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CHAPTER 57 SPECIES INTERACTIONS Brenda Leady, University of Toledo
Prepared by Brenda Leady, University of Toledo Copyright (c) The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
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Competition Intraspecific – between individuals of the same species
Interspecific – between individuals of different species Resource competition – organisms compete indirectly through the consumption of a limited resource Interference competition – individuals interact directly with one another by physical force or intimidation
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Amensalism – one species has a strong effect on another, but the reverse effect is negligible
-/0 relationship Allelopathy Roots excreting chemicals to kill off competitors Penicillin inhibiting the growth of bacteria
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Connell’s Experiments with Barnacle Species Show That One Species Can Competitively Exclude Another in a Natural Setting 1954, Joseph Connel worked with 2 barnacles along the Scottish coastline Determine range of Chthamalus stellatus in the absence of competition from Semibalanus balanoides (formerly known as Balanus balanoides) Potential distribution (the fundamental niche) of Chthamalus extends over the entire intertidal zone, its actual distribution (the realized niche) is restricted to the upper zone Among the first to show that, in a natural environment, one species can actually outcompete another, affecting its distribution within a habitat
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Outcomes of competition can vary
Late 1940s, Thomas Park examined competition between two flour beetles, Tribolium confusum and Tribolium castaneum Presence of parasite (biotic factor) can alter the outcome of competition Random variation, stochasticity, evident Also influenced by temperature and moisture Mechanism of competition is predation of larva by other species
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Field studies 1983 review competition found in 55% of 215 species surveyed, demonstrating that it is indeed frequent in nature Generally in studies of single pairs of species utilizing the same resource, competition is almost always reported (90%) Percent competition varies with the number of species involved Plants showed a high degree of competition Marine organisms tended to compete more than terrestrial ones
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Competitive exclusion hypothesis
Gause worked with 3 protists – Paramecium aurelia, Paramecium bursaria, and Paramecium caudatum Grown separately all 3 grew logistically When P. caudatum and P. aurelia grown together, P. caudatum went extinct When P. caudatum and P. bursaria grown together, neither went extinct Concluded that 2 species with exactly the same requirements cannot live together in the same place and use the same resources, that is, occupy the same niche Competitive exclusion hypothesis – complete competitors cannot coexist
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Robert MacArthur examined coexistence between five species of warblers feeding within spruce trees in New England Found that the species occupied different heights and portions in the tree and thus each probably fed on a different range of insects Resource partitioning – differentiation of niches, both in space and time, that enables similar species to coexist in a community
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Morphological differences allow coexistence
Some partial level of competition between complete competitors can exist if it is not severe enough to drive one competitor extinct G. Evelyn Hutchinson compared size differences in feeding apparatus between sympatric (same geographic area) and allopatric (different geographic areas) species When species were sympatric, feeding apparatus size changed to specialize on different food Size differences did not occur in allopatric species Character displacement – tendency for 2 species to diverge in morphology and resource use due to competition Galapagos finch example
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Predation Categories of predation can be classified according to how lethal they are for the prey and how close is the association between predator and prey
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Antipredator strategies
Chemical defense Bombardier beetle ejects hot spray Aposematic coloration, or warning coloration, which advertises an organism’s unpalatable taste Many tropical frogs have bright coloration to advertise their skin’s lethality Cryptic coloration Camouflage Stick insects mimic branches Sea horses adopt body coloration to mimic habitat
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Displays of intimidation
Mimicry Resemblance of mimic to another organism (model) Mullerian mimicry – noxious species converge to reinforce warning, black and yellow stripes of bees and wasps Batesian mimicry – palatable mimic resembles unpalatable model, scarlet king snake and coral snake Displays of intimidation Porcupine fish inflates itself Deceive predator about ease of eating prey Fighting Horns and antlers can be used in defense
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Escape Grasshoppers powerful jumping ability Armor Shells of turtles provide strong defense Beetle exoskeleton Masting Synchronous production of progeny satiate predators and allow some young to survive Seed herbivory Periodical cicadas Most common antipredator mechanisms in arthropods were chemical defenses and aposematic coloration
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Predators still affect prey densities
Importance of predation depends on if it is Donor-controlled system, prey supply is determined by factors other than predation, such as food supply, so that removal of predators has no effect on prey density Predator-controlled system, action of predator feeding reduces supply of prey, so removal of predator results in increase in prey
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Invasive species do not have their natural predators
Brown tree snake introduced to Guam By 1980’s 8 of 11 of island’s forest birds extinct Birds had no defense against snake they didn’t evolve with Systematic decline in whales of various species in response to human whaling industry
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Plant/herbivore arms race
