Soil Organisms

• Classification schemes for organisms

• Basis for classification
• How classifications relate to each other
• What classification says about an
organism’s role in environmental
processes and vice versa
 How are life forms and their
activity in soil categorized?
• Physical identification of organisms, their
description and naming
• Biochemical characterization of organisms
and naming (microbes)
• Identification of soil processes and
organisms’ role(s) in these
 Categories

• Ecosystem function
– producers, consumers
• Trophic (physiological) group
– e.g., aerobe, anaerobe, denitrifiers, etc.
• Phylogenetic/taxonomic identification
– e.g., Proteobacteria, Pseudomonas, etc.
Ecosystem function

Organic material

Carbon dioxide, inorganic materials

 Ecological classification:
Producers
– Generate organic materials that consumers
and decomposers feed on
• Plants
• Green algae, Cynaobacteria (Blue-green algae)
• Mineral-oxidizing bacteria (lithotrophs)
 Ecological classification:
Consumers
– Feed directly on plants or on other
consumers
– Animals including
• Macrofauna (moles, snakes)
• Mesofauna (ants, worms)
• Microfauna (nematodes, protozoa)
 Ecological classification:
Decomposers
Microflora
• Bacteria (including actinomycetes)
• Fungi (including yeasts)
 Trophic categories
• Defined by:
– Source of carbon for cell synthesis
– mechanisms used to generate energy
• Define majority of transformations that
comprise element cycles
 Carbon source categories
Autotrophs
(Producers)

Carbon dioxide Organic carbon

Heterotrophs
(Consumers & Decomposers)
 Energy generation
• Energy harnessed during the movement
of electrons (e-)
– Requires a reduced e- donor & oxidized e-
acceptor
• Dissimilatory reaction
– Energy is extracted
– Materials are released (oxidized e- donor,
reduced e- acceptor)
Energy generation categories:
Electron donor types
• Organotroph
– Organic compounds
• Consumers and decomposers
• Invariably use organics for cell synthesis
Energy generation categories:
Electron donor types
• Lithotroph
– Inorganic compounds
• Producers
• Invariably use carbon dioxide for cell synthesis
Energy generation categories:
Electron donor types
• Phototroph
– Light
• Producers
• Invariably use CO2 for cell synthesis
 Trophic category summary
• Organoheterophs
– Often abbreviated to “heterotroph”
• Photoautotrophs
– Often abbreviated to “phototroph”
• Lithoautotrophs
– Often abbreviated to “autotroph”
Energy generation categories:
Electron acceptor types
• Oxygen
• Oxidized forms of:
– nitrogen
– sulfur
– metals
• Halogenated organic compounds
 Energy generation categories
based on electron acceptor use

• Aerobes
• Facultative anaerobes
• Anaerobes
 Aerobes
• Use of oxygen only
• “obligate” or “strict”
– Includes:
• Macro-, meso- , micro-fauna
• Microflora: most fungi, many bacteria including
all actinomycetes
 Facultative anaerobes
• Use O2 if available
• If O2 is absent, use alternate acceptor or
ferment
• Alternate electron acceptors include
– Nitrate (most common)
– Iron/manganese/sulfate (less common)
• Includes:
– Bacteria
– Eukaryotes (capable of limited fermentation)
 Anaerobes
• Unable to use O2 as electron acceptor
• Use:
– oxidized metals (Fe, Mn, others), S
– CO2
– Halogenated organics
• Lack protection against toxic oxygen species
• “obligate” or “strict”
• Include Bacteria and Archaea
 Soil organisms & ecosystem
function: Bottom-up
MAMMALS/
PREDATORY MITES BIRDS

