Ecology and Evolution Flashcards

(783 cards)

1
Q

Ecological Networks

A
How well did you know this?
1
Not at all
2
3
4
5
Perfectly
2
Q

Indirect Effects within Networks

A

Trophic cascades, keystone species and apparent competition.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
3
Q

Darwin’s Entangled Bank

A

species don’t exist in isolation, but in complex networks of antagonistic or mutualistic interactions.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
4
Q

What dynamics are described by antagonistic networks?

A

Food webs, predator-prey, host-parasitoid, herbivore-plant and pathogen-host.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
5
Q

Mutualistic Networks

A

Pollinators and flowers. Seed dispersers and fruits.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
6
Q

The First Food Web

A

Produced by Charles Elton after an expedition to Bear Island, Spitsbergen. However, some of the connected boxes represented one species whilst others represented multiple species.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
7
Q

How are interaction strength networks produced?

A

By manipulation experiments whereby a species is removed from an ecosystem and the effects are measured.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
8
Q

Degrees of Separation in a Network

A

How many connections are required to link two nodes on a network. A small world property.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
9
Q

How many degrees of separation in ecological networks?

A

2 degrees of separation with >95% within 3 links of each other.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
10
Q

What is the significance of 2 degrees of separation?

A

Dynamics within ecosystems can be highly inter-connected. Changes to the abundance of one species will propagate rapidly. Biodiversity loss, over-harvesting and species invasions may affect more species than previously thought.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
11
Q

Community Structure

A

Which species are rare, common or absent.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
12
Q

Why study networks?

A

They’re more realistic than studying a few interacting species with the Lotka-Volterra model. they summarise the complexity of community interactions. They aid our understanding of community structure and dynamics and what happens if humans interfere.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
13
Q

S

A

The number of species in the web.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
14
Q

L

A

The number of links/connections.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
15
Q

C

A

Connectance: the fraction of possible links in the web that actually occur. C= Actual Links/Potential Links

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
16
Q

Formula for Connectance

A

C= L/[S(S-1)/2]

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
17
Q

How are possible links represented in food web diagrams?

A

Dashed lines.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
18
Q

How can food webs be studied?

A

Observations, modelling and experiments.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
19
Q

Omnivory

A

Feeding at more than one trophic level.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
20
Q

Why are lots of patterns found in past food webs artefacts?

A

They overlooked omnivory and underestimated rare links. There was a taxonomic bias whereby some species were lumped into a single unit, e.g., plankton.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
21
Q

Observation

A

Analysing patterns in published, purpose-built webs.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
22
Q

Example of a pattern observed in food webs:

A

Most food chains are short– 3-4 trophic levels.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
23
Q

Why are food chains short?

A

Partly the Energy Attenuation Hypothesis and partly the Trophodynamics Hypothesis.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
24
Q

The Energy Attenuation Hypothesis

A

Energy is wasted at each trophic level through heat, excretion/egestion and respiration due to the 2nd law of thermodynamics. There’s only a finite amount of energy that is input at the bottom of the food chain as primary productivity. Eventually, there isn’t enough energy left to support another trophic level.

