Ecosystems 1) Feedback system involving interrelated parts which function to maintain themselves in a steady state, or equilibrium a.

A portion of the output of the system is used as the input of the system b. Control of temperature in warm-blooded animals sensory system is the skin, hippocampus is triggered, and panting or sweating works to decrease body temperature through release of body heat c. Requirements of all systems i. All systems must have a source of energy (in many ecosystems, this is the sun) ii. Must have a means of transforming this into usable forms, loss of energy at each level energy transformation iii. Must have a built-in ability to vary with conditions in order to return to equilibrium (allows for reactions to perturbations to the system, can react to different levels of stress), some trial and error as it returns to steady state, dynamic equilibrium iv. Inherent limits, exceeding these limits will lead to a break-down in the system d. Ecosystem is an open system composed of many parts working cooperatively, and a steady state will be conserved if the parts continue to work in harmony despite perturbation i. Never have perfect equilibrium, always responding to perturbations, always attempting to return to equilibrium, otherwise the system will break down ii. Stability is the end product of millennia of parts competing against one another to reach equilibrium e. Ecosystem is a model of a self-regulating system 2) Components of an Ecosystem a. Biotic - living b. Abiotic non-living (solar energy, oxygen, carbon, hydrogen, nitrogen, phosphorus, and potassium), energy is passed through the ecosystem through the trophic system i. Autotrophic plants (photosynthesizers), get energy directly from the sun ii. Heterotrophic animals (primary consumers, secondary consumers, scavengers, parasites, decomposers), loss of energy at each level, chemical are recycled by the system 1. Primary consumers expend a great deal less energy to obtain nutrition than secondary consumers (Eltonian Pyramid illustrates

decreasing total biomass, and usually density, at each successive level) a. Only 3% of the total energy available from the sun is incorporated into the system, only 10% of the remaining energy is captured by the next level, if any level became 100% efficient, the level below would be eliminated, and the top level would follow b. 14 wolves studied, ate 70 herbivores in one year, each wolf weighed approximately 70 lbs, each herbivore that was eaten was approximately 1000 lbs, they ate 6,000,000 lbs of plants c. A limit being placed on any one level will lead to limitations of subsequently higher levels c. Food Chain the sequence of events by which energy moves up the tropic level, from plants to herbivores to carnivores to top predator d. Food Web much more complex system of interactions wherein a single element may play a role at many tropic levels Trophic Interactions 1) Biomes ecosystems can be split into six primary biomes (desert, grassland, tropical forest, deciduous forest, taiga, tundra) a. The first three biomes are found primarily within 20 degrees of the equator, high annual temperature, daily temperature variation is greater than seasonal variation, amount of rainfall differs b. Desert succulents and cacti, bloom during a short period right after rainfall which is infrequent, tends to occur in bursts, most plants and animals are small, mammals and birds are rare (difficult to maintain constant body temperature), difficult to maintain water, non-human primates do exist on the fringes (baboons, patas monkeys, some lemurs), no apes nor New World monkeys c. Grassland exists nears the tropics as well as outside, intermittent and erratic rainfall, 10-40 inches per year, more species in terrestrial habitats than in any other biome, many primates are specially adapted (gelada baboon, patas monkeys, ring-tailed lemurs, macaques, gorillas and chimpanzees), but no New World monkeys d. Tropical Forest torrential rainfall, may have very short dry season, growth is year-round (perennial), most diverse

