BIOS 6150: Ecology •  

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BIOS 6150: Ecology

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

2. Decomposers and detritivores:

•   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

3. Resources include:

•   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

4. Resource decomposition rates:

•   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

5. Differences from other consumers:

•   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

6. A continuous model of detritivory:

•   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

7. Size classification and biomass of

detritivores:

•   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

8. Distributions of detritivore size

classes:

•   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

11. Mutualism:

•   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

12. Mutualism and symbiosis:

•   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

13. Abundance of 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

14. Nonsymbiotic

mutualisms:

•   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

15. Nonsymbiotic mutualisms:

•   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

16. Nonsymbiotic

mutualisms:

•   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

17. Agricultural/domestic mutualists:

•   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

19. Symbiotic mutualisms:

Degrees of symbiotic association.

•   Fig. 13.10.

Such a range of association

dependence implies that closer

associations might benefit the

interactants:

•   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

20. Gut 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

21. Termite and aphid mutualists:

Termites are “insect deer” with their

own microflora that digest cellulose:

•   Figs 13.11 & 13.14

of termite gut flora.

Aphids also have tightly associated

symbiotic mutualists:

•   Fig. 13.12.

Slide - 21 BIOS 6150: Ecology - Dr. S. Malcolm. Week 9: Decomposers, detritivores & mutualists

22. Mycorrhizae:

•   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

23. Nitrogen fixation:

•   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

24. Postscript on mutualisms:

•   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

Figure 11.15 (4

th

). African dung beetle.

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

Figure 13.21 (3

rd

). Lichen structure.

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

Figure 13.7

(3

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

Figure 13.15 (4

th

). Sheathing

ectomycorrhiza of pine roots.

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

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