Biology 313 Notes

What is Competition? An interactions between individuals for limited resources that reduces survival, growth, and/or fertility Inevitable consequence of increasing population size + limited resources (sets boundaries) o Leads to evolutionary change (i.e. those failing to compete are not selected for in evolutionary terms) Competition is extremely common in nature almost every interaction is competitive

Intra- vs. Interspecific Competition Intraspecific Competition: Competition between individuals of the same species o More likely to be more intense due to similar dietary/lifestyle requirements and needs for resources Interspecific Competition: Competition between individuals of different species o When two species are phylogenetically similar, this type of competition can then be strong too o Important in determining: Natural Selection Fixation of adaptive traits i.e. what we see now is a result of competition in the past Phenotypic divergence between species 2 species with similar traits will look different over time due to competition Abundance and distribution in species Species coexistence

Interference vs. Exploitive Competition Interference = DIRECT Exploitive = INDIRECT o Interact via a lower trophic level o The two individuals dont need to physically interact with one another, yet indirectly effect
eachothers fitness

Contest vs. Scramble Competition Contest Competition Monopolization of resources o Requires a resource that can be defended o Some individuals are able to monopolize a particular resource in short supply o Often associated with ritualized behaviours or combat often resulting in injuries/fatalities o Results in a very uneven distribution of resources Many individuals may not get any resources, while some get a lot Scramble Competition Free-for-all o All individuals have access to a limiting resource and acquisition is a free-for-all o Resource distribution may still be unequal due to individual competitive ability not being identical o Monopolization not possible Sometimes competition can start out as scramble and result in contest competition o Ex) Plants start growing = scramble.those that outgrow the smaller individuals monopolize the sun as a resource as well as nutrients with longer roots and this this is turned into contest competition o Ex) Some pea-beetles choose to engage in scramble competition in large peas because the space is less monopolized, while in smaller peas contest strategies take place because space is small enough that it is monopolized

Comparing Plant and Animal Competition Plants are sedentary (fixed position and have a relatively smaller distance over which competition occurs) A plants immediate neighbours are more important than the entire population due to its inability to move Some sedentary animals (i.e. barnacles) are unable to move and thus exhibit similar patterns as plant competition Plants have NON-SUBSTITUTIBLE requirements for life (nitrogen, oxygen, carbon dioxide, light, etc) and thus INTERspecific competition is COMMON over a wide range of plant species

Lab Models of Competition Indefinite Coexistence only happens at reduced population densities and abundance < K for each species Single Species Extinction intense competition One Species Leaves Area of Overlap mobile organisms choose to leave an area by assessing risks at the site due to pre-colonization of another species at the time

Competitive Exclusion Principle o Exclusion of one species often occurred between closely related species in highly simplified environments where resource partitioning not possible, no emigration/immigration o One species of these two similar species will gain an edge and compete out the other species (on goes to K, the other goes to zero) thus when seeing coexistence, one must infer that the species are somehow using the environment differently

Mathematical Models of Competition Lotka-Volterra model of interspecific competition Allows us to predict outcomes based on different combinations of K and There are FOUR graphical possibilities Competition coefficients () they give you a measure of the individual of another species on the members of your focal species o If 12 = 1, then each individual of Species 2 has the same effect on an individual of Species 1 o If 12 > 1 Species 2 is a STRONG competitor o If 12 < 1 Species 2 is a weak competitor Solving for outcomes in 2 manners: o Given r, N, and for BOTH species and running computer simulations to find an outcome o Given N and for BOTH species, plot the zero isoclines on a state-space graph (plotting abundance of species 1 on x-axis and species 2 on y-axis) and infer an outcome based on relative position of each line (More likely for the exam)

Graphical Representations of Models Any point on the plot represents the abundance of each species at t = 0 The zero isoclines go through points that represent combinations of abundances that represent no growth of that particular species There are three outcomes to competition: o Species 1 grows to K1, species 2 goes extinct o Species 2 grows to K2, species 1 goes extinct o Both species coexist and N1 < K1 and N2 < K2

