Transcript Competition and character displacement

Character Displacement
Character displacement occurs when interspecific competition
leads to the selection of different characteristics (usually
morphological) in the zone of geographical overlap of species.
What shifts is the morphology of some aspect of the species
important in interactions. The classic example is in Darwin’s
Galapagos finches:
Geospiza fortis
Geospiza fulginosa
Since food resources are commonly the limiting resource over
which competition occurs, the morphology of feeding
structures is the feature frequently 'displaced'.
Grant (1968) drew comparisons of wing size and bill size
among closely related species on islands and made parallel
comparisons between some of the same species when both
were present in mainland habitats.
The differences found clearly indicate both greater difference
between species in their bill sizes than their wing lengths
(competition is for food), and that differences are greater on
islands than on nearby mainland.
Here is some of Grant’s data extracted from summary tables:
% differences in characteristics between congeneric species
Species
Mainland
Island
wing
bill
wing
bill
Parus
4.0
28.2
2.7
55.9
atricapillus/hudsonicus
Hylocichla
5.8
9.2
8.4
13.0
guttata/ustulata
Centurus
24.2
47.7
33.5 63.9
aurifrons/pygmaeus
Platypsaris
9.4
8.3
11.8
17.7
aglaiae/bidentata
Cynanthus
11.6
9.5
32.6
28.1
latirostris/A.rutila
These species are all present on Pacific islands near and on mainland of
California and Mexico.
What happens if the survey is across a wider range of
geographic areas? Answer: the same basic pattern…
Island
Species
% Difference in:
wing
bill
Norfolk
Zosterops
8.6
16.7
lateralis/albogularis
Lord Howe Zosterops
9.8
36.4
strenua/lateralis
Tristan de
Nesospiza
16.3
33.3
Cunha
wilkinsi /acunhae
Ongea levu Mayrornis
10.1
19.1
lessoni/ versicolor
Kauai
Laxops
13.4
50.0
parva/ virens
Socotra
Onchyognathus
7.2
20.2
frater/ blythii
Island
Species
Celebes
(Sulawesi)
Andaman
Dicrurus
montanus/hottentottus
Dicrurus
19.8
paradiseus/andamanensis
Acanthiza
0.0
pusilla/ewingi
Zosterops
6.8
mauritiana/curvirostris
Tasmania
Mauritius
% Difference in:
wing
bill
16.4
22.6
19.0
16.2
17.1
These differences, plus many others that could be cited,
demonstrate that food is frequently a limiting resource, and
that the world is not so green as Hairston et.al. claimed.
Kauai
Sulawesi
Norfolk
Mauritius
Tristan de Cunha
Lord Howe
Tasmania
Food as a Limiting Resource
Recognizing that limits to the food resource can limit
population size is straightforward. The question remains - Can
food regulate populations?
The problem with most field studies of 'regulation' of
populations by their food resources is lack of a proper control.
To establish regulation in the proper sense, population density
must tend toward an equilibrium from both above and below
as food resources change.
There is one field study which attempts to demonstrate that - a
study in which both the density of Lymnaea elodes and food
resources were manipulated.
In the field, all measurements must be taken relative to an
unmanipulated control treatment. Eisenberg did that,
fencing off areas at the margin of a pond in southeastern
Michigan, and maintaining three kinds of areas:
1) control,
2) areas in which snail density was reduced to 1/5 of control,
and
3) areas where density was increased to 5x control.
4) In additional replicate pens 10 oz. packages of frozen
spinach every 4 days.
The life history of the snails:
Lymnaea produces only 1 generation per year, laying eggs in
gelatinous egg masses on the surfaces of vegetation.
The eggs hatch during the summer and grow to 1-5 mm shell
diameter.
Overwintering (estivation) occurs as small juveniles by
burrowing into the mud along pond margins. In the spring
they emerge and grow to reproductive adulthood as the
pond thaws and spring runoff submerges the margins where
estivation occurred.
Reproduction occurs through June an July.
That means there are only two basic size classes in the pond at
any time, young of the year and adults from the previous year,
which are easily separated.