PREDATORY PREDATORY PREDATORY

COLLEMBOLA COLLEMBOLA INSECTS
tertiary
PREDATORY
NEMATODES

secondary
COLLEMBOLA MITES NEMATODES PROTOZOANS OLIGOCHAETES

PLANT-FEEDING
NEMATODES FUNGI BACTERIA primary

PLANT ROOTS PLANT DEBRIS ROOT DEPOSITIONS

 Autotroph Heterotroph

Producer Producer Consumer Decomposer

Phototroph Lithotroph Organotroph Organotroph

Aerobe Aerobe Aerobe Aerobe

Bacteria Bacteria Eukarya Bacteria P
Eukarya Archaea Eukarya

Fac. Anerobe Fac. Anerobe Fac. Anerobe

Bacteria Bacteria Bacteria
Eukarya
Anaerobe Archaea
Anaerobe
Bacteria Bacteria
Archaea Anaerobe
Bacteria
Archaea
 Integrated functional and taxonomic classification road maps link food chain, trophic, and

biogeochemical functions with common and genomic names of biological community members

BIOLOGICAL COMMUNITY

Organisms producing organic C for biomass by reducing Organisms using organic C for biomass ( heterotrophs ) , and mineralizing it to

-
-
CO ( autotrophs ) with inorganic e donors ( lithotrophs )
2
CO as organic e donor ( organotrophs ) for chemical energy ( chemotrophs )
2

Organic Matter Producers CO Regenerators

2

Organisms using light as primary

Organisms using chemical

energy ( phototrophs ), and

energy ( chemototrophs ) with Food-ingesting

Predigested food-
supplemental chemical energy with
-
inorganic e donors ( lithotrophs ) organisms absorbing organisms
-
endogenous org or inorg e donors

PRIMARY CHEMO
MIXOTROPHS CONSUMERS DECOMPOSERS

PRODUCERS
PRODUCERS

1
Animals

Plants, Algae, Bacteria Bacteria Bacteria Fungi, Yeasts Actinomycetes , Bacteria

OBLIGATE AEROBES OBLIGATE AEROBES FACULTATIVE ANAEROBES OBLIGATE AEROBES OBLIGATE AEROBES

* Photosynthetic:
*H , CO oxidizers * Photo & nonphotosynthetic * Org C oxidizers * Org C oxidizers
2

Oxygenic Water oxidizers *O reducers. Bacteria: Acidovorax , Anoxygenic Org C, CO, H , *O reducers *O reducers. Actinomycetes . Bacteria:
2 2 2 2

0
* Nonphotosynthetic : Hydrogenophaga , Paracoccus , Sulfide, S oxidizers * Fungi. Eukarya : Actinomadura , Micromonospora ,

Org C oxidizers Variovorax * Nonphotosynthetic : * Oomycetes : Pythium , Nocardia , Rhodococcus , Streptomyces ,

0
*O reducers
* Sulfide, S oxidizers O reducers if O present; Phytophthora , Frankja
2
2 2

* Plants. Eukarya : *O reducers Org C , Heavy metal (Se, Saprolegnia * Org C, H oxidizers
2 2

2
* Green algae. Eukarya : * Bacteria : Beggiatoa , Bosea , Te) anion reducers if no O * Zygomycetes : *O reducers. Bacteria : Acidovorax ,
2 2

Chlamydomonas , Thiobacillus * Purple non S bacteria Glomus , Alcaligenes , Agrobacterium , Agromyces ,

Chlorella
* Archaea : Acidianus * Bacteria: Mucor , Rhizopus Arthrobacter , Cellulomonas , Cytophoga ,

* Diatoms. Eukarya : * Ferrous, Manganous oxidizers Rhodobacter , * Ascomycetes / Deinobacter , Flexibacter , Haloferax ,

* Cyanobacteria. Bacteria: *O reducers. Bacteria : Gallionella , Rhodopseudomonas , Deuteromycetes : Myxococcus , Paracoccus ,

2

Anabaena, Calothrix , Leptothrix , Planctomyces , Rhodospirillum , Acremonium , Pedomicrobium , Planctomyces ,

Gloeocapsa , Microcystis ,
Spharotilus , Thiobacillus Rhodovibrio Alternaria , Pseudomonas, Xanthomonas

