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5/27/2010



                                                                                                                           Soil macrofauna as ecosystems engineers
 Soil macrofauna Biodiversity, Soil structure and Organic
      Resource Management in East and West African                                                                                                                                             Earthworms and termites are important
                    cropping systems                                                                                                                                                         ecosystem engineers (Jones et al. 1994).
                                                                                                                                                                                               Their role in sustainable crop
Fredrick Ayukea*, b, Mirjam Pullemana, Lijbert Brussaarda, Johan Sixc, Bernard                                                                                                               production includes:
                                   Vanlauweb
                                                                                                                                                                                                   Improve soil structure & water
                                                                                                                                                                                                 retention
                                                                                                                                                                              Earthworms

                                                                                                                                                    Lavelle et al. 1998



                                                                                                                               Microbial
                                                                                                                                                  Soil water
                                                                                                                                                  content
                                                                                                                                                                              OM
                                                                                                                                                                          degradation             Incorporate organic material into soil
                                                                                                                               activities


                                                                                                                                                                              Improved             Improve soil fertility through organic
                                                                                                                                                                                                 matter decomposition and nutrient
                                                                                                                                                                              Water
                                                                                                                                                                              Infiltration
                                                                                                                                                                               rate

                                                                                                                                                                                                 release
           *a   Department of Soil Quality, Wageningen University, P.O Box 47, NL-6700 AA Wageningen, The Netherlands                                                         Nutrient
                                                                                                                                                                              uptake
                                                                                                                                Clay transfer &
                   b   Tropical Soil Biology and Fertility (TSBF) Institute of CIAT, P.O Box 30677-00100, Nairobi, Kenya        transformation
                                                                                                                                                                              N


                               c Department of Plant Sciences, University of California, Davis, CA 95616, USA              Termites (Courtesy of Soulemayne Konate)




                Objectives & Hypotheses                                                                                     Objectives continued:
This study aimed at: 1. Assessing effects of climate and agricultural
    management on earthworm and termite biodiversity across SSA                                                            2. Explore relations between soil macrofauna (earthworms
                                                                                                                              and termites), soil aggregation and SOM dynamics in
Hypotheses                                                                                                                    differently managed agro-ecosystems”

Biodiversity of earthworms and termites will:                                                                              Specifically:
                                                                                                                                  a) the extent to which soil macrofauna explains
1. Decrease with increasing temperature and decreasing                                                                       differences in aggregation across a wide range of climatic
    precipitation                                                                                                            and soil conditions in SSA, and
                                                                                                                                  b) compare these relationships between arable and
2. Be lower in agricultural than in fallow systems                                                                           fallow systems representing different levels of
                                                                                                                             management intensity.
3. Be higher under long-term agricultural management that had led
    to high-C soils than where it had led to low-C soils




                                 Materials & Methods                                                                                              Sub-humid AEZ                                                       Semi-arid AEZ
                                                                  Study sites
                                                                    Long-term trials across the
                                                                  sub-humid to semi-arid
                                                                  agroecological zones of East &                           Fallow
                                                                  West Africa
                                                                                                                                                            Shrubland                                 Bush               Shrubland

                                                                     The trials differed in management
                                                                  of:
                                                                       Organic resources
                                                                       Mineral N                                           High-C
                                                                       Rotation                                                                   T. candida-maize                              S. siamea+NPK Millet-cowpea
                                                                       Tillage




                                                                                                                           Low-C
                                                                                                                                                    Maize, no input                             Maize, no input         Millet, no input
                                                                                                                                                             KENYA                                 NIGERIA                   NIGER




                                                                                                                                                                                                                                            1
5/27/2010



                                           Results                                                                                                 Earthworms




                    Soil macrofauna biodiversity
                                                                                                                    Millsonia inermis, Saria, Burkina Faso

                                                                                                                                 20 earthworm taxa in 3 families:
                                                                                                                                      Ocnerodrilidae (4 taxa)
                                                                                                                                      Acanthodrilidae (8 taxa)
                                                                                                                                      Eudrilidae (8 taxa)