How well did you know this?
1
Not at all
2
3
4
5
Perfectly
25
What does the Energy Attenuation Hypothesis predict?
Communities with more efficient energy transfer should have longer food chains. Communities with higher primary productivity ought to have longer food chains.
26
What evidence supports the energy Attenuation Hypothesis?
Ectothermic food chains tend to be longer than endothermic food chains as less energy is expended regulating their temperature.
27
To what extent is the 2nd prediction (communities with higher productivity ought to have longer food chains) correct?
At low productivity, adding more energy equates to more trophic levels, but there's a limit. Only 3-4 trophic levels are found, typically.
28
Proponents of the Energy Attenuation Hypothesis
MacArthur and Elton
29
Proponents of the Trophodynamics Hypothesis
Pimms and Lawton
30
The Trophodynamics Hypothesis
Longer food chains are less stable. Chance variations in population size are amplified up the food chain leading to unpredictable dynamics for the top predator.
31
Evidence for the Trophodynamics Hypothesis
Environmental variability is correlated with food chain length. In more predictable environments, food chains will be longer.
32
Modelling using 16 real networks.
Jennifer Dunne. Removing species with few connections tends to have little effect. There are critical thresholds whereby after a certain number of individuals of a species have been removed, there are marked effects on the food web. Robustness increases with connectance.
33
Robustness
Secondary extinctions that followed simulated extinctions.
34
Dynamic Models
May, Lawton and Pimm extended the Lotka-volterra models to sets of interacting species.
35
Modelling using simulated networks.
Create artificial food webs. Manipulate the properties, and examine the stability. E.g., food webs with different numbers of trophic levels, but similar values for S and C.
36
What did May's models predict?
Food chains would be short, and network complexity reduces stability. Both greater S and C decreased stability.
37
When did May's model find webs were stable?
If β(SC)^1/2 < 1.
38
May's Model
Randomly assembled food webs with different numbers of species, connectances and interaction strengths (β).
39
β
Interaction strength: the impact of the changing the abundance of a species on another species.
40
Survivorship Bias
The food webs observable in nature are not a random subset of all food webs. They are the stable ones. May's model could be attributed to survivorship bias.
41
Field Experiments
Labourious, but informative. Manipulations can be used to identify keystone species and indirect effects.
42
Robert Paine
Excluding the starfish Pisaster from rocky shores reduced species diversity and shifted food web structure. Ethically challenging to manipulate real ecosystems.
43
Sharon Lawler
Lab experiments with aquatic micro-ecosystems. Corroborated Bob May. Fine control over the composition and conditions of these micro-ecosystems. With 2 species, 2.5% of populations went extinct. With 8 species, 27.5% of populations went extinct.
44
Weak Interactions
Species are connected, but a change in the abundance of one species doesn't propagate to the other species. More important than complexity. A highly stabilising factor.
45
Indirect Interactions
Populations of two different species are dynamically linked, but not through a feeding interaction.
46
Apparent Competition
the hare and the rabbit aren't directly competing, but they share a predator. If the abundance of the hare increases, the abundance of eagles increases, so predation on the rabbit increases.
47
A Trophic Cascade with a Twist
Cats reduce the mice population, which increases the bee population. The plant population then increases also due to a mutualistic relationship with the bee pollinators.
48
Apparent Competition: Squirrels
Grey squirrels appeared to outcompete red squirrels, but actually they share a common pathogen (the squirrelpox virus) to which grey squirrels are more resistant. When they interact, red squirrels are less resistant, so suffer more from the virus.
49
Sea Otters
A keystone species in the otter-urchin-kelp cascade.
50
Keystone Species
A species whose impacts on the community or ecosystem are much larger than expected based on its abundance or biomass. Often predators, but not always. Key to conservation.
51
Wolves
A keystone species that went extinct in the Yellowstone National Park, but were restored in the late 20th century.
52
What were the effects of the restoration of wolves?
Their prey (ungulates and elks) decreased, so herbivory and browsing also decreased. Aspen, cotton wood and willow increased in abundance, so beavers also increased.
53
Beavers
Keystone species/ecosystem engineers whose dams provide a habitat.
54
Fig Trees
Keystone species as all frugivores in rainforests depend on them. Fig trees, in turn, depend on specialised pollinators (fig wasps), thus fog wasps are arguable also keystone species.
55
Giant Tortoises
Keystone herbivores (not a trophic cascade, however, as there are only 2 levels). By grazing, the tortoises reduce vegetation from trees to short grass on Aldabra island.
56
Leaf-cutting Ants
Create large openings in the vegetation that affect plant regeneration in neotropical rainforests.
57
Why is the keystone concept simplistic?
Not all ecosystems have a keystone, and keystones can be more easily identified after extinction. As species are removed, the stability of the ecosystem decreases. Different species within the ecosystem can be differentially important.
58
Nodes
Species
59
Edges
Feeding Links
60
61
Interactions Between Populations I: Predator-Prey & Host-Parasite Interactions
62
63
Trophic Interactions
Feeding interactions in which one species benefits and the other doesn't.
64
Antagonistic Interactions
Interactions where one species benefits from the other.
65
Antagonistic Trophic Interactions
One species gains nutrition from another.
66
Natural Enemies
Parasites, predators, parasitoids and herbivores.
67
Equation for Determing the Size of a Population Over time
Nt+1 = Nt + Births - Deaths
68
When is a population at carrying capacity, K?
When births = deaths.
69
What influences the number of births and deaths?
Resources and competition for them (inter- & intra-specific) and natural enemies.
70
Features of Parasitic Interactions
The host is always harmed by the parasite living in or on it, only part of the host is consumed, the parsite tends to be much smaller than the host and only 1 of very few hosts are consumed per parasite.
71
Features of Parasitoid Interactions
The host is always killed by the parasitoid which lives in or on it. The parasitoid kills one host in its lifetime. The parasitoid and the host are similar sizes.
72
Most parasitoids are wasps or flies.
A parasitoid wasp lays its eggs in aphids that then hatch and eat the aphid. Another parasitoid wasp larvae eats ladybirds then tunnels itself out to form a coccoon.
73
Herbivory
The resource individual is a producer. The herbivore consumes only part of each resource individual.
74
Weevils
Lay their eggs in a seed. The larvae then eat the seed.
75
Kelp
An ecosystem engineer that provides habitats for many marine organisms. A primary producer.
76
How do natural enemies influence abundance of a species?
When the lionfish was introduced into the Caribbean sea, they were invasive and non-native, so reduced the populations and carrying capacities of native fish.
77
Example of a Trophic Cascade
Over-hunting of sea otters in the Pacific increased the populations of urchins and decreased the populations of kelp as well as kelp-dependent marine organisms.
78
Example of Biological Control
The Opuntia prickly pear was well adapted to arid conditions, so became overgrown and invasive in Australia. Cactoblastis cactorum moth was found in Mexico (the native habitat of Opuntia) to act as a biological control agent and reduce the carrying capacity of the Opuntia population.
79
How can you calculate zero isoclines?
Solve for dN/dt= 0 using the Lotka-Volterra equations.
80
Prey Zero Isocline
Combinations of predator and prey that lead to an unchanging prey population. r/a.
81
Predator Zero Isocline
Combinations of predator and prey that lead to an unchanging predator population. q/fa.
82
Why are coupled predator-prey interactions unusual in nature?
In more complex food webs, there are many predators for each prey and vice versa that act as a buffer.
83
Why was the lynx-snowshoe hare predator-prey cycle so tightly coupled?
The Arctic environment has a simple food web with the hare having few other predators and the lynx having few other prey.
84
Functional Response
The intake rate of a conusmer as a function of food density.
85
What does the Lotka-Volterra Model assume?
Type I functional response. Assumes there is a cobstant, linear relationship between the prey density and the number of prey consumed. The model can be adjusted to be more realistic by adding in functional response graphs.
86
Not just predators exhibit these functional responses.
Parasites, herbivores and pathogens can too.
87
Parasites may limit populations in a density-dependent way.
Cliff swallows nest colonially. The number of parasites (usually ectoparasites) increases with the size of the colony.
88
Janzen-Connell Hypothesis
Pest pressure from specialised herbivores or pathogens causes density-dependent survival of seeds and seedlings, promoting diversity via a negative feedback system. A relatively rare species can be released from the affects of natural enemies, and then increase in abundance.
89
Larger-bodied Predators
More vulnerable to extinction.
90
Lago Guri Reservoir
Created by damming a river. This flooded a rainforest, creating islands. Such habitat fragmentation can affect trophic cascades.
91
What happened on the small islands of Lago Guri?
Lack of predators of vertebrates as these predators are most vulnerable to extinction. Densities of rodents, howler monkeys, iguanas and leaf-cutting ants were 10-100x higher than on nearby mainland. Densities of seedlings and saplings reduced dramatically. Severe trophic cascade due to the lack of top-down regulation by predators.
92
93
Population Ecology I & II
94
95
What is ecology?
Concerned with the distribution and abundance of organisms.
96
Evidence for density-dependence:
Bacterial growth curves. Wildebeest recovering from Rinderpest.
97
UK Birds in Decline
All groups of birds have declined from 1970-2016. Farmland birds have declined the most by 90%. From 2018-2023, they declined again by 90%.
98
Features of populations models aim to capture:
Populations should be able to recover from low-density. Eventually, environmental limitations will halt population increase. Populations also flutuate apparently randomly.
99
Continuous vs. Discrete Time Logistic Models
Bacteria divide continuously, all are out of sync with each other= continuous time. Wildebeest have a single reproductive event each year, all calve within a week of each other; synchronised reproductiv events= discrete time. One is more useful for a given population.
100
Life-History Trait
A trait that influences demography (population dynamics).
101
r
Maximum per capita population growth rate. A life-history trait.
102
What determines r?
The typical number of offspring per clutch or brood, number of clutches/broods per year, mortality rate through reproductive life, age at first reproduction and reproductive lifespan.
103
K
The maximum number of individuals an environment can support-- carrying capacity. A property of the environment, not a life-history trait.
104
When is there no net growth?
When N=K, where n is population size.
105
What is the difference between exponential and logistic growth?
When r takes the same positive value regardless of population size, it's exponential growth. Logistic growth occurs when r decreases as the population increases towards the carrying capacity.
106
Maximum population is at growth rate N= K/2, but…
…this is an oversimplification. This affected fishing quotas as as it was assumed that fish stocks were at K, but really they were much lower, so fishing to K/2 almost drove certain fish to extinction.
107
Absolute Growth Rate (in the Discrete-Time Logistic)
The change in population size in a single growth step.
108
Why do wood mice and bank mole populations fluctuate massively?
Environemntal stochasticity. The availability of acorns fluctuates due to the weather.
109
If the wood mice populations fluctuate, why don't the tawny owl populations fluctuate much?
The tawny owls have a smaller value of r than mice.
110
Why is density dependence difficult to detect in real populations?
You will mostly observe fluctuations in K, once equilibrium/carrying capacity is reached.
111
How can environmental stochasticity be shown in a model?
By varying K by pulling random numbers from a normal distribution.
112
K varies.
There is a still a mean K, but K fluctuates. If one year is good, this indicates nothing about the next year.
113
The smaller the value of r for a population, …
… the less responsive the population is to changes in the environment.
114
When r is larger, the fluctuations are greater. Why?
The population is more responsive to changes in K.
115
Occam's Razor
Choose the simpler model when confronted by a simpler and a more complex model. However, simple models/toy models may be able to reproduce simple patterns, but are not entirely trustworthy.
116
r determines time to recovery after a population crash.
Larger values of r mean the population recovers faster. Smaller values of r mean the population recovers more slowly, and it's seldom at its carrying capacity.
117
Why have humpback whales still not recovered from whaling?
Humpbacks are still hit by ships. Humpback whales produce 1 calf every 2-3 years. It takes 5-10 years for a calf to reach sexual maturity. It has a low value of r.
118
r is under selection.
The r/K continuum is difficult to estimate in real populations and only goes so far. r can evolve.
119
K Selection
At K, there is lots of competition as resources are scarce. If the population is often at K, there's selection to produce few, larger offspring. Competition favours large body size, late maturation and having few, large offspring. E.g., humpback whales.
120
r Selection
r selected species are selected to have higher values of r as they live in more hazardous environments. They can recover easily from low density. Large litters, early sexual maturity, frequent broods, small body size and small offspring. E.g., rabbits.
121
Average Population Size
Mean value of K.
122
Grey heron populations crash periodically.
This is due to harsh winters. If the water is frozen, herons can't reach any fish. This can be modelled as a discrete-time logistic with hazards that kill members of the population randomly.
123
What is recovery time proportional to?
1/r, hence species with higher r values recovery faster.
124
European rabbits have a very high r.
Litter size: 1-14. Can produce one litter per month at low density. 13 rabbits were introduced to Australia in 1859. In 1866, hunters shot 14,000 rabbits. In 1940, there were an estimated 60 million rabbits across Australia.
125
Species can be pushed into positive feedback loops.
E.g., a hazardous environment might mean selection on r, leading to earlier maturation and smaller body size. This makes individuals more susceptible to hazards, which increases selection on r.
126
127
Population Ecology III: Modelling Single-Species Populations
128
129
Deterministic Formulation
Always adds or subtracts the same number of individuals, but this isn't realistic. No stochasticity is factored in.
130
Issue with the discrete-time logistic:
r is a real number, so in the model, fractional individuals are possible, which isn't the case in real life.
131
How can the fractional-individuals-issue with the discrete-time logistic be resolved?
Truncate, or make the actual number of new individuals a random variable and use the Poisson distribution.
132
Demograohic Stochasticity
Randomness inherent to populations as a result of randomness in the birth and death processes. Only potent in small populations.
133
For a Poisson distribution, variance=
mean. Variance cannot be changed.
134
Fluctuations at Equilibrium
Due to environmental or demographic stochasticisty. You cannot tell which. For a small population, fluctuations due to stochasticity at equilibrium are greater.
135
r is determined by both births and deaths.
The actual number that die or are born isn't constant. It includes a component of randomness.