Nostoc * Ammonium to nitrite oxidizers * Photosynthetic (only Aspergillus , Azoarcus , Azospirillum , Azotobacter ,

*O reducers. Bacteria : anaerobic) Blastomyces , Bradyrhizobium , Rhizobium

2

FACULTATIVE ANAEROBES Nitrosococcus , Nitrosomonas, Anoxygenic Org C oxidizers Chaetomium , * Org C oxidizers
1

3
* Photosynthetic (only anaerobic): Nitrosospira * Nonphotosynthetic : Cladosporium , * O reducers. Bacteria : Methylobacillus
2

0
Anoxygenic H , Sulfide, S * Nitrite oxidizers Org C oxidizers Coccidioides ,
2

oxidizers *O reducers. Bacteria : Nitrobacter , * O reducers if O present; Colletotrichum , FACULTATIVE ANAEROBES

2 2 2

* Nonphotosynthetic : Nitrospira Org C reducers if no O Fusarium , * Org C, H oxidizers

2 2

H , Org C oxidizers
2
* Filamentous green bact Histoplasma , * O reducers if O present;
2 2

* O reducers if O present; FACULTATIVE ANAEROBES * Bacteria: Paecilomyces , * Nitrate denitrifiers i f no O .

2 2 2

4
Org C reducers if no O *H oxidizers Chloroflexus , Penicillium , * Bacteria : Acidovorax , Bacillus,
2 2

* Purple S bacteria. Bacteria: * O reducers if O present; Heliothrix Rhizoctonia , Haloferax , Paracoccus

2 2

Chromatium , Thiocapsa
Nitrate denitrifiers if no O . Sclerotium , * Archaea : Haloferax
2

* Bacteria : Bacillus , Paracoccus OBLIGATE ANAEROBES Trichoderma * Nitrate denitrifiers , Ferric, Manganic , Org C

0
OBLIGATE ANAEROBES * Sulfide, S oxidizers * Photo & nonphotosynthetic * Basidiomycetes: reducers i f no O . Bacteria: Shewanella
2

* Photosynthetic: * O reducers if O present; Anoxygenic Org C oxidizers Mushrooms, Lignin * Org C oxidizers ± H producers
2 2 2

0
Anoxygenic H , Sulfide, S
Nitrate denitrifiers if no O . * Nonphotosynthetic degraders * O reducers if O present;
2
2 2 2

oxidizers * Bacteria : Thiobacillus , * Org C reducers * Yeasts. Eukarya : Nitrate, Org C reducers i f no O
2

5
* Nonphotosynthetic : Thiomicrospira * Bacteria with bchl g C andida, Rhodotorula * Bacteria : Erwinia , Escherichia,

H , Org C oxidizers * Bacteria: Pantoea , Proteus, Prosthecobacter ,

2

* Org C reducers
OBLIGATE ANAEROBES Heliobacillus FACULTATIVE ANAEROBES Serratia , Verrumicrobium ,

* Green S bacteria. Bacteria: * H oxidizers * Org C oxidizers Aeromonas , Bacillus, Klebsiella

2

0
Chorobium , Pelodictyon * Thiosulfate ,S reducers * O reducers if O present;
2 2

OBLIGATE ANAEROBES
* Bacteria : Desulfurococcus Org C reducers if no O
2

* Org C oxidizers ± H producers

2
* Archaea : Archeoglobus , * Yeasts. Eukarya :

* Org C, Org Cl , Org N, Nitrate, Proton

Pyrobaculum Saccharomyces
Color coding
reducers. Bacteria: Acidobacterium ,
* CO reducing methanogens
2
Green: C assimilators, and
Clostridium, Dehalobacter ,
* Archaea : Methanobacterium ,

Dehalococcoides , Dehalospirillum ,
N fixing organisms
2 Methanococcus , Methanosaeta ,