habitat for plants, rarely have a single dominant species, Central Africa, South and Southeast Asia, Central and South America (Amazon), being reduced rapidly, usually continuous canopy, may have multiple layers, usually cuts off rain and sunlight from ground level, high shade tolerance in many plants, many microhabitats for animals, specific adaptations, most primates evolved in tropical rainforests e. Deciduous Forest most abundant found outside of the tropics, cold winters and warm summers, seasonal temperatures vary more than seasonal, 30-40 inches of rain, seasonal, growing season is discontinuous, trees are farther apart, fewer species represented, many primate species, cold deciduous forests of China and Japan are the furthest from the equators (macaques and langurs) f. Taiga and Tundra found at the far northern latitudes, no primates occur in these biomes 2) Given certain characteristics and environmental conditions, we can predict which of these habitat types will dominate a. Abiotic components are fairly constant in any area b. In similar habitats, structure and physiology of plants and animals may be similar, but taxonomically separate c. Climax community is achieved through the process of succession i. Ordinary change from one community to the next until it reaches a predictable equilibrium point, distinct stages 1. Grassland to shrubs to pine forest to hardwood forest (maples, birch, and hemlock) 2. Animal communities will change in conjunction with the plant community succession 3. Natural succession is mimicked by human induced changes, some species are specially adapted to this first stage of succession ii. Occurs because each species alters the environment in such a way that it will no longer be able to succeed as well as another species, more shade-tolerance in subsequent seedlings 1. Climax community replaces individuals of the same species, death rate equals germination success rate, equilibrium is reached at this level 2. Disruption will lead to a reversion back to early stages iii. Man is keeping much of the earths surface in a state of disclimax, where there should be grassland, we have desert, and where we should have forest, we have grassland by over-utilizing resources and

altering chemical concentrations 3) Leibigs Law of the Minimum the material (chemical, water, heat, space) which is most limited in supply will be the factor limiting a species population size and distribution, an organism is no stronger than the weakest like in the ecological chain of materials a. Carrying capacity of an environment depends on these limitations 4) Shelfords Law of Tolerance not only a minimum of some factor can limit a population, but also a maximum of a factor (heat, light, water) a. Stenothermal a population with a narrow tolerance for temperature, more specialized, may have greater efficiency at expense of adaptability, do not thrive is variable environments b. Eurythermal a population with a wide tolerance for temperature, more generalized 5) Interactions at different trophic levels a. Neutralism two species do not effect each other (0,0) b. Competition each population is adversely effected (-,-) c. Mutualism the growth and survival are dependent on the other (+,+) d. Protocooperation both populations benefit, but relationship is not obligatory (+,+) e. Commensalism one species benefits while the other is unaffected f. Ammensalism one species in inhibited while another is not helped directly (0,-) g. Parasitism one species benefits by directly hurting another (+,-) h. Predation one species feeds directly on another species (+,-) Interactions Among Animals 1) Different Trophic Levels (Predator-Prey) a. In a stable ecosystem, predator and prey populations are in equilibrium, if predator overkills or if prey gets too good at escaping predation, could lead to the extinction of both b. Usually not the limiting factor for prey populations, normally limited by food resources, though predators may be limited by prey i. Lemming migration diminishes through suicide, the predator population fluctuates in response c. Prey develop methods of escaping predators, prevents predators from overeating, coevolution may occur as mutual adaptations aid both the predator and the prey in their arms race

i. Detection prey must know that a predator is there, once they are detected, predator may move on to new prey, energetically most efficient for both predator and prey ii. Escape didnt detect them in time, means of escape, energetic loss for both predator and prey iii. Defense after pursuit d. Three operational mechanisms for reducing predation i. Exploiting the limits of a predators motor or sensory capacities 1. Dispersal outrun, outfly, or outmaneuver, active a. Patas monkey is fastest primate, male is always on the lookout, when predator is spotted, performs ritualized display to attract attention away from the females and young, then outrun 2. Camouflage blend into surroundings, passive a. Patas females and infants blend into the tall grasses of their environment ii. Exploiting the limits of a predators learning abilities 1. Imitate species that a predator is disinclined to eat (mimicry), leaves or toxic animals 2. Intimidate using large canines or other defense mechanism a. Primates with large canines will yawn upon seeing a predator to scare them off 3. Mobbing by a large group of small animals a. Lemurs, Macaque iii. Direct protection against predators 1. Passive poison or toxins a. Lorises smell bad because they are toxic 2. Active spines, grouping behavior, vertebral spines a. Prey species would defend themselves against predators that were smaller than three times their own mass b. All diurnal primates are group-living species, apparently in response to predators 3. Combination a. Small nocturnal prosimians tend to be solitary, cryptic, quiet b. Lorises develop camouflage, scapular shield because of their slow moving lifestyle