Scenario 1: The first scenario is one in which the isocline for species 1 is above and to the right of the isocline for species two. For any point in the lower left corner of the graph (i.e., any combination of species abundances), both populations are below their respective isoclines and both increase. For any point in the upper right corner of the graph, both species are above their respective isoclines and both decrease. For any point in between the two isoclines, species 1 is still below its isocline and increases, while species 2 is above its isocline and decreases. The joint movement of the two populations (thick black arrows) is down and to the right, so species 2 is driven to extinction and species 1 increases until it reaches carrying capacity (K1). The open circle at this point represents a stable equilibrium. In this scenario, species 1 always outcompetes species 2, and is referred to as the competitive exclusion of species 2 by species 1. Scenario 2: The opposite of the first; the isocline of species 2 is above and to the right of the isocline for species 1. This graph can be interpreted in much the same way as the previous one, except that the joint trajectory of the two populations when starting in between the isoclines is up and to the left. In this case species 2 always outcompetes species 1, and species 1 is competitively excluded by species 2.

Scenario 3: the isoclines of the two species cross one another. Here, the carrying capacity of species 1 (K1) is higher than the carrying capacity of species 2 divided by the competition coefficient (K2/a21), and the carrying capacity of species 2 (K2) is higher than the carrying capacity of species 1 divided by the competition coefficient (K1/a12). Below both isoclines and above both isoclines the populations increase or decrease as in the first two scenarios, and there is an unstable equilibrium point (closed circle) where the isoclines intersect. For points above the dashed pink line (species 2 isocline) and below the solid yellow line (species 1 isocline), the outcome is the same as in the first scenario: competitive exclusion of species 2 by species 1. On the other hand, for points above the solid yellow line (species 1 isocline) and below the dashed pink line (species 2 isocline), the outcome is the same as in the second scenario: competitive exclusion of species 1 by species 2. The two stable equilibrium points are again represented by open circles. In this scenario, the outcome depends on the initial abundances of the two species. Scenario 4: The isoclines cross one another, but in this case both species' carrying capacities are lower than the other's carrying capacity divided by the competition coefficient. Again, below both isoclines the populations increase and above both isoclines the populations decrease. In this case, however, when the populations of the two species are between the isoclines their joint trajectories always head toward the intersection of the isoclines. Rather than outcompeting one another, the two species are able to coexist at this stable equilibrium point (open circle). This is the outcome regardless of the initial abundances.

Ecological Niche Naturalists and ecologists collected info on habitats and feeding relationships of animals and plants Over time patterns were recognized in terms of how organisms differed in requirements for space, food, light, etc. Descriptions of where you could find the organisms, their requirements for growth Grinnell o described a niche as the sum of the habitat requirements allowing a species to persist Elton o described a niche as being an organisms role in the environment o Both Grinnell and Eltons descriptions were qualitative rather than quantitative Hutchinson o Said that you could look at all of the variables contributing to organisms ability to persist pH, salinity, other abiotic factors, species interactions o He took the variables and described them graphically in an n-dimensional hypervolume with n-variables

The realized niche will always be smaller than the fundamental niche, and will generally be seen in the wild since it is unlikely to have no species interactions in a natural environment

Evolution of Competitive Ability Competitive ability does in fact change over time The difference that we see among various organisms can be accredited to evolution in their competitive ability over time o Drosophila example Populations kept at high and low densities Low density populations showed less competitive ability High density populations showed greater competitive ability Showed that over 128 generations that competitive ability can in fact change through evolutionary mechanisms over time

Minimizing Competition MacArthur was concerned with how certain similar species managed to coexist He found various warblers: o Were similar in body size o Tended to eat the same species of insects o Resources appeared to be limiting o Often within the same tree foraging This was found to be occurring by resource partitioning competition was minimized here by dividing the tree into relatively distinct feeding zones

It is important to examine HOW species manage to coexist over time and how competition is thus minimized over time Ecological Character Displacement - the phenomenon where differences among similar species whose distributions overlap geographically are accentuated in regions where the species co-occur but are minimized or lost where the species distributions do not overlap Two populations destined to become two different species must be very similar initially, and then become more different over time As long as there are some reasons present, the relative fitness of each species genotype will change over time o It is not good to be the initial similar genotype of the similar species due to competition the intensity of competition depends on your relative trait value (what species you belong to) o Selection over time favors the two extremes