                                                                                                              Earthworm and termite taxonomic richness and functional groups
                                            Termites                                                          based on monolith and transect methods
                                                                                                                           Earthworm taxa                                              Termite taxa
                                                                                                                                                 Functional                                                Functional Food
                                                                                                              Taxonomic group                                 Taxonomic group
                                                                                                                                                 group a                                                   group b      type c
                                                                                                              Ocnerodrilidae                                  Rhinotermitidae-Rhinotermitinae
                                                                                                              Nematogenia lacuum                 Endogeic     Captotermes intermedias                      I            WLG
                                                                                                              Gordiodrilus robustus              Endogeic     Schedorhinotermes lamanianus                 I            WLG
                                                                                                              Gordiodrilus wemanus               Endogeic     Rhinotermitidae-Psammotermitinae
                                                                                                              Gordiodrilus marcusi               Endogeic     Psammotermes hybostoma                       I            WLG
                                                                                                              Acanthodrilidae                                 Termitidae-Nasutitermitinae
                                                                                                              Millsonia inermis                  Endogeic     Nasutitermes spec                            II           WLG
                                                                                                                                                 Endogeic     Trinevitermes spec                           II           LG
                                                20 termite taxa in two families:
                                                                                                              Millsonia guttata
                                                                                                              Dichogaster (Dt.) saliens          Epigeic      Termitidae-Macrotermitinae
                                                                                                              Dichogaster (Dt.) affinis          Epigeic      Ancistrotermes cavithorax                    II           WLG
                                                                                                                                                 Epigeic                                                   II           FWLG
                                                      Termitidae (17 taxa)
                                                                                                              Dichogaster (Dt.) bolaui                        Macrotermes nr. Vitrialatus
                                                                                                              Dichogaster (Dt.) modiglianii
                                                                                                              Di h      t (Dt )      di li ii    Epigeic
                                                                                                                                                 E i i        Macrotermes subhyalinus
                                                                                                                                                              M     t         bh li                        II           FWLG
                                                                                                              Dichogaster (Dt.) spec nov 1       Epigeic      Macrotermes herus                            II           FWLG
                                                                                                              Dichogaster (Dt.) spec nov 2       Epigeic      Microtermes pusillas                         II           FWLG
                                                      Rhinotermitidae (3 taxa)                                Eudrilidae
                                                                                                              Polytoreutus annulatus             Epigeic
                                                                                                                                                              Macrotermes spp.
                                                                                                                                                              Microtermes spp.
                                                                                                                                                                                                           II
                                                                                                                                                                                                           II
                                                                                                                                                                                                                        FWLG
                                                                                                                                                                                                                        FWLG
                                                                                                              Hyperiodrilus africanus            Epigeic      Odontotermes magdalenae                      II           FWLG
                                                                                                              Hyperiodrilus spec nov             Epigeic      Odontotermes spp.                            II           FWLG
                                                                                                              Eudrilus buettneri                 Epigeic      Pseudacanthotermes spiniger                  II           FLG
                                                                                                              Ephyriodrilus afroccidentalis      Epigeic      Pseudacanthotermes spp                       II           FLSD
                                                                                                              Eminoscolex violaceus              Epigeic      Termitidae-Termitinae
                                                                                                              Stuhlamannia spec nov              Epigeic      Amitermes-Amitermes stephensoni              II           WLSD
M. bellicosus, Tamale (Ghana)                                                                                 Lavellea spec nov                  Epigeic      Microcerotermes parvulus                     II           WLG
                                                                                                                                                              Termes-Termes baculi                         III          WS
                                                                                                                                                              Cubitermes- Tubeculitermes spec              IV           S
                                                                                                              a
                                                                                                                Based on classification by Swift & Bignell (2001). (W=wood; L=leaf litter; S=soil; D=Dung/manure; F=fungus
                                                                                                              grower; G=dead/dry grass. bBased on classification by Eggleton et al. (2002). c based on field notes/observation.
                                                                                                              c
                                                                                                                based on observations by Kooyman & Onck (1987).