136
If the mean population size becomes very low, what can happen to the species?
The species can go extinct just by chance. For demographic stochasticity to cause extinction, K<50 roughly: the population size must be very small.
137
The combination of environemntal and demographic stochasticity means some populations may never reach K.
This puts a huge selection on r.
138
Black-footed Ferrets
Once baundant in U.S. prairies, the population plummeted when the bubonic plague was introduced. In one population, only males were born in one year. This led to extinction due to stochasticity in the sex ratio.
139
Genetic diversity can be lost from a population when it shrinks in size…
… by genetic drift, especially if the population remains small for a long time.
140
Florida Panthers
Had undergone genetic erosion due to drift in tehir small population, so the subspecies couldn't be saved. Instead, panthers from Texas were introduced in 1955.
141
Red Kites
Weren't able to expand their range and were restricted to south Wales, until Swedish red kites were introduced.
142
Weak Allee Effect
The population growth rate decreases as populations get smaller.
143
Strong Allee Effect
Population growth rate becomes negative below a critical threshold in population size.
144
Musk ox form defensive rings around their offspring to fend off wolves,
If the herd is too small, all the offspring are eaten as they can't defend themselves against wolves. They would suffer an Allee effect.
145
African wild dogs experienced the Allee effect. Why?
They hunt in packs, and their population shrank due to canine distemper and rabies.
146
Kakapo males form a lek, and attract females with a booming call.
There could be an Allee effect as a female is attracted to several males' calls.
147
The Kakapo: the ultimate K-selected animal.
4kg-- the heaviest parrot. Flightless and nocturnal with whiskers to feel its way in the dark. Breed slower than any other bird (every 2-7 years, when high-quality food is available). Can sometimes live for more than 100 years!. Low basal metabolic rate allows it to subsist on low-quality food. once hugely abundant, but cannot recover easily.
148
What doesn't New Zealand have?
Any native mammals.
149
What caused Kakapos to decline?
When the Maori arrived ~1200 years ago, they brought dogs, intrdouced Pacific rats and hunted kakapos. Europeans in the late 18th century brought more mammals-- mustelids that predate ground-nesting birds.
150
Mustelids
Stoats, ferrets and weasels.
151
In 1977, there were 100-200 kakapos on Stewart Island.
This island had no mustelids, but did have cats and ongoing predation.
152
Low Point in the Kakapo Population
51 birds. There are more males than females.
153
From 1980 to 1997, birds were translocated to four small islands.
An intensive kakpo recovery programme began.
154
Tracking
Every kakapo bird is fitted with a smart transmitter and tracked.
155
Health Check
Kakapo birds are caught annually and inspected.
156
Nest Monitoring
Some kakpo nests are monitored remotely; others have a dedicated human.
157
Constant fear of predator arrival.
Strict biosecurity protocols and traps.
158
Supplementary Feeding
Feeding stations are re-filled throughout the island.
159
Artificial Incubation and Hand-Rearing
69 have been incubated and hand-reared so far, including removing chicks when the food supply fails. This eliminated some of the demographic stochasticity.
160
Kakapo aspergillosis outbreak on Codfish island in 2019.
21 cases. Twelve recovered after treatment. Two adult females and seven chicks/juveniles died. Birds were helicoptered to the mainland for intensive treatment. Associated with low immunity, likely due to genetic erosion.
161
Median Minimum Viable Population
4000 individuals.
162
Why does minimum viable population size need to be cautiously set?
Due to potential Allee effects.
163
The effect of in a discrete-time logistic:
Low r = smooth curve to equilibrium. Higher values of r= dampened oscillations followed by limit cycles. Very high values of r= chaos-- entirely unpredictable.
164
Limit Cycles
Population size just fluctuates below and above K by the same amount repeatedly in a stochasticity-free discrete-time logistic.
165
Chaos
Not stochastic, but highly susceptible to initial conditions.
166
Populations are not chaotic.
St. Kilda's wild soay sheep appeared chaotic due to tehir large fluctuations. In reality, many sheep give birth to twins at the same time. They have a high r value, so can recover post-winter.
167
How are some populations able to exceed K?
High r values. In a single generation, the population can add a large number of recruits.
168
Overcompensating Density Dependence
Associated with exceeding K. For high values of Nt, the population size of the next generation will actually be lower than the current population size.
169
Overcompensating density dependence is extreme for very high values of r.
this means once the population booms, it can crash back down again. The addition of a 1:1 line on a Nt against Nt+1 graph illustrates this clearly.
170
When r is very high, what is the only population size at which the population is stable?
When Nt= K, but with chaotic dynamics, this will never be reached.
171
More attention should be paid to declining populations.
Intervention should be earlier, before the population has reached its minimum viable population size.
172
173
Comparative Method
174
175
Comparative Method
Taking pre-gathered data on one species to compare with another species.
176
Polygyny
Single male, multiple females.
177
Polyandry
Single female, multiple males.
178
Darwin, 1871.
Sexual selection. Why do male peacoacks have such tails?
179
Monogamous Birds
Tend to have paired systems where both parents care for the offspring. In polygamous mtaing systems, the males leave, so females have to raise the offspring.
180
John Crook, 1964:
Red-headed weaver: monogamous with dispersed territories. Southern masked weaver: polygynous, and nests in colonies. Village weaver: polygamous and nests in colonies.
181
Correlation between food and mating system:
Correlation between monogamy and insect-eating, polygamy and seed-eating. Pairs can catch insects to feed their offspring, but it's easier to find lots of seeds to support a large colony.
182
Monogamy vs. Polygamy
Polygamous groups tend to be colonial. Monogamous birds tend to be solitary. Polygamy increases genetic diversity and variation.
183
Closely-related Species
Don't represent independent data points.
184
Issues with John Crook's Conclusions
Comparative studies can't prove something is the result of something else-- correlation not causation. Crook didn't quantify his ecological variables. Closely-related species don't represent independent data points. Causalty could be in the other direction with sociality determining what birds eat.
185
Comparing Sexual Dimorphism in Primates
Indri, macaque, baboons and mandrils.
186
Indri
Largest out of 80 different types of lemur. Males and females tend to be the same size.
187
Macaque
Females are just slightly smaller than males.
188
Baboons and Mandrils
Males are roughly double the size of females.
189
Clutton-Brock and Harvey, 1977: Sexual Dimorphism in Primates.
Quantified ecological variables: sexual dimorphism= difference in weights, plotted against sex ratio in the breeding groups.
190
Clutton-Brock and Harvey's Hypotheses: Niche Differentiation to Reduce Competition or Sexual Selection.
When there's high sexual selection and competition between males, males may be larger to compete with one another. The higher the sexual dimorphism, the higher the number of females per male.
191
Limitations to Clutton-Brock and Harvey's Primate Comparison
There may be more untested hypotheses. Are species independent? This hasn't been tested. Pan and gorillas are more closely related than indri and lemurs, so some data points rae more independent than others.
192
Why not use species in the comparative method?
Statistical analyses assume data points are independent. Closely-related species tend to be similar because they share traits through common descent, rather than independent evolution.
193
Pseudo-Replication
Doing an experiment, but thinking you sample size is larger than it is.
194
What does plotting the contrasts do?
Partitions out the effects of the shared relationship/the phylogeny. Finds the differences between datapoints. Accounts for the statistical non-independnece of the data.
195
Fig Wasps
The pollen-laden female wasp lays her eggs inside the fig. Pollinates the flowers on teh fig tree. Male wasps fertilise the females. Some females lose their wings, and spend the rest of tehir lives their. Others don't.
196
Pollinating Fig Wasps
Both sexes are smaller than their non-pollinating counterparts. Some non-pollinators can lay their eggs without even entering the tree.
197
Non-Pollinating vs. Pollinating Male Fig Wasps
Non-pollinating males are armoured, often live their whole lives in fig trees and some can be violent.Pollinator males don't tend to be violent. This is just a correlation as they are closely-related species.
198
West et al., 2001. Fig Wasps
The higher the female density, the lower the level of male injury. Blending phylogeny with quantified data by plotting the contrasts between species instead of the species as datapoints themselves.
199
200
Major Evolutionary Transitions
201
202
The ancestor of most animals was probably like the mole.
Solitary. Only a few times in spring does it emerge to find a mate.
203
Colony of Guillemots
They pack together on a cliff to rear their young, but the breeding pairs seldom interact.
204
Shoal of Fish
They avoid predators in a group. Each individual benefits directly from being in a group. There is no cost.
205
Bacteria cooperate in a colony.
They all produce a molecule to break down a food source, so they can take it up.
206
Social Behaviour
Any behaviour that impacts another organism.
207
Humans are less social than slime moulds.
Generally, slime moulds are single cells that break down wood. When the food source dries up, they aggregate together to form a fruiting body that produces spores. Spores disperse to find a new patch of food. Slime moulds are semisocial.
208
Reproductive Division of Labour in Slime Moulds
The stalk cells are sterile or may have died in some species. Their reproductive potential is over, but the cells in the fruiting body can still reproduce.
209
Portuguese Man of War
Made up of zooids with an air sac on top. As juveniles, they can live independently, but as adults they can no longer live alone and have specialised roles.
210
The termite queen is huge.
After the termite king and queen make a nuptial flight, they burrow, mate and the queen produces the first batch of eggs. The workers help her produce more eggs. The queen can live for 50-60 years! All the workers are sterile.
211
John Maynard Smith
Signalling theory, game theory and major evolutionary transitions.
212
Major Evolutionary Transition
An interruption of evolution as a series of gradual changes. Instead of a continuum, it's a series of steps. Revolutions.
213
Language
Major transition in how information is passed on.
214
The evolution of chromosomes and genomes: major evolutionary transition.
Evolution of information transfer by DNA. Bits of genome have clustered together to form chromosomes that have come together to form a genome.
215
The evolution of the eukaryotic cell: major evolutionary transition.
Endosymbiosis. All eukaryotes descend from an ancestor with an alpha-proteobacterial symbiont. Two individuals from different domains coming together.
216
Evolution of sexual reproduction: major evolutionary transition.
Ancestors would have been asexual. Producing gametes relies on another individual to transfer information to the next generation.
217
Evolution of multicellularity: major evolutionary transition.
Individual cells form groups then become a multicellular organism. A unicellular alga formed groups with reproductive division of labour. Some algae were soma cells, whilst others reproduced. The transition to multicellularity occurred multiple times.
218
The evolution of eusociality: major evolutionary transition.
Extreme form of a social group with total reproductive division of labour between different castes.
219
Solitary Cockroaches
Most recent common ancestor of colonial termites.
220
Transition in Individuality
Type of major evolutionary transition. When individuals come together to form a 'higher-level' individual, so that individuals can no longer reproduce separately, just as this new individual.
221
Transitions in Individuality: Examples
Replicating molecules becoming populations of molecules, prokaryotes to eukaryotes, protists to animals, plants and fungi and solitary individuals to colonies.
222
Major evolutionary transitions that are NOT transitions in individuality: examples.
Independent replicators to chromosomes, RNA to DNA, asexual clones to sexual populations and primate societies to human societies, incl. language.
223
What stage occurs between solitary individuals and a 'higher-level' group?
A cooperative group.
224
Why would individuals form a cooperative group?
High relatedness is important. Low promiscuity means an individual is 0.5 to its offspring and its siblings. Cooperation can evolve, then be lost again, if promiscuity increases.
225
What is required to transition from a cooperative group to a new, 'higher-level' individual?
No selection or conflict within the group.
226
How can conflict be eliminated from a cooperative group?
Aligning genetic interest by increasing relatedness through monogamy (in sexual organisms) or clonality. Repression of competition.
227
Workers invest in repressing conflict in haplodiploid honeybee colonies.
Workers don't mate, but can produce male eggs. Males arise from unfertilised eggs. Worker policing: workers eat the eggs laid by other workers.
228
In multicellular organisms, all cells are genetically identical/clonal, so it doesn't matter which reproduce.
No competition between cells as they are all the same. There's a division of labour: some cells reproduce whilst others don't. If cells with one genotype becom ethe germline and cells with another genotype become the soma, there would be conflict as soma cells do all the work. This is avoided in a clonal system, but where cells ahve different genomes, this is an example of conflict, e.g., chimeric genomes.
229
Why do mitochondria nd host cells have the same selection across both individuals?
They are dependent on each other to reproduce.
230
Subsocial (offspring)
r(offspring) = r(siblings). When the queen has mated only once, relatedness to the siblings and offspring is the same. High relatedness in the group.
231
Semisocial (siblings)
r(nephews and nieces) < r(offspring). Multiple queens in a colony. Relatedness to potential offspring is greater than the offspring they help to rear, because they'll be your nephews and nieces instead of siblings.
232
Comparative method: is monogamy required for eusociality to evolve.
Monogamy was ancestral to all the independent origins of multicellularity. Polygamy has only evolved after eusociality evolved. Major evolutionary transitions are irreversible.
233
Obligate Multicellularity
Only evolves from ancestors with clonal group formation (discovered by Bertie Fisher).
234
Facultative Multicellularity
Cells aggregate together, but haven't undergone a transition as they can still separate. Can evolve when groups are non-clonal.
235
Multicellular group formation can also be sub- or semisocial.
Subsocial: where a single cell produces many clones that stay with the parent, r=1. Semisocial: cells aggregate together; they're not clonal.
236
What does complex, obligate multicellularity require?
A subsocial route with a single cell stage. Organisms begin from a single-celled egg-- a single cell bottleneck. This removes competition as all cells are clonal.
237
Major Transitions Between Species
Egalitarian, e.g., mitochondria and host cells. Competition is removed as genes are transmitted together.
238
Major Transitions Within Species
Fraternal.
239
How does reproductive division of labour evolve in bacterial colonies?
In monocultures, the density of altruistic helpers and reproducing bacteria is greater than that of cheats. In a mixed culture, the cell density of cheats increases.
240
Rhizobia and other symbionts can jump between hosts.
Thus, their evolutionary lineages are not completely matched to their hosts', and a transition will not occur.
241
242