Predator-Prey Dynamics Predation when one organism feeds on another, thereby killing and consuming some portion of it (+/- interaction) Prey/Victim individual that is killed/consumed Herbivory the loss of tissue in plants due to grazing from herbivores typically plant isnt killed, only grazed The number of prey is affected by number of predators, and vice versa Prey numbers negatively impacted by predation as predators need to consume prey to survive and reproduce Prey often have adaptations to reduce the predators profitability of predation through biological defenses Predator-prey cycles can be observed through time o Not particularly stable numbers, yet the effects of the relationship on each populations fluctuations can be observed o The peaks and troughs of the predator occur AFTER peaks and troughs of prey o Predators are usually physically larger than prey and thus the number of prey in the cycle is observed to be consistently higher over time than the number of predators Food Availability how it effects cycles o Near peak densities of populations, shortages of food are observed and the quality of food also decreases o In certain plant species, some choose to facultatively increase defense mechanisms to prevent herbivory, especially in short growing seasons Predation how it effects cycles o At peak densities, 60-90% of mortality is due to predators a great deal of a crash in the cycle is accredited to predation rather than food availability

o Generalist predators even choose to turn into specialist predators due to a large influx in prey populations contributing to the great decline in prey populations Lotka-Volterra Predator-Prey Model

Change a biological parameter, and the model fits what we would expect to see (above).

Change a non-biological parameter, and the model does NOT fit what we would expect to see (above). Changing P0 and V0 should have not resulted in drastic changes therefore there is a problem with the model.

Base model does not incorporate competition among prey or predators (intraspecific)

Include a term in the model that invokes densitydependence and you will have dampened oscillations in the prey shows a greater degree of realism and a stabilizing effect in the population.

Need to consider things like handling time for predators of prey, and search time too (both of which the model neglects). Capture rate ultimately levels off at a certain value as prey number increases due to these parameters (Type II Functional Response).

Density-dependence in prey + Type II Functional Response in predator = stable limit cycles

The Prey Capture Process 1) Encounter o Predators need to be physically close to prey to capture them o Prey can evade predation by being active at different times of day, or by leaving/avoiding areas of vulnerability 2) Detection o Predators need to detect their prey o Prey can reduce their ability to be detected by moving unpredictably, confusing predators by being in large groups, etc 3) Identification o Predators need to recognize that the individual they have encountered and detected is in fact consumable prey o This is prevented by the prey through i. Crypsis ii. Aposematism iii. Mimicry 4) Approach o Predator now must move in to capture prey o Prey can outrun predator or retreat to refuge 5) Subduing Prey o When prey is captured, it must gain control/kill it to consume it o Prey may be able to escape predator still, detach body parts to escape, or rely on chemical substances to deter predation

Crypsis Blending into the environment physically (body shape, body color) Crypsis doesnt mean that an individual is monochromatic, yet it can be that they can have dynamic colorations/body shapes main message here is that it really depends on the environment the organism is persisting within

Mimicry The attempt to RESEMBLE a distasteful species Mllerian: both the copycat and original species distasteful o Most likely to evolve when both species are equally unpalatable and equally common o If you have a rare unpalatable signal, you will most likely still be sampled by predators and thus it makes sense to follow the status quo Batesian: one species is palatable (mimic) and the other is not (original)

Aposematism Opposite of the cryptic strategy Rather than blending in, individuals choose to stand out relative to their environment The prey has toxic compounds that make it unpalatable These prey honestly advertise their toxicity via bright colors/striking color patterns

Some question whether or not aposematic lineages more successful than nonaposematic sister groups It is true that aposematism is associated with greater species richness There is reduced predation risk on these individuals and this appears to be associated with increased speciation rates and/or reduced extinction rates in aposematic lineages This doesnt ALWAYS occur since crypsis can be very effective as well, and the proper chance mutations must occur for it to be beneficial thus it is not ubiquitous

Herbivory Herbivores derive energy and nutrients from the plants they graze on Grazing on plants puts selective pressure on these plants then leading to increased specialization within herbivores Plants have an impressive arsenal of techniques to prevent herbivory Herbivory is generally not good for the plant plants invest a great deal of energy into producing tissues only to be consumed by the herbivores An example of a plant defense is trichomes o Hooked small plant hairs o Implication as a successful defense strategy to defend against larvae species o The species with the trichomes was found to be the most abundant within the genus due to the defensive strategy present o Certain larvae species were able to compete with the plants defensive strategy by creating silk scaffolding allowing them to prevent injury from trichomes while still benefiting from herbivorous behaviour The latex resin production defense strategy used by plants to kill herbivores attempting to consume the plant overcome by the use of petiole constriction by herbivores (to prevent latex resin flow allowing for successful herbivory)