- Fibrobacter , Lactobacillus, Sarcina ,

Red: e donor oxidizers Methanosarcina ,

Clostridium
-
* CO reducing acetogens
Blue: e acceptor reducers 2
0
* Sulfate, S reducers
* Bacteria: Acetobacterium ,
6
* Bacteria Desulfotomaculum ,
Clostridium, Sporomusa

Desulfovibrio ,

* Org Cl reducers. Bacteria :

Desulfitobacterium , Desulfomile ,

Desulfuromonas ,

* Archaea : Archeoglobus , Pyrobaculum ,

1
Sulfolobus
Animals : OBLIGATE AEROBES, Org C oxidizers, O reducers. Eukarya .
2

* Org C oxidizers
Macrofauna : Groundhogs, Moles, Snakes
1

* Org C reducing methanogens

Mesofauna : Ants, Centipedes, Grubs, Millipedes, Mites, Spiders, Springtails, Worms
1ox

7
* Archaea : Methanobacterium ,
Microfauna : Nematodes:

Methanosaeta , Methanosarcina
Protozoa: Small flagellates, Naked amoebae, Slime molds - Dictyostelium , Physarum , Ciliates, Testacea

2
* Acetate splitting methanogens
Also, Acetobacter , Flavobacterium , Hyphomicrobium , Kocuria , Leptohtrix , Pirella , Sphaerotillus , Xanthomonas

3
* Archaea : Methanosarcina
Also, Methylobacter , Methylobacterium , Methylocystis , Methylomonas , Methylophaga , Methylosinus

4
Also, Blastobacter , Brachymonas , Bradyrhizobium , Halomonas , Hyphomicrobium , Jonesia , Roseobacter , Thiomicrospira

5
Also, Enterobacter ,

6
Also, Desulfobacterium , Desulfobulbus , Desulfohalobium , Desulfonatrum , Desulfurella , Desulfurococcus , Desulfuromonas

7
Also, Methanococcus , Methanocorpusculum , Methanoculleus , Methanospirillum ,
How might organo(hetero)trophic soil microbes obtain carbon?

Passive acquisition from by-products

Active
acquisition
 How might decomposers obtain
carbon from producers?
• Passivelly from dead biomass, excretions
and secretions
– Saprotrophs
• Actively from living biomass
– Necrotrophs, attack and decay biomass
– Biotrophs, obtain carbon from plant in
exchange for a microbe-supplied nutrient
 Saprotrophs
• Consume inanimate organic material
– Plant: litter, exudates
– Animal: bodies, excretions
– Microbial: cells, exudates
 Necrotrophs
• Plant pathogens
• Attack and decompose living plant parts
• Fungi (Pythium)
• Bacteria (Erwinia)
 Biotrophs
• Plant symbionts
• Bacteria = Nitrogen fixation
– Rhizobia (legumes)
– Actinorhiza (trees)
• Fungi = Phosphorus uptake
– Mycorrhizal fungi (woody and non-woody
plants)
 Learning Goals
• Classification schemes for organisms
• Basis for classification
• How classifications relate to each other
• What classification says about an
organism’s role in environmental
processes and vice versa
• What can we infer from element’s redox
transitions in biogeochemical cycles about
the microbial driving force underlying the
individual transformations?
– What do the microbes get out of reaction?
– What ecological/ tropic categories of would
mediate such a reaction?
Coupled Redox Reactions and Oxidation Numbers

•Dissimlatory processes: Oxidation of one element is coupled to reduction

of another.

•Recognizing which element is oxidized and which is reduced allows

identification of trophic classification.

•Knowing the trophic classification is key to understanding the microbiology

and biochemical basis of any transformation.