RDA biplot on earthworm species abundance and environmental variables                                       RDA biplot on termite species abundance and environmental variables




    Axis 1: temperature/altitudinal gradient
                                                                                                                                                              Axis 1: precipitation/latitudinal gradient
    Axis 2: Soil C/latitudinal gradient
                                                                                                                                                              Axis 2: temperature/texture/ longitudinal gradient.
    Axes partition sites that are cooler and are at high altitudes from lower relatively hotter altitudes
                                                                                                                                                              More taxa occur on warmer/drier sides
    Earthworm taxa are less abundant on the higher, cooler altitudes characteristic of East African sites
                                                                                                                                                              Axes mainly separate East & West Africa.




                                                                                                                                                                                                                                  2
5/27/2010




Shannon diversity indices across fallow and arable systems

                                       Treatment

                                          Arable system
                                                                                      Soil Macrofauna and ecosystem functioning

                            Fallow      High-C     Low-C          P-value

 Earthworm diversity        0.13a       0.07b      0.02c          <0.001
 (Hâ€Č)
 Termite diversity (Hâ€Č)     0.08a       0.06a      0.03b          0.008




Factor analysis and factor pattern after varimax rotation
                                                                                 Possible attributes: factor & multiple regression analysis
                                               Treatment
                                                                                                                 Fallow                               Arable system
                                Fallow system
                                                                                                   Factor I          Factor II              Factor I          Factor II
                                    (n=36)             Arable system (n=72)                                                                                   Termite
Variable                    Factor I Factor II         Factor I    Factor II                       SOM,                                                       (abundance &
Termite Shannon index         −0.1         0.5          −0.1          0.2                          Texture,                                                   biomass),
Earthworm Shannon index        0.5        −0.4          −0.1          0.7                          Climate,         Termite                                   Earthworm
Termite abundance              0.2         0.6           0.2          0.7                          Earthworm        (Shannon                                  (Shannon
                                                                                                   (Shannon         diversity,                                diversity,
Earthworm abundance            0.6        −0.3           0.0          0.7        Possible          diversity &      abundance &             SOM, Texture,     abundance and
Termite biomass                0.3         0.7           0.3          0.6        attributes        abundance)       biomass)                Climate           biomass)
Earthworm biomass             −0.1       −0. 8          −0.2          0.6
Average t
A        temperature
                t             −0.9
                                09          02
                                          −0.2          −0.9
                                                          09            01
                                                                      −0.1
Precipitation                  0.7         0.1           0.7           0.0       Multiple regression analysis with aggregate fractions and factors with loading coefficients >0.05
Total organic C                1.0         0.1           1.0           0.0       (%variance accounted for by factors)
                                                                                 Total
Total organic N                1.0         0.1           1.0           0.0       macroaggregates
Clay                           0.9         0.3           0.9          −0.1       (sand corrected)       88.5              4.1                       90.4                   7.6
Sand                          −0.9        −0.1          −0.9          0.1        Microaggregates
                                                                                 within
 Factor eigenvalues            5.6        2.0              5.1             2.4   macroaggregates
                                                                                 (mM)                   82.6              4.9                       91.3                   8.7
 Explained variance (%)       45.9       17.8              42.6           20.0




 Conclusions                                                                      Conclusions continued:
                                                                                   Macrofauna, especially earthworms, and to a lesser extent
     Differences in climate coincide with differences in soil
     types and geographical region. These factors are                              termites, are important drivers of stable soil aggregation, in
     associated with earthworm and termite diversity                               conjunction with climate, soil organic C content and soil
                                                                                   texture in Sub-Saharan agroecosystems
     Agriculture has negative effects on earthworm and
     termite diversity as compared to long-term fallow                             The beneficial impact of earthworms and termites on soil
                                                                                   aggregation is reduced with increasing management intensity
     Under continuous crop production, agricultural                                (e.g. soil disturbance due to cultivation)
     management that resulted in low-C soils had lower
     earthworm and termite diversity than agricultural
     management that had resulted in high-C soils                                  Our results are important for designing agricultural
                                                                                   management systems aimed at increasing long-term soil
                                                                                   fertility in Sub-Saharan Africa.