Density manipulations revealed rapid
regulation. It is evident in egg laying by
adults in the summer following density
treatment. It's evident both in the total
numbers of eggs in pens, and in the
number of eggs per egg mass. The total
number of eggs produced per pen shows
no relationship to adult density, but the
number of egg masses, and particularly
the number of eggs per egg mass show
essentially perfect density compensation.
Adult and juvenile survivorship thus
show little or no response, but mx or
fecundity showed a strong response. This
response does not implicate food.
The key to demonstrating that food was the regulating factor
is the effect of adding only 10 oz of spinach every 4 days to a
1x5 m pen. In those pens there was a 25x increase in the
number of eggs laid, with increases evident in both the
number and size of egg masses.
Pens are 'filled' with aquatic vegetation, therefore it wasn't
food quantity, but quality which was the limiting factor. Does
adult density the following year increase 25x? No, lx in 'fed'
pens also changed, so that the number of juveniles estivating
was increased only by 2x.
Hairston et al.'s hypothesis that 'the world is green' is not
warranted from the herbivores' point of view. There are other
reasons why plant quality may suggest food limitation. They
include the various forms plant defenses can take.
Chemical Defenses
There are 3 broad classes into which we can pigeonhole
chemical defenses:
1) toxic substances,
2) substances which reduce the digestibility or effective
quality of consumed food, and
3) pharmaco-active substances.
Most chemical defenses (other than tannin production) are
offshoots of normal metabolic pathways in the leaves. An
outline of 'normal' and secondary product metabolism reveals
that there is a close metabolic relationship among many toxic,
pharmacoactive and inhibitory defenses.
Sugar metabolism
Amino acid metabolism
acetate
phenolics  tannins
quinones
quinones
waxes
alkaloids
terpenes
glycosides
steroids
glucosinolates
Many plant chemical defenses against insect herbivores have
their origins as simple oxidations (or other modifications) of
ordinary phenolics, for example:
the active ingredient in cinnamon - trans cinnamic acid
the active ingredient in aspirin - salicyclic acid
the active ingredient in coffee - caffeic acid
and a related flavenoid:
the active ingredient in vanilla - vanillic acid and
a quinone produced from simple phenols:
don't plant things under a walnut tree - juglone
Vanilla planifolia
And raw vanilla beans
Cinnamomum zeylandicum
and its bark
Salix babylonicum
and aspirin
Toxic Defenses
Are there characteristic kinds of plants or plant tissues in
which different kinds of plant chemical defenses predominate?
Cates (1977, 1980) suggests there are general patterns:
Toxins are more frequently characteristic of ephemeral tissues
of plants (e.g. flowers, young leaves) which the plant cannot
afford to lose. These tissues are, however, less predictable in
time and space since they are ephemeral.
As a result, herbivores successful in utilizing plants or tissues
protected by toxins are likely to be specialists who have
evolved the ability to detoxify the protection, and related
plants are likely to have individualized (by species) defenses.
Toxins directly affect metabolic processes in the herbivores, are
easily absorbed by animal target systems, and are active in low
concentrations.
They represent generally < 2% of plant tissue dry weight.
If the plant, or the endangered tissue, is unpredictable in time
and/or space, then damage from the specialist herbivore is
limited due to difficulty in finding this plant or tissue.
Ephemerality selects for generalist herbivores, in that they are
not searching, but feeding on encounter.
Since detoxification (and the evolution of the capability) is
expensive and 'difficult', toxic defenses will be most effective
against generalists, particularly if the defenses deviate widely
from the already extant or ‘average' pattern present in the
community.
From the point of view of the herbivore - the generalist
herbivore feeds from a wide variety of plants as a strategy. In
doing that, while avoiding eating intensively from any single
food source, the limited detoxification capabilities which these
herbivores have is not overwhelmed by any single defensive
chemical.
For the apparent/persistent plant tissue the direct defense
needed is one against a specialist herbivore, which, having
encountered the tissue, would feed heavily on it. Each species
of plant evolves independently, but convergence has been
shown to occur. Within plant taxa many species have patterns
of digestibility-reducing defenses in common.
The costs and amounts of defensive chemical may be high.
There is an evolutionary race between toxin evolution and the
evolution of detoxification capability. Liver microsomal
enzymes are responsible for much of the detoxification.
