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55 Cards in this Set

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Why might defenses angiosperms not have evolved more effective against nectar robbers?
Nectar robbing results in only a slight fitness cost to the plant – weak natural selection for greater defense

Defenses against nectar robbers might negatively affect pollinators also

Allocation costs of constructing such defenses
Direct negative fitness consequences of nectar robbing?
Energy Costs-some contain amino acids

Water lost in dry conditions

Nectar Robbers Damage Reproductive parts
Indirect negative fitness consequences of nectar robbing
– reduce visitation of efficacy of legitimate animal pollinators
Extent to which nectar robbers will reduce pollinator efficacy depends on
- ability of pollinators to detect nectar reduction / depletion from afar

- amount of nectar removed by the nectar robber
Reasons why nectar robbers may have slight, if any negative effects on plant fitness
Bees make longer flight distances between stops after visiting “low” nectar flowers – reduces inter-breeding with close relatives

Bees visit fewer flowers on the same inflorescence when to encounter “low” nectar flowers – reduced self-pollination

Nectar robbers transfer some pollen
Fitness advantages of seed dispersal to plants
Departure-related benefits: benefits that result from escaping the surroundings of the parent plant
What are some examples of departure-related benefits?
-Avoid highly competitive environment due to overshading and sibling competiton

less herbivory by specialist herbivores

more competitive seeds get disperesed to new microsites (leptokurtic seed dispersal

escape from pathogens

chemical influence of parent, alleopathy of parent
Arrival related benefits:
benefits that result from arrival at a specific location that is particularly conducive to survival, growth and reproduction

1. Colonization of unpredictable, vacant, resource-rich habitats (recently disturbed sites)

2. Directed dispersal – movement to predictably favorable sites for germination and growth (bell birds, Wenney and Levy)
Herrera’s three modes of vertebrate-mediated seed dispersal
Imperfect harvesting: granivores cache seeds for later consumption, but those cached seeds that are never re-located and germinate are dispersed

External “contamination” by seeds: seed with hooks or barbs adhere to the outside of an animal’s body

Interal “contamination” by seeds: seeds are consumed by the animal and are dispersed when the animal spits, regurgitates or defecates out the seed
Binary seed dispersal
“two-stage” seed dispersal; seed dispersal initially away from the parent is by one dispersal mechanism then a second dispersal mechanism moves the seed from its original deposition point
Primary seed dispersal
process that moves the seed from the parent plant to a new location
Secondary seed dispersal
process that moves a seed that has already been removed from the parent plant
Ecological Patterns in Vertebrate-mediated Seed Dispersal
Vertebrate-mediated seed dispersal is more common in:
Plant species with large seeds
Woody plants (trees and shrubs) than herbaceous plants
Woodlands, shrublands and forests than in grasslands
More productive (NPP) ecosystems, within a ecosystem type
Hypotheses for why there is a positive relationship between ecosystem productivity and the proportion of plant species that are vertebrate-dispersed?
Westoby et al. 1990 tested these two hypotheses

In more productive communities, seeds must be larger to contain large endosperm and cotyledons required in low light environments; large seed are only effectively dispersed by vertebrates

In more productive communities, parent plants are larger and, therefore, the seeds must be dispersed further to receive “departure-related” benefits

Neither proved very good.
Eliasome
A food body on a seed which ants like, fatty
Additional hypotheses for the positive relationship between vertebrate seed dispersal and ecosystem productivity
3. Dispersal structures required to attract vertebrates are energetically expensive and the resources to produce them are only adequate in productive ecosystems

4. In low productivity sites, vertebrate populations are too small to efficiently disperse seeds
Do vertebrate seed dispersers act as agents of natural selection that determine plant fruiting phenology?
Overall, no not really...

But, fruiting might coincide with avail. of seed dispersers

Could produce either staggard or synchronous fruiting.
What might be an advantage of “staggered” or “over-dispersed” fruit production by plant species in a community?
-Reduces competition for animal seed dispersers

staggard decreases satiation, all species benefit... (but doesn't hold very true, frugivores aren't specialists)
Advantages of synchronized fruit production by plant species in a community
synchronized, large bursts of fruit production will attract large numbers of vertebrate frugivores / seed dispersers
Factors other than phenology of frugivore / seed disperser abundance that might acts as agents of natural selection on fruiting phenology
Avoidance of frugivores that do not disperse seeds

Avoidance of microbes that might infest fruits making them less appealing to frugivore / seed dispersers

Seasonal availability of abiotic resources required for reproduction, including fruit production

Timing of fruit production may be constrained by selection acting on the timing of flowering
Color evolves in response to frugivores?
Seed dispersal by bird frugivores often involved red or black fruits

Mammals-fruit is pretty drab

Birds discriminate among fruits based upon color, birds chose red over orange (Gervais)

Color selection might act as selective force.
Do larger-bodied vertebrate seed dispersers explain larger fruit sizes in the Old World tropics than in the New World tropics?
Mack 1993 shows trends suggesting yes

Might be ecological filtering... only those frugivores with correct gape size survive and colonate.

