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Chapter 6 1. Habitat selection - selection of a particular area in which to live 2. Territoriality - defending your habitat 3. Dispersal, migration and homing - when suitable habitat is not available nearby 4. Nest Construction – making own habitat 1. Observation: if you go out in nature, one finds specific animals usually occur only in specific locations. If you compare species of animals in an open field to those found in forests, you would find most are different. 2. If you allow animals to make choices about the type of area they live in, one finds that even single celled animals are discriminating choosing to live in particular locations and not others European songbirds: coal titmice - live in pinewoods (pine trees) blue titmice - oak woodlands She hand reared baby coal tits and blue tits until they could fly And placed young birds in a cage with a "composite" branch oak and pine portions of same branch % Time 100 50 Blue Wild Coal Blue Coal Hand-Reared 1. an animal may be best adapted (anatomically or physiologically) to feed or reproduce in a particular area - selection favors animals that also prefer to stay in these areas factors restricting habitat use: a. climate, physical +/or chemical factors, competition, predation, location of food, etc. Ex. Polistes nests - picked off out in open Ex. Competitive exclusion: barnacles Chthamalus stallanus - high on rocky shores Balanus balanoides - lower on rocky shores Remove Balanus, Chthamalus moves in. Balanus smothers and crushes Chthamalus (Connell 1961) Cottonwood aphids - Utah Tom Whitham In spring, eggs (laid previous winter) hatch and from them females about 0.5mm long emerge Females walk along trunk and branches of tree to a location where a batch of leaves is just starting to form. Female chooses a leaf and induces the formation of a hollow ball of plant tissue (gall) to form along the midrib. Aphid lives in gall - first batch of eggs are female. Larger the leaf - the more female offspring produced and the greater their average weight One would expect females to select only large leaves when given a choice (assuming that their behavior has been shaped by natural selection) Is this what they do? If so, what happens if there is competition for the good leaves? 1. Average Density = 35 aphids / 100 leaves - about 2% leaves = large, 33% = small, the rest (65%) were in between - none of the small leaves were utilized Conclusion: -Aphids use large leaves disproportionately more than small. They are able to tell which leaves are the best and choose those that will be the most productive and support the most offspring. -But what happens if a large leaf is not available? 2. some will double up or triple up on the same leaf; Late comers end up farther out on the leaf and therefore don't get as much of the flow as the first female No. of Galls per Leaf Mean Leaf Size (cm) Mean Number of Progeny Produced Basal Female Third Female Second Female 1 10.2 80 2 12.3 95 74 3 14.6 138 75 29 ½ workers & old queen swarm Scout bees search for new home ◦ Each dances to direct others to potential site One site chosen Everyone leaves together & flies to the new nest site Preferences: ◦ ◦ ◦ ◦ Small entrance – 12 cm2 Entrance – south facing Ht of nest – 5 m min Cavity volume – 40 liters Scouts inspect entrance, walk around interior ◦ Distance walked relative to the cavity entrance was experimentally shown to be basis of bee’s estimate of cavity volume Tom Seeley set up box choices on treeless island Transported swarm and studied decision process ◦ Scouts recruit only for a short time – so voting for a site can fade away if new recruiting scouts aren’t picked up ◦ Best site is not always picked Neutral bees become supporters for a site in proportion to the number of bees dancing for that site ◦ Scouts do not stop dancing for their site after visiting another site ◦ They just stop recruiting after a while & become quiet ◦ This loss of motivation allows a less supported site to fade out of the “debate” Regular - even spacing of individuals in an area (territorial) Random - rare Clumped - social groups, family groups, mating clusters, etc. An area or space, which is defended through aggression or less overt behavior against other organisms. 1. What is defended? May be a piece of land (Cardinals, Red Winged Blackbirds), a resource (Acorn Woodpeckers), or simply space around animal of same species (stallion / herd) - or more unusual space around another species: ex. Bitterling (small fish) territory is centered around moving clam 2. Defense may be through active fighting or aggressive displays – antlered flies or by means of chemical markers which are not as obvious - dogs, cats 3. Territory may exclude animals of same species or other species, including plants. ex. Acacia ants - live in thorns, they defend their homes and the nectaries the plants provide from other ants, grazing animals and also from competing plants in the area Using Optimality Theory – should occur when benefits outweigh costs The cost of defending the area: if around a resource ◦ How many competitors? ◦ How dispersed is the resource? ◦ How predictable is resource? The benefits of defending the area ◦ What is gain in resources? ◦ Does it help in attracting a mate and improve reproductive success? 1. When there is a resource worth defending: ◦ ◦ ◦ ◦ Predictable in time and space Defendable given the number of competitors. Concentrated or rich resource Benefit is greater than cost of defending 2. Territory may be required for reproduction (high reproductive benefit) ◦ wide range of species - typically males set up small display territory (known as a lek) where they can display themselves, females choose among them Some defend permanent territories where food is predictable and abundant. They defend both sides of a river and collect food washing up on the bank. - If area is very rich, a partner or young may be allowed to share, but chased off if resources dwindle. - Interestingly, some of same species feed on transient, patchy food supplies - when this is the case, they feed in flocks ex. Chipmunks - J. Brown - Nevada 2 sp.: Eutamias dorsalis - open terrain, territorial E. umbrinus - forest, no territories - Where they occur alone, there is no difference in habitat elevation. Where they overlap, E. umbrinus is at higher elevations. There appears to be competitive exclusion. Experiment: placed peanuts out in areas of near overlap In open terrain - E. d. does better - aggressively defends food dish from others, can see them approach from far