot much biomass accumulates – peak standing biomass -> equilibrium or ‘climax’this was an important model, but it’s only dealing with relatively short-term changes – does the equilibrium state persists over longer time scales?
to answer questions like this you need soil age gradients much longer than what Lake Michigan can give youone of the most well-known system are the Hawaiian islandshot spot that leads to volcanic eruptionsas islands move away from the hotspot, volcanic eruption ceaseshave not been glaciated for a long time, no major disturbancetherefore you can find soils and ecosystems from very young (a few 100’s years) to very old (4 million years)Sequence has been widely used to improve our understanding of nutrient cycling and nutrient limitation in terrestrial ecosystems
you don’t need to go as far as Hawaii to find similar long-term soil age sequencesin fact you’re sitting on one right nowall across the Swan Coastal Plain you find systems of dune that range from very young to very old, around 2 million yearsthis is what I use in my current research
there are a number of other long-term sequences like this around the world, around 10in different biomes and climatesforest biomass -- does it follow Odum’s model?
causes are pretty clear and are due to nutrient limitationyou start with low nitrogen because that comes from the atmospherehowever you start with all the phosphorus you’ll ever have because phosphorus comes mostly from minerals