Single-Species Populations IV: Metapopulations and Other Spatially-Structured Populations
243
244
Frequent Features of Populations of Individual Species
Often small, extinction-prone and patchily distributed in space.
245
Populations can grow exponentially at first, then..
… density dependence kicks in, and the population size approaches carrying capacity.
246
Small populations are especially prone to…
… extinction.
247
Demographic Stochasticisty
Variation in population size due to variation in births, deaths and sex ratio around the mean. Works alongside environmental stochasticity.
248
Fragmented Habitats
Often have small, extinction-prone populations within them. Anthropoengenic activities can lead to patchy and fragmented landscapes.
249
Why are small populations inherently more extinction-prone?
Stochastic events can cause local population sizes to fall to zero.
250
Allee Effect
The tendency for population growth rate to become very small or even negative when the population size is very small.
251
In a fragmented landscape, patches supporting individual populations are reduced in area.
Populations can become trapped in small patches of habitat that tend to lack the resources to support them long-term. These populations are extinction-prone.
252
Semi-Natural Fragmentation
E.g., insects in clusters of acacia trees in Kenya, or populations in ponds (like inverse islands).
253
Challenges for Small Populations
Demographic and environemntal stochasticity, genetic effects and Allee effects.
254
What may occur in patches where populations go extinct locally?
They may be re-colonised by immigrants from other populations.
255
Dispersal may be key to persistence.
The dispersal ability of a species relative to the level of habitat fragmnetation will determine how its population is structured and whether it can persist in fragmented landscapes.
256
Metapopulation
Population of populations. Species can persist as a balance between extinction and colonisation events.
257
At any one time, some fraction of the patches will be occupied.
The number of patches occupied at any one time depends on the balance between rates of colonisation and rates of extinction.
258
Metapopulation Example: Melitaea cinxia.
Areas of meadow (suitable habitat for the butterfly) surrounded by sea and forest (unsuitable habitat) in part of Finland. Local extinction will be balanced by butterflies colonising previously uncolonised areas.
259
Levins Population Model
First determined by Richard Levins in the 1960s. Rate of change of occupied patches= colonisation (c ) - extinction (e).
260
The higher the rate of colonisation and the lower the rate of extinction, …
…the greater the proportion of patches occupied.
261
When is colonisation rate highest?
When there's an intermediate number of patches occupied. There are lots of empty patches available to be colonised, as well as lots of occupied patches to generate colonists.
262
When does colonisation rate decline?
As the patches are saturated.
263
What happens when colonisation rate balances extinction rate?
This gives the average proportion of patches occupied in a given landscape in a given time.
264
Distance Colonisation
Patches are isolated from each other in a landscape, so it's harder for individuals to disperse to colonise other patches.
265
What conditions give the smallest proportion of patches occupied?
When patches are far apart, and when populations are small in the metapopulation. High extinction rate, and low rate of colonisation.
266
Correlated Population Dynamics
Across different populations in a metapopulation. Increases the chance of landscape-wide extinction.
267
Uncorrelated Population Dynamics
Across different populations in a metapopulation. Decreases the probability that all local patches will go extinct simultaneously. Dispersal events allow patches where local populations have gone extinct to be re-colonised. However, lots of drivers of environmental stochasticity, e.g., weather, tend to affect populations in that environment similarly.
268
Conservation Insights from Metapopulation Dynamics
Minimising extinction rates, and maximising colonisation rates. Conserving empty habitat patches. Reducing population synchrony. Most of this can be achieved through habitat mangament, e.g., ensuring some populations do better in wet years and others do better in dry years, or by ensuring patches are well-connected.
269
Examples of Spatially-Structured Populations
Metapopulations, patchy populations, mainland-island dynamics and source-sink dynamics.
270
Patchy Population
Monitored change in occupancy of holly leaf miners in these patches (holly trees in a woodland over time). Every holly plant has leaf miners on it every year. The populations appears patchy, but the flies that lay the leaf-mining larvae are mobile enough that it is effectively one habitat. best treated as one population.
271
Mainland-Island Dynamics
The mainland, core population is large enough that it effectively never goes extinct. It should be protected. The satellite populations are small, so extinction prone, and are reliant on re-colonisation from the mainalnd population.
272
Source-Sink Dynamics
The source is a large, high-quality habitat-- a next exporter of individuals. The sink only persists due to immigration from the source, otherwise it would become extinct. Without high immigration, the right hand side of the equation (Nt+1= Nt + births - deaths + immigrants - emigrants) would be negative, and teh population would decline nnntowards extinction.
273
274
Community Ecology I & II
275
276
Ecological Community
A group of species that occur together in space and/or time, and compete for limiting resources. Inherently competitive.
277
Resource
Something that is consumed, and used up for growth.
278
A resource can only be limiting when it's depleted.
E.g., CO2 cannot be limiting for plants because it's so availble; whereas nitrates in the soil can be depleted, so can be limiting.
279
Trophic Guilds
Species in the same trophic guild have similar feeding roles in an ecosystem.
280
Hyperdiversity
In one hectare of the the tropical rainforest in Brazil, there are 400 different species. Other examples include tropical coral reefs.
281
How many tree species in total across the U.S. and Canada?
~700.
282
Natural Monocultures
Low diversity. E.g., Coco de mer is one species that dominates the palm forests in the Seychelles.
283
Gause's Principle of Competitive Exclusion
Two species cannot exist on one limiting resource, if other ecological factors remain constant.
284
What consequences does Gause's principle of competitive exclusion have?
If there's only one limiting resource and no other meaningful ecological variation, we only expect to observe one species, because it will be better adapted, and outcompete the other species.
285
Connell's Barnacles: example of competitive exclusion.
Cthamalus is completely excluded from the lower shore by Balanus, but persist in the upper shore because it is more tolerant of desiccation. Balanus was less well-adapted to the upper shore, so were killed there. Cthamalus larvae were still found on the lower shore when dispersed, but as adults they were killed due to increased predation. As Cthamalus is less aggressive, it was outcompeted by Balanus.
286
Cthamalus
The planktonic larvae swims around, then fixes to the rock by secreting CaCO3. The barancales open their paltes to feed-- filter feeders. They clsoe their plates to prevent desiccation.
287
Paramecium bursaria
Has symbiotic alage termed chlorella.
288
Gause's Parameciums
Both P. bursaria and P. aurelia eat bacteria. P. aurelia outcompetes P. bursaria in aerobic conditions, but P. bursaria can persist in anoxic conditions where P. aurelia cannot.
289
What happens when the Parameciums are grown together in a well-stirred mixture?
P. aurelia outcompetes P. bursaria.
290
What happens, if the Paramecium mixture is not stirred?
Both species persist, but P. bursaria is mostly confined to the bottom of the beaker. This is because anoxia occurs at the bottom of the beaker, but P. bursaria can geneerate O2 internally via its photosynthetic symbiont.
291
Fenchel's Snails (Hydrobia)
The 2 species didn't occur together, even though the distribution of their food (algae) was the same, and they were almost indistinguishable.
292
What happened when H. ventrosa and H. ulvae (Fenchel's snails) were sympatric?
There was character dispalcement: H. ulvae fed on larger food particles and became significantly larger than H. ventrosa.
293
Tilman's Diatoms, 1981.
Photosynthetic, planktonic organisms. Diatoms are contained within a unique silica-frustule cell wall. Can be grown in culture such that silica is the limiting factor.
294
R*
The level to which the concentration of a limiting resource is reduced by an equilibrial monoculture of a species.
295
R* Value Explained
When the concentration of silica is high, the population of the diatoms increases. Their growth depletes the silica, hence their population growth rate slows. R* value: concentration of silica at equilibrium.
296
R* in Different Species
Different species have different R* values, but each R* value is unique and constant for that species.
297
Labile
Highly mobile.
298
A theory of competition for a limiting resource:
R*.
299
Assumptions of the R* Theory
1. Species compete for a single, limiting resource. 2. The resource is labile, and the mixture is well-mixed.
300
Alternative Definition of R*
The minimum resource concentration a species requires for positive population growth. E.g., the diatoms cannot take up silica from the water when silica concentration is below the R* value for that species.
301
What does the R* theory predict?
The species with the lowest R* for the limiting resource is predicted to be the best competitor.
302
Prairie Grasses
Often C4. Compete for limiting nitrogen.
303
Nitrates vs. Ammonia.
Nitrates are labile. NH3 is not.
304
R* Theory: Prairie Grasses in Minnesota.
Assumptions of the R* theory were met. 4 species. After 5 years, they measured the biomass and R* values of species in monocultures. It wasn't the species ebst able to accrue biomass that were most abundant. It was the species with the lowest R* values.
305
What happened when the seeds of Agropyron and Agrostis (prairie grasses) were grown together?
Agropyron outcompeted Agrostis. R* theory predicts this, but monoculture biomass doesn't.
306
Stable Coexistence
Two species persist together forever.
307
What are the possible outcomes when you put two species together?
Species 1 outcompetes and excludes species 2, or vice versa. Stable coexistence. Sometimes the dominant species cannot be predicted because this depends on the initial conditions: there are two alternative equilibria, e.g., the species that arrives first may dominate.
308
Alpha Values
Competition coefficients by which interspecific competitors are weighted.
309
If alpha is less than 1, …
… adding an individual of species 2 depletes the resources less than adding an individual of species 1, and vice versa, if alpha is greater than 1.
310
If the carrying capacities of two species are similar (k1=K2), what is required for them to co-exist?
Both competition coefficients must be less than 1. the worst kind of competitor should be a member of the same species.
311
What conditions are required for coexistence generally?
K1 > K2α12 AND K2 > K1α21.
312
When is interspecific competition likley to be weaker than intraspecific competition?
When each species has its own ecological niche.
313
When do multiple species coexist?
If the environment is more complex than one ecological condition being limiting.
314
According to Gause's principle, if species occupy the same niche, …
… they cannot coexist.
315
Robert MacArthur
Observed five different species of warblers in coniferous forests.
316
Niche Separation in Warblers
Each warbler fed on a different part of the tree. E.g., Cape May warblers fed only on the tops of trees. If any one of these species' populations increased in size, intensified intraspecific competition would mean the species whose population had just increased would suffer the most. Population rescue also occurs as when a species is rare, competition is released, so the species' population increases.
317
There is still some interspecific competition between warblers.
If one of the species went extinct, the other species' populations would still increase, but there may be a delay as the other species are not as sensitive.
318
When species coexist, competition will be most intense among…
… conspecifics.
319
Feeding niches create strong density dependence.
If species have different niches, then competition is concentrated within a species, and species have a greater impact on themsleves than they do on others.
320
Dynamical Effect
When the population density of a given species is high, there are high levels of competition that tend to cause the population growth rate to decline. Vice versa when population density of a given species is low.
321
Life-History Trade-Offs
Constrain species, so they cannot be good at everything. Competition between species through evolutionary time forces them to specialise.
322
Life-History Trade-Off: Growth Rate vs. Defence in Plants.
Cacti have spines and photosynthetic stems, but they grow slowly. Whereas, bamboo is defenceless and fast-growing because it grows in montane forests here resources are abundant, so bamboo must grow fast enough to reach the light. Plants with hairy leaves represent a middle-ground as they do have some defence, but the hairs do affect light penetration and, thus, growth rate. The Darwinian demon would be a hairy-leaved bamboo, but this is impossible as resources are limiting.
323
Resources invested in one trait aren't available to be invested in another as resources tend to be limiting.
The resulting trade-offs offer potential niche axes along which species can be differentiated.
324
Pioneer Trees
Trees that grow quickly in the gaps due to fallen trees, but not well in the shade. Cast a light shade.
325
Shade-Tolerators
At the other end of the growth-survival trade-off to pioneer trees. Can't grow quickly in the gaps, but can survive well in the shade. Cast a dense shade.
326
1% of Daylight
Reaches the forest floor.
327
Birch
Pioneer trees. Cannot regenerate in the shade. Produce lots of small seeds. Short-lived.
328
Beech
Shade-tolerant. Larger than birch. Grows slower than birch because the wood is higher density.
329
A simple model of forest dynamics:
When a shade-tolerant tree dies, a large gaps facilitates lots of light. The pioneer tree produces lots of seeds, so it's likely that the pioneer tree will fill the gap. When a pioneer tree dies, the gap is smaller, but shade-tolerant saplings were already growing underneath it in the less dense shade. Though pioneer trees will attempt to outgrow them, the shade-tolerant trees were already there, so will fill the gap.
330
Forests that experience hurricane damage
Tend to have a greater proportion of pioneer trees.
331
Why are shade-tolerant trees typically more common in forests than pioneer trees?
Shade-tolerant trees live longer.
332
How can co-existence in the forest model be achieved?
1. When a shade-tolerant tree dies, it creates a gap, which is captured by a pioneer tree. 2. When a pioneer dies, its place is atken by a shade-tolerant tree.
333
Shade-tolerators and pioneers create heterogeneity in the light environment.
This allows the different species to specialise, so they can co-exist. Other forms of heterogeneity, e.g., different soil types, can also promote the co-existence of plant species.
334
When pioneer trees are rare…
… there are lots of gaps available, so the pioneer population increases.
335
When shade-tolerant trees are rare…
… there are lots of pioneer trees, which creates the ideal environment for shade-tolerant trees to grow, increasing their population.
336
Anolis lizard
Colonised the Greater Antilles from mainland South America.
337
Species of Anolis lizard evolved different morphologies and feeding habits to avoid competition-- an adpative radiation.
E.g., in Cuba, there are 6 main ecomorphs that feed on different parts of the habitat, e.g., crown-giant, grass-bush etc.. The larger islands support more ecotypes. Classified by Losos based on habitat.
338
How does the phylogenentic tree demonstrate that the radiation of Anolis lizards on each island is independent?
The order each morph evolved on each island differs. Their evolution is repeatable.
339
What drives specialisation?
Competition between species.
340
