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What about the predator? • Again, seem to forget the predator in all this. • What adaptations do they have to try and make attack successful? Anatomy…. • Obvious that they have the tools • Specific to each species as to which are used more • But what about the behaviors? • Are there things they can do to improve chances? Obvious ones • Cursorial vs ambush • Solitary vs group or pack here Attack points • Some prefer the back end (usually cursorial ones) • http://www.youtube.com/watch?v=CT_3QiW Qh8M Attack points • Some (stalkers) prefer the back or head region • http://www.metacafe.com/watch/yt7HogWIpfLjc/cougar_and_cub_hunt_a_deer/ Aerial predators? • Owls tend to swoop in low • http://animal.discovery.com/videos/killerclips-birds-of-prey-owl-mouse.html • Hawks tend to hover higher • http://video.yahoo.com/watch/4577360/1225 5461 More • Falcons • http://www.youtube.com/watch?v=j3mTPEuF cWk • Each species has specific attack style. Summary • Two player (often more!) evolutionary game where behavior is the game plan! • Again, a prey’s behavior will revolve around trying NOT to get eaten. • A predator’s around trying TO eat the prey Why doesn’t the predator WIN? If responses by prey are countered by evolutionary responses of the predator. Eventually, predator could win! • But remember, which has more to lose? • A predator loses a meal • A prey loses its life! • Selection pressure is greater for the prey! What next? • The stage: Habitat is where it all happens • Talked about use of habitat in reference to foraging theory, landscape of fear. • But again in more theoretical aspects • Now look at some of the more detailed behaviors associated with habitat use. • Chapter 13 What First? • Had talked about animals selecting habitat for resources, for safety, etc. • On the landscape scale, ample data indicate individual animals will use the same area repeatedly. Example • Coyote locations over 9 weeks • This we commonly refer to as the Home range Burt 1943 • First proposed the concept and that an animal did not roam randomly around. • “Home range then is the area, usually around a home site, over which the animal normally travels in search of food.” • “Even then I would restrict the home range to that area traversed by the individual in its normal activities of food gathering, mating, and caring for young.” Home range behavior • So animal is purposely restricting its movements. • Reasons? • 1) Nest site: becomes a central forager, efficiency of food gathering decreases with distance • 2) becomes familiar with where food is distributed, phenology of food abundance. But still… • If don’t have central nest site (many animals don’t), why restrict your activity to a given area? • Would possibly lead to over use?? • What other reason? Predation risk • Again, trade off between foraging and safety. • Might be advantageous to move randomly over a large area • But, with each new area, unfamiliar with the landscape of fear! • Where are the safe and risky areas? • Best to stay in the “hood” than wander. How big should the “hood” be?? • Ok, if your going to stay in one area, be it for familiarity with resources or predation risk… • How big of an area should it be? • Age old question: • Some say related to size of animal: Regressions of various mammals Looks good but…. • Feral pigs: size difference in a species adds variation. Some say it is density! • Same study found inverse relationship between home range size and density. Others? • • • • Some say it is latitude Some say it is productivity of habitat Both related to either mass or density so…. But doesn’t get at the basic question of why is it as big as it is! • Indicates maybe resource levels but does not hold as well. • How about fear!! profiles • Please email your profiles to me at: [email protected] • I have easier access to this address. here Predictions • 1) If it is fear, then home ranges of each prey individual should have similar range in makeup of risky vs safe areas. • 2) across species should also see relationship • Should be trying to carve out an area that provides sufficient safe areas or refuges. • To my knowledge this has yet to be tested. What about the predators? • Best to learn the landscape of opportunity. • Again, not where prey most abundant but where they are most vulnerable. • As with prey, home ranges of each should have sufficient successful habitat • Prediction: home range size will be a function of quantity of successful hunting habitat. This we tested! • Puma edge talk. So that is large scale use • Within home range we are saying that animals are selecting certain habitats more than others. • How do we know they are selecting? • What is our reference point?? What should we use as a model? • IF we say that habitat use is resource based • That is, a habitat with a higher level resource should be used by more individuals • The question becomes what would the distribution rule be? • Ideal Free Distribution (IDF) suggested model What is IDF model? • Simplest is two