•The oxidation number indicates the relative oxidation state of an element and
can be used to identify electron donor/electron acceptor pairs
 Classification according to
growth characteristics
• Two general pools of organic carbon in soil

• Resistant to degradation
– Soil organic matter (humus, humic materials)
– Amounts are relatively constant

• Labile to degradation
– Fresh inputs(root exudates, soluble components released
from dead cells)
– Amounts are highly variable (spatially and temporally)

• Soil organisms may adapt to exploit one or the other of

these pools
 Zymogenous v. autochthonous
Winogradski (~1900)
•zymogenous: “ferment-producers”

•Attack readily available carbon sources

•Population densities fluctuate widely

•Large (1-3 um diam), spore-forming bacilli

•autochthonous: “self-feeders of the earth”, “humus

degraders”

•Attack more resistant materials

•Population densities are relatively stable.

• Small (1-1.5 um diam), cocci

 Oligotrophs and copiotrophs
(Sources?)
•Copiotrophs (aka, eutrophs): “abundant [resource]
feeders”.
•Activity adapted for the presence of abundant, readily-
available carbon sources.

•Oligotrophs: “scant [resource] feeders”.

•Activity adapted for the presence of low amounts of
readily-degradable carbon sources or compounds not
readily-degradable.
 Characteristics of an oligotroph
•Aerobic, but lack catalase and/or superoxid dismutase,
• lack full protection against toxic oxygen species

•High nutrient uptake capacity

• high surface/volume (S/V) ratio (small diameter)
• high density of transporter sites

•Efficient nutrient management to allow growth

• formation of reserve polymers (poly--hydroxybutyrate)
• biosynthesis regulation
Arthrobacter: a possible soil oligotroph

Morphology: coccoid cells (high S/V) under low nutrient

conditions, rod-shaped (lower S/V) under high nutrient levels.

Uptake systems favor organic acids, amino acids. Reduces

competition for sugars

Endogenous metabolism of starving cells 10% of that of

starving copiotrophs (e.g., E. coli, Pseudomonas)

Cells are resistant to dessication (% survival similar to Bacillus

endospores)
Morphology transition in Arthrobacter
 r- and K-strategists

• Mathematical model developed from studies on population

dynamics of animals and plants

• r = rate of population increase, K = carrying capacity of the

environment (carrying capacity = density of organisms that can
be supported

•K -strategists: optimized to function at high population sizes

(high K), but low growth rate (low r).

•r-selected: optimized to function at low population sizes (low

K), but high growth rates (high r)
 Organism type r value K value
r-strategist high low
K-strategist low high

r wins K wins
rI
competitive cross-over density
rate of
population rII
growth

KI KII
low population density
high
high resource density (availability)
low
r-strategist favored when resources are high relative to population
K-strategist favored when resources are low relative to population
Growth types: Equivalent terms

Zymogenous = copiotroph = r-strategist

Autochthonous = oligotroph = K-strategist

 Integrated functional and taxonomic classification road maps link food chain, trophic, and

biogeochemical functions with common and genomic names of biological community members

BIOLOGICAL COMMUNITY

Organisms producing organic C for biomass by reducing Organisms using organic C for biomass ( heterotrophs ) , and mineralizing it to

-
-
CO ( autotrophs ) with inorganic e donors ( lithotrophs )
2
CO as organic e donor ( organotrophs ) for chemical energy ( chemotrophs )
2

Organic Matter Producers CO Regenerators

2

Organisms using light as primary

Organisms using chemical

energy ( phototrophs ), and

energy ( chemototrophs ) with Food-ingesting

Predigested food-
supplemental chemical energy with
-
inorganic e donors ( lithotrophs ) organisms absorbing organisms
-
endogenous org or inorg e donors

PRIMARY CHEMO
MIXOTROPHS CONSUMERS DECOMPOSERS

PRODUCERS
PRODUCERS

1
Animals

Plants, Algae, Bacteria Bacteria Bacteria Fungi, Yeasts Actinomycetes , Bacteria

OBLIGATE AEROBES OBLIGATE AEROBES FACULTATIVE ANAEROBES OBLIGATE AEROBES OBLIGATE AEROBES

* Photosynthetic:
*H , CO oxidizers * Photo & nonphotosynthetic * Org C oxidizers * Org C oxidizers
2