Generalists vs. Specialists Debate of whether generalists give rise to specialists and once a specialist always a specialist? This part of the lecture was not really a core concept thus, dont worry about it very much

Invasion of Exotic Plants Are plants not native to a place that they are newly introduced to successful since they have no native herbivores to control their growth? Example: Two alien species (P. aduncum and P. umbellatum) o It was found that results didnt follow expectations that the two invasive species would have less herbivory by caterpillars due to a lack of native herbivores o Rather, herbivore rates on the alien species was high o It was found that the herbivores fed on a single family of plants and thus aliens were not immune to attack hypothesis put forward (first point) proved to be incorrect o P. aduncum were indistinguishable from the native plants of the area and dominated by a single species, while P. umbellatum dominated by multiple species (thus not all alien species the same in a new environment)

Parasitism Successful attacks of parasitoids should kill the hosts Hosts creates defenses against attacks o These two points lead to coevolution of the host and parasitoid For parasitoids to proliferate, they must have a means of attaching to hosts for a long period of time, must be able to overcome defenses of hosts, or simply go under the radar of the host Even though parasites are typically the enemy, they have their own predators (hyperparasitoids) Parasites have been considered to be useful control agents to control pest host populations because they typically specialize on the host species Being a parasitoid is the one of the MOST successful modes of life o This is determined scientifically by how often they have evolved and by currently looking at how many parasitic species are currently in existence

What is a Parasitoid? Insects using other insects for development of their larvae Larvae usually develop within/on host = usually death of host, survival of parasitic larvae Parasitoids of parasitoids are called HYPERPARASITOIDS
Hyperparasitoid Pupa
Parasitoid to...

Parasitoid Pupa

Parasitoid to...

Moth Larva

Nicholson-Bailey Model Model to examine coexistence of parasitoids (P) and hosts (H) Assumptions of BASE MODEL o No density-dependence in population growth of P or H o Every P equally successful at attacking/laying eggs in H o All attacked H DIE and give rise to new P Only H that have ZERO ENCOUNTERS with P can reproduce

Base model produces unstable large amplitude oscillations. Without any new hosts, there are no new parasitoids at t = 22. The base model doesnt model the coexistence we see in real life.why?

When considering the following more biologically realistic conditions stated below, the model is functional: o MUTUAL INTERFERENCE COMPETITION within parasitoid populations causes one or both parasitoids to leave population invoking a sense of density-dependence in P o FUNCTIONAL RESPONSE IN PARASITOIDS: parasites cant attack endlessly as model dictates o SPATIAL COMPONENTS: equidistribution as the model predicts is not likely and therefore we must consider the possibility of clumped distributions of organisms

When altering base model and incorporating various variables, the model yields more realistic results. Interference competition between parasitoids is a huge factor (because it causes D-D in P).

When thinking about the spatial components, it is important to think about subsequent generations in patches of space o Many H = Many P next generation has Fewer H = Fewer P o Although there is local fluctuations, there is stable coexistence across over an entire landscape as the simulation below displays with local hotspots/coldspots

Parasite Transmission Direct Transmission o Transferred from one host to another DIRECTLY (ex. rhinovirus cold virus in humans) Transmission Through Intermediates (VECTORS) o Vectors are not usually adversely affected by the parasite
Definitive Host Parasitoid Vector(s) Definitive Host

Life cycles of many parasitoids are linked to life cycles of one or more hosts The DEFINITIVE HOST is the host in which the parasitoid will produce (final step in the line of transmission) Intermediate hosts (IH) = vectors and are found in the habitat where definitive host (DH) is usually found o IH must allow P to find its way back to DH o IH is usually prey to the DH, or at least found nearby the food the DH will consume o Parasite may alter the behaviour of the IH in an attempt to perpetuate cycle

Coevolution of Hosts and Parasites There is fairly good evidence a gene-for-gene system with H and P o The synthesized proteins by P are foreign to H, and selection will FAVOR the hosts with the ability to detect and resist P over the course of time

Virulence The additional per capita mortality in host caused by infection If something is highly virulent, host populations will have lower potential fitness, higher mortality rates, etc. Virulent strains have greatest host-to-host transmission produce more copies of itself per unit time
The more virulent strain (in this case the teal strain) is more likely to be passed on to a mosquito, for instance, because when consuming blood it is more likely to take with it the more numerous teal parasitoids). This correlates to higher fitness in the teal strain since it is more numerous (more virulent) = further proliferation of strain in hosts.