Occasionally the process turns on the herbivore:
If caterpillars were classified by the number of plant families
on which they fed into monophagous (1 plant family),
moderately polyphagous (2-10 families) and highly
polyphagous (11 or more families), the activities of
microsomal oxidase activites paralleled diet breadth.
Activities were measured by the conversion of dieldrin to
aldrin, and were 20.4, 90.7 and 294.4 nanograms processed
per milligram of enzyme per minute respectively.
The oxidation of certain toxins, dieldrin among them,
produces a 'de-toxified‘ molecule which, in many animals, is
more toxic than the environmental chemical. In other cases,
e.g. Brassica (cabbages), a diversity of herbivores and their
predators are attracted to the plants by the mustard oil present
in leaves as an anti-herbivore chemical toxin.
An example of diversity in toxic defenses in related species is
alkaloids in Lupinus species of the southwestern U.S.
Species Alkaloids A B C D E F G H
Lupinus arboreus
L. chamissonis
L. arbifrons
L. excubitus
L. niveus
L. latifolius
3
2
1
1
2
1
1
1
1
1
1
2
1
1
1
1
1
1
1
Digestibility Reducing Defenses
Digestibility reducing substances are generally protein or
carbohydrate complexing agents that prevent or reduce
digestion of food materials from plant tissues. They are
characteristic of mature tissues of perennials, particularly
woody plants. In those tissues a surprisingly large fraction of
biomass can be defensive chemicals. 80% of woody dicot
perennials contain tannins, while only 15% of annuals
and non-woody dicot perennials contain tannins.
Digestibility reducers are generalized compounds, both in
terms of the breadth of the targeted herbivore spectrum and in
terms of the variety of plant species using similar or identical
chemicals.
Compare alkaloid distribution to the distributions of different
tannins in the leaves of maple (Acer) species…
Species
Tannins A B C D E F G H
Acer ginnale
A. tartaricum
+ + + + +
A. saccharinum
A. platanoides
A. campestre
A. rubrum
A. pennsylvanicum
A. spicatum
A. rotundilobum
A. griseum
A. saccharum
A. pseudoplatanus
+ +
+
Common names of (some) of those maple species:
A. tartaricum – tartar maple
A. saccharinum – silver maple
A. platanoides – Norway maple
A. campestre – field maple
A. rubrum – red maple
A. pennsylvanicum – striped maple
A. spicatum – mountain maple
A. griseum – paperbark maple
A. saccharum – sugar maple
A. pseudoplatanus – sycamore maple
Thus far chemical defenses have been considered as
constitutive, that is defenses prepared and maintained whether
the herbivore is present or not. There is also evidence of
inducible defenses, i.e. those synthesized only in response to
'initial' herbivore damage.
The existence of an inducible defense has been intensively
studied in arctic birch (personal interest here!), which is a
favorite forage of caribou (reindeer) in northern Europe. We’re
not interested in reindeer.
The herbivore that can defoliate entire trees of some strains of
northern birch is the autumnal moth Oporina (now called
Epirrita) in outbreaks.
Eppirita autumnata – the
defoliating moth
Betula pendula – the birch common
in northern Scandanavia
How do birch trees respond to herbivory? There are a number
of responses, including induction of defensive chemicals.
Responses include:
a) By escape in time. The tree is perennial; it certainly
suffers from herbivory, but even defoliation in a single year
does not result in certain mortality. Instead, there are a number
of responses evident in the following year. There is delay and
increased variability in the date of leafing out the following
spring. Measured as standard deviations in the date of leaf
burst, in control plants (not defoliated) the s.d. was .32 days,
but in defoliated plants it was .88 days.
Larval hatching from overwintering eggs had s.d.'s of .73
or less, decreasing as temperature increased. The difference in
variability makes larval starvation more likely in years
following defoliation.
b) By inducible chemical defense. Mechanical damage to
leaves, for example by larval chewing, increases the amount
of phenol (the particular chemical is called platyphylloside) in
the leaves.
Induction occurs in less than 48 hours. Damaged leaves,
gathered 48 hours after damage occurred, had 9% more
phenol than undamaged leaves on the same plant.
