Predation, Herbivory, and Parasitism Types of Species Interactions When two species interact, the effects for each species can be positive, negative, or neutral. Competition -/Amensalism -/0 Commensalism +/0 Mutualism +/+ Exploitation +/- Types of Species Interactions Exploitative include: Predation Herbivory Parasitism interactions Parasitism Parasites live in or on their host's body and often spend most or all their lives eating tissues or body fluids of just one host individual. Sometimes multiple generations of parasites live on the same host. Because parasites depend on their hosts for continued feeding, they do not generally kill their hosts (at least not immediately). Parasitism Most parasites associated with a single host species have a free-living life stage during which they are not attached to a host. A great many other parasites, though, have multiple hosts with different life stages adapted to each host (and possibly free-living stages as well). Endoparasites The inside of an organism is a much more stable and protected environment than the outside, and endoparasites (e ndo = inner) take advantage of this by living and feeding inside their hosts Ectoparasites Although parasites tend to be tiny and hard to see, ectoparasites (ecto=outer), which live on the outside surface of their host, are often easier to observe. Ectoparasites By living outside their host, ectoparasites avoid having to defend themselves against the host's immune system. The trade-off, however, is that they are exposed to predation and a sometimes harsh exterior environment. Some species of predators specialize on ectoparasites. Ectoparasites The cleaner wrasse is a type of fish that lives in coral reefs. Other fish will wait patiently while a cleaner fish picks off parasites from their scales, mouth, and gills. Parasitoids and Hyperparasites Most of the parasites discussed so far do not directly kill their hosts but, parasitoids do. Parasitoids develop inside their host and essentially eat it from the inside out. When the host is completely consumed, the parasitoid transforms into an adult and crawls out to find new hosts for its offspring. Parasitoids and Hyperparasites Interestingly, the parasitoid wasp has a parasitoid of its own, known as a secondary parasitoid, or hyperparasitoid. Hyperparasitoid wasps find aphids with internal parasitoid larvae and lay eggs inside the larvae. Ecological Impacts of Parasites Parasites can have broad ecological impacts. These effects begin at the individual level; because parasites rob their hosts of resources, host survival and/or reproduction can be reduced even when hosts are not killed by parasites directly. Effects of Parasites on Individual Hosts A protozoan causes rats to become attracted to cats. A nematode turns the bellies of Amazonian ants red, attracting berry-eating birds. A fluke makes ants climb grass stems so they will be eaten by sheep. A trematode causes killifish to swim closer to the surface of the water, making them easy targets for birds. Herbivores Grazers are herbivores that specialize on herbaceous plants (grasses, forbs, and herbs), while browsers eat the leaves, bark, and twigs of woody plants. Herbivores that specialize on seeds are granivores, while those specializing on fruits are frugivores. Plant Defenses Against Herbivory Herbivory is generally not a positive experience for a plant, so plants have evolved forms of self-defense. Mechanical - Developing structures like thorns that make it harder for animals to eat them. Chemical - Producing chemicals that are noxious or poisonous to herbivores. Nutritional - Growing structures that are less nutritious for grazers (have less N and P). Tolerance - Adaptions to regrow quickly after being grazed. Impacts of Herbivory on Plant Communities Herbivory can reduce the overall number of plants and can also have a profound impact on the composition of a plant community. Impacts of Herbivory on Plant Communities When plants have evolved without selective pressure from herbivory, they may not be very well-defended, and the influence of herbivores can be even more drastic. This is a problem when new herbivores are introduced. Predation Lynx are fast, but lack endurance, so they don't chase hares over long distances. Instead, they stalk hares, hiding behind trees and brush until they can get close enough to pounce on a hare. Predation Stalking is one of a variety of strategies used by predators for catching mobile prey. Others include pursuit, where the predator chases prey over a distance; ambush, where the predator hides and waits in one spot until prey comes along; and random encounter, where the predator and prey meet by chance. Animal Defenses Against Predation Just as plants have evolved defense mechanisms to combat herbivory, animals have evolved ways to defend themselves from predation. Chemical - Producing chemicals that are noxious or poisonous to predators. Physical - Developing physical barriers to predation (e.g., shells). Behavioral - Behaving in ways that minimize risk from predation. Animal Defenses Against Predation Aposematism - Warning colors, sounds, or other characteristics to alert predators that this prey will not be tasty. Crypsis - Camouflaged colors, shapes, and other ways of hiding from predators. Mimicry - Looking, sounding, or in other ways mimicking a species that predators avoid. Predator-Prey Dynamics Lynx & Snowshoe hare population size records available back to 1800s. Fur traders Populations of lynx and hare seemed to follow an interesting cyclic pattern. Both species peak and then sharply decline about every ten years, at slightly non-overlapping intervals. Prey Dynamics: Modeling Hare Scientists often use computer models to investigate how different underlying mechanisms might produce particular dynamics in systems. Cycling and Extinction As the prey population size drops from predation, the predators have less and less food to eat, causing the predator population size to drop. With fewer predators, the prey population starts growing again, and the cycle repeats. Interactions among Trophic Levels Predators do not act alone in determining whether prey populations can survive. Prey can also be strongly affected by the availability and quality of their food. Deterministic vs. Stochastic The deterministic Lotka-Volterra predatorprey model predicts regular, even cycles of predator and prey populations, with predator cycles following prey cycles. More realistic models that include chance (stochastic) events show variation in both the period and amplitude of predatorprey cycles. Predator-Prey Dynamics in a Complex World Large