Selection and Virulence Is there evidence that highly virulent interactions are selected for over time? It was traditionally thought that any instance of a Host-Parasite interactions where P was highly virulent must be of recent origin, proceeding to a state of commensalism o This was thought of being a faulty manner of thought It is NOW known that natural selection can actively favor highly virulent P over long periods of time The following factors influence the level of virulence observed in a H-P interaction o Characteristics of the vector in transmission o Mode of transmission Host dies before it can pass on disease through o Degree of population structure in host direct transmission to other host/before
High Virulence Intermediate Virulence Low Virulence another mosquito can further transmit disease Favored balance of virulence titre and ability to transmit disease successfully Host comes in contact with other hosts, but doesnt manage to pass on disease due to low virulence titre

Infection

Modes of Transmission Horizontal Transmission: P move from one H to another H within the SAME generation o Implies MORE VIRULENT due to the strain killing host before it can reproduce Vertical Transmission: P moves from host parents to their offspring o Implies LESS VIRULENT since the parasite can allow the host to reproduce before killing them Think of these two modes of transmission in terms of a family tree:
Parent Parent
Blue offspring created before parental infection and spread within own generation (horizontal transmission). Red parent transmitted disease to a third individual through vertical transmission. Offspring Offspring Offspring

Mutualism Interaction that is mutually beneficial for parties involved Can still be best thought of as mutual exploitation rather than cooperation
Symbiotic Relationships Parasitism Mutualism

Commensalism

Resource-Resource

Resource-Service

Service-Service

One resource traded for another interaction based upon resource need/availability (ex. N-fixing bacteria and legumes legumes get N, bacteria get carbohydrates).

One resource is traded for a resource (ex. Nectar and pollen traded for pollen dispersal). Zoochory is the dispersal of seeds by animals usually fleshy nutritious fruits of plants desired by animals.

Services exchanged (RARE). (ex. clownfish in anemones clownfish receive protection, anemones defended by clownfish that would otherwise be attacked by butterflyfish.

Symbiosis: Long term interaction between two species (parasitism, commensalism, mutualism) Commensalism: An organism benefits, the other is not benefitting or being harmed Sometimes there can be occasional cheating within a mutualistic relationship o Ex) Cleaner wrasses occasionally feed on tissue of host fish they help clean o Ex) Orchids that mimic nectar-producing orchids to attract insects for pollination, but dont produce nectar (mimicry) o Ex) Nectar robbers that chew through flower corollas to receive nectar without pollinating flower Mutualism is believed to have evolved from an initially negative interaction, and mutualism evolves when the benefits outweigh the costs

Interesting Example of Mutualistic Relationship Domatia and Predatory Insects o Domatia = small hairs or depressions on the bottoms of plant leaves that were thought to be refuges for predatory insects that plants want to attract o The domatia allowed for predatory insects to ward off herbivorous insects protecting the plant from herbivory Ant Guard Example (refer to notes)

Introduction and Species Abundance Communities o An assemblage of interacting species in a defined area o Consist of many species (10-20) impacted by various types of interactions o Defined by the features of the environment to an extent Intertidal zones is an example of an area that is spatially defined where these interactions could occur Community Structure includes: number, kinds, and relative abundances of species Species Abundance o It is equivalent to (the relative abundance of species 1) (n in the focal community) o When considering this, determine if there is a dominant/more abundant species o Dominance in terms of abundance can be defined by the amount of encounters made with that species in the community o Lognormal Distribution is used to quantify species using relative abundances Obtained a normal curve on a log scale Species Diversity o A combination of the absolute number of species and their relative abundance o A measure of variation in the species that are found in a community o Species Richness (s) Number of species in a community A community with s = 23 is less diverse than one with s = 59 species o Species Evenness Relative abundance of species If there is no species dominance in a community, then there is species evenness o Shannon-Wiener Index Measures species diversity Minimum value = 0 when s = 1, and increases with species richness and evenness