patches (H1 and H2), each with different resource levels (R1 and R2) • R1 is five food units/minute • R2 is three food units/minute • If N individuals can move freely between the two patches, how should they distribute themselves? (how many in H1 and How many in H2?) The prediction • IFD predicts that: equilibrium distribution of individuals into patches should be that distribution at which, if any individual moved to the patch she was not in, she would suffer a reduced payoff! • In this case, it is if any individual moved from H1 to H2 or visa versa, it would obtain fewer resources as a result of its move. Visually what does this mean? • Imagine animals moving around at will. The more enter one patch, the less resource/individual/time is available. • If one moves from H1 to H2 and If it can do better in H2, it will stay. If not, no reason to move! • Means they all assess their relative harvest rate compared to each patch and don’t move anymore when they can’t do better. Mathematically? • Equilibrium is when R1/N1 = R2/N2 • When per capita intake rate in both patches is equal • Example: simplest is 5/5 = 3/3, Does it apply? • Laboratory situations where resource level only selection criteria: fairly good. Results • Separated out as predicted AND changed when ratio changed In the field? • Again, under controlled conditions. Results • When food was 1:1, ducks were 1:1, when food was 2:1 the ducks were 2:1 Conclusion • So, when resource levels are the only consideration, seems to work • Conclude that ALL ELSE BEING EQUAL, ideal free distribution will predict distribution of individuals over landscape: they will be distributed according to the relative abundance of food resources in each patch. Is all else being equal reasonable? • Again, as with many models, this is the starting point. • IF you do find a IFD, then all else is equal and resources are major factor • In fact, rarely is all else equal and IF you do not find an IFD, that indicates other factors than resources are important. What other factors? • Microclimate: can affect foraging strategies • Resource types: remember each species has ranking of food types so gross resource availability may not be appropriate • Predation risk? One last one • Territoriality: you may not have a choice of where to go! • What is it? Occupation and defense of a particular area or areas within a habitat. • How does it differ from home range? • In some species it is the same! • They defend their whole home range area Territoriality • • • • • In many, it is a subset of the home range In many, it is not evident. How do we distinguish? Home ranges can overlap extensively Territories do not. Advantage? • Home range is to better know the environment you live in: resources, predation risk • Territory is to defend those resources! • Function of a territory is to provide owner with exclusive access to food, mates, safe habitat, etc. Costs of territoriality? • Once you choose to defend an area for its benefits, it will cost you! • First, it doesn’t come easy. • If you figure it is worth defending, others will too! • So defense of a territory has to be vigorous, no half-hearted measures, if so, you will lose! Time commitment • Territorial defense is a 24/7/365 time commitment. • You have to expend a lot of time non-contact time making sure no one is trying to take it over! Here Is it worth it? • No brainer: must be, if not would not see this behavior. • Real answer is: it depends on what you are defending. • If the resource is mating opportunities, then YES it is often worth it. • Majority of territorial behavior is related to sex Mating territories • Mentioned before, males of many species stake out courting and breeding territories. • Leks to song birds all actively defended “space” or territories • Male mountain lions • Size defended depends on mating systems Resource territories • Here it becomes less evident. • Many herbivores, resources are not restrictive and so see little resource territoriality. • Will see a little more in predators. • Here they may not be defending the resource but the habitat in which to catch the resource. Predatory resource territoriality • If a predator’s ability to catch its prey declines as it enters a patch (prey become more aware), needs to use its area in a manner where areas “rest” long enough for prey to drop their guard. If others use the patch? • If other predator visits patch, becomes more difficult. • Prey will have a higher level of vigilance than predicted. • So defend use of patches to “manage” fear in prey. Summary • Defending an area (territory) has its advantages. • Costly but if resource worth it, will be done. • Usually most common related to reproduction. Establishing a territory • Ok, territory is good but how do you decide what area to defend? • Limit ourselves to none breeding territories • Evidence indicates that Learning behavior figures in. Learning and territory establishment • Example is juvenile lizards (Anolis aeneus) • Found that juveniles established territories based not on food resources