Differences in fruit sizes between the Old World and New World tropics cannot be explained only based upon differences in the plant families present
Palatability-defense trade-off hypothesis
plant species whose fruits are dispersed less quickly should have higher levels of secondary compounds in the fruit pulp
Nutritional challenges of eating fruit
Extensive nutrient dilution

Lots of water/carbs not much protein/fat

Nutritional imbalance – low in N content

Secondary compounds

Strong temporal variation in fruit availability, including strong seasonality in fruit production
Advantages/Disadvantages of secondary compounds in mature fruits.
Disadvantages:
-deter see disperesers, fewer fruits consumed before leaving
-reduce number of species as dispersers
-induce a learned avoidance response

Advantages:
-Deters parasitic frugivores (insects)
-secondary compounds may be antimicrobial
-inhibit premature seed germination
-compount regulates rate of gut passage in frugivore
-not bad to eat fewer fruits, more likely to move on before deficating
Behavioral, Anatomical and Biochemical Adaptations in Frugivores to deal with Nutrient Dilution
Regurgitation, rapid gut passage of seeds

Long small intestines, seasonal adjustment of intestine lengths

Fast gut passage rates – more fruit consumed per unit time

4. Adjustment of activity of carbohydrate assimilating enzymes

-Gut retention time is a function of amt of fruit consumed, not body mass
Adaptations that aid Frugivores in dealing with Nutrient Imbalance in Fruit Pulp
Generalized diet – many frugivores consume some animal matter also

Consumption of a wide range of fruits from different plant species

Compensatory consumption – amount of fruit consumed may depend on the protein or lipid content of the fruit

Physiologically many frugivores are highly efficient at extracting N from low protein food
Adaptations that aid Frugivores in dealing with High Levels of Secondary Compounds in Fruit Pulp
Consume fruits of a variety of plant species during individual foraging bouts (recall Toxin Dilution Hypothesis)

Anatomical and physiological adaptations for enhanced detoxifying ability – frugivorous species have larger livers than non-frugivorous species
Adaptations in Frugivores to deal with Pronounced Seasonality of Fruit Production
Expand home range size in seasons of low fruit production

Seasonal shifts in habitat to track seasonality of fruit production (altitudinal migration)

Broaden diet to include animal or non-fruit plant material

Reproductive cycles synchronized with seasonal peaks in fruit production
-Tamarans in Peru... all births occur in corresponance to wet season when most fruit is available.
Coevolution
evolution in two or more species brought about by reciprocal selective effects between the entities
Specific coevolution
two species interact closely with one another and changes in one species select for adaptive changes in the other species and vice versa
Diffuse coevolution
groups of species interact with groups of other species leading to adaptations that cannot be identified as arising from specific pairwise interactions between species
How to quantify the degree of specificity in species interactions?
Proportion of interactions in the community that are strictly pairwise (meaning one-on-one)

Mean number of species with which each species interacts (e.g. mean number of pollinators with which each plant interacts)
Connectance
the proportion of potential interactions between species pairs in a community that actually occur; greater connectance indicates less specificity in interactions

Generally, plant pollination systems have less connectance than Seed Systems

5 host plant species X 7 pollinator species = 35 potential species pairs interactions
11 species pairs interactions actually occur
Connectance = 11 / 35 = 31.4%
Why is there less specificity in vertebrate seed disperser – host plant interactions than in animal pollinator – plant interactions?
Vertebrate dispersal of seeds limits opportunities for local genetic divergence and specialization

Opportunities for plant specialization are limited by the absence of a specific target site

Temporal variability in fruit availability limits opportunities for specialization on a single host plant species by frugivores
-Natural selection favors a generalist diet
-In different years, vertebrate frugivores are likely to be exposed to conflicting selection pressure
Myrmecochory
seed dispersal by ants; occurs in up to 3000 plants species from 80 families
Ecological distribution of myrmechochory
Southern hemisphere: low nutrient, fire-prone shrublands of South Africa and Australia (2700 plant species). Generally secondar dist by ants.
Life form: small shrubs, canopies usually <2m diameter, not serotinous


Northern hemisphere: temperate deciduous forests (300 plant species). Primary distribution by ants.
Life form: understory herbaceous plants, weight of seed bends the shrub to the ground where ant picks it.
Diaspore
seed plus the attached elaisome
Why do ants eat elaisomes?
-Most ants are carnivores, so many of the rules we consider with herbivore nutrion do not apply

--Hughes 94, eliasome close to fatty acid composition of insects more than seeds

diglyceride 1,2 diolein is strongest ant attracter, very prevalent in insects preyed upon by ants

-ant collects the entire seed, takes back to nest, eats elaisome, and discards the seed in the nest. This puts the seed in optimal position for germination.