What suggests that the same ecological forces operate in different places?
Convergent evolution of similar ecological strategies.
341
342
Co-Evolution of Antimicrobial Resistance
343
344
Why does the UK have increased rates of sepsis?
Due to E. coli.
345
How many people have died due to Covid-19 since November 2019?
7 million.
346
How many people die due to anti-microbial resistance each year?
1.5 million at least as there is a lack of data on low-income countries.
347
Between now and 2050, it is estimated that how many people may die from anti-microbial resistance?
50-60 million.
348
Where is anti-microbial resistance most prevalent?
Sub-Saharan Africa. It is also prevalent in South Asia. Particularly in low-income countries.
349
How many strains of E. coli are found in the gut?
5
350
Klebsiella pneumoniae
One of the biggest killers in low-income countries as it causes neonatal sepsis. It absorbs DNA easily.
351
Coevolution/Collateral Damage
When an antibiotic is given that not only selects for a resistance mechanis against that antibiotic itself, but also selects for a resistance mechanism against another antibiotic (or resiatnce to multiple antibiotics).
352
How does co-evolution of anti-microbial resistance work?
Genes are clustered together, and move in those clusters. Thus, using antibiotic A also selects for resistance to antibiotics B, C and D due to the cluster.
353
What would happen, if doctors switched from administering antibiotic A to antibiotic B, when resistance to both A and B are in the same cluster?
The bacteria would be unaffected by the switch. This may result in clinical failure.
354
It's not just antibiotics that select for anti-microbial resistance.
Biocides, such as those used to clean surfaces in hospitals, and the use of nutrients such as copper and zinc in the farming sector also select for anti-microbial resistance.
355
The more foreign DNA a bacterium has, …
… the more foreign DNA it can acquire by homologous recombination. This means the more resistance genes a bacterium has, the more it can acquire.
356
Inflation in much of Africa has increased the price of medicines.
The cost of antibiotics in Nigeria has risen fourfold since last June.
357
A hospital in Dessie, Ethiopia, that was ransacked by militants.
5-6 newborn babies were sharing a cot and an incubator. The supply chain across many African countries for antibiotics was so poor that there were no available antibiotics to treat Klebsiella pneumoniae in the hospital. Mothers of newborns bought the drug meropenem, and sahred it around the unit to treat sepsis. Neither the mothers nor the hospital could afford it long-term.
358
3 Global Health Organisations
WHO, FAO (Food and Agriculture Organisation) and WHO for animals.
359
How many bacteria are released into the environment every time a human defecates?
10^14 bacteria.
360
Running Water in Hospitals in Low-Income Countries:
In Bo in Sierra Leone, there is no running water or soap in birthing suites, so infection is high and can spread quickly. There are no hospitals with running water in Ethiopia.
361
Why does the lecturer critique the One Health approach to anti-microbial resistance?
He views hospitals as the battle-ground for infections because bacteria are on and in humans.
362
ISCR Elements
Successful at moving large quantities of DNA at no cost to the bacterium.
363
Enviornmental Stimuli
Can cause DNA to be moved, e.g., plastics.
364
Toxin-antitoxin ssytems
Genes on the plasmid that hold the bacterium hostage.
365
The same drugs have been used in humans as in animals since…
… 1953.
366
Usage of Antibiotics in Agriculture
Exceeds 200,000 tons per year. Usage is projected to increase by 8-11% by 2030.
367
Global Sales of Antibiotics
$7 billion. Projected to reach $11 billion by 2033.
368
How much is spent on tackling anti-microbial resistance compared to on weapons?
Barely $1 billion is used to tcakle anti-microbial resistance each year, whereas $2.4 trillion is spent on international arms, even though less than 1 million people have died in conflict since 1985.
369
3 Groups of Antibiotics according to the WHO:
Access, Watch and Reserve.
370
Access
Antibiotics you should be able to get relativley freely.
371
Watch
Antibiotics you should be able to get, but the use of which should be monitored.
372
Reserve
Antibiotics that should only be used for life-threatening infections.
373
Ampicillin
Used as a growth promoter in animals, but confers resistance to the antibiotic group carbapenems, which is used in humans to treat serious infections.
374
Chicken Feed in Vietnam
Contains colistin, which is used to treat serious infections in humans.
375
Conjugation Frequency
1 in 10^3-10^4.
376
Classical Collateral Damage
Plasmid (120-250 kb) have resistance genes arranged in clusters. There's a transfer integron, which contains a cluster of genes, incl. aadA1 (confers resistance to aminoglycosides) and VIM-1 (Confers resistance to carbepenems). If this cluster is transferred, be it onto the chromosome or another plasmid in another bacterium, not just one gene is received, but a series of genes.If treating a patient with carbapenems, VIM-1 confers resistance to it, and aadA1 is transferred as well.
377
Integrons
Have an integrase (a recombinase)-encoding gene, a recombination site and a promoter, as well as a gene casette, which may contain anti-microbial resistanec genes.
378
NDM5
New Delhi Metallo β-Lactamase. Discovered in 2010. Confers resistance to carbapenems.
379
IncX3 Plasmid
In E. coli in eastern China, this plasmid was associated with the resistance gene NDM5. NDM5 confers resistance to carbapenems, but China doesn't use carbapenems. The NDM5 genes were being transferred due to the use of ampicillin in chicken and pigs.
380
NDM-5 genes drive resistance.
The IncX3 plasmid with NDM5 genes appeared in gram positive bacteria fter mating with plasmid-containing gram negative bacteria. This method was culture-independent, so allowed factors affecting the transfer of plasmids between bacteria and the environment to be observed.
381
Do plastics affect the speed of movement of plasmids?
Plotted concentration of plastics (x-axis) against transfer frequency (y-axis). The presence of plastics increases the transfer frequency of plasmids. This links environmental degradation with anti-microbial resistance.
382
Presence of the antibiotic doesn't drive the transmission of plasmids necessarily, but it does select for resistance.
In the gut of chickens, IncX3 with NDM-5 is transferred between bacteria without the antibiotic being applied.
383
Why was tigecycline resistance emerging in China, when China doesn't use tigecycline?
The low-grade antibiotic tetracycline (used in fish, chickens and pigs) is driving resistance to tigecycline via the tetX4 resistance gene carried on a plasmid. ISCRs are resposnisble for the dissemination of tigecycline resistance in E. coli in China.
384
MCR Gene
Mobile Colistin Resistance. Modifies the lipid A portion of the membrane in E. coli and other bacteria.
385
Why did the MCR gene increase in frequency in 2008?
Colistin was used in farms.
386
Why, when culturing strains, was the presence of the MCR genes much lower than in detection by presence of DNA?
The strains were sick. MCR genes massively affect bacteria by modifying their outer membranes, which is toxic to bacteria.
387
Most resistance genes in bacteria don't affect the bacteria themselves.
E.g., Klebsiella pneumoniae can survive with 10% of its genome being foreign DNA.
388
Some resistance mechanisms are a balance between survival and resistance to the antibiotic.
The toxicity of the MCR genes was tested by using a promoter to over-express the mcr-1 gene, which caused E. coli to burst, and become non-functional.
389
When was colistin banned as a growth promoter?
In April 2017 in China. The European Medicine Agency and America later followed suit.
390
Colistin resistance dropped significantly and quickly after the ban.
This wouldn't have worked so effectively with any other antibiotic. This drop was due to the fitness costs of MCR genes.
391
mcr-1
Index case. MCR genes are present in many plasmids.
392
Plasmids could come togetehr to make a super plasmid.
It would be full of antibiotic resistance genes and jumping genes. Any bacterium in possession of it would be resistant to all antibiotics on the planet, except the experimental ones.
393
394
Reproductive Value and Life History Theory
395
396
Darwinian Demon
An organism that does everything well. It lives forever, reproduces frequently and successfully and develops into an adult quickly. If it existed, it would have outcompeted all other species.
397
3 Drivers of Fitness
Survival, reproduction and development.
398
Blooms of Duckweed
close to being a Darwinian Demon, but it doesn't live long.
399
How does natural selection optimise fitness?
By optimising trade-offs.
400
Life History theory
Study of how an organism's traits influence fitness at different stages of its life, from birth to death. Founded by George Williams et al..
401
Life History Traits:
Size at birth, growth pattern, age at maturity, number and size of offspring, life span and sex of offspring.
402
Size at Birth
Joeys live in their mother's pouch for months-- tiny and defneceless. Altricial species. Whereas, the Ugandan Kob produces offspring that are abel to run, graze and socialise-- almost independent of each other. Precocial.
403
Precocial Species
Produce ready-to-go offspring. Opposite of altricial. Altricial to precocial is a continuum.
404
Growth Pattern
Determinate and indeterminate growth patterns. The sit-and-wait growth pattern. Some species even have a growth pattern of climbing to grow as tall as possible, e.g., Lianas.
405
Determinate Growth
Determinate growers finish development and growth at maturity.
406
Indeterminate Growth
Indeterminate growers don't stop growing at maturity, e.g., corals.
407
Sit-and-wait Growth Patterns
Seedling-looking plants can be 50 years old as they are opportunits waiting for a tree to fall, so light becomes available.
408
Age at Maturity
Aphids give brith to pregnant daughters, so the age at maturity is in utero. Wheras, the agave century cactus (not a real cactus) can wait 100 years before it reproduces.
409
Semel Parity
The strategy in which organisms only reproduce once, then they die.
410
Number and Size of Offspring
Orchids are the plant equivalents of clams with many offspring. Whereas, kiwis produce 1 large offspring in a clutch.
411
Sex determination is temperature-controlled in reptiles.
In Crocodilians, this follows a normal distribution, which can be male- or female-biased.
412
What is the shortest vertebrate lifespan?
The seven figure pygmy goby that lives for just 3 weeks.
413
Bristle Cone Pine Tree
One in the U.S. is 5000 years old. Dated using tree rings.
414
Silene stenophylla
Has the longest lifespan, if counting from when the propagule (here, the seed) was set. Seeds obtained from the last glacial 32,000 years ago germinated to produce viable plants.
415
Philopatric
An organism that always returns to breed in the same place.
416
Life Tables
Can be used to calculate fitness. Individual life history strategies determine population growth rates.
417
Average Reproductive Lifetime Success
Sum of lxmx across age classes, i.e., the sum of the R column in a life history table is net reproductive success.
418
Reproductive Value
The average contribution to the ancestry of future generations made per individual for each age class.
419
dx=
Dx/Sx
420
mx
Offspring per capita.
421
lx=
Sx for a particular year/ Sx in year 0. Survival to a particular age.
422
R0
Like with Covid-19, the contribution of an organism to a population throughout its lifetime. How amny daughters a female has in her lifetime.
423
R0 = 1
The population is stable.
424
R0 <1
The population is decreasing.
425
R0 > 1
The population is increasing.
426
R. A. Fisher: future reproduction.
Traits of individuals can be used to predict the contribution of individuals to a population in terms of reproduction. These ideas were misused in eugenics.
427
When fishing, which age groups are targeted?
Age groups where the fish have a low Rv.
428
In a diploid, sexually-reproducing population, how does the genetic value of males and females compare?
The genetic value of males and females is equal because the sex ratio is equal.
429
The Lack Clutch Size
When Rv (reproductive value) is maximised. A balance between egg size and egg number.
430
The action of natural selection is proportional to…
Rv (reproductive value).
431
Natural selection should select for older individuals in the clutch according to the Rv curve.
Parent birds tend to sleectively feed the older offspring in tough conditions as they have a higher Rv than the younger offspring.
432
Humans have evolved to provide…
… parental care.
433
Average Rv for Newborn Giant Clams
Much lower than for humans, so an individual offspring is less valuable to its parents.
434
Some clams have evolved a fih-like structure.
The structure is 'preyed upon' by fish, so the clam larvae becomes trapped in the fish gills.
435
The strength of selection peaks in early adulthood.
Deleterious mutations that are expressed at this age are more likely to be selected out as this is when Rv is highest.
436
Menopause
Human females continue to live beyond the point at which Rv drops to 0.
437
The Grandmother Hypothesis
Whenever the grandmother lived in the same area as the mother, the mother's fecundity and the number of grandchildren increased.
438
Integrative Fitness
The fitness of an individual increases, if the number of its grandchildren increases. Also applies to cooperative breeding, e.g., in wasps.
439
440
Molecular Evolution I and II
441
442
Genetic Drift
Stochastic changes in allele frequency (due to chance). Faster in smaller populations.
443
Positive Selection
Causes alleles that confer a beneficial phenotype to increase in frequency, e.g., selection for low body armour in freshwater sticklebacks.
444
Negative/Purifying Slection
Causes deleterious alleles to decrease in frequency.
445
Selection
Deterministic loss of low-fitness alleles.
446
Both slection and genetic drift…
… decrease genetic diversity. Mutation increases genetic diversity.
447
Population Genetics in the 1950s:
Most mutations were found to eb deleterious, low levels of genetic diversity in populations and occasional rapid selective sweeps by beneficial mutations.
448
Migration Rate (in protein electrophoresis):
A molecular phenotype of a protein as aminoa cid variation causes proteins to migrate at different speeds.
449
Allozyme Electrophoresis
First used to examine mutations in natural populations, but used proteins instead of DNA. Different charges of proteins affect the distance they move.
450
Heterozygotes
Have two bands in allozyme electrophoresis.
451
Limitation of Allozyme Electrophoresis
Not all mutations change the charge of the protein.
452
What did allozyme electrophoresis surveys reveal?
High levels of genetic diversity in populations, e.g., the alcohol dehydrogenase F allele.
453
Clock-like Divergence between Species
Conserved sequences of amino acids indicate descent from a common ancestor. Proteins exhibit a clock-like divergence over time due to mutation. There's a strong correlation between molecular divergence time and fossil divergence time.
454
Dayhoff Distances
Indicate the similarity of a protein from another species to the human version.
455
What does the neutral theory of molecular evolution aim to explain?
The divergence between species, and the polymorphisms within species.
456
The Neutral Theory of Molecular Evolution
Variation within species and divergence between species is chiefly the result of neutral mutations. Genetic drift drives molecular evolution. Selection at a molecular level is principally a purifying force removing deleterious alleles. Beneficial mutations exist, but they are so rare that they can be ignored.
457
Mutation Rate
µ/allele/generation.
458
What are the possible fates of every allele?
Every allele either goes extinct (0) or to fixation (1) due to genetic drift over time.
459
Probability of Fixation=
Allele frequency.
460
What is the likelihood that new mutations are fixed by drift?
A new mutation starts out as a single copy. Population size= 2N (2 because it's diploid). Frequency of the new mutation= 1/2N. Popbability of fixation by drift = 1/2N. Hence, almost all new mutations are lost.
461
Time to Fixation of a New Mutation
4N
462
Why is diversity greater in larger populations under the neutral model?
Slower drift.
463