                                                                                                                                                                                     3
5/27/2010



Recommendations from this study:                                    General recommendations:

The agroecological conditions studied, i.e:                        As mineral and organic fertilizers are often limited in quantity and quality,
                                                                   soil fertility research has to focus on developing integrated management
  Long-term application of manure in combination with fertilizer   strategies to address soil fertility decline (Integrated Soil Fertility
  Conservational tillage plus maize stover residue application;    Management (ISFM) and Conservation Agriculture (CA))
  Hand-hoeing plus manure
affect faunal activities in different ways. Hence:                 As the soil biota are responsible for the key ecosystem functions of
                                                                   decomposition and nutrient cycling, soil organic matter synthesis and
recommendation of these practices should be tailored to            mineralization, soil structural modification, aggregate stabilization, ISFM
                                                                                                       f                                   S
meet the circumstances of target farmers                           & CA have to maximize beneficial soil biota.

                                                                   Although there is pressure on land in many parts of Sub-Saharan Africa
                                                                   fallowing (or similar practices) has to be an integral part of land
                                                                   management in view of the conservation of biodiversity for enhanced
                                                                   ecosystem functioning




                                                                                  Acknowledgements
                                                                   WOTRO/NWO through Wageningen University for awarding the
                                                                   Scholarship and Research funds and Norman E. Borlaug LEAP
                                                                   through UC-Davis and the International Atomic Energy Agency
                                                                   (IAEA) for co-funding the research

                                                                   Collaborating Institutions
                                                                       TSBF-CIAT, KEFRI, Kenyatta University (Nairobi, Kenya)
                                                                       INERA,
                                                                       INERA IFDC (Burkina Faso)
                                                                       Others: IITA-Nigeria, SARI-Ghana, ICRISAT-Niger, KARI-Kenya,
                                                                       CHITEDZE Agricultural Inst.
                      It was a teamwork
                                                                   for cooperating during periods of data collection
                                                                   Special thanks to all my promoters and in particular Prof. Lijbert
                                                                   Brussaard for sending me off on the long journey.




                                                                                                                                               4

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PK12:Soil macrofauna Biodiversity, Soil structure and Organic Resource Management in East and West African cropping systems