predator populations cannot be supported by small prey populations. There is a feedback loop: when prey populations are small, predator populations shrink, and when predator populations shrink, prey populations grow. Predator-Prey Dynamics in a Complex World The equations in the Lotka-Volterra predator-prey model predict stable population cycling. However, because this is a deterministic model, there are no fluctuations in the cycles. Each cycle is exactly the same as the one before it. Predator-Prey Dynamics in a Complex World By simulating a simple predator-prey system that included some stochasticity (randomness), we see that irregular population cycling can lead to extinction. Predator-Prey Dynamics in a Complex World Predator-prey systems in nature involve many complicated dynamics that are thought to contribute to irregular patterns of cycling. Predator-Prey Dynamics in a Complex World Researchers studying lemmings (prey) and stoats (predators) in Greenland believe that the irregularities of cycling in their data is due in part to the fact that predators besides stoats prey on lemmings (e.g., arctic fox, snowy owl, and longtailed skua). Unrealistic Assumptions of Lotka-Volterra There are no effects of crowding for either the prey or predator (i.e., there is no density dependence in growth rates). All predator and prey individuals are equally likely to meet any of the others (i.e., the populations are evenly spread out and well-mixed). The prey species is the only food source for the predator. The predator is the only significant cause of death for the prey. Each predator individual can catch and eat prey individuals instantaneously (there is no handling time). There is no immigration or emigration of either prey or predators. Density Dependence Density-dependence prey and predator populations tends to stabilize the populations and reduce cycling. in Refuges Refuges and metapopulations tend to stabilize predator-prey systems and reduce cycling. Functional Responses to Exploitation The functional response, describes how an individual predator's feeding rate depends on prey density. The question is, given a certain density of prey, how fast will a predator eat them? Refuges & Metapopulations Patchiness in populations can promote stability in predator-prey systems and help avoid extinctions. This was shown in a famous laboratory experiment done by Carl Huffaker using two mite species, one of which was a predator on the other. The prey mites can live on oranges, so Huffaker set up a variety of trays with different arrays of oranges. Refuges & Metapopulations When all oranges were easily accessible from their neighbors, the predator mites quickly ate all the prey mites, and one or both populations went extinct. But when Huffaker isolated the oranges by adding vaseline "walls" throughout the tray that made it harder for mites—especially predator mites—to move from orange to orange, he isolated groups of prey from their predators. In essence, he created refuges for prey, and the populations started cycling. Refuges & Metapopulations Some of the prey mites arrived at an orange without any predators, and their population would increase. Eventually, the predators would find the orange, but by that point some of the prey mites had moved on to another orange. In this way, although individual populations did not survive for long on any one orange, they did persist as a metapopulation across the whole landscape of oranges. Predicting Value of Models Models allow ecologists to predict which factors might be important in determining predator-prey dynamics. Functional Response In a type I functional response, predation rate increases linearly with increasing prey density. In a type II functional response, predation rate increases with prey density, but the rate of increase slows down as prey density gets higher, and the predation rate eventually levels off. A type III functional response shows an S-shaped curve, with predation rate increasing more slowly at low prey density, then more quickly at high prey density. Like a type II response, it gradually levels off at a maximum predation rate. Functional Response The type of functional response and the shape of the functional response curve depend on characteristics of the predator, the prey, and their environment. The Evolutionary Arms Race Environments are dynamic. When a population's environment changes, different characteristics in that population become more or less advantageous. A population's environment includes not only physical factors, but also biological ones, including interactions with other species. Exploitation can favor genetic diversity in the species involved. The Evolutionary Arms Race When one species exploits another, traits that increase the success of the exploiter will be selectively favored, while traits that decrease the vulnerability of the exploited species will also be favored. Exploitation can thus create an "evolutionary arms race" between two species. Coevolution Predator-Prey Coevolution Newts produce a highly poisonous substance known as tetrodotoxin, or TTX. Small amounts of this toxin can kill most potential predators, but garter snakes have evolved ways to neutralize the toxin, allowing them to eat newts. Predator-Prey Coevolution As snakes become resistant to the newt toxin, this selectively pressures the newts to elevate production of the toxin, which again selects for snakes with higher resistance. As the graph at right shows, this reciprocal selection creates a tight match between newt toxicity and snake resistance for co-occuring populations. Trade-offs Coevolution may come with a price. All traits, including those under selective pressure from coevolution, generally involve trade-offs. Some alleles that benefit an individual's ability to exploit (or avoid exploitation) may reduce growth, survival, or reproduction, and the allele's advantageousness may be offset by other disadvantages. Red Queen Hypothesis If one species evolves much slower than the other, the slower evolver suffers a disadvantage and would likely go extinct. Species that evolve fast enough to keep up with (or outpace) evolution in their enemies will generally persist longer than those that evolve more slowly. Like the Red Queen, a species and its enemies continually evolve "to keep in the same place." Red Queen Hypothesis Sexual reproduction is energetically costly. Sexual reproduction, through recombination, speeds up the rate of evolution, improving its practitioners' ability to survive in a Red-Queen world. The Red Queen hypothesis is a likely explanation for the evolution of sexual reproduction.