Factors Affecting Diversity Environmental Complexity o Consider two environments of equal geographic extent/volume, but with one being more complex than the other (more mountainous vs. flat surface) o The environment with greater complexity = higher species diversity o More complexity = greater ranges of abiotic factors Ex) Forests with greater foliage height tend to support greater bird species diversity Disturbance o Intermediate Disturbance Hypothesis Proposes that disturbance influences species diversity in communities In undisturbed environments, conditions tend not to change, and a dominant species will overtake the environment over time In disturbed environments, conditions tend to change preventing a dominant species to overtake other species Ex) Forest fires

o An example of how disturbances effect species richness comes from the boulder idea Small rocks in the tide zone were turned over frequently, killing all on it Large rocks in the tide zone were infrequently turned over allowing dominant species to thrive Medium rocks were turned over at a rate that allowed high diversity preventing a dominant species from being apparent in the community SPECIES RICHNESS LINKED WITH DISTURBANCES

Species Interactions o Coexistence can only occur for many species if intraspecific competition is greater than interspecific competition o Typically the competitive interactions of certain species will greatly affect the diversity within the community, especially when the needs for life overlap to some extent

Community and Related Structure What is a Community? o Contains all of the organisms living within a defined geographical area o Includes those of same/different species o Involves the direct and indirect interactions between these species What is an Ecosystem? o Essentially encompasses multiple communities while acknowledging the abiotic factors that influence the abundance and distribution of organisms

Communities Elton first documented feeding relationships in communities revealing a flow of energy Feeding relationships are links in food webs Lindeman organized the relationships into tropic levels o Separation of PP, C, SC, TC, etc. o Noticed that the amount of energy in upper trophic levels < energy of lower trophic levels general inefficiency of energy transfer (laws of thermodynamics)

Trophodynamics

Bottom-Up Regulation
Abundance of primary producers are believed to influence the amount of primary consumers in a community Is proven to be true to an extent when looking at the proportional relationship between algae and herbivores

Top-Down Regulation
Belief that biomass at most trophic levels maintained by predation from the top Green World Hypothesis predators control herbivory allowing plants to flourish

Both ultimately function in shaping communities and regulating trophic level dynamics, and this effect is best seen when seeing effects of a predator in a community before and after its introduction

Keystone Species Certain species that exert strong effects despite having low biomasses and relative abundances Not to be confused with species, that when removed from the system, have a large effect on the dynamics of said system simply because they are largely abundant need to look for the individuals of low biomass that when removed = huge effects Dominant Species = species with the largest abundance in a community with great effects on community structure by virtue of large biomass Keystone species arent always predators o Consider the cleaner wrasse fish that removed parasites from certain species in an interaction that is mutualistic o When cleaner wrasses removed from system, diversity of fish species decreased

Trophic Cascades The concept of how top-down control in a community can cause variable cascading and fluctuating effects on species in lower trophic levels This concept holds true in systems with complex dynamics

Food Web Matrices Lists the potential predators and prey Communities can be summarized under this notation Prey = row entries, predators = column entries Higher # in matrix corresponds to being in a higher trophic level 1 in the matrix indicates the predator in that column feeds on prey of that row 0 = no interaction 0s along diagonal unless cannibalism is present Basal Species = one that is prey for most species and does not eat other species Top Species = one that feeds on one of more species but is not prey for others

Influences on Ecosystem Size and Food Chain Length Two Non-Mutually Exclusive Hypotheses

Productivity Hypothesis

Ecosystem-Size Hypothesis

- Mean chain length is proportional to the amount - Food-chain lengths should be greater in areas that of energy at primary producer level cover greater areas - (i.e. more energy = more trophic levels) - (i.e. larger ecosystem = more trophic levels) Both are true in different cases, but it tends to be the Ecosystem-Size Hypothesis that is more often true Dynamic Stability Hypothesis (DSH) Proposes that long food chains are less stable than short food chains This is because population fluctuations at the bottom of longer chains may result in the extinction of top predators Combines aspects of both the productivity and ecosystem-size hypotheses