2) Same Trophic Level a. Competition i. Gauses Law of the Struggle for Existence, placed two species of paramecium (P. aurelia and P. caudatum) in a petri dish with fixed food resources, one species grew at the expense of the other, no direct attack, one was a better competitor (utilized resources and reproduced most quickly) 1. No two species can coexist in the same environment if they have a resource in common that is in limited supply 2. Misinterpretation: no two closely related species can coexist in the same niche ii. When a different species (P. bursaria) was added to P. caudatum, the two species reached an equilibrium, occupied different parts of the environment, not sharing the same limiting factor 1. Habitat selection one species lived on the flat surface while the other lived on the rim 2. Niche Definitions a. Everything an animal does (functional or behavioral) b. Where an animal does the things they do c. Quantitative value of the measurements of the resources available versus the resources that a particular species uses, percentage of time spent in a given part of the habitat, overlap at each of these parameters, unused niches i. Among 8 South American primate species, dietary overlap ranged from 24 to 97%, cant use limited resources, but can use other resources ii. Populations must shift (within environment or in size) if limiting resource is being competed for iii. Surviving in the same ecosystem 1. When resources are not limited, more resources than animals needed, manage their populations through epideictic factors such as fecundity, balance with the environment (more offspring in good environment) 2. Severe climate keeps population densities low, widely dispersed so that individual will have enough resources

3. Differential habitat use, forest strata, adapted to different resources iv. Character Displacement determine if characteristics developed because of interaction or were they preadapted b. Coevolution evolution of two or more taxa that have a close ecological relationship in which reciprocal selective pressures operate to make the evolution of each taxon partially dependent upon that of the other i. Plants and animals coevolve in a similar manner as predator-prey relationships, plants may protect themselves by thorns, corky bark, hard seed coats, they may also evolve behaviorally, different flowering times, poisonous secondary compounds 1. Arms race between the coevolving species through detoxification of secondary compounds, other alterations to the digestive tract to enable use of these resources, exploit these protective mechanisms 2. Plants use animals to pollinate other members of the species, this also leads to coevolution, most of these animals can fly, looser relationships exist with seed dispersers to move seeds outside the seed shadow a. Birds, bats, and primates are the primary seed dispersers in rainforest environments 3. Diffuse coevolution no a one-to-one relationship between plant and animal, but occurs between a community of plant and animal species Primate Taxonomy 1) Taxonomy classification based on the Linnean Hierarchy a. Kingdom (Animalia), Phylum (Chordata), Class (Mammalia), Order (Primates), Family (idae), Subfamily (inae), Genus, Species b. The natural scale is an inaccurate representation of the evolutionary process, prosimians and humans have been evolving for the same amount of time c. Tree shrews are good representations of the animals that eventually evolved into the modern primates d. Prosimii i. Lemuriformes Madagascar, these may have existed elsewhere, but were outcompeted by primates in other parts of the world 1. Lemuridae - diurnal, monkey-like, most are

primarily arboreal, but ring-tailed lemurs are terrestrial adapted, walk bipedally 2. Lepilemuridae nocturnal, folivores 3. Cheirogaliidae most similar to primitive primates, nocturnal frugivores 4. Indriidae vertical clingers and leapers, folivorous, two diurnal, one nocturnal species 5. Daubentoniidae hard fruit and insects in bark, previously thought to be a rat (Aye-aye) ii. Lorisiformes 1. Lorisidae solitary but social, nest and interact, except when hunting a. Lorisinae slow-moving, look like nocturnal lemurs, frugivorous, no tail, cryptic, Africa and Asia b. Galaginae (galagos) nocturnal, fastmoving, fruits and insects, long tails and large ears, Africa iii. Tarsiiformes only found in Asia, only primate 100% faunivorous, nocturnal iv. Platyrrini (New World Monkeys) only primates with prehensile tails, though only some have it, flat-nosed 1. Callitrichidae a. Callitrichidae tamarins and marmosets, small, readapted claws instead of nails (17-21 species), gum-feeding, may cause injuries themselves, marmosets are more adapted to gnawing into trees, communal breeders, all members care for young 2. Cebidae a. Cebinae monkey-like, capuchin, squirrel monkeys, omnivore, lives in large groups, diurnal, some have prehensile tails b. Aotinae Aodis (only nocturnal Anthropoid) 3. Atelidae a. Atelinae spider monkeys, long arms and legs, prehensile tails, suspensory locomotion, frugovores, cannot walk quadrupedally well b. Alouattinae howler monkeys, foliverous, slower, prehensile tails, but not suspensory, quadrupedal, spend a lot of time resting c. Pitheciinae sake and wooly monkeys, seed-predators, some have dichromatism