Neutral evolution should occur at a constant rate= µ.
Rate of evolution due to drift = rate of appearance of mutations x probability of fixation = 2N*µ * 1/2N= µ
464
Rate of Divergence between Species
Neutral mutation rate x time (in generations).
465
Why don't deleterious mutations affect divergence between species?
Deleterious mutations are removed before they cn contribute to the diversity of the population.
466
Selectionist Interpretation of Evolution
Fixed differences and polymorphisms reflect positive sleection for beneficial mutations.
467
Neutralist Interpretation of Evolution
The vast majority of polymorphisms and fixed differences reflect genetic drift of neutral mutations.
468
The Molecular Revolution
Allozyme electrophoresis (demonstrated high diversity in populations) and clock-like divergence between species.
469
Global Surveillance of the Influenza Virus
Indicate mutations in the hemagglutinin gene sweeping to fixation because these mutations allow the virus to escape human antibodies.
470
Functional Characterisation of Mutations
Enables the identification of beneficial mutations.
471
The spike protein mutation D614G alters SARS-CoV-2 fitness.
The SARS-CoV-2 genome was modified to have this mutation, then grown on plates to measure replication in the mutated strain compared to a non-mutant. An example of functional characterisation of mutations.
472
Parallel Evolution
The same mutation is fixed in different populations. More parallel evolution than expected by chance alone (as genetic drift fixes mutations and is stochastic) is indicative of positive selection.
473
Parallel Evolution Experiment
A single clone of E. coli was grown in a tube deficient in nutrients for 24 hours. The some of the E.c oli is trasnferred to the next tube. This process repeats. There have been 12 separate repeats of this whole experiment since 1988. Some of the E. coli was cryogenically frozen, so could be sequneced later.
474
How did the E. coli experiment involve parallel evolution?
The same genes with different utations had similar consequences. E.g., the ribose gene was knocked out due to different mutations in each population as there was no ribose in the medium for E. coli to catabolise.
475
Parallel Evolution of Resistance to Milkweed Plant Toxins among Insects
The exact same mutation in the monarch butterfly, red milkweed beetle, large milkweed beetle and Oleander aphid that affects the Na+/K+ pump confers resistance to the milkweed toxins. It's not just the same gene; it's the same nucleotide substitution across the 4 different species.
476
Limitations of Parallel Evolution
Requires a statistically null hypothesis-- how much is due to random genetic drift? It's good at picking up genes that are under strong selection, but doesn't capture a lot of adaptation.
477
Synonymous Mutations
Typically in the 3rd position in the codon. Don't change the amino acid. Likely to be neutral. Indicate the effects of random genetic drift.
478
Non-synonymous Mutations
Change the amino acid and protein. May be neutral, deleterious or beneficial. Indicate the effects of selection.
479
dN
Rate of non-synonymous substitutions.
480
dS
Rate of synonymous substitutions.
481
No Selection
dN/dS = 1
482
Purifying Selection
dN/dS < 1
483
Positive Selection
dN/dS > 1
484
Drosophila yakuba and Drosophila simulans
6 million years of divergence. 45% of non-synonymous differences are due to positive selection. 270,000 selected differences with one selective sweep every 45 years.
485
There have been at least 8 million years of divergence between humans and gorillas.
30-1% of non-synonymous differences are due to positive selection.
486
Why is there more positive selection between more closely-related species than more distantly-related species?
Over time, as the less closely-related species have diverged longer ago, the dN will be swamped by the dS as there have been so many synonymous mutations.
487
Limitation of comparing rates of evolution at synonymous and non-synonymous sites:
It only picks up genes under recurrent selection.
488
Neutrality provides a null hypothesis:
What would genetic variation look like without selection?
489
Innovation
New traits that allow organisms to exploit their environment in new ways. Important in the context of major transitions and individuality.
490
Enzyme Promiscuity
Proteins are dynamic with conformational flexibility. Enzymes have a primary reaction that they catalyse, as well as another ligand that can bind to it in a less-preferred conformation.
491
Why is enzyme promiscuity important?
A mutation can mean that the enzyme switches, so that its less-favoured reaction becomes the primary reaction that it catalyses.
492
Directed Protein Evolution
Mutagenic PCR of a gene. Artificially select for E. coli that best produce the protein encoded by the gene of interest. Repeat the PCR amplification.
493
Regulatory Re-Wiring
A gene (FleQ) was deleted. This meant the FleQ regulator wasn't produced, so the flagellar operon wasn't transcribed. Within 1 week, they had re-evolved the ability to grow a flagellum. Mutations switched the NtrC binding, so the Tf could bind to and regulate both the flagellar structural genes and the nitrogen assimilation genes. Eventually, it evolved only to regulate flagellar structural genes.
494
Why was there such a strong sleection pressure for the bacteria to regian motility, after the FleQ was deleted?
They couldn't swim, so used up their resources.
495
How are new genes acquired?
HGT. Plasmids are semi-autonomous DNA molecules that are replicated and transferred across the conjugation pilus. Plasmid genes can be incorporated into the bacterial chromosome.
496
HGT is common in bacteria.
Up to 90% of some bacterial genomes come from HGT. the lac operon genes were inly recently acquired by E. coli via HGT.
497
How does HGT drive lots of antibiotic resistance?
Plasmids carry antibiotic resistance genes from environemntal bacteria into gut bacteria in the gut microbiome.
498
Bdelloid Rotifers
Have acquired genes for protection against fungal pathogens.
499
Genes Involved in Eukaryotic Immune Systems
Acquired from bacteria, liekly very early in eukaryotic history.
500
EPAS1 Allele
Hypoxia-induced. Enables adaptation to low-oxygen on the Tibetan plateau. This allele was acquired from the Denisovans via hybridisation.
501
Where do new genes come from?
The origin of new genes is thought to be gene duplication. Duplications occur just as frequently as mutations at a single site do.
502
Most duplicates are inactivated by mutation, but sometimes there is neofunctionalisation.
Duplication relieves the selective pressure on the gene. There is amutation in one of the duplicated genes. Duplication means the fucntion of the original gene is retained, and a new function is acquired.
503
What have been important for body plan regualtor genes sucha s Hawk's genes?
Duplication and divergence.
504
505
Climate and Life on Earth: the Terrestrial biosphere
506
507
Climate plays a role in…
… determining the distribution and cahracteristics of ecological communities at the global scale.
508
Annual Average Temperatures
Higher at the equator. Lower near the poles. Variation within a latitudinal band due to topology and elevation.
509
Precipitation is less tied to latitude.
The wettest areas on Earth areWest africa, tropical rainforests in South America and Southeast Asia.
510
Primary Influencers of Biome Dsitribution
Temperature and precipitation.
511
Biomes
Zones dominated by plants with characteristic shapes, forms and physiological processes.
512
Where is Mediterranean vegetation found beyond the Mediterranean?
The U.S. west coast, the tip of South Africa and parts of Australia due to similar cliamte with high light levels and propensity to drought. E.g., Garrigue in southern France, Chapparal in California and Fynbos in South Africa.
513
Whittaker Biome Classification
Plots different biomes, e.g., tundra, on a graph with annual precipitation (y-axis) against average annual temperature (x-axis).
514
Tundra
Low shrub vegetation at high latitudes. Relatively species poor.
515
Mount Kilimajaro
At the equator in Tanzania. From the lowest to the highest elevation: civilisation, rainforest, moorland, alpine desert and eternal ice.
516
Primary Productivity
Rate of energy production per unit area. Can also be measured as mass of carbon, or dry organic matter.
517
Secondary Productivity
The biomass produced by consumers.
518
Gross Primary Productivity (GPP)
The total fixation of energy by consumers. Some of the GPP will be lost as heat due to respiration by the autotroph, but that which isn't lost is Net Primary Productivity (NPP).
519
What is the largest biome by global surface area?
The ocean.
520
How much of global NPP is contributed by tropical rainforest?
Almost 20%.
521
Algal Beds and Coral Reefs
Have higher annual NPP than tropical rainforests, but cover a smaller surface area.
522
Where is NPP highest?
Tropical rainforests and surrounding Savannah. It’s also quite high in temperate zones, too.
523
What limits NPP in terrestrial ecosystems?
The intensity of solar radiation received, the availability of water, tempearture and the availability of nutrients.
524
What % of potential NPP have humans appropriated?
23.8%, of which 53% is from harvesting (this includes plats for feeding livestock, as well as harvesting crops and timber), 40% is from land-use-induced productivity changes and 7% by human-induced fires.
525
Why do different regions of Earth receive different amounts of solar radiation?
Earth's tilting. Seasonal variation.
526
Where is photosyntehtic efficiency highest?
Coniferous forests. Higher than in deciduous forests or deserts. Plants in different regions have different degrees of efficiency in using this solar radiation.
527
Where are effects of seasonality increased?
Away from the equator.
528
Winter at Very Northern Latitudes
Water is inaccessible as it's locked up in snow and ice. There's also less light in winter. Productivity is only possible when plants can photosynthesise.
529
What tend to be limiting nutrients for plant growth?
Phosphorous and nitrogen.
530
Why is phosphorous limiting in rainforests and/or tropical regions?
The soil has been weathered, and iron-oxide in the soil tends to bind to phosphorous, limiting its availability.
531
Limiting Nutrient Distribution
Nitrogen is limiting, or both nitrogen and phoshporous are co-limiting, except for in tropical regions and/or rainforests where phosphorous is limiting.
532
533
Primary Production in the Oceans
534
535
Continental Shelves
Have shallow gradients and high productivity. Impacted greatly by humans.
536
Abyssal Plain
4-5 km down. Little is known about it.
537
Mid-Ocean Ridges
Seafloor spreading at a constructive plate boundary. Volcanically active areas. Can include depp-sea vents.
538
Deep-Sea Vents
Driven by the heat and chemical gradients of Earth instead of by sunlight. Source of critical minerals.
539
How do seamounts form?
When the plume of magma is separated from the vent, it sinks to form a seamount.
540
Hotspots of Volcanic Activity
Form islands.
541
Mariana Trench
7km deep.
542
Thermocline
Transition layer between the warmer surface waters and the coller waters below. Sharp reduction in temperature down to 1000m deep. It then remains that temperature until the abyss. Established in the tropics (permanent feature here) and in summer at temperate latitudes. Less-dense warm water prevents vertical mixing, leading to nutrient depletion in surface waters.
543
How much less O2 is there in a litre of water than in a litre of air?
40x less. It also diffuses ~1,000x slower.
544
Oxygen Minimum Zones
~200m -1,500m. Respiration exceeds photosynthesis and diffusion of O2 from the air. O2 is less oluble in water than CO2.
545
Why is there more vertical mixing in coastal waters generally?
Due to waves that can mix the entire water as the water is shallower.
546
Euphotic/Sunlight Zone
Sea level-200m deep. Only zone in which photosynthesis occurs, and most of it in the top 50m, as sunlight seldom penetrates beyond this zone. Home to tuna.
547
Dysphotic/Twilight Zone
200m-1000m deep. Sunlight decreases rapidly with depth. Photosynthesis is not possible here. Home to shrimp, hatchet fish and swordfish.
548
Aphotic/Midnight Zone
Below 1000m deep. No light penetration at all. Home to giant squid, angler fish and tripod fish.
549
Which wavelengths of light penetrate deepest?
Blue-green.
550
Why doesn't light penetrate as deep in coastal waters?
More turbidity due to river mouths.
551
How much of human emissions since the industrial revolution have been absorbed by the oceans?
~1/3. It is estimated that ~1/2 of current emissions continue to be absorbed.
552
Oceans are still at ~pH 8, but they are less alkaline due to acidification.
Seawater reacts with water to produce carbonic acid that dissociates into bicarbonate ions and H+ ions. These additional H+ ions react with existing carbonate ions in the water forming further bicarbonate.
553
Why does the removal of carbonate ions from the water by additional H+ ions pose a problem for organisms with CaCO3 skeletons?
They rely on saturating concentrations of carbonate ions. At a pH below 7.5, this becomes critical.
554
What creates currents?
Temperature gradients and wind.
555
The Gulf Stream
Carries warmer water from the Caribbean to the UK.
556
Ice Formation
Creates dense, hypersaline water as the ice is pure, salt-free water.
557
The Great Ocean Conveyor Belt
Redistributes nutrients. Slow-moving currents. As deep water flows, it picks up nutrients from the bottom of the ocean, bringing them to the surface. This fuels hotspots of productivity.
558
Large Areas of Tropical Oceans
Virtual deserts. Lack of primary productivity due to the permanent thermocline under the constant sun.
559
Roughly half of the world's primary production takes place in the global ocean:
50 billion tonnes of crabon are fixed every year.
560
2 Main Producers in the Open Ocean
1. Bacterioplankton (30-50%): cyanobacteria. 2. Phytoplankton. 3 major groups: the diatoms, the dinoflagellates and the coccolithophores.
561
What comprises the 70 Gt of carbon in the deep sub-surface of the Earth?
Bacteria and archaea.
562
Biomass of consumers in the marine environment…
… 5x the biomass of producers.
563
Although the majority of the biomass is in the terrestrial biosphere, what is the marine biomass made up of?
2 Gt of protists, 2 Gt of animals, 0.5 Gt of plants and 1.5 Gt of bacteria.
564
Marine Food /chains
Typically longer than terrestrial food chains due to more efficient energy transfer. Food chain length can vary hugely, e.g., upwelling that support large phyotoplankton communities enable lots of grazing by fish etc..
565
Where is productivity highest in the oceans?
The coast/continental shelves, and the Arctic. Hence why lots of animals migrate to the Arctic.
566
Microbial Loop
Recycles dissolved organic matter (DOM) in the surface waters.
567
Why can dissolved organic matter be so massive?
The prevalence of viruses in seawater-- up to 10 million per ml of sewater.
568
Zooplankton carry out the planet's largest migration every day.
1000m every 24 hours. They transfer carbon from the surface to the bottom of the ocean by moving up to eat and down to defecate. This is part of why human-emitted CO2 is dissolved in the oceans. The zooplankton excrete tiny faecal pellets that form particuklate organic matter (POM) that sinks, but nutrients are rapidly returned by heterotrophic bacteria.
569
What makes temperate and polar waters nutrient-rich?
Absence of a thermocline in winter. Strong vertical mixing.
570
Temperate latitudes have high amrine productivity, but when do the productivity peaks occur?
A major peaks occurs in spring, often followed by a second smaller peak in autumn.
571
What limits ocean productivity?
Rarely CO2. Often light in temperate and polar regions.
572
Peruvian Anchoveta
Most heavily exploited fish worldwide.
573
How do upwellings form?
Longshore winds on the western edges of continents draw up the cold, nutrient-rich water from the deep.
574
How are hotspots of high productivity created in coastal waters?
Upwellings of nutrient-rich water. Common along the west coasts of continents, e.g., on the west coasts of California, Chile and Peru.
575
What % of the human population lives within 100 km of the coast?
~ 60% of the human population, i.e., 4 billion people.
576
What causes upwellings to shut off?
El Niño bringing warm water.
577
Coastal Seas