-ants disperse seeds over short distances
Some advantages to plants of ant-mediated seed dispersal
Departure-related benefits, such as reduced parent-offspring and sibling competition

Transfer of seeds to nutrient rich sites (middens in nests) – an arrival-related benefit (directed dispersal)

Escape from granivorous rodents and granivorous ants – ant-mediated seed dispersal is one of the surest ways to get a seed belowground (directed dispersal)
Why does myrmecochory occur in the plant community types and with the plant life-forms that it does?
Plant body sizes are relatively small – short distances that ants disperse seeds are adequate to escape the parent

Serotiny may incompatible with ant-disperal because long-term storage in a sealed cone may cause the eliasome to deteriorate

Burial of seeds is advantageous in fire-prone ecosystems because the soil insulates from the heat of the fire
Patterns in seed selectivity by harvester (granivorous) ants
Often harvest seeds of the most common plant species in the community

Correlation between ant body size and the size of seeds consumed
Ecological consequences of ant granivory (harvester ants)
Ant removal increases the densities of an equal number of winter annuals as does rodent removal, but rodents have larger indirect effects
Ants reduce densities of more summer annual species than do rodents (Samson Et. Al)

Ants can structure desert rodent communities through interspecific competition for seeds (brown 79)
Myrmecophyte
plant that is defended by ants
Adaptations of myrmecophytes to attract ants
Extrafloral nectaries

Food bodies produced from epidermal tissue that are harvested by ants (e.g. Beltian bodies)

Domatia-spines that can house ants.. plants can regulate size of ant which can use it.
Relationship of Ants, plants, and herbivore, simple.
Ants can have a net fitness advantage.
Ants can have a net fitness advantage.
What are the advantages to ants of tending myrmecophilous herbivores, like hompoterans and some lepidopteran larvae?
Aphids are phloem suckers, lepidopterns eat plant and secrete sweet substance. Ants then eat some byproduct of the herbivore.

This makes the relationship more complex.


Ants often tend some herbivores on myrmecophytes while deterring other herbivores

Ant guarding with aphids present actually increases damage by leaf-chewing herbivores (OLiver 07), Ant guarding with aphids present decreases host plant fitness
What are the advantages to the myrmecophilous herbivores of being tended?
Ants deter predators and parasitoids

Ants reduce pathogen contamination

Ants shelter the herbivores by carrying them into domatia

Ants move herbivores around on the plant to higher quality tissue patches
If ants often guard homopterans or lepidopteran larvae on host plants and the presence of ant guards decreases plant fitness then why are extrafloral nectaries still present?
No genetic variation to lose extrafloral nectaries

Conflicting selection pressures across years

3. Extrafloral nectaries are adaptive because they attract non-ant predators and parasitoids

Extrafloral nectaries draw ants away from flowers where they could deter pollinators

5. Distraction hypothesis – extrafloral nectaries provide an alternative for ants to tending herbivores on plants, may prevent ants from guarding herbivores
Common effects of ant nest building on physical properties of soil
Reduce soil bulk density

Move smaller-sized soil particles to the soil surface

Homogenize soil horizons
Common effects of ant nest-building on chemical properties of soil
Increase quantities of N, P and K

Increase organic matter in soil

Sometimes alter soil pH, usually making both acidic and alkaline soils more neutral
How is decomp. rate altered?
-Actinomycse are less abundant, they are involed in later stages of decomp/hummus breakdown

Microbes involved in early decomp are more abundant
metapleural gland (ants)
produces antibodies secretion to prevent fungal infections (spread all over nest and on larvae) maybe this stuff ends up in refuse piles... making the pile good for seeds

also involved in chemical defense, territory marking and pheremones
Why are grasses more abundant than forbs on mounds than off?
Soil temp on the nest is higher than off, Favors the grasses

Maybe... rate of ant import, and maybe diff. rate of germination due to soil antibodies.
Refuse Piles (ants)
More nutritionally rich than surrounding forest floor. When in this soil during a study, fewer seeds died due to fungal infection (perhaps from secretions of ants), more seeds germinate.