like prosimians v. Catarrhini not flat-nosed 1. Cercopithecoidae (Old World Monkeys) a. Cercopithecidae live in large groups i. Cercopithicinae grass-adapted, baboons (savannas), mandrils (forest), cercopithecines (arboreal), found primarily in Africa, except for macaques ii. Colobinae primarily in Asia, except for colobus, folivorous vi. Hoinoidea 1. Hylobatidae (Lesser Apes) gibbons and siamangs, no prehensile tails, hooked hands without prominent thumbs, move extremely quickly, frugivorous, walk bipedally arboreally and terrestrially 2. Pongidae only in Southeast Asia a. Ponginae (orangutan) - arboreal b. Homininae birth interval due to long period of gestation and care, no tails i. Gorilla largest living primate, adapted for upright posture, but center of gravity is so high that they knuckle walk on the ground, leather-like pads on hands ii. Pan (chimpanzees and bonobos) structured like gorillas, but smaller iii. Homo 2) What is a Primate? a. Other orders within Mammalia have developed special adaptations as they moved away from their primitive mammalian ancestor b. Morphology i. Pentadactyly five independently mobile fingers and toes ii. Flattened nails and sensitive pads on hands and feet iii. Retained generalized primitive mammalian bone structure (collarbone, radius and ulna) iv. Retained generalized dental structure (4 types), may have lost canines or molars depending on their diets 1. Incisors have uses in biting into fruits 2. Canines are used for ripping and cutting into fruits and meats, showing in display 3. Premolars rip/cut and chew, may shift towards one or the other depending on diet

v.

vi.

vii. viii.

ix. x.

4. Primitive mammals have tritubercular molars with three cusps, in most primates, a quadritubercular system dominates, apes and humans exhibit a Y5 pattern (more complex tooth, more plant material) 5. Primitive mammals will have a 3:1:4:3 dental structure with a total of 44 teeth, many New World Monkeys have 2:1:3:3 (36) pattern, Old World Monkeys, Apes, and humans have 2:1:2:3 (32), some lemurs have variations on these patterns a. Indri lemur has 2:1:2:3 on top and 2:0:2:3 on bottom, Aye-Aye has 1:0:1:3 on top and 1:0:0:3 on bottom for gnawing and chewing Visual powers elaborated (better than many other mammals) 1. Frontation movement of the eyes to the front of the face, rather than sideways 2. Convergence moving towards each other, enables over-lapping vision, towards stereoscopic vision 3. Hemi-decussation rather than each eye going to the opposite side of the brain, the left side of both eyes goes to the right side of the brain and the right side of both eyes goes to the left side of the brain Reduction of smell 1. Not necessarily the case for all primates, lemurs combine olfactory and visual displays, as have many other prosimians in New and Old World monkeys, the increase in eye size has led to a reduction of the olfactory system (cattirhine) Progressive development of the brain 1. Relative to body size, primates have some of the largest brains, though not the largest Infant dependency last for longer 1. Long period over which an infant develops into adult, during which it stays closely tied to its parent(s) Trend towards sociality 1. Nearly all diurnal, and some nocturnal species are very social animals R.D. Martin said that all of these traits are descriptive of primates, but not necessarily unique to them (may apply to other mammals)

