Dr. Stephen Malcolm, Department of Biological Sciences
• Week 9: Decomposers,
Detritivores & Mutualists.
• Lecture summary:
• Decomposition &
detritivory:
• Examples & resources.
• Comparisons.
• Model of detritivory.
• Mutualism:
• Non-symbiotic.
• Symbiotic.
Slide - 1 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Decomposers are saprobes like bacteria and
fungi that feed on dead or dying plant and
animal tissues.
• Detritivores feed on the same material once
it has been fragmented and processed to
varying extents by both these
decomposers and physical events.
• Interactions tend to be very general.
• Taxonomic origin usually unimportant.
• Result in release of nutrients (Fig. 11.2).
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 2
• 1. Dead bodies of animals:
• carrion
(Fig. 11.18)
• 2. Feces & other excreted products
(Fig. 11.15)
• Australia was nearly covered with sheep/cow feces because of a lack of dung beetles!
• 3. Dead plant material:
• Trees, roots, stems, leaves as standing material
• Litter, and ripe fruit separated from the parent
• Fig. 11.11.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 3
• Resistance of resources to decomposition
increases in the order:
• sugars < starch < hemicelluloses < pectins and proteins < cellulose < lignins < suberins < cutins
• Shown partially in Fig.11.2
for 2 different ecosystems.
• Cellulose is difficult to break down:
• Cellulose catabolism (cellulolysis) requires cellulase
enzymes which most animals don’t have:
• 1 cockroach & a few termites.
• Complex mechanisms have evolved as in Fig. 11.12.
Slide - 4 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Although predators and herbivores also eat
dead food after they have caught and
killed it, the primary distinction between
these consumers and decomposers/
detritivores is that the latter do not affect
the rate at which their resources are
produced, but of course predators and
herbivores do.
• In addition, while mutualists may increase
resource availability, decomposers and
detritivores do not have an influence.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 5
• Represent resource ( R ) renewal as F(R).
• P as the number of predators.
•
• a as the efficiency with which individuals find and capture
their food resource.
For exploiters , such as predators, herbivores and parasites , the rate of resource renewal dR/dt is: dR/dt = F(R) - aP
• for mutualists , where M is the number of mutualists and
is a measure of mutual benefit dR/dt is:
δ dR/dt = F(R) + δ M
• for decomposers and detritivores that have no influence
on resource renewal, dR/dt is: dR/dt = F(R)
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 6
• Detritivores and microbivores (tiny
detritivores that feed on bacteria and
fungi rather than larger particulate
detritus - but their food is often alive!).
• Taxonomically diverse and can be classified
by size from:
• microfauna (<100 µ m) through,
• mesofauna (100 µ m-2mm) to,
• macrofauna (2-20 mm) (Fig. 11.3).
Slide - 7 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• The relative distribution of micro- meso- and
macro-detritivores among biomes related to
temperature, rainfall and latitude is shown in
Fig 11.4
:
• Most macrofauna in tropics.
• Most microfauna in cold regions.
• Mesofauna dominant in temperate zones.
• Darwin (1888) estimated that earthworms near his
house formed new soil layers at the rate of 18
cm/30 years and bring up 5.1Kg soil/m 2 .
Slide - 8 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
9. Diversity & abundance of detritivores:
• In 1m 2 of temperate woodland soil there could be:
• 10 million nematodes and protozoans.
• 100,000 springtails (Collembola) and mites
(Acari).
• 50,000 other invertebrates.
• In woodlands, microbial decomposition is
highest (Fig. 11.7), but larger detritivores can
enhance microbial respiration and so the
different species function as a connected
community (Fig. 11.8).
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 9
10. Diversity & abundance of detritivores:
• In freshwater ecosystems, detritivores are
also diverse.
• Different “guilds” according to feeding
methods:
• “shredders”, “collecto-gatherers”, “grazer-
scrapers”, and “collecto-filterers” (Fig. 11.5).
• Together this community breaks down detritus in
a stream (Fig. 11.6).
Slide - 10 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Mutualism is an interaction in which both
partners benefit:
• “ … individuals in a population of each
mutualist species grow, survive and
reproduce at higher rates when in presence
of individuals of the other species”
• Note: it is not a “cosy” relationship - each
species acts completely selfishly.
Slide - 11 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Symbiosis just means "living together" in
close association (excluding parasitism).
• Mutualism is a special kind of symbiosis,
but mutualists don’t have to be symbionts
to benefit each other.
• So mutualisms can be either symbiotic or non-symbiotic.
Slide - 12 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Most of the world's biomass is made of
mutualists:
• Most plants, coral reefs, pollinators.
• Nonsymbiotic mutualisms are common:
• E.g.
cleaner fish, ants tending aphids, or pollinator-flower interactions.
• Symbiotic mutualisms also common:
• Include fungus-alga associations in lichens
(Fig. 13.21), fungus-plant associations in mycorrhizae, or animal-alga associations such as the flatworm Convoluta roscoffensis .
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 13
• Bull's horn acacia and
Pseudomyrmex ants:
• Figs. 13.2 & 13.3 (4 th )
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 14
• shrimps and gobiid fish
• Fig. 13.3 (3 rd )
• clown fish & anemones
• cleaner fish & customers
• honey guide and ratel
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 15
• Defense mutualisms:
• E.g.