Those phenols have important effects. Measurement of
inhibition of trypsin, a key enzyme in protein digestion,
indicate undamaged leaf extract reduces trypsin activity by
23% (even undamaged leaves carry some chemical defenses),
but induced leaves reduce activity by 32%, or an increase in
trypsin inhibition by 40%. Thus, the ability to digest
protein has been significantly reduced.
This inhibition slows the growth of larvae both following
initial induction and in the following year. The platyphylloside
is effective in inhibiting digestion down to a concentration of
0.8% in leaf tissue. That is a concentration lower than is
observed in even undamaged leaves.
c) These defensive adaptations don't disappear rapidly after
coming into play. The relaxation time for these defenses may
be as long as 3-4 years. Larvae reared in 1979 on trees
artificially defoliated in 1975-6 were still significantly lighter
(smaller due to slower growth, presumably due to inhibitory
defenses) than those grown on leaves from control plants.
This picture may not be as general as European studies of
arctic birches might suggest…
Chapin et al.(1985) searched for parallel responses in the
woody plants of Alaska and found none. Simulated
herbivory did not induce increased production of phenols,
resins, or tannins in Alaskan plants including willows,
poplars, aspens, or birch.
Even the basic pattern of digestibility-reduction by tannins
has been found to have exceptions.
a) Some arctic ruminants have salivary proteins that bind
(complex) tannins as leaves are initially chewed, so that the
defensive function of the tannins is circumvented.
b) Bernays (1980) found growth and survivorship of locusts
differs among species exposed over evolutionary time to
diets including different frequencies and abundance of
tannins. She studied 15 locust species ranging from
monophagous to highly polyphagous species.
Most of the locust species feed on grasses in nature, and
were maintained in the laboratory and tested using a grass,
wheat. For tests of tannic acid effects the wheat leaves were
dipped in tannic acid solution. Other species feed on dicots
in nature; they were maintained and tested using lettuce
leaves instead of wheat. Four groups of locust species were
distinguished in terms of their responses to tannic acid:
Group 1) Tannic acid caused complete mortality during early
instars. In these locusts protein complexing by tannins is
unimportant to its effect. Instead, tannic acid had dramatic
histological consequences; lesions were rapidly apparent in
the insects' midgut. Tannic acid then entered the open
circulatory system and acted as a systemic poison.
As a group these locusts are characteristically grass feeders;
grasses rarely contain tannins naturally, and the locusts are
unlikely to have been exposed to tannins in their natural diets
Group 2) Mortality is delayed to later instars, and a few
individuals molt successfully to adulthood. These species are
also generally grass-feeders and inexperienced with tannins.
Their weights as late instar larvae are severely reduced,
probably reflecting protein complexing by dietary tannins.
One example is Acrida conica. 50 first instar larvae
constituted each group. Survival and weight (mg) of
survivors were:
Treated
Control
Male
Female
Male
Female
weight
146 + 4
396
216 + 6
623 + 10
number surviving 10
2
1
19
There must be a miscopied number
here. There is no SD with only 1
survivor.
Group 3) These species showed no significant effect on
survival. They are generally polyphagous, and therefore had
been exposed to at least a moderate proportion of dicots
containing tannins in their natural diets. Digestibility, digestive
efficiency and weight increase also seemed unaffected by
tannins.
A good example of this category is the plague locust,
Schistocerca gregaria:
Treated
Control
Male
Female
Male
Female
weight (mg) 1093 + 24 1343 + 20 1110 + 16 1319 + 15
number surviving 31
28
29
28
digestibility (%)
34 + 4
34 + 3
digestive efficiency 38 + 6
34 + 7
% wt.inc. (last instar) 79 + 5
69 + 8
Group 4) A few species (two in these studies) do better on
tannins than without. Both species are specialist feeders; their
normal diets consist of the leaves of tropical acacias, which
are loaded with just the sort of tannin used in these
experiments. At least for these two species there is the
suggestion of enzymatic hydrolysis of tannins (and possibly
other related phenolics) into compounds useful in normal
metabolic pathways. Example: Anacridia melandorhodum
Treated
Control
Male
Female
Male
Female
weight
1517 + 30 2124 + 37
1314 + 42 1813 + 67
survival
22
20
8
11
digestibility
52 + 2
40 + 4
efficiency
25 + 2
20 + 2%
wt. inc. last instar 116 + 6
103 + 5