but on temperature AND safety from predation. • How did they determine an area had these requirements? Observed others? • Prediction was that juveniles observed other older lizards and learned which areas were more suitable. • Experiment: allowed them to watch others in territories and see another area where there was no lizard. • Removed occupant and then gave test animal opportunity to select. Results? • Arrived sooner to pre-occupied areas • And used them more So learning from conspecifics! • Strong visual component: IF test animal did not see occupant before, no preference. • Referred to as: Conspecific cueing. Sharing your territory?? • • • • Seems contradictory, it’s yours, why share? Obvious male-female sharing Other times? IF you have enough resources to share, you might allow an intruder IF it shares defense duties. • Case study: Pied wagtails Satellites • Defends a circuit where it looks for food. • IF food abundant enough, allows satellite individuals. Flexibility • What this demonstrates is just how flexible territorial behavior is. • Can be confusing at times because of the variation among AND within species! • Add to this the size variation • How to best visualize it? Territoriality is a matter of scale • • • • Looking at social behavior expanded Personal space to territorial space No species is indeed then truly solitary Live imbedded in a social system established by interactions among members. • Could be daily interactions as in truly social species • Could be occasional interactions among territorial occupants. • Pack is territorial behavior condensed • Solitary behavior is social behavior expanded. • What determines if you have a large or small “territory”? • Resource abundance, Resource availability, predation risk. • What is required to extract a living AND stay alive! Migratory behavior • Now lets contrast home range and territorial behavior with Migratory behavior • Though many species do have home ranges and even territories, they don’t stay in them all year. • May be a relatively short distance (deer moving up or down slope) or from pole to pole (arctic tern) Birds and more • Most striking are the annual bird migrations. • Waterfowl most obvious but MOST birds in northern hemisphere migrate. • Many mammals do it but not as extensive, unless we include the oceans! • Fish!! In and out of the oceans: salmon • Insects: butterflies. Birds as an example • Early naturalists thought migratory birds just hibernated! • 5 billion land birds migrate from Europe and Asia for Africa and similar numbers leave North America! First Patterns • In general movements are intercontinental or at least multi-national movements northsouth Examples of migration patterns How do we know? • • • - Banding programs in winter and summer ranges Doppler radar shows flocks of birds moving Feather chemistry Carbon 13-12 ratios in feathers Will reflect ratio of food source where grew feathers - Compare summer and winter areas, match feathers to sources of origin. General patterns • There are certain migratory routes that birds will use for generation after generation • In North America, they primarily follow mountain ranges and coast lines/Mississippi river: waterfowl follow major flyways. • In Europe and Asia, strong east west segments because of e-w mountains and Mediterranean sea. North American Flyways Western Hemisphere routes General patterns • Many fly over oceans and so routes vary. • Characteristics of routes taken - Easiest flying (not crossing mountains, bypassing large bodies of water, more thermals on land) - Easiest: leads to bottlenecks: narrow areas over water, Straits of Gibraltar. - Dependent on individual species General patterns • Daily timing of flights - General pattern is to chose times when travel is safest and most rapid. - Depending on species: lot migrate at night - Why? Safer for sure, also air can be more stable, cooler (heat loss) - Altitudes: low mostly (700-800 m) some high to avoid turbulence Up to 9000 m recorded. Altitude of flight Migration distances • Vary in length relative to how far they need to go to find the right conditions (some individuals of a species DON’T migrate) • Many go 100’s, even 1,000’s of kilometers - Arctic shorebirds > 13,000 km to South America - Bar-tailed Godwit 11,000 km nonstop! - Blackpoll Warblers nonstop (> 80 hrs) over Atlantic from East coast to South America Migration Distances • Migrants from Europe fly 1,100 km across Mediterranean Stop and rest and then fly nonstop 1,600 across Sahara Desert! • Besides trying to impress you, what do these migration distances tell us? • VERY energy intensive • So why do they do it?? Why migrate? • Go back to beginning of class: • Need to consider two absolutes: - The day (24 hours or 199 hours) still time from sunrise to sunrise (or any other point) - remember marking time artificial: hours ! - The year : 365.25 days to complete trip around sun. Annual Cycles • So, the two absolute time frames are Cyclic! • Start at one point and you eventually get back to it. • Time may be linear