c. Unique features of Primates i. Arboreal inhabitants of tropical and sub-tropical forests ii. Extremities are adapted for prehension and arboreal support 1. Grasping, nails and tactile pads with enhanced sensitivity iii. Hind-limb domination push off with their hind-limbs rather than pulling with their fore-limbs, lateral footfall pattern (monkey gait, left foot to right hand, right foot to left hand, diagonal sequence) iv. Tarsi-fulcrumation elongated heel bone allows for primates to leap, different balance v. Large eyes in relation to skull size, post-orbital bar (eyeball completely enclosed by bone), frontation and convergence leading to overlapping vision vi. Petrosal bulla the bulla is the bone that covers inner ear, only in primates (and plesiadopoids) use the petrosal as this bone vii. Specialized sulcal pattern (troughs in the grey matters of the brain) viii. Long gestation period (slow fetal and pre-natal growth), late maturity, and long life-span relative to mothers body size ix. Small litters of precocial (not able to live independently of parents) infants x. Dentition maximum of two incisors, molars and premolars are relatively unspecialized, cusps on molars, lower molars have a grinding basin (talonid) xi. Gregarious and social 3) Evolution of Primates a. Reptiles (300 MYA) Mammals (200 MYA) Pre-Archontan (insectivorous, tree shrew like) Mammals (120 MYA) Archonta (Plesiadaformes) (60-65 MYA) Euprimates (35-60 MYA) b. Archonta contains the orders Dermatptera (flying lemurs), Chrioptera (bats), Scandentia (tree shrews), plesiadaformes, euprimates c. What major adaptive shift led to this set of adaptations? Why did primates of modern aspect evolve? d. Currently 4 theories concerning primate origins i. Classic Arboreal Theory (Grafton Elliot Smith, Wood Jones, W.E. Le Gros Clark) 1. Shift from ground insectivore to trees became a successful lineage by depending more on vision and touch rather than smell

2. Led to hand-eye coordination, larger brains 3. To acquire an arboreal way of life demands prehensile functions of the limbs, a high degree of visual acuity, and the accurate control and coordination of muscular activity by a welldeveloped brain ii. Plesiadapoids were the first primates (Fred Szalay) 1. Marker traits petrosal auditory bulla 2. Features of the molar teeth, moved into trees and began eating fruits 3. Features of the ankles and wrists 4. Not like primates claws and nails, long snout, laterally facing eyes, basically insectivore adaptations iii. Visually-oriented predators (Matt Cartmill) 1. Plesiadapoids are not primates, many did not have primate-like dentition, features of ankles/wrists disputed, some dont have petrosal bulla, some evolve to flying lemurs or other mammals 2. Problem of monophyly plesiadapoids evolved into a number of different modern mammals, have to make a taxonomic shift 3. The last common ancestor of all members of a taxon must also be member of that taxon 4. If exclusively arboreal mammals exist and are successful without primate adaptations, something is wrong with the arboreal theory of primate evolution 5. Of 13 orders of terrestrial mammals, 9 have arboreal forms, may of which do not have primate adaptations a. Squirrels are not at a disadvantage despite lack of primate adaptations, retain primitive mammalian bone structure, clearly another way to exploit arboreal habitat 6. Comparative method a. Claws highway branches (larger than their body size), take food to large horizontal branches, often go to ground to retrieve food, descend face down b. Fingers branches smaller than body size, use small terminal branches to feed 7. Primates are adapted to use terminal branches of dense bush layer of the forest

8. What is the adaptive significant of orbital convergence, frontation, and stereoscopy? a. If aiming towards agility in trees, would expect closely related animals to have greater convergence in tree forms, faster, more acrobatic forms would have more convergence b. Neither prediction held true: in marsupials, squirrels, tree shrews, and carnivores, no greater convergence in arboreal forms, slowest primates have the greatest convergence c. Like cats and owls, earliest primates were visually-orientated predators of the small branch milieu of the bushes and trees, moved into a new niche 9. Problems with the theory a. Only a very few of the 100 prosimians eat over 50% insects, most are omnivorous b. Omnivory is the primitive condition of primates morphologically (dentition and gut tract) c. Most convergent eyes found in slow lorises (90% of diet is fruit and gums, of the insects they do eat, they are slowmoving, smelly, and conspicuous) d. Hearing is the main sense used by other fast moving nocturnal primates and insects mostly ambushed on the ground e. Cats use hearing mainly and eye anatomy is only superficially similar to that of primates f. Many strictly plant-eating arboreal mammals have convergent eyes (primates, sloths, koalas, phalangerid marsupials) and many predatory birds g. The animals with the visual system most similar to that of primates are megachiroptera (fruit bats) iv. Angiosperm coevolution (Robert Sussman) 1. What was happening on earth 60-120 MYA, when the earliest primates were evolving? 2. What adaptive niche were the earliest primates exploiting? 3. Frugivorous birds, bats, and primates are important because the pollinate flowers and,

4.

5. 6.

7.