  • 1. 5/27/2010 Soil macrofauna as ecosystems engineers Soil macrofauna Biodiversity, Soil structure and Organic Resource Management in East and West African Earthworms and termites are important cropping systems ecosystem engineers (Jones et al. 1994). Their role in sustainable crop Fredrick Ayukea*, b, Mirjam Pullemana, Lijbert Brussaarda, Johan Sixc, Bernard production includes: Vanlauweb Improve soil structure & water retention Earthworms Lavelle et al. 1998 Microbial Soil water content OM degradation Incorporate organic material into soil activities Improved Improve soil fertility through organic matter decomposition and nutrient Water Infiltration rate release *a Department of Soil Quality, Wageningen University, P.O Box 47, NL-6700 AA Wageningen, The Netherlands Nutrient uptake Clay transfer & b Tropical Soil Biology and Fertility (TSBF) Institute of CIAT, P.O Box 30677-00100, Nairobi, Kenya transformation N c Department of Plant Sciences, University of California, Davis, CA 95616, USA Termites (Courtesy of Soulemayne Konate) Objectives & Hypotheses Objectives continued: This study aimed at: 1. Assessing effects of climate and agricultural management on earthworm and termite biodiversity across SSA 2. Explore relations between soil macrofauna (earthworms and termites), soil aggregation and SOM dynamics in Hypotheses differently managed agro-ecosystems” Biodiversity of earthworms and termites will: Specifically: a) the extent to which soil macrofauna explains 1. Decrease with increasing temperature and decreasing differences in aggregation across a wide range of climatic precipitation and soil conditions in SSA, and b) compare these relationships between arable and 2. Be lower in agricultural than in fallow systems fallow systems representing different levels of management intensity. 3. Be higher under long-term agricultural management that had led to high-C soils than where it had led to low-C soils Materials & Methods Sub-humid AEZ Semi-arid AEZ Study sites Long-term trials across the sub-humid to semi-arid agroecological zones of East & Fallow West Africa Shrubland Bush Shrubland The trials differed in management of: Organic resources Mineral N High-C Rotation T. candida-maize S. siamea+NPK Millet-cowpea Tillage Low-C Maize, no input Maize, no input Millet, no input KENYA NIGERIA NIGER 1
  • 2. 5/27/2010 Results Earthworms Soil macrofauna biodiversity Millsonia inermis, Saria, Burkina Faso 20 earthworm taxa in 3 families: Ocnerodrilidae (4 taxa) Acanthodrilidae (8 taxa) Eudrilidae (8 taxa) Earthworm and termite taxonomic richness and functional groups Termites based on monolith and transect methods Earthworm taxa Termite taxa Functional Functional Food Taxonomic group Taxonomic group group a group b type c Ocnerodrilidae Rhinotermitidae-Rhinotermitinae Nematogenia lacuum Endogeic Captotermes intermedias I WLG Gordiodrilus robustus Endogeic Schedorhinotermes lamanianus I WLG Gordiodrilus wemanus Endogeic Rhinotermitidae-Psammotermitinae Gordiodrilus marcusi Endogeic Psammotermes hybostoma I WLG Acanthodrilidae Termitidae-Nasutitermitinae Millsonia inermis Endogeic Nasutitermes spec II WLG Endogeic Trinevitermes spec II LG 20 termite taxa in two families: Millsonia guttata Dichogaster (Dt.) saliens Epigeic Termitidae-Macrotermitinae Dichogaster (Dt.) affinis Epigeic Ancistrotermes cavithorax II WLG Epigeic II FWLG Termitidae (17 taxa) Dichogaster (Dt.) bolaui Macrotermes nr. Vitrialatus Dichogaster (Dt.) modiglianii Di h t (Dt ) di li ii Epigeic E i i Macrotermes subhyalinus M t bh li II FWLG Dichogaster (Dt.) spec nov 1 Epigeic Macrotermes herus II FWLG Dichogaster (Dt.) spec nov 2 Epigeic Microtermes pusillas II FWLG Rhinotermitidae (3 taxa) Eudrilidae Polytoreutus annulatus Epigeic Macrotermes spp. Microtermes spp. II II FWLG FWLG Hyperiodrilus africanus Epigeic Odontotermes magdalenae II FWLG Hyperiodrilus spec nov Epigeic Odontotermes spp. II FWLG Eudrilus buettneri Epigeic Pseudacanthotermes spiniger II FLG Ephyriodrilus afroccidentalis Epigeic Pseudacanthotermes spp II FLSD Eminoscolex violaceus Epigeic Termitidae-Termitinae Stuhlamannia spec nov Epigeic Amitermes-Amitermes stephensoni II WLSD M. bellicosus, Tamale (Ghana) Lavellea spec nov Epigeic Microcerotermes parvulus II WLG Termes-Termes baculi III WS Cubitermes- Tubeculitermes spec IV S a Based on classification by Swift & Bignell (2001). (W=wood; L=leaf litter; S=soil; D=Dung/manure; F=fungus grower; G=dead/dry grass. bBased on classification by Eggleton et al. (2002). c based on field notes/observation. c based on observations by Kooyman & Onck (1987). RDA biplot on earthworm species abundance and environmental variables RDA biplot on termite species abundance and environmental variables Axis 1: temperature/altitudinal gradient Axis 1: precipitation/latitudinal gradient Axis 2: Soil C/latitudinal gradient Axis 2: temperature/texture/ longitudinal gradient. Axes partition sites that are cooler and are at high altitudes from lower relatively hotter altitudes More taxa occur on warmer/drier sides Earthworm taxa are less abundant on the higher, cooler altitudes characteristic of East African sites Axes mainly separate East & West Africa. 2