but it passes in a cyclic fashion. Annual Cycles • Annual cycles involved in responding to annual changes in environment because earth is a ball traveling around the sun. Annual Cycles • Review: position in orbit imparts specific conditions Result • Again, related to physics of earth placement and movement. • Some parts of globe freeze!! • Most animals have to adjust • Migration is a way to take advantage of these changes. • Many species, whole life is spent going from “winter” to “summer” territories. Annual cycles • Related to light length • Even in tropics, but here it is food availability. Thursday 4:00 Illick 5 • The ecology of Fear: A unifying paradigm in ecology?? • Dr. John W. Laundré Annual Cycles: how do they know?? • Two main changes: 1) Day to night (24 hour) Circadian rhythm 2) Length of day (Annual or annual rhythm) - here All organisms, plant and animal have physiological controls for changes that occur. These controls are influenced by circadian and annual rhythms: we have “biological clocks that sense changes in rhythms. Biological clocks – where? • Pineal gland on top of brain in birds • Well developed gland with photosensitive cells. • Direct production of Melatonin, regulatory chemical. • Most pronounced is Circadian responses - Daily light level, twilight, triggers switch in physiology. Circadian Rhythms • Intrinsic rhythm of 23 hours! Oops! • Need to synchronize with external cues Zeitgebers or “time keepers” • Experiments show value of Zeitgebers, if not, lose synchrony. Annual rhythms • Endogenous rhythms of approximately one year • Photoperiod (length of day) • Photoperiodic control systems use 2 types of info: 1) Internal circadian cycles help bird measure day length 2) External environmental-light stimulates neural receptors. Sensing annual changes • Circadian rhythms have limited period of photosensitivity per day. - Longer day length means more overlap of light with this period, which can be used to measure day length. • Also have receptors in hypothalamus in brain, pineal gland, and retina! - Longer day lengths are sensed and produce changes. How produce changes? • Neurohormones! Sounds like science fiction! • Light stimulates cells to produce and secrete them. • Master hormones mostly located in pituitary gland. Known Hormones Why Migrate? • So the why is related to food and temperature, escaping cold temperatures in north….but • Why go north to begin with?? Why not stay south?? • So is not as simple as we thought! • What factors do they use to decide? Why Migrate • Still some doubt but there are the ideas. • Based on the obvious that benefits of migration must offset risks and costs • Main idea is that birds aren’t heading south to avoid cold • Tropical birds heading north to take advantage of long light/abundant insects. Why Migrate? • Large expanses of land in the north so less competition- waiting list in tropics! • Reduced predation: If a predator has prey species year round, can specialize. If only for a short period of time, will be less predators and less specialized in a particular migratory species. Why migrate? • IF you are a northern bird (some are yearround residents in some locations), actually spend less energy migrating south (energy of migration vs thermoregulation • Key is “some locations”: If winters severe enough, migration is better, if not, might be worth staying. Sexual differences Differential migration • • • • • Young migrate greater distance than adults Males less distance than females Why? Again balance of needs In Dark-eyed Juncos, Young males least, adult females greatest: Sex advantage for young males- return first for territories. Why??? • Resource based but not a simple answer. • More animals taking advantage of abundant summer resources than avoiding harsh winters • They would consider themselves southern residents who vacation in the north rather than visa versa. How do they do it? Navigation • How do they know where to go? • Again use birds: • Can return to the same marsh/tree etc. after a trip of 1,000s of km! • Holds for both ways! • What cues do they use? Navigation • • • • • What do they need to know? Compass direction: what is N/S/E/W Where they are relative to goal How would we do it? Direction: compass easy Latitude –stars (what a sextant is for). Angle we are relative to north star changes as go N-S • Longitude – a little more difficult need to know time of day! Navigation • So how do birds do it? • Actually a mixture of methods, a “complementary and interactive toolkit”. • Includes a variety of mechanisms and abilities • Consider each. Navigation • First is “visual landmarks” • Animals are no dummies as we have seen. So, like us, when traveling overland, use landmarks. • Why waterfowl and many others have specific routes, coastlines, rivers, mountains. • Even modern railways/highways seem to be used. Navigation • However, does not explain how they get over water or on foggy days/nights! • Sun compass use position of sun to determine direction. • Experiments in circular cages showed birds were directional