Energy Flow

Exploitation Efficiency

Assimilation Efficiency

Net Production Efficiency

Nutrient/Biogeochemical Cycles Carbon, nitrogen, sulfur, phosphorus and collectively called essential nutrients Earth is a closed system with respect to essential nutrients Large pools of nutrients are called reservoirs Flux o The rate of movement of one component of the biogeochemical cycle to another o Measured in amount of nutrient per unit time Hydrological Cycle o Largest reservoir = oceans o Greatest flux = oceans

Phosphorus Cycle o Largest reservoir = marine sediments o Largest flux = on land between organisms and soil Carbon Cycle o Largest reservoir = carbonate rocks o Largest flux = increasingly becoming the addition of CO2 from fossil fuel use into atmosphere Nitrogen Cycle o Depends critically on microorganisms that convert one form to another o Ammonification = organisms derive energy from this reaction o Largest flux = oceans Ecotron Experiments o Found that reduction in species diversity may negatively impact ecosystem function o Ecosystems with greater amount of species = increased PP o Provides further reason to adding to conservation efforts of species because this may lead to irreversible environmental change.

Landscape Ecology The study of relationship between spatial patterns and ecological processes over a large range of scales Landscape heterogeneous area consisting of distinctive patches of landscape elements organized in a mosaic-like pattern Landscape Elements forests, meadows, ponds, etc Considers humans and their impacts on the environment Involve effects of organisms on environment (beaver), fire, and glaciers

Glaciers 100,000 year cycles o Glaciations = very cold periods (60-90 K years) last was 20,000 years ago (Wisconin) o Interglacials = warmer periods (10-40 K years) Different ice sheets retreated in different ways effecting populations Glacial Refugia = areas where populations persist during glaciation o May result in phylogeographic lineages that are genetically distinct o Refugia have consequences on the genetic composition of lineages

Landscape Structure Measured through analyzing o % cover of patches o Relative shapes of patches amidst a continuous matrix background Organisms prefer more circular patches (do not like edges of patches) The effects of fragmentation of the environment have a greater effect on small mammals than organisms like birds that are highly mobile o Being small + need for mobility to forage in fragmented patches = more energy lost + mobility increases chances of being consumed by predator = increase mortality o Increased fragmentation = increase chance for extinction

Island Area and Species Richness Increase in island size = increase in species richness due to possibility for increased geographic diversity There are limits to this relationship there are boundaries that limit species richness on a large island such as climate/productivity, etc. Mountains can be thought of as islands o Patches of island vegetation act as island-like habitat patches o Once continuous forests became scattered and dispersed habitat patches as a result of climate change forming montane habitats o Increased area of montane habitats = an increase in species richness Lakes can be thought of as islands o An increase in lake area = increase in species richness o Lakes that had similar maximal depths and lake bed substrates shared this relationship

Island Isolation and Species Richness Increase in island isolation = decrease in species richness This relationship is relative to distance to the nearest mainland, which will be the source of immigrants For different organisms, dispersal ability comes into play (i.e. their ability to travel to distant islands from a mainland to populate) It was found that when comparing birds with ferns, that distant islands correlated with a decreased species richness of birds when taking island size into consideration too Conversely, it was found that certain fern species were unaffected by island distance thus indicating highly effective dispersal ability compared to bird species

Equilibrium Model of Island Biogeography Species diversity on a given island is a balance between immigration of new species and extinction of existing species This rate of immigration = fast on a bare island, declining as species accumulate because fewer and fewer species will be new species Rate of extinction believed to rise with number of species present because o More species = larger pool of potential extinctions o As # species on island increases, population size of each decreases o As # species on an island increases, potential for competition increases Equilibrium point must exist between extinction and immigration because they are opposing factors The number of species is a delicate balance between immigration and extinction Isolation effects immigration, while island size effects extinction

Latitudinal Biodiversity Gradient When moving from poles to equatorial regions, biodiversity said to increase There is a strong relationship for marine and terrestrial organisms, and slightly less strong for freshwater organisms Strength of latitudinal gradient strongest for species in higher trophic levels, and for larger organisms in general Carnivores were found to generally be more strongly effected than, say, autotrophs Mechanisms promoting this o The tropics are older than other places (i.e. not effected by glaciations) Therefore possibly less extinction in the tropics o More geographic area in the tropics than extratropics = more species richness o There is a greater spread of temperature being steady across the -25/25 degree latitudes that encompass the tropical areas leading to Decreased generation times A larger range in species and larger possible population size