8.

more importantly, disperse fruits Birds were few and insignificant 65 MYA, some very primitive mammals, reptiles dominated landscape and most niches, most trees were gymnosperms, angiosperms prominent in edge regions, important after disturbance Dinosaurs kept angiosperms in check, despite very rapid reproduction, never reaches tree height, maintained for millions of years With extinction of dinosaurs came fall of gymnosperms, either killed by the same thing or angiosperms were no longer kept in check a. Allowed more dense growth of angiosperms, more shade tolerance seeds, larger seeds stored more resources, allowed for taller growth, slower reproductive turnover i. Movement from r-selection to kselection, fewer large seeds, rather than many small b. Rapid evolution of these new angiosperms during this Paleocene period Mammals and birds no longer have to compete with the dinosaurs, and plenty of niches are open with emergence of these large angiosperms a. Mammals could go after the large seeds, so plants develop nutritious covers for the seeds, animal takes seed and deposits away from mother tree b. Mammals can move into the trees to take on an arboreal lifestyle, pre-cursors of primates c. Radiation of tropical rainforest emerges toward the end of the Paleocene i. Plesiadapoids are a good example of movement of mammals moving into these arboreal niches, one of these probably evolved into modern primates ii. Begin as insectivores, but frugivores emerge within the radiation, some are completely folivorous The Eocene is the period during which this adaptive radiation come to fruition, now large plants with large seeds dominate the landscape,

modern taxa are present in modern rainforests, modern families and genera of angiosperms are present a. Major avian radiation to fill niches provided by large fruits b. Major mammalian (rodents, rabbits, and hoofed animals) radiation, bats and batadapted fruit c. First euprimates develop during early Eocene d. Animals are better able to exploit the terminal branches of these angiosperms (gums, fruits, leaves, insects that eat these things) e. Primates, with their grasping appendages, can exploit this new niche very effectively f. Frontation and Convergence develop in a number of mammal groups, some of which feed solely on foliage, evolve to see through clutter in the vegetation i. 2 forward facing eyes can see 2 to 8 times further in a highly cluttered environment than one lone eye ii. Strong correlation between leafiness and convergence relative to body mass g. Most likely hypothesis based on the time frame in which grasping limbs, primate dentition, and orbital convergence evolved v. Cartmill: If the first euprimates had grasping feet and blunt teeth adapted for eating fruit, but retained small divergent orbits, the terminal branch feeding hypothesis would be supported, thus disproving his own theory 1. Carpolestes simpsonii (plesiadapoid) fossil discovered basically has primate grasping hind foot, spent little time on large diameter supports, occupied small branch niche, lowcrowned molar teeth, primarily fruit eaters, also exhibits small divergent orbits vi. The question remains where we should make the cutoff for primates in the taxonomy, is Carpolestes the common ancestor for more than just the primate phylogeny e. Several of the previously recognized unique

characteristics of living primates were acquired incrementally during the course of primate-ancestor evolution f. Consistent with the idea that primates acquired their suite of diagnostic features through diffuse coevolution with angiosperms during the Paleocene-Eocene transition Lorisiformes 1) Taxonomy has been the one consistently inconsistent aspect of primatology, many more species of lorisiformes are now recognized than decades ago, in nocturnal primates, similar looking animals turned out to be different species, mating vocalizations are now the most accurate way to distinguish a. Now, for conservation purposes, new species are being made out of small populations discovered in new systems or members of a single population that appear slightly different 2) Using primarily Charles-Dominques study of these five species in Gabon a. Determines the type of habitat each species used i. Discontinuous study 836 first sightings (42 months over 8 years), notes the height in the forest, orientation of the branches they used, the diameter of these branches, and nature of the support b. Determines the diet i. Shot animals outside of his study site (201 specimen of all different species, sizes, times of year, and times of day), divided into animal, fruit or gum 3) Galaginae (fast-moving African galagos) a. Synterritoriality found in some galagos, lemurs, and tarsiers, one male and one female home ranges overlap (corresponding home ranges), may have multiple females sharing one home range, do not move around together at night, may sleep together in the same nest b. Galagoides demidovii (Demidoffs Dwarf) 60 grams i. Canopy dwellers, most of the time from 10-20 meters in primary forest, 64% were less than 1 cm, most of the rest of the time on branches less than 5 cm, 22% on horizontal, 30% on oblique, and only 48% on vertical branches (or so Charles-Dominque thought, he was actually studying primarily thomasi) ii. 70% of the diet was animal, 19% was fruit, and 10% was gum (insectivore) iii. Galagoides thomasi 60 grams 1. They were found in edge forest, from 0-10 meters in secondary forest c. Sciurochieirus gabonensis (Gabon squirrel) 260 grams i. Spent most of the time in the understory, 0-2 meters

ii. iii.