Oxygenic Water oxidizers *O reducers. Bacteria: Acidovorax , Anoxygenic Org C, CO, H , *O reducers *O reducers. Actinomycetes . Bacteria:
2 2 2 2

0
* Nonphotosynthetic : Hydrogenophaga , Paracoccus , Sulfide, S oxidizers * Fungi. Eukarya : Actinomadura , Micromonospora ,

Org C oxidizers Variovorax * Nonphotosynthetic : * Oomycetes : Pythium , Nocardia , Rhodococcus , Streptomyces ,

0
*O reducers
* Sulfide, S oxidizers O reducers if O present; Phytophthora , Frankja
2
2 2

* Plants. Eukarya : *O reducers Org C , Heavy metal (Se, Saprolegnia * Org C, H oxidizers
2 2

2
* Green algae. Eukarya : * Bacteria : Beggiatoa , Bosea , Te) anion reducers if no O * Zygomycetes : *O reducers. Bacteria : Acidovorax ,
2 2

Chlamydomonas , Thiobacillus * Purple non S bacteria Glomus , Alcaligenes , Agrobacterium , Agromyces ,

Chlorella
* Archaea : Acidianus * Bacteria: Mucor , Rhizopus Arthrobacter , Cellulomonas , Cytophoga ,

* Diatoms. Eukarya : * Ferrous, Manganous oxidizers Rhodobacter , * Ascomycetes / Deinobacter , Flexibacter , Haloferax ,

* Cyanobacteria. Bacteria: *O reducers. Bacteria : Gallionella , Rhodopseudomonas , Deuteromycetes : Myxococcus , Paracoccus ,

2

Anabaena, Calothrix , Leptothrix , Planctomyces , Rhodospirillum , Acremonium , Pedomicrobium , Planctomyces ,

Gloeocapsa , Microcystis ,
Spharotilus , Thiobacillus Rhodovibrio Alternaria , Pseudomonas, Xanthomonas

Nostoc * Ammonium to nitrite oxidizers * Photosynthetic (only Aspergillus , Azoarcus , Azospirillum , Azotobacter ,

*O reducers. Bacteria : anaerobic) Blastomyces , Bradyrhizobium , Rhizobium

2

FACULTATIVE ANAEROBES Nitrosococcus , Nitrosomonas, Anoxygenic Org C oxidizers Chaetomium , * Org C oxidizers
1

3
* Photosynthetic (only anaerobic): Nitrosospira * Nonphotosynthetic : Cladosporium , * O reducers. Bacteria : Methylobacillus
2

0
Anoxygenic H , Sulfide, S * Nitrite oxidizers Org C oxidizers Coccidioides ,
2

oxidizers *O reducers. Bacteria : Nitrobacter , * O reducers if O present; Colletotrichum , FACULTATIVE ANAEROBES

2 2 2

* Nonphotosynthetic : Nitrospira Org C reducers if no O Fusarium , * Org C, H oxidizers

2 2

H , Org C oxidizers
2
* Filamentous green bact Histoplasma , * O reducers if O present;
2 2

* O reducers if O present; FACULTATIVE ANAEROBES * Bacteria: Paecilomyces , * Nitrate denitrifiers i f no O .

2 2 2

4
Org C reducers if no O *H oxidizers Chloroflexus , Penicillium , * Bacteria : Acidovorax , Bacillus,
2 2

* Purple S bacteria. Bacteria: * O reducers if O present; Heliothrix Rhizoctonia , Haloferax , Paracoccus

2 2

Chromatium , Thiocapsa
Nitrate denitrifiers if no O . Sclerotium , * Archaea : Haloferax
2

* Bacteria : Bacillus , Paracoccus OBLIGATE ANAEROBES Trichoderma * Nitrate denitrifiers , Ferric, Manganic , Org C