Lie above the continental shelf, highly productive due to strong vertical mixing, heavily exploited and heavily polluted, e.g, agricultural runoff creates dead zones that expand each year, and oil spills are a constant threat.
578
Where are kelp forests abundant?
At mid-latitudes. They are ecologically unstable, and sensitive to harvesting of key animals.
579
Kelp is economically important.
It is harvested directly, and supports important fish, such as rockfish in the U.S. and shellfish such as European lobsters.
580
Brown Algae
Not true plants.
581
Red Algae
True plants.
582
Features of Kelp
A multicellular, brown algae with no true vascular system. Often annual.
583
Dominant Kelp Genera
Laminaria and Macrocystis.
584
Kelp can differentiate into multiple cell types:
A holdfast, stipe and baldes.
585
How many species can live within just a single kelp holdfast?
30-70!
586
Kelp can be highly productive.
Macrocystis can grow up to 30cm per day to a height of 30m, forming a dense underwater forest.
587
The Kelp Stipe
Supports epiphytes, e.g., red algae.
588
Red Algae Communities Beneath Kelp
40-180 species around the UK. Supports many invertebrate species such as brittle stars, anemones, crabs and jellyfish.
589
Why do only a few animals graze directly on the living kelp?
It has very high C:N ratio, and conatins compounds that deter grazers.
590
Blue-Rayed Limpet
Specialist grazer of kelp. Found at low tide in the UK.
591
Urchins
Grazers of kelp that specialise on eating their holdfasts. If present in large enough numbers, they can prevent kelp from regnerating after a disturbance.
592
Giant Kelp
Macrocystis pyrifera: supports important crab, lobster, abalone, snail fisheries and kelp harvesting.
593
Kelp Forest
A major, largely temperate coastal ecosystem.
594
Study on a kelp forest in Mexico:
Constructed a food web based on 10 years of monitoring of 40 different functional groups. Bottom-heavy biomass pyramid. Estimated throughput: 5477.6 t.km^-2. year^-1.
595
Urchin Barren
High population density of urchins converts a kelp forest into an urchin barren.
596
What can cause urchin populations to boom?
They have very high reproductive rates, so can increase in population density quickly. Often top-down controlled.
597
Sea Otters
Enhydra lutris. Feed on shellfish and echinoderms. Hunted fo fur until 1911, when hunting of sea otters was banned,a nd their population had plummeted to just 2000 individuals.
598
Sea otters are a keystone species.
As their population recovered and their range expanded, sea otters controlled the urchin population, allowing kelp forests to recover.
599
Why have kelp forests collpased in the past 20 years in northern California?
Warmer waters, strong El Niño events that blocked coastal upwellings (this was an issue as kelp requires a constant supply of nutrients from deep water-- bottom-up control) and a disease of the sunflower sea star.
600
Sunflower Seastars
Target medium-sized urchins. Even when the seastar population collapsed, otters helped to control the urchin population.
601
602
Primary Production in the Oceans: the Tropics
603
604
3 Major Types of Coastal Tropical Ecosystems
Mangroves, seagrass meadows and coral reefs.
605
What do mangroves and seagrass meadows have in common?
They are both dominated by angiosperms. Both have high biomass of primary producers, but not many producer species, and actively sequester huge amounts of carbon.
606
Seagrass meadows support high biodiversity.
They have both bottom-up and top-down controls on their productivity.
607
Threats to Coral Reef Biodiversity
Overfishing, coastal pollution and rising sea temperatures.
608
Key Difference between Coral Reefs and the Other Two Coastal Tropical Ecosystems
Coral reefs have a much higher number of producer species.
609
Mangrove Trees
Angiosperms that grow in coastal brackish or saline habitats.
610
True mangroves belong to what genus?
Rhizophora.
611
Mangrove trees trap sediment around their roots, creating anoxic conditions. How are the trees adapted to cope?
They have lenticels (pores for gas exchange) or pneumatophores (act as conduits to provide O2 to the roots).
612
How do mangroves cope with salt?
Salt is mostly excluded by the roots. Some species have the ability to excrete salt via specialised glands in their leaves.
613
Where are seagrass meadows found?
In both temperate and tropical regions. Temperate bed are typically species-poor, and often dominated by eelgrass (Zostera marina). Tropical beds are more species-rich.
614
How many species of seagrass globally?
72 species.
615
How does seagrass differ from algae?
Alage have a holdfast to attach to a surface, whereas seagrass have roots and a vascular system, so can take up nutrinets from the sediment.
616
How does seagrass spread rapidly?
Rhizomes enable asexual reproduction.
617
Features of Seagrass
No stomata, but athin cuticle. They can produce pollen and seeds, and they can flower. Flowers are pollinated by Crustaceans.
618
What directly grazes seagrass?
Sirenians (dugongs and manatees) and green sea turtles, which have green flesh as all they eat is seagrass.
619
At what depths does seagrass grow?
Up to 50m deep, but mostly < 30m.
620
Small-bodied Seagrass
Have a guerilla strategy: they can colonise new areas as they are fast-growing, and have a seed bank to disperse seeds.
621
Large-bodied Seagrass
Phalanx strategy. Fewer, larger seeds. Grow more slowly, and have larger leaves, so they're better competitors, and better at holding down space.
622
Tropical Seagrass Meadows
High biomass systems with high productivity. Important nursery grounds for commercially crucial fish species. Grow in coastal areas, incl. the intertidal zone. Can cope with brackish water and some water turbidity.
623
Why do seagrass meadows often grow alongside reefs?
They protect the reefs by trapping sediment, and gain protection from wave action as reefs absorb the energy from the waves.
624
Mangroves and Coral Reefs
Mangroves filter and trap sediment, so light can reach coral reefs.
625
What happens, if segarass meadows take up extra nutrients from eutrophication?
The water column stays free.
626
What indicates seagrass are bottom-up controlled?
If they respond to addition of nutrients.
627
What indicates seagrass is top-down controlled?
If herbivores affect the seagrass biomass.
628
Seagrass biomass is both bottom-up and top-down regulated.
Bottom-up: if only seagrass was present, then seagrass biomass increased with nutrient addition. Top-down: if there were no predators, so only seagrass and herbivores, then the response to nutrient addition was dramatically reduced. If all three were present (full food web), predators controlled teh herbivores, allowing seagrass to mop up excess nutrients.
629
Procedural Control: Cage Control.
Caging can be sued to kepe predators or herbivores out of the seagrass ecosystem.
630
Van der Zee et al., (2016) examined the effects of non-trophic interactions on seagrass ecosystems via a successional sequence.
Successional sequence: 1. Bare areas with a coarse, sandy substrate, often brought by an estuary. 2. The seagrass starts to colonsie the bare area, and traps silt and sediment. 3. After >40 years, established areas of seagrass with 90cm-thick silt and pools created by burrowing crabs. Climax community: seagrass and crabs.
631
Primary and Secondary Habitat Modifiers in the Successional Sequence
Primary Habitat Modifier: seagrass. Secondary Habitat Modifier: the burrowing crabs. Removing habitat modifiers impacted species richness at differents tages of the successional sequence.
632
Coastal Wetlands
Mangroves and seagrass meadows.
633
What % of coastal wetland carbon is found in the soil?
50-90%, depending on vegetation type.
634
How do coastal wetlands sequester carbon?
Tidal inundation keeps the soil wet or submerged, inhibiting microbial action and slowing decomposition. The roots penetrate deep into the substrate, trapping carbon such as in the form of decaying matter. The fine particles of silt create an anoxic environment in which organic matter doesn't decompose. Destroying coastal wetlands would release lots of CO2 into the atmosphere.
635
Coral, Kelp and Phytoplankton
Can't store carbon as long-term as coastal wetlands.
636
Deep Water Reefs
Different to tropical coral reefs. Found in cooler waters.
637
Why don't coral reefs grow in areas with major river outflows?
Coral is sensitive to feshwater and silt.
638
Where are coral reefs found?
The Indo-Pacific and tropical west Atlantic. The Indo-pacific has ~10x the biodiversity of the west Atlantic.
639
What conditions do corals require?
high light intensity, so low turbidity and oligotrophic waters. Clear waters, temperatures 18-30°C and no sudden changes in salinity.
640
Fringing Reef
Coral reef that starts growing on the shore of a high volcanic island in shallow water.
641
Barrier Reef
Forms from a fringing reef that is pushed upwards and outwards by tehd eposition of sediment, as the island begins to erode and shrink.
642
How does a coral atoll form?
As the volcanic island continues to erode, the lagoon between the barrier reef and the volcanic island becomes wider and deeper. The peak ultimatley sinks below the water, leaving only the encircling reef behind.
643
Reef-building Corals
Scleractinian corals within the class Anthozoa within the phylum Cnidaria. The ectoderm contains cnidocytes (stinging cells), the endoderm often contains photosyntehtic algae and in-between is the mesoglea.
644
Lots of marine calcifying organisms are photosynthetic.
Photosynthesis may aid the calcification process.
645
Why do corals have cnidocytes?
Corals compete for space, so corals attcak each other with their stinging tendrils at night.
646
Corallite
Protective CaCO3 skeleton.
647
Coeonsarc
Tissue that connects a coral colony. Coral colonies can grow for years, or even decades. Coral can also exist as polyps.
648
Zooxanthellae
Photosynthetic dinoflagellate algae belonging to the genus Symbiodinium.
649
Symbiosomes
Specialised membranous structures that separate the symbiont celsl from the host cytoplasm.
650
What % of the biomass of the polyp does Symbiodium constitute?
Up to 30%. Can slo be found free-living at low density in tropical waters.
651
What does the coral provide the Symbiodinium?
Protection and CO2 from respiration.
652
What does the Symbiodium provide the coral?
Much of its carbon and nitrogen (when not enough is obtained from heterotrophy). It also aids the calcification process.
653
How does the Symbiodium provide the coral with fixed nitrogen?
The symbiont produces amino acids from dissolved inorganic nitrogen (DIN) form the water column and nitrogenous waste produced by the coral.
654
When its feeeding rates are low, what % of nitrogen does the coral acquire from the symbiont?
More than 80%.
655
Diazotrophs
Nitrogen-fixing bacteria involved with coral. Its role is unknown as yet.
656
How much of the carbon fixed by their symbionts is channeled by the coral into its own growth?
~90%.
657
Algae on reefs provide a turf for grazing fish.
Many different types of algae are found on reefs such as coralline red algae, fleshy green algae and brown algae. Graizng fish are herbivorous, e.g., sturgeonfish, which, in turn, support predatory fish.
658
Coral reefs take up minimal space, but are highly biodiverse.
Despite taking up only 1% of the ocean floor, they support 25% of marine life.
659
In 99 countries, what % of fisher-people fish on reefs?
25%.
660
Fishing on Reefs
Starts by targeting top predators, such as groupers and sharks, but these are easily overfished. Then fishing down the food chain begins: smaller, herbivorous fish such as parrotfish are targeted.
661
What destroyed lots of coral reefs in the Caribbean?
Chronic overfishing, sea urchin boom, hurricane damage and sea urchin die-off.
662
1. Chronic Overfishing
Began by targeting predatory fish, then moved in to herbivorous fish, incl. parrotfish. Parrotfish tend to feed on fleshy algae that grows in-between the coral. The algae and coral compete for space, so if the algae grow tall, they will block the light from the coral.
663
2. Sea-Urchin Boom
Massive numbers of long-spined urchins, Diadema antillarum, kept the fleshy alage in check, depsite the loss of herbivorous fish.
664
3. Hurricane Damage
Destroyed much of the Acropora cervicornis species in 1980. There was a short-lived coral recovery afterwards.
665
4. Sea-Urchin Die-Off
1982-84. A pathogen caused the death of 995 of urchins. No herbivorous fish or sea urchins to keep the algae in check. Huge nutrient run-off as well. The benthic algae grew out of control, so the coral couldn't recover.
666
Coral Bleaching
The reaction of coral to sudden but prolonged high temperatures. Corals expel their symbionts. UV also contributes. Corals don't always die due to bleaching, but if the warming is prolonged, they cannot recover. Some algae grows on the surafce of the bleached coral.
667
When was the first recorded bleaching event?
1998
668
Why are symbionts expelled?
Unclear as yet. It could be because the symbionts become selfish by stopping exchanging nutrients with the coral, or could be due to oxidative stress as the Symbiodium produces reactive oxygen and nitrogen species.
669
A phase-shift occurred in Caribbean reefs.
Reefs can exist in one of two stable states (e.g., reef-building coral or fleshy algae, hard coral or macroalgae).
670
What is required for reef recovery in the Caribbean?
Stronger management of fisheries, and reduced nutrient run-off. Caribbean reefs have still not recovered.
671
1998 Seychelles Bleaching Event
Destroyed reefs around the inner islands. They have not recovered.
672
Bleaching events are linked to El Niño/La Niña.
Part of a large-scale climate phenomenon that occurs every 3-8 years, and takes months to spread across oceans. In addition, ocean temperatures are rising.
673
Bleaching of the Great Barrier Reef in Australia
Half of the Great Barrier Reef has been bleached to death since 1995. the average interavl between bleaching events has halved between 1980 and 2016. There were bleaching events in 2017 and 2020.
674
Aldabra Atoll
UNESCO World Heritage Site. Zero commercial fishing. Some corals are adapted to warming and are resistant to bleaching as it's already a thermally intense environment. Reduced coral bleaching, and faster recovery. Some coral that have experinced bleaching appaear to be more resistant to subsequent bleaching.
675
676
Large-Scale Patterns in Diversity
677
678
What are large-scale patterns in diversity?
Trends in the distribution of species across Earth's surface. Macroecology: species curves and latutudinal gradients.
679
Species Curves
How the number of species recorded increases as the area studied increases.
680
Latitudinal Gradients
The trend towards increasing diversity as you move from the poles towards the tropics.
681
The increase in the number of species as area increases is not linear.
It tends to follow a power function.
682
Typically, a tenfold increase in area…
… leads to a doubling in the number of species recorded.
683
Schoener, 1976.
Species area curves (log S= z log A + log c) is one of the few genuine laws of ecology.
684
Where do species area curves apply?
On islands of different sizes within an archipelago, for pieces within a single biome, e.g., states within the U.S. and for interprovincial areas, e.g., continents.
685
Who discovered species area curves?
H. C. Watson in 1859.
686
3 main mechanisms to explain the species area curve pattern:
1. The habitat diversity hypothesis. 2. The passive sampling hypothesis. 3. The equilibrium hypothesis. These are not mutually exclusive, but they do emphasise different processes.
687
The Habitat Diversity Hypothesis
If there is a wider range of habitat types, more species can live in the area. Larger areas have more habitats, and species idversity increases with habitat diversity.
688
Evidence for the habitat diversity hypothesis can be found where area and habitat diversity can be separated.
Keep habitat constant, and vary the area within sampling sites, or keep area constant, and vary habitat within sampling sites.
689
Mangroves-- a homogenous habitat.
Larger mangrive islands still have more species than smaller mangrove islands, even though habitat diversity is the same. Thus, habitat diversity doesn't always explain this pattern.
690
The Passive Sampling Hypothesis
Often used as a 'null model'. Assumes the probability a species is present on a island is proprtional to the island's area, and that islands sample individuals randomly and independently.
691
Limitations of the Passive Sampling Hypothesis:
