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Long-term ecosystem development
       and belowground controls over
          terrestrial plant diversity




Etienne Laliberté
School of Plant Biology, UWA
ENVT3363 Ecological Processes
Sept 11, 2012
Organisms   Climate    Parent material        Topography   Time




                                                             Ecosystem
                                                             processes
    Soils       Soil abiotic         Soil biotic
                properties           properties




                                                     Community
                                                     processes
                          Terrestrial plant
                              diversity
Vegetation succession on Lake Michigan dunes




                             Cowles (1899) Botanical Gazette
Classical vegetation succession model
                                         ‘Climax’




                    Johnson & Miyanishi (2008) Ecology Letters
Eugene P.
Odum
(1913-2002)




         Odum (1969) The strategy of ecosystem development. Science 164:262-270
Hawaiian 4.1 million-year island
          sequence




                         Crews et al. (1995) Ecology
Jurien Bay >2-million-year dune   Jurien
                                     Bay
           chronosequence
                                     Perth


0-7 ky


120-500 ky


         >2000 ky
Wardle et al (2004) Science
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
Long-term soil chronosequences
Build-up (progressive) phase      Maximal phase                   Decline (retrogressive) phase




                                     Soil age
                               Peltzer et al (2010) Ecol Monogr
What causes ecosystem decline?
          Total P
                    Total N




               Soil age




                              10 mg kg-1
Pedogenesis – Jurien Bay dunes
                     Ecosystem progression
 Very young dune
(10’s—100’s years)

A


C


        Very low N
        High P
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
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
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’
Implications for Australia


               Most
               ecologists
               work here          Mt Michaud, Lesueur National Park
Productivity




                                              Most of Australian
                                          terrestrial ecosystems
                                                         are here




                            Soil age
Plant strategies

Soil ‘available’ P     Leaf P concentration
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
Plant diversity along soil
   chronosequences




                   Graham Zemunik




                       Laliberté et al (in preparation)
Nutrient availability and stoichiometry


                                                   Time



                               Pedogenic stage




                Nutrient
            availability and
             stoichiometry

 resource-ratio
 model, productivity-
 diversity (+/-)
                                         Plant
                                       diversity
‘Humped-back’ model

             • Low diversity at
               high fertility
             • Low diversity at
               very low fertility
             • Highest
               diversity at
               intermediate
               fertility

                     Grime (1973) Nature
Jurien Bay

Fertility increases to a
peak around 1000’s years
and then declines in older
soils
High diversity at low productivity in old soils
  Low diversity at high
          productivity




Low diversity at
low productivity in young soils
Multiple resource limitation and
            diversity




         Harpole & Tilman (2007) Nature
Multiple resource limitation and
            diversity




         Harpole & Tilman (2007) Nature
High diversity under strong P limitation
                                                                        Strong P
        N limitation    Co-limitation   Co-limitation   P limitation   limitation




Laliberté et al. (2012) J Ecol
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
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 (+)
Nitrogen uptake and partitioning




                                            Bever et al (2010) TREE




                 Hill et al (2011) Nature
                          Climate Change
Phosphorus-acquisition strategies
                                        P ‘miners’ = non-mycorrhizal/cluster roots

    P ‘scavengers’ = AM fungi




Lambers et al (2008) Trends Ecol Evol
Turner (2008) J Ecol
Resource partitioning

                                                   Time



                           Pedogenic stage




Diversity
of N and
P forms                                        Perhaps, but no data yet!

                                     Plant
       resource                    diversity
     partitioning (+)
Soil spatial heterogeneity

                          Time



      Pedogenic stage




         Soil spatial
        heterogeneity




                Plant
              diversity
More niches, more species




homogeneous
soil conditions     calcrete
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
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
Belowground heterotrophs

                                         Time



      Pedogenic stage




                          Belowground
                          heterotrophs




                Plant
              diversity
Plant-soil feedback

                  Janzen-Connell
                      hypothesis




                  Host-specific pathogen
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
Barro Colorado Island, Panama




Photo: STRI




              Mangan et al (2010) Nature
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
Species pool hypothesis

                                   Abiotic        environmental
                        Time
                                  conditions        filtering (-)


    Pedogenic stage                              Stage-
                                                specific
                                                species
                                                pool size




              Plant
            diversity           species pool
                               hypothesis (+)
Siskiyou Mountains, Oregon, USA




Grace et al (2011) Ecology
Carbonate dunes
(Quindalup, stage 2: 100s-1000 years?)




       pH > 8
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 (+)
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 (+)
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
Honours, PhD?
etienne.laliberte@uwa.edu.au

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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
  • 3. Vegetation succession on Lake Michigan dunes Cowles (1899) Botanical Gazette
  • 4. Classical vegetation succession model ‘Climax’ Johnson & Miyanishi (2008) Ecology Letters
  • 5. Eugene P. Odum (1913-2002) Odum (1969) The strategy of ecosystem development. Science 164:262-270
  • 6. Hawaiian 4.1 million-year island sequence Crews et al. (1995) Ecology
  • 7. Jurien Bay >2-million-year dune Jurien Bay chronosequence Perth 0-7 ky 120-500 ky >2000 ky
  • 8. Wardle et al (2004) Science
  • 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
  • 10. Long-term soil chronosequences Build-up (progressive) phase Maximal phase Decline (retrogressive) phase Soil age Peltzer et al (2010) Ecol Monogr
  • 11. What causes ecosystem decline? Total P Total N Soil age 10 mg kg-1
  • 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
  • 17. Plant strategies Soil ‘available’ P Leaf P concentration
  • 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
  • 22. Jurien Bay Fertility increases to a peak around 1000’s years and then declines in older soils
  • 23. High diversity at low productivity in old soils Low diversity at high productivity Low diversity at low productivity in young soils
  • 24. Multiple resource limitation and diversity Harpole & Tilman (2007) Nature
  • 25. Multiple resource limitation and diversity Harpole & Tilman (2007) 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
  • 33. Resource partitioning Time Pedogenic stage Diversity of N and P forms Perhaps, but no data yet! Plant resource diversity partitioning (+)
  • 34. Soil spatial heterogeneity Time Pedogenic stage Soil spatial heterogeneity Plant diversity
  • 35. More niches, more species homogeneous soil conditions calcrete
  • 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
  • 38. Belowground heterotrophs Time Pedogenic stage Belowground heterotrophs Plant diversity
  • 39. Plant-soil feedback Janzen-Connell hypothesis Host-specific pathogen
  • 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
  • 41. Barro Colorado Island, Panama Photo: STRI Mangan et al (2010) Nature
  • 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 (+)
  • 44. Siskiyou Mountains, Oregon, USA Grace et al (2011) Ecology
  • 45. Carbonate dunes (Quindalup, stage 2: 100s-1000 years?) pH > 8
  • 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

Notes de l'éditeur

  1. 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
  2. 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
  3. 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’
  4. Equilibrium‘Self-perpetuating’
  5. 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?
  6. 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
  7. 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
  8. 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?
  9. 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