in their movements when could see sun. When could not, did poorly. Sun compass • • • • Use of sun requires knowing time. Do they? Test with starlings showed Yes! Could trick them by using “sun” that did not move! - Also worked when changed internal clock. - So yes can tell time! Navigation • Star Compass: If flying at night can use stars • Do they? • Planetarium experiments demonstrated YES. -familiar with most constellations. Navigation • Geomagnetism Sensing the geomagnetic fields. • Can they do it? Experiments with magnets tied to heads demonstrated that yes can do it. • Respond to angle of inclination of magnetic field, not just north vs south!! • Respond to changes in intensity of magnetic field!! Navigation • Odors: Chemical odors can be detected by some. (sensitive sailors could “smell” land). • Twilight cues: Use setting sun to determine west. But also use polarized light that runs NS to determine direction. Zugunruhe • The onset of migratory activity is characterized by a restlessness commonly referred to as Zugunruhe • Hormonal in nature as we saw and does seem to be genetically controlled. • Demonstrated via selective breeding experiments • Could alter when this occurred. Summary • Many environment is changeable over most of the planet • Many species respond by moving from one area to another and back again, Migratory behavior • Energy intensive but benefits can be substantial. • Report due Monday 27 • Email to [email protected] • You can email extra credit to the same address • Name format: A Student EFB480 profile (extra credit) • Indicate notification of opening email here Summary • For birds, benefits seem to be related to accessing a seasonally abundant food supply, especially for reproduction • Hormonal control under the influence of day length. • Migration involves navigational behavior • All forms used: visual to magnetic fields. • Constitutes a significant part of their lives. Dispersal behavior • • • • Other form of movement. Usually unidirectional Usually young but old may exhibit it also. So usually from natal area to new home range or territory • Most often males Dispersal behavior • Reasons? • Resource based: greener pastures! From area of high density to lower. • Sexually based. • Some feel it is an innate aversion to inbreeding (young males avoiding mating with their mothers or sisters) • However often find males mating with their daughters Inbreeding avoidance? • Genetically equal so hard to imagine a young male resisting inbreeding but then engaging in it as an adult. • Rare that young male will have more will power than older male! • More of how we WANT to view the system than how it really is! Conflict avoidance • Another proposed hypothesis for male dominated dispersal is that young males are forced out by their fathers or other males. • Under this hypothesis would predict high amounts of male-male conflicts, especially father-son • This type of aggression is common, sexual competition is the norm among males Mechanism • But why is it the young males that disperse? • Under the conflict hypothesis, the young males are….younger! • No match with father, unless father really old! • Even if a match for father, other males coming into area are older, better suited to defeat father and son! Resulting dispersal patterns • Thus young male leaves and continues to move as long as he gets beat up! • Either by territorial holders or by other dispersers over disputed vacant territories. • Until… enough experience/strength, becomes the winner. Dispersal distances in cougar males 0.5 Idaho Data New Mexico data (approximate) Frequency 0.4 0.3 0.2 0.1 0.0 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Home range Diameters >22 Summary: Dispersal • • • • • Many young leave natal home range Predominance of males. For many it is resource based For many others it is for reproductive reasons Most likely driven from home range area by fathers or larger conspecifics. • Is a mechanism for colonizing new areas. Behavioral Conservation • Last topic to cover is the use of behavioral ecology in conservation biology • How do we use what we know about the behavior of animals in efforts to conserve their populations. • Can be an extensive consideration • Here only touch on some aspects to give an appreciation as to how important behavior is in conservation efforts. Significance of behavior to conservation • Conservation biology is relatively recent • Though conservation was advocated by early naturalists, e.g. Muir, Thoreau, etc. • American opted for management rather than conservation. • Management: “wise” use of selective species • Still lead to abuses and declines of undesirable species Birth of conservation biology • Began to realize with management we were saving the desired species but losing the structure and function of the systems that supported them! • Conservation biology began in the 1980’s, Soulé a main figure. • Argued we needed to conserve all species of the systems. Focus of con bio • Multi-disciplinary: ecology, population biology, population