Fails to explain the diversity on small islands. Nothing limits diversity: given time all species should be present on an island. Doesn't predict the turnover of species over time.
692
Turnover
Some species go extinct, some species arrive, hence the composition of species changes over time, even if still in equilibrium.
693
The Equilibrium Hypothesis a.k.a. The Equilibrium Theory of Island Biogeography
Closer islands will have more species than more isolated islands. Larger islands will have more species than smaller islands. MacArthur and wilson (1963, 1967). The number of species on an island would be a fucntion of its isolation and area.
694
Assumptions of the Equilibrium Theory of Island Biogeography
Evolution doesn't influence species richness. Isolated islands are less likely to be colonsied. Large islands have large populations. Large populations are less likely to become extinct. Colonisation and extinction are independent of the species composition of the island.
695
The equilibrium hypothesis assume evolution doesn't influence species richness. What is the issue with this assumption?
On a very isoalted oceanic island, e.g., the Galapagos where independent radiations within the archipelago have occurred, divergence leading to speciation may occur.
696
What does the Equilibrium Hypothesis predict?
1. There should be substantial turnover in species composition, but species number should remain relatively constant over time. 2. All else being equal, the number of species on an island should decrease with increasing isolation.
697
Birds on the Channel Islands in California
As per the Equilibrium Hypothesis, species richness was stable, but there was significant turnover in species composition.
698
Simberloff and Wilson (1969)
Defaunation: they killed off the invertebrates living in mangroves. Invertebrate species tehn recolonised from other islands. ~2 years later, the number of species reached the pre-defaunation number. This demonstrates hoe the number of species reaches a plateau.
699
Species-area relationships have been used for over 30 years in conservation.
Used to predict how many extinctions will occur following area reduction, and to suggest the best way to slow the loss of spceis by reserve design.
700
Assumptions of Species-area Relationship Use in Conservation:
Species richness is the main object of conservation, and that varaition in area is the main cause of species richness.
701
Z Value
The gardient of the line for a species area curve on log axes. The steeper the slope, the more rpaid the cahneg in species richness. For a given reduction in area, the graph indicates the corresponding change in species richness.
702
Species-area relationships have been accurate in predictions for forest birds in eastern North America.
52% of forest in eastern North America have been cleared in the past 200 years. Species-area relationships predicted a loss of 4.8 endemic species. 4 have gone extinct with a fifth species close to extinction.
703
Designing Nature Reserves/Protected Areas
Size matters: larger areas have more species. The SLOSS controversy: should you have a single large reserve or several smaller reserves with a similar area?
704
Alpha Diversity
The number of species on an individual island.
705
Gamma Diversity
The number of species across the whole archipelago.
706
Beta Diversity
Gamma diversity / alpha diversity. The turnover of species in space (previously, it was turnover over time). A measure of the differentiation of communities.
707
Solving the SLOSS controversy depends on beta diversity:
The degree to which species overalp between different areas. Many smaller islands or reserves can collectively have more sepcies than a larger, equivalent area.
708
Latitude of 0
At the equator.
709
Latitude of 90.
At the North Pole and -90 is the South Pole.
710
The habitats with the greatest biodiversity both on land and in the sea…
… are tropical.
711
Tropical Rainforests
Cover 7% of the Earth's surface. Account for >50% of Earth's biodiversity.
712
The universal ecological pattern of latitudinal fgradients holds for…
… different levels of the taxonomic hierarchy, e.g., genera, family etc..
713
Exceptions to the Latitudinal Gradient Pattern
Penguins (have higher species richness at the poles) and ichneumoid wasps.
714
Competing Explanations for latitudinal Gradients
Biotic explanations, e.g., productivity, competition and predation. Abiotic explanations, e.g., time and stability.
715
Explanations for latitudinal gradients, categorised by resource availability and usage.
A greater arnge of possible resources. Each species is more specilaised. More overlap in reosurce use. Resources are more fully exploited.
716
Why is explaining the latitudinal gradient so difficult?
Many of the explanations are circular, many explanations make similar, qualitative predictions, the data is often messy (from many sources), our sample size is small (1 planet) and doing experiments is difficult.
717
A greater range of posisble resources.
Productivity or Species Energy Hypothesis. The hottest, most humid places have the greatest net primary productivity. More resoucrsx can support more posisble species.
718
Issues with the Greater Range of Possible Resources Hypothesis
Tropical seas have low productivity, but high richness. Eutrophic lakes have high productivity, but low richness. High productivity doesn't always lead to high abundance or biomass.
719
Each species is more specialsied.
Similar resources, but each species is more specialised. If niches are narrower, more species can be packed into an environment. Constant climate and intense competition in the tropics leads to niche narrowing. Most species grow towards tehir carrying capacity.
720
Issues with the Each Species Being More Specilaised Hypothesis:
Comparative analysis of strength of competition and niche breadth are difficult.
721
More overlap in resource use.
Due to less intense interspecific competition. More intense mortality in the tropics due to natural enemies, instead of predators. Predator-mediated co-existence: population sizes of tehir prey are kept low, which reduces competition. This allows overlap in resource use, prevents any one species becoming locally dominant. Thus, many species can co-exist.
722
Resources are more fully exploited.
Communities diversify over time, and temperate regions are younger due to glaciation. Tropics have been relatively unchanged for at least 1.5 x 10^8, and display low seasonality. There are empty niches outside the tropics, but there has been no ecological time for species to re-invade, or evolutionary time for species to evolve to fill them.
723
Issues with the Resources Are More Fully Exploited Hypothesis:
Many exceptions where stable habitats have low diversity, e.g., tropical mountain tops. The deep ocean is a constant ocean with a low number of species. Many tropical environments have seasonal fluctuating environments.
724
Species-area relationships may partly explain latitudinal gradients.
At this scale, the number of species is a balance between speciation rates and extinction rates. Species present = species evolving - species going extinct.
725
Tropics may have lower extinction rates and increased speciation rates.
Lower extinction rates due to their larger populations and refuges they provide agianst environmental change. Increased speciation rates as geographic barriers facilitate allopatric speciation, and the greater chance of a rare mutation leads to more rapid evolution.
726
727
Freshwater Ecology
728
729
Freshwater habitats are rare.
Only 0.8% of Earth's water is a suitable freshwater habitat. Only 2% of the Earth's surface is freshwater, but much of that is locked up in permafrost and glaciers.
730
Freshwater habitats are physically diverse.
Lake Baikal in Russi is the deepest freshwater lake, and is home for the only species of freshwater seal. By contrast, hot springs, e.g., Hengill in Iceland are small and can be up to 100°C.
731
Freshwater habitats are hotspots for biodiversity.
Incl. microbes such as diatoms. Freshwater habitats are home to 25% of described mollusc species. 55% of fish species reside in freshwater for at least part of their lifecycle.
732
How many species of fish are found in the UK?
Native: 38. Introduced: 12.
733
How many fish species are found in Lake Malwai?
Up to 1,000.
734
Freshwater vs. Terrestrial Habitats
For animals, the number of species per km^2 in freshwater habitats is more than double that of terrestrial habitats. Phylogenetic diversity of terrestrial species also exceeds terrestrial species.
735
Freshwater habitats are threatened.
Since 1970, the population decline of freshwater vertebrates is 84% compared to 30-40% in marine and terrestrial ecosystems.
736
Ecosystem Services
Benefits to humans from the natural environment. Freshwater habitats provide drinking water, fish protein and flood protection.
737
What lives in freshwater?
Thousands of insect and bird species. 6 orders of mammals.
738
6 Orders of Freshwater Mammals
Sirenia: manatees, the freshwater manatee is the smallest species of manatee and lives in the Amazon. Cetartiodactyla: dolphins and hippos. Carnivora: otters, minks and seals. Rodentia: beavers, capybara and voles. Eulipotyphyla: shrews. Monotremata: platypus.
739
Living in water has evolved several times in mammals. How do we know?
These orders are spread out across the phylogenetic tree.
740
Freshwater Thermophiles
Bacteria, viruses and fungi. Live in hot springs. E.g., Hydrogenobaculum and brine flies.
741
Freshwater hot springs can be acific.
Such as those in Yellowstone that are more acidic than a car battery.
742
Challenges of Living in Caves
Darkness, little food and low oxygen.
743
Astyanax mexicanus
No scales, for efficient swimming, and no eyes in cave morphs, unlike the surface-dwelling morphs.
744
What do comparisons between cave-dwelling and surface-dwelling morphs reveal?
Cave dwellers have lower metabolism and increased red blood cell development.
745
How have some species adapated to survive in ephemereal (temporary) streams or ponds?
4 Ds: dormancy, diapause, die young (live fast) and dispersal.
746
Dormancy
Adult rotifers can have an inert body form with almost no body water. When rehydrated, they resume activity within a few hours.
747
Diapause
In crustacean zooplankton (e.g., cyclopoid copepods), they can produce a different kind of eggs that can undergo diapause when they sense a stress. The eggs can survive for up to 200 years in this state. When rehydrated, the eggs hatch.
748
Dispersal
Many species have aquatic larval and terrestrial adult stages. E.g., mayflies and dragonflies. The insects use the temporary pools for their larvae, which then emerge as flying adults.
749
Die Young (Live Fast)
The Turquoise killifish is the fastest-living fish. Lives in the Zimbabwean Savannah. Adults live for 3-9 months and reach sexual maturity as quickly as two weeks-- the fastest of any vertebrate.
750
Killifish Lifecycle
Once maturity is reached, females lay 20-120 eggs per day, which are then fertilised by a male. When the pool dries up, adults die, but the dormant embryos are left behind. When the rainy season begins, they rapidly hatch, growing from 5 to 30-50mm in a few weeks.
751
Waterfalls
Present an issue for migratory fish.
752
The rock-climbing goby in Hawaii:
Uses a sucker mouth to climb waterfalls.
753
What causes the high rates of population decline in freshwater vertebrates?
Freshwater habitats occupy the lowest point in the environment. Thus, whatever occurs in the surrounding catchment accumulates in the freshwater environment.
754
Chemicals
they can have ripple effects on food chains, and result in biomagnification as toxins are concentrated in tissues when travelling up the food chain. Biocides incl. insecticides, herbicides, pesticides and fungicides as well as antibiotics often end up in freshwater, especially in the lowland catchments and floodplain's of the world's major rivers, which are often densely-populated.
755
Heavy Metal-Based Fungicides
Also poisonous to fish.
756
Fertiliser is a diffuse source of pollution.
Because it's spread out across fields then enters the river at multiple points.
757
1858 Great Stink in London
Heatwaves and raw sewage combined to increase the spread of cholera.
758
Organic Pollution
Pollution from nutrients. Two main sources: sewage and fertiliser.
759
Eutrophication
Elevated nutreients due to organic pollution promote algal growth. As the alage decompose, microbes deplete the O2 in the water. Invertebrates and fish perish, or leave the affected area.
760
Dams cause habitat fragmentation.
They also impair sediment transfera nd migratory fish.
761
What % of rivers longer than 1,000 km remain free-flowing (dam-free)?
37%. Uninterrupted rivers are restricted to the Arctic, the Amazon and Congo basins.
762
How does water abstraction lead to water scarcity?
If demand exceeds supply. The combined effect of climate change and increased demand leads to dry rivers.
763
How are freshwater habitats exploited?
By water usage and fisheries.
764
Biological invasion is increasing with…
… globalisation.
765
Freshwater ecosystems can be affected by invasions in the water body itself as well as by invasions of plants on the banks.
Leaf litter that falls into streams is an important food source for aquatic invertebrates. Leaves from innvasive plants may be less palatable. E.g. rhododendrons don't provide sufficient food for native species.
766
Invasive Crayfish
Are outcompeting native crayfish by bringing disease. In the UK, there's 1 native and 6 invasive species of crayfish.
767
What do elevated temperatures in freshwater habitats mean?
Some sites are no longer suitable for species with a narrow thermal niche. Fragmented habitats mean individuals often cannot move.
768
Climate change-associated changes in rainfall lead to flooding or drought.
Flooding changes nutrient dynamics, availability of seasonal resources and habitat availability.
769
Heatwaves
Some evidence suggest fish activity decreases with warming.
770
3 possible outcomes when stressors interact:
Mitigate, amplify or additive effects.
771
Additive Effects
The sum of the mortality of each of the stressors.
772
Mitigate
Antagonistic interaction between stressors mean the overal effect is less than the sum of its parts.
773
Amplify
Synergistic interaction between stressors. The overall effect is more than the sum of its parts.
774
What do controlled mesocosm experiments suggest?
Non-additive effects are common.
775
Lake Victoria is affected by multiple stressors.
Fishing pressure, invasions and pollution. Nutrient pollution has promoted the spread of invasibe floating hyacinth. Native fish have lower fitness from pollution, which means they are more likely to get caught in the growing fishery.
776
The effect of warming on pollutants:
Warming can increase the potentcy of some pollutants, but can aso decrease the effects of pollutants by breaking them down faster.
777
A stressor increases or decreases the sensitivity...
… of a species to subsequent stressors.
778
Negative co-tolerance to stressors is additive.
One species will only be lost in instances of plant invasions. The other species will only be lost when fish invade. Different parts of the food web are affected, so the effect of the stressors can be summed. This si because stressors are quite different.
779
Positive co-tolerance is less than additive.
One species isn't sensitive to either invasion (both invasions are by fish, but different species). The other species is senitive to both invasions. Antagonistic effects: its' non-additive. Less than the sum of its parts. For the affected spcies, it doesn't matter if it;s one spcies of both invading.
780
What does co-tolerance depend upon?
Stressor similarity. Stressor similarity can be used to predict whether the effects of the interacting stressors will be additive or non-additive.
781
The stressors in co-tolerance scenarios don't have to be invasions.
They could be insecticide and an invasive insectivorous fish, for example.
782
Non-additive effects can also be caused by food web interactions.
Altered interactions between species, e.g., competition, parasitism and predation. Perch has restructured the Lake Victoria food web by causing extinctions. After the effects of the null perch, stressors are starting from a different baseline.
783
In one study in Europe, 4 pressures (hydrology, morphology, water quality and connectivity) were investigated.
~90% of all lowland rivers in the study area were imapcted bya combination of all 4 pressures.