Herbivory can be lethal to small species Usually not lethal to larger species Generalist herbivores can feed on many plant species Specialist herbivores restricted to 1 or 2 host plants 2 proposals explain why every plant isn’t consumed Predators and parasites keep herbivore numbers low Plant defenses make a difference
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Plant defenses Array of unusual and powerful chemicals
2 classes – nitrogen containing (alkaloids) and lacing nitrogen (phenolics and terpenoids) Secondary metabolites – not part of primary energy generating metabolic pathway Over 25,00 compounds Also mechanical defenses like thorns and spines
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Herbivores can overcome plant defenses
Detoxify using 2 pathways Oxidation – catalysis of secondary metabolite to corresponding alcohol by mixed-function oxidases (MFOs) Conjugation – unites results of oxidation with another molecule to create inactive and readily excreted product Certain chemicals that are toxic to generalist herbivores actually increase the growth rates of adapted specialist species
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Parasitism One organism feeds off another, but does not normally kill it outright Predatory organism is termed a parasite and the prey a host
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Parasitic flowering plants
Holoparasites lack chlorophyll and are totally dependent on the host plant for their water and nutrients Rafflesia arnoldii, which lives most of its life within the body of its host with only the flower developing externally (largest known flower in the world) Hemiparasites generally do photosynthesize, but they lack a root system to draw water and thus depend on their hosts for that function Mistletoe (Viscum album)
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Host range Monophagous – feed on one or two closely related hosts Polyphagous – feed on many hosts Size Microparasites – multiply within hosts, usually within cells (bacteria and viruses) Macroparasites – live in host but release juvenile stages outside host’s body Site Ectoparasites – live on the outside of the host body (fleas and ticks) Endoparasites – live inside the host body (bacteria and tapeworms)
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Parasites may outnumber free-living species by four to one
Most plant and animal species harbor many parasites Few cases of experimental removal of parasites confirm that parasites can reduce host population densities Blue tit and parasitic blowfly larvae example
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Introduced parasite species an opportunity to examine parasitism as a mortality factor
Plants Chestnut blight virtually eliminated chestnut trees in North America Dutch elm disease eliminated elms in Europe and North America Canker eliminated cypress in Italy Animals Canine distemper caused a large decline in the last remaining black-footed ferrets in the US 1/3 of lions wiped out by canine distemper in Serengeti National Park Ethiopian wolf has been vaccinated against rabies Mountain gorillas vaccinated against measles
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The Fight Against Plant Diseases
Sudden oak death is a recently recognized disease killing tens of thousands of oak trees and other plants in California Usually a forest disease but known to infect woody ornamentals like rhododendrons Mapped genome of disease-carrying fungus to help develop specific tests for disease Some have suggested limited, cautious transfer of resistant genes to threatened species Using traditional breeding technologies to cross susceptible plant with resistant species can later affect appearance of original plant
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Mutualism Close associations are known between species in which both species benefit Leaf-cutting ants and fungus Ants chew up leaves to feed to fungus they care for Fungus produces gongylidia as ant food Ants circumvent chemical defenses of leaves
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Defensive mutualism Dispersive mutualism
Animal defends a plant or herbivore Ants protect aphid and aphid secretes honeydew Facultative mutualism since they can live apart Ants nesting in acacia trees defend the tree and trim away competing plants Obligatory mutualism since neither can live without the other Dispersive mutualism Involve pollination and seed dispersal Plant would like pollinator with high fidelity to one species that moves quickly among individuals Animal wants to be a generalist to obtain as much food from many flowers in a small area to reduce energy expenditure Although mutualisms are beneficial, optimal needs of each party can be different
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Commensalism One member derives a benefit while the other neither benefits nor is harmed Epiphytes growing in trees do not harm the trees Cattle egrets benefit from cattle stirring up insects Phoresy – one organism uses another for transportation Flower-inhabiting mites use hummingbird nostrils Cheating Grass-pink orchid produces no nectar, but it mimics the nectar-producing rose pogonia and is therefore still visited by bees Plants cheat seed-dispersal agent out of meal with barbs or hooks on seeds
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Conceptual models Bottom-up model – food limitation controls population density Plants regulate the population size of all other species that rely on them as the source of energy Nitrogen-limitation hypothesis – increasing supply of nitrogen to plants will increase herbivore population size, survivorship, growth and fecundity Top-down model – natural enemies control population densities Predators control prey populations Herbivores control plant populations Evidence from studies where natural enemies are introduced to control exotic pest species Small cactus moth brought in to control prickly pear cactus growth in Australia
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Modern models suggest both important
Recently, different models have been proposed that take into account the effects of both natural enemies and limited resources on species Ecosystem exploitation hypothesis – strength of mortality factors varies with availability of plant biomass involved Simple systems with limited plants resource limited At the other end, when plants are abundant, they support herbivores and carnivores, which compete
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