genetics, systematics, etc. • Develop conservation models, plans, actions to conserve species, which then conserves main focus; biodiversity • PVAs, MVPs, metapopulations: focus on main reason for biodiversity loss, fragmentation. Actions • Dealing with small populations, trying to recuperate them. (conservation biology is the biology of small populations; Ne becomes important) • Land protection: Parks, reserves, etc. • Reintroductions • Captive breeding • Corridors • Etc. Where’s the behavior? • Unfortunately, in things such as PVAs, MVPs, metapopulation dynamics, and corridors, behavior is often left out. • Why and how is behavior important? • Look at several behavioral categories we have covered and see their link to conservation biology Small populations • In all cases, dealing with small populations • Individual behavior and behaviors among individuals then become important What your trying to do is increase flow of energy through the species. Two levels: individual; you want the individual to get enough energy to survive Population: you want the individual to reproduce: successfully! Individual energy gains • Foraging behavior becomes important • Area has to provide adequate answers to the four questions: what, where, how long, and where to next Diet choice • Saw earlier diet choice is a balance between gains and costs • Obviously range of what a species eats is important • Generalists vs specialists: Behavioral choices based on gains and costs • Usually in conservation biology dealing with specialists: animals with narrow diet range In either case • Knowledge of diet AND whether area supplies these food items becomes important and is direct result of diet choice. • Diet switching behavior also important: What are the “rules” the species of concern follows? • Type II curve, what is its shape, when will it switch to second choice? • Remember it is the abundance of the preferred food item that determines when to switch. Significance of this? • 1) abundance of primary food will determine abundance of species. • Why? Lower food items are lower because they provide less gain at greater cost. • IF secondary food is abundant and primary food low, will not have as great of a potential in increasing the population Patch selection/residence • Now know that distribution of food as important as abundance! • What is patch structure of area? • Does it provide adequate choices? • What are relative patch qualities? • Juxtaposition becomes important • Travel times • Not as simple as providing “adequate” food resources. Intra specific behaviors • Individuals need to interact and how they do this can impact success of conservation program. • Need to know something about competitive behavior between males. • For mates: outcomes of male-male competition has impacts on who will breed Male-male competition • Who will breed affects: Ne , genetic composition of population (Possible loss of genetic material could be high) • Example: Bison in the Badlands National Park • Aggression determines which males breed • Lineage of more timid males, less able to defend females is declining • Knowing this, adjustments can be made (e.g. separation of two groups) Other examples • Infanticide by males: killing offspring sired by other males: can affect reproductive output • Hybridization between two similar species: • Just a few factors about male competition that would need to be considered Female choice • Now know female choices and is an attempt to increase fitness of offspring • In small populations, there is less to choose from! • This is especially true in captive populations where we often do the choosing for females! • Many time why captive breeding can fail or is less of a success as we hope! Symmetry • Saw earlier that morphological symmetry plays a role in mate choice. • Environmental disturbances could affect this symmetry and affect female choice • Leg bands! In captive breeding, females may prefer not to mate with a male that has one but no the other! Courtship • As with food resources, areas for courtship rituals to play out have to be considered • Very species specific behaviors! • May have adequate food but if there are no lek areas, will not have the species there. here Mating systems • Often small populations can have skewed sex ratios • Ne will be affected depending on mating system: monogamous; Ne will be set by lowest number sex • Polygynous: Lek system may not be sustained if there are not enough males OR females! Parental care • Again, in species where both parents provide care, a female may mate (extra pair mating with male of other female), but chances of survival of offspring low. • Lions: one male may do the most mating but other males (subordinate siblings) needed to protect from other males; not enough protective males, system breaks down. Sex allocation • • • • Many species have biased birth sex ratios This has to be taken into consideration Example: Wild ass: Middle aged females give birth primarily to males, younger and older to females. • Need females to increase the population • Released primarily middle aged females! • Population grew very slowly until middle aged became older and started producing females! Dispersal behavior • Obviously you want population to grow and increase in range • This requires dispersal of young, male and female! • In many species males primary dispersers • So may need to plan on helping females! Dispersal behavior • Many species have distinct requirements for dispersal habitat • Dispersal routes become important • Bighorn sheep, reintroductions can fail because don’t know how to get off the mountain in the fall! Dispersal distances • Again, some species will more readily disperse over longer distances than others. • This has to be taken into consideration Inbreeding avoidance • Dispersal becomes very important in small populations where you want new individuals to come in to reduce effects of inbreeding. • DON’T assume there is an innate avoidance of inbreeding! Interspecific considerations • Predator-prey relations • Antipredatory behavior • The landscape of fear/opportunity Landscape of fear and conservation • More important than resources, risk involved in getting these resources • How can landscape of fear affect conservation efforts? • Mentioned before, landscape will consist of differing percents of risky/safe habitats. • If area is primarily risky, reintroductions will fail What do we need? • Need that balance between risky and safe that insures species has enough safe areas to persist. • Predator? Often conservation efforts are for predators • Landscape of opportunity. How do you modify? • Behavior specific! • Need to identify risky/successful vs safe/unsuccessful habitat and then plan habitat modifications that will tip the balance the way you want. • Example: desert bighorn and shade. Ecosystem conservation • Newer emphasis in conservation biology • Realize can’t conserve species if we don’t conserve ecosystems that support them • Two things: structure: obvious, if we don’t have the habitat, we don’t have the species • Function: not as obvious, need to conserve the function of ecosystems: energy flow Ecosystem function • How do we maintain energy flow? • Depends on behaviors of individuals! • One of the most important is the predatorprey interaction. • Reason predators are keystone species. • The prey killed by predators is the “cost of operation” of the ecosystem • This is lethal effects Non-lethal effects • Beyond killing prey, predators have an overwhelming non-lethal influence • Landscape of fear again • Predators “keep everyone in their place”! • Cascading effects Predation and Conservation • The role of predatory behavior and conservation of ecosystem processes • Saw fear affects behavior and habitat use. Elk Level of vigilance of females with calves before and after The reintroduction of wolves in Yellowstone. 70 Females with calves in area with wolves 60 % Vigilance Females with calves in areas originally without wolves 50 40 30 20 10 0 1 2 3 4 Number of years after wolf reintroduction 5 6 3 Area with wolves Y = 1.9 - 0.002 X Area without wolves a 2 Number of fecal groups/10m2 r = 0.66, P = 0.001 2 1 0 NS 0 100 200 300 400 Distance from forest edge Elk: Habitat use patterns 500 600 Deer and Pumas. GUDs for deer in two types of forest. 1.5 1.4 1.3 1.2 1.1 Open areas Forest edges Forest areas 1.0 GUD (kg) 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 Douglas Fir Mountain Mahogany Habitat type Fecal groups/10 m2 6 Open Edge Forest Average # fecal groups/ 10 m2 5 4 3 2 1 0 Year 1 Year 2 Year 1 Year 2 Juniper Fir Deer: Habitat use. Year 1 Year 2 Mahogany Year 1 Year 2 Aspen Produces landscapes of fear • Where animals are afraid to go and if they do, are less efficient in foraging. October 2008 VAN DER MERWE AND BROWN—MAPPING THE LANDSCAPE OF FEAR 1167 FIG. 3.—Open Colony study site. A) Landscape of fear superimposed on map of feeding stations and burrow entrances. Contour lines represent areas of similar quitting harvest rates. Units are in joules per minute. Differences between contour lines indicate differences in predation cost of foraging. B) Landscape of fear superimposed on map of tree positions. The species code in the key is: A.m. for Acacia mellifera. Contour lines represent areas of similar quitting harvest rates. Units are in joules per minute. Differences between contour lines indicate differences in predation cost of foraging. Effect on vegetation? • Herbivores are plant predators • The less predation, the more survival, the higher the population density Ripple and Beschta. 2004. BioScience. 54: 755-766 What next? • Cascading effects • More plants, more production for other species. • Other species provide resources for still more. • Etc. etc. Best example • Yellowstone National Park More? • Coyote example: • Mesopredators return. What next? • Enough! • Still a lot more on how behavior effects conservation efforts, it is the key! The end • Hope you have enjoyed the course • Hope you learned something! • At the very least, that behavior IS important • Final: May 5; 2:45-4:45, 146 Baker