0
OBLIGATE ANAEROBES * Sulfide, S oxidizers * Photo & nonphotosynthetic * Basidiomycetes: reducers i f no O . Bacteria: Shewanella
2

* Photosynthetic: * O reducers if O present; Anoxygenic Org C oxidizers Mushrooms, Lignin * Org C oxidizers ± H producers
2 2 2

0
Anoxygenic H , Sulfide, S
Nitrate denitrifiers if no O . * Nonphotosynthetic degraders * O reducers if O present;
2
2 2 2

oxidizers * Bacteria : Thiobacillus , * Org C reducers * Yeasts. Eukarya : Nitrate, Org C reducers i f no O
2

5
* Nonphotosynthetic : Thiomicrospira * Bacteria with bchl g C andida, Rhodotorula * Bacteria : Erwinia , Escherichia,

H , Org C oxidizers * Bacteria: Pantoea , Proteus, Prosthecobacter ,

2

* Org C reducers
OBLIGATE ANAEROBES Heliobacillus FACULTATIVE ANAEROBES Serratia , Verrumicrobium ,

* Green S bacteria. Bacteria: * H oxidizers * Org C oxidizers Aeromonas , Bacillus, Klebsiella

2

0
Chorobium , Pelodictyon * Thiosulfate ,S reducers * O reducers if O present;
2 2

OBLIGATE ANAEROBES
* Bacteria : Desulfurococcus Org C reducers if no O
2

* Org C oxidizers ± H producers

2
* Archaea : Archeoglobus , * Yeasts. Eukarya :

* Org C, Org Cl , Org N, Nitrate, Proton

Pyrobaculum Saccharomyces
Color coding
reducers. Bacteria: Acidobacterium ,
* CO reducing methanogens
2
Green: C assimilators, and
Clostridium, Dehalobacter ,
* Archaea : Methanobacterium ,

Dehalococcoides , Dehalospirillum ,
N fixing organisms
2 Methanococcus , Methanosaeta ,

- Fibrobacter , Lactobacillus, Sarcina ,

Red: e donor oxidizers Methanosarcina ,

Clostridium
-
* CO reducing acetogens
Blue: e acceptor reducers 2
0
* Sulfate, S reducers
* Bacteria: Acetobacterium ,
6
* Bacteria Desulfotomaculum ,
Clostridium, Sporomusa

Desulfovibrio ,

* Org Cl reducers. Bacteria :

Desulfitobacterium , Desulfomile ,

Desulfuromonas ,

* Archaea : Archeoglobus , Pyrobaculum ,

1
Sulfolobus
Animals : OBLIGATE AEROBES, Org C oxidizers, O reducers. Eukarya .
2

* Org C oxidizers
Macrofauna : Groundhogs, Moles, Snakes
1

* Org C reducing methanogens

Mesofauna : Ants, Centipedes, Grubs, Millipedes, Mites, Spiders, Springtails, Worms
1ox

7
* Archaea : Methanobacterium ,
Microfauna : Nematodes:

Methanosaeta , Methanosarcina
Protozoa: Small flagellates, Naked amoebae, Slime molds - Dictyostelium , Physarum , Ciliates, Testacea

2
* Acetate splitting methanogens
Also, Acetobacter , Flavobacterium , Hyphomicrobium , Kocuria , Leptohtrix , Pirella , Sphaerotillus , Xanthomonas

3
* Archaea : Methanosarcina
Also, Methylobacter , Methylobacterium , Methylocystis , Methylomonas , Methylophaga , Methylosinus

4
Also, Blastobacter , Brachymonas , Bradyrhizobium , Halomonas , Hyphomicrobium , Jonesia , Roseobacter , Thiomicrospira

5
Also, Enterobacter ,

6
Also, Desulfobacterium , Desulfobulbus , Desulfohalobium , Desulfonatrum , Desulfurella , Desulfurococcus , Desulfuromonas

7
Also, Methanococcus , Methanocorpusculum , Methanoculleus , Methanospirillum ,