Müllerian mimicry in heliconiid butterflies:
• Eltringham (1916), from cover of
Futuyma & Slatkin
(1983) Coevolution .
• Group defense:
• E.g.
musk oxen or sawflies.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 16
• Are domestic crops and domestic animals
examples of mutualisms with man?
• Is your dog or cat a mutualist?
• If there are many more of a species than
there would have been without the
association it must be a mutualism!
• “Farming” also occurs in termites and ants
where they “tend” aphids and butterfly
larvae and fungus gardens (Fig. 13.5).
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 17
18. Fruit dispersal and
pollination:
• Fruit dispersal is mostly a generalist
phenomenon Fig. 13.7.
• Pollination:
• Charles Darwin was fascinated by pollination and he
described the specialized floral structure of the
Madagascar star orchid in 1859 with nectar tubes
approx. 30 cm long.
• He suggested that a pollinator must exist with an
appropriately long proboscis and 40 years later a
hawkmoth with a 25cm proboscis was found:
• see Fig. 8.5 Howe & Westley 1988 for floral diversity in relation to pollinators & Fig. 7.7 for fig wasp mutualism.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 18
• Fig. 13.10.
• Greater stability for the symbiont or the
opportunity to control environmental
conditions through the association.
Slide - 19 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Gut inhabitants in plant-feeding vertebrates and
invertebrates:
• Problem is coping with cellulose - a very hard to digest polysaccharide.
• Many animals have solved the problem by hosting
bacterial microcosms in their guts.
• Ruminants (deer, cattle and antelope) have a 4chambered stomach of which the rumen harbors a complete ecological community of protozoa and bacteria that compete, predate and cooperate through mutualisms, all driven by cellulose.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 20
• Figs 13.11 & 13.14
of termite gut flora.
• Fig. 13.12.
Slide - 21 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Sheathing ectomycorrhizae and vesicular arbuscular
(VA) endomycorrhizae:
• Found in majority of plant species (Figs 13.15
& 13.17).
• Clearly benefit plants (Fig. 13.18) by providing P, N & Ca.
• Ectomycorrhizae:
• Occur as a sheath most often on roots of trees.
• Can be cultured in isolation from their hosts
• Require soluble carbohydrates as carbon resource from
host (not cellulose like free-living fungi).
• VA endomycorrhizae:
• Extremely widespread and penetrate host cells.
• Makes them very difficult to culture.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 22
• Bacteria associated with roots of some
plants can fix atmospheric nitrogen and
make it available to the plant:
• Fig. 13.19.
• Benefits plant and influences outcome of
other ecological processes:
• Fig. 13.21.
Slide - 23 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
• Ultimate symbiotic mutualism may be
evolution of the eukaryotic cell.
• “tit-for-tat”
• Axelrod & Hamilton demonstrated
theoretically the increased stability
generated from cooperation.
• Sex may have evolved through parasitism
that lead to cooperation and mutualism
because of mutual benefit.
Slide - 24 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
Figure 11.2 (3 rd ). Release of resources
during decomposition of oak leaves.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 25
Figure 11.18 (4 th ). Mouse burial by
Necrophorus beetles.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 26
th
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 27
Figure 11.11 (4 th ). Detritus decomposition rates.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 28
Figure 11.12 (4 th ). Mechanisms of
cellulose digestion.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 29
Figure 11.3 (4 th ). Sizes
of decomposers.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 30
Figure 11.4 (4 th ). Faunal variation of
decomposers among biomes.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 31
Figure 11.5 (4 th ). Freshwater
invertebrate feeding guilds.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 32
Figure 11.6 (4 th ). Energy flow in a stream.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 33
Figure 11.7 (4 th ). Forest litter decomposition.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 34
Figure 11.8 (3 rd ). Influence of isopods
on microbial breakdown of leaf litter.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 35
rd
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 36
Figure 13.5 (3 rd ). Feeding by Atta ants
at their fungus garden.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 37
rd
• Diet breadth of fruit feeders in Malaysia.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 38
Figure 8.5 (Howe & Westley, 1988).
Floral diversity and pollinators.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 39
Figure 7.7 (Howe & Westley, 1988).
Fig wasp mutualism.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 40
Figure 13.10
(3 rd ).
• Degrees of symbiotic association.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 41
Figs 13.11 (4 th ) &13.14 (3 rd ). Symbiotic
mutualists in termite intestines.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists
E – endospore-forming
bacteria
S – spirochaetes
Others are anaerobic,
flagellate protozoa
Figure 13.12 (4 th ). Congruent phylogenies of
aphids and their bacterial endosymbionts.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 43
th
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 44
Figure 13.17 (3 rd ). Vesicular
arbuscular mycorrhiza.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 45
Figure 13.18 (3 rd ). Effect of mycorrhizae on
phosphate concentration in leek roots.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 46
Figure 13.19 (4 th ). Rhizobial bacteria in
root nodules of legume roots.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 47
Figure 13.21 (4 th ). Influence of rhizobia on
plant performance and competition.
BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists Slide - 48