away In forest – E.u. does better. Aggression spent in vain, while chasing off one, another dashes in and feeds, with dense brush, several can get very close to dish a. wide range of species do this - typically males set up small territory (known as a lek) where they can display themselves, females choose among them Lekking species include: birds, frogs, antelopes, bats, dragonflies, and many species of flies. ex. sage grouse - males stand out in area, individual stamping ground, puff feathers, and call or stamp feet - display areas are often traditional - typically in a lek central males mate with the most females (80% by 3 central males in group of 10?) - female preference, forces males to compete for these locations and thus the females automatically get the strongest male. a. Territories exclude others. - If resources or nesting areas and /or mates can be monopolized by one or a few individuals, they will survive and reproduce more successfully than those without territories Why do excluded individuals tolerate exclusion? - If exclusion means loss of mates or food, why are most territorial disputes relatively harmless. - most exclusions are temporary - one breeding season, one area of food abundance. In insects with just one reproductive season, - small males may use alternative strategies. - territorial holders have alot at stake in dispute will tend to fight much harder than newcomer - newcomer will do better if as soon as it is fairly clear that s/he might get hurt in the fight, s/he runs away. If s/he stays and fights, s/he risks injury and potentially total loss of reproductive success. - what one finds is that usually surplus males and females = young individuals - waiting their turn tits, sparrows, impala, elephant seals, others Dispersal: Often young disperse ◦ Often males (Belding ground squirrels), may be female (Florida scrub jay), maybe both Costs may be high – energy, exposure to predators, finding resources in a new place Benefits are high ◦ Avoid inbreeding depression (lowered fitness from mating with closely related individuals) ◦ Avoid competition with parents ◦ Avoid aggression from older same sex individuals Ex. Migratory locust (Schistocerca gregaria) of N. Africa and Middle East Two races: 1. normally live in a solitary fashion, cryptically colored 2. swarm forming, long-distance movement, brightly colored, eats anything High density or If given diet of other locust tend to get 2cd race At intervals of several years, fluctuations in weather lead to a reduction in the size of the population in favorable areas. When such a reduction follows a period of population buildup, the locusts are concentrated and the gregarious, dispersing race is produced. Once dispersal is in full swing - swarm may be as many as 10,000 million individuals, travel a total of 2000 mi (tend to move downwind). - to sustain flight locusts may consume their body weight in plant material per day. Large swarms may weigh up to 50,000 tons or more. Latin equivalent of locust means "a burned place" a. Why? 1. Avoid bad climate during winter or dry season 2. To maintain a source of food - decrease due to onset of winter or dry season - buildup of population density = dispersal _ egg laying site is different from foraging area a. Why not just stay in tropical zone (warm and food avail)? 1. Temperate zone - spring and summer - large increase in plant and insect pop growth = excellent resource if taken advantage of. Same can be said of an area that has wet and dry seasons 2. Stiff competition in wet tropics, great species diversity 3. Compromise: overwinter in acceptable area, breed in rich area with lots of space for breeding 1. Arctic tern - 25,000 mi round trip each year (longest known) - flies 8 mon/yr a. Breed in Arctic summer (long days, lots of food), overwinter on islands in the Antarctic Sea 2. Many mammals: wildebeest, caribou, seals, whales - same reasons Ex. Buffalo = migrated between Canada and Mexico in large herds - between 1870-1875 12.5 million were shot (tongue = delicacy, carcasses were left while indians starved) - by 1889 pop. = 1091 today = 30,000 Insects - Monarch butterflies - Northern breeding areas, southern overwintering area - overwinter in very specific small areas - coast of California, LA to Monterey, and just north of Mexico city at an elevation of 9000 ft. a. None - travel with prevailing winds ex. Migratory locust b. Topographic features - visual and chemical cues Ex. Salmon - born in mountain streams, swim to ocean, live several years - return to natal stream to spawn and die - learn smell of river as fry and learn olfactory characteristics of currents near river mouth Ex. Digger wasp (Tinbergen) 1. Orientation alone - no biological clock involved Ex. northern butterflies (Painted Ladies) fly at an angle to sun only during limited hours of the day even so path looks like: - sun as a compass heading in conjunction with clock - angle travelled is with respect to the sun, but that angle is corrected for the movement of the sun Ex. pigeons - clock allows the birds to fly in a straight line Ex. Honeybees – communicate location of food with respect to the location of the sun Ex. indigo buntings (Emlen) - learn star pattern - orient with respect to the star around which others rotate: northern star Ex. birds and bees - magnets attached to (pigeon) heads alters their orientation. Ex. Heart and Dart Moth (Agrotis exclamationis) (Robin Baker,1987) moves across country - Moth used the geomagnetic field to calibrate a moon compass. Directional preference changed such that it tracked the change in the moon’s azimuth. 1) Orients to geomagnetic field; 2) Sees the direction of the moon; 3) Rest of night flies according to the moon. a. some bird nests are primitive "structures" - slight depression in dirt b. Some termite nests are so elaborate, no one fully understands how the builders can coordinate their building efforts to construct it. Ex. Macrotermes bellicosus Compass termite nests get up to 5m high and 3m long -broad sides face east and west -narrow surface = north and south, so get little exposure at mid day -wide sides = maximum surface for sun warming early and late in the day -how are nests oriented? they are sensitive to earth's magnetic field, Example of behavioral radiation and the effects of predation on shaping that behavior (Jeanne 1975) - much variability a. two types - open and enclosed b. major predators = ants, which once they find brood can recruit many helpers and decimate a wasp nest c. ant guard Stimergy – inherited building program Learning - Weaverbird nests