  • 3. 5/27/2010 Shannon diversity indices across fallow and arable systems Treatment Arable system Soil Macrofauna and ecosystem functioning Fallow High-C Low-C P-value Earthworm diversity 0.13a 0.07b 0.02c <0.001 (Hâ€Č) Termite diversity (Hâ€Č) 0.08a 0.06a 0.03b 0.008 Factor analysis and factor pattern after varimax rotation Possible attributes: factor & multiple regression analysis Treatment Fallow Arable system Fallow system Factor I Factor II Factor I Factor II (n=36) Arable system (n=72) Termite Variable Factor I Factor II Factor I Factor II SOM, (abundance & Termite Shannon index −0.1 0.5 −0.1 0.2 Texture, biomass), Earthworm Shannon index 0.5 −0.4 −0.1 0.7 Climate, Termite Earthworm Termite abundance 0.2 0.6 0.2 0.7 Earthworm (Shannon (Shannon (Shannon diversity, diversity, Earthworm abundance 0.6 −0.3 0.0 0.7 Possible diversity & abundance & SOM, Texture, abundance and Termite biomass 0.3 0.7 0.3 0.6 attributes abundance) biomass) Climate biomass) Earthworm biomass −0.1 −0. 8 −0.2 0.6 Average t A temperature t −0.9 09 02 −0.2 −0.9 09 01 −0.1 Precipitation 0.7 0.1 0.7 0.0 Multiple regression analysis with aggregate fractions and factors with loading coefficients >0.05 Total organic C 1.0 0.1 1.0 0.0 (%variance accounted for by factors) Total Total organic N 1.0 0.1 1.0 0.0 macroaggregates Clay 0.9 0.3 0.9 −0.1 (sand corrected) 88.5 4.1 90.4 7.6 Sand −0.9 −0.1 −0.9 0.1 Microaggregates within Factor eigenvalues 5.6 2.0 5.1 2.4 macroaggregates (mM) 82.6 4.9 91.3 8.7 Explained variance (%) 45.9 17.8 42.6 20.0 Conclusions Conclusions continued: Macrofauna, especially earthworms, and to a lesser extent Differences in climate coincide with differences in soil types and geographical region. These factors are termites, are important drivers of stable soil aggregation, in associated with earthworm and termite diversity conjunction with climate, soil organic C content and soil texture in Sub-Saharan agroecosystems Agriculture has negative effects on earthworm and termite diversity as compared to long-term fallow The beneficial impact of earthworms and termites on soil aggregation is reduced with increasing management intensity Under continuous crop production, agricultural (e.g. soil disturbance due to cultivation) management that resulted in low-C soils had lower earthworm and termite diversity than agricultural management that had resulted in high-C soils Our results are important for designing agricultural management systems aimed at increasing long-term soil fertility in Sub-Saharan Africa. 3
  • 4. 5/27/2010 Recommendations from this study: General recommendations: The agroecological conditions studied, i.e: As mineral and organic fertilizers are often limited in quantity and quality, soil fertility research has to focus on developing integrated management Long-term application of manure in combination with fertilizer strategies to address soil fertility decline (Integrated Soil Fertility Conservational tillage plus maize stover residue application; Management (ISFM) and Conservation Agriculture (CA)) Hand-hoeing plus manure affect faunal activities in different ways. Hence: As the soil biota are responsible for the key ecosystem functions of decomposition and nutrient cycling, soil organic matter synthesis and recommendation of these practices should be tailored to mineralization, soil structural modification, aggregate stabilization, ISFM f S meet the circumstances of target farmers & CA have to maximize beneficial soil biota. Although there is pressure on land in many parts of Sub-Saharan Africa fallowing (or similar practices) has to be an integral part of land management in view of the conservation of biodiversity for enhanced ecosystem functioning Acknowledgements WOTRO/NWO through Wageningen University for awarding the Scholarship and Research funds and Norman E. Borlaug LEAP through UC-Davis and the International Atomic Energy Agency (IAEA) for co-funding the research Collaborating Institutions TSBF-CIAT, KEFRI, Kenyatta University (Nairobi, Kenya) INERA, INERA IFDC (Burkina Faso) Others: IITA-Nigeria, SARI-Ghana, ICRISAT-Niger, KARI-Kenya, CHITEDZE Agricultural Inst. It was a teamwork for cooperating during periods of data collection Special thanks to all my promoters and in particular Prof. Lijbert Brussaard for sending me off on the long journey. 4