iv.

v. vi. vii. viii. ix.

x.

xi.

in primary forest, 60% of the time on small branches (under 5 cm), 6% on horizontal, 13% on oblique, and only 81% on vertical branches, specialized for the vertical branches 25% of the diet was animal, 73% was fruit, and 5% was gum (frugivore), mostly fallen to the ground, already ripe, not being fed upon by other potto Most generalized form of sociality in nocturnal prosimians, females have fairly small home ranges, that overlap quite a bit, and multiple females will share home range and avoid females form other home ranges, will form nesting groups within (1 to 5 individuals) Males have fairly large home ranges (30 hectares vs. 10 in females, overlap more than one female group, males avoid one another, do not form nesting groups, will mate with females in their home range, if two males overlap with a female home range, females will potentially mate with both males Smaller males are forced to the periphery or may have very irregular ranges Reproductive harems develop whereby one male tends to mate with more than one female, interact throughout the year Evolution towards one-male group in harsh environments because population relies on females, one male can mate with many females Males are territorial, females are less territorial, though do try to avoid each other Female offspring stays with mother long after weaning, leads to matriarchal groups, though males are forced out after puberty, young are left in nests by mothers 1. Some species have more tolerance for males, may have home ranges within the ranges of other males Urine markings for social, territorial purposes, CharlesDominque found that females move throughout the whole range, spent more time urine marking around nests and 3.7 times per hour in overlap zones (with other females), communicated where they were so they could avoid each other, male can also tell mating condition (estrous) by marking in overlap zone between male and female Two types of calls: alarm is never answered so that they can remain hidden, croaking call is always

answered for communication purposes, attracts males and tells females to stay away (8 times more likely in overlap zones) d. Euoticus elegantulus (Southern needle-clawed) 300 grams i. Found from the ground all the way up to 50 meters, moves primarily vertically, use nails as a secondary adaptation to claws to move along highway branches, 22% on horizontal, 51% on oblique, and only 27% on vertical branches, specialized claws and dentition allows them to fit into a unique niche between the two other species ii. 21% of the diet was animal, 10% was fruit, and 75% was gum iii. 78% of insect diet are beetles and moths (fast moving insects) iv. Lives in the canopy with potto, both eat gums, during dry season, potto eats only fruits and insects while elegantulus focuses on gums, uses canine-like incisors to get to it e. Protection against predation - very fast moving, can usually outrun or outmaneuver predators, warning calls, mobbing behavior 4) Perodicticinae (African slow lorises) a. Perodicticus potto (Western potto) 1100 grams i. Remains in one place without moving when faced with a predator, faces the predator, adaptations to hands and feet (maximizes blood flow) that allow it to stick to the branch for a long time, spines on its neck that allow it to bend its head forward, revealing only these spines, butts the predator, when faced with a snake, will drop to the bottom of the branch and falls to the ground, then moves away relatively quickly once on the forest floor (7 out of 10 times), difficult to find them afterwards 1. Researchers have never found a potto in a predators stomach contents ii. Canopy dweller, 10-30 meters in the primary forest, in the secondary forest it was 10-15 meters up, locomotion was climbing rather than jumping, branches went from 1-30 cm in diameter, found 39% on horizontal, 35% on oblique, and only 26% on vertical branches iii. 10% of the diet was animal, 65% was fruit, and 21% was gum 1. Actually consumed approximately the same amount of insect mass as Archtocebus, both

were going after the same thing, but because potto was much larger, had to supplement insect with fruit 2. Eats primarily ants (formic acid), centipedes (iodine), and crickets, all toxic, adaptations to detoxify 3. Tested whether these species prefer toxic insects or just eat them because they are available, actually prefer moths without their wings, but cannot catch them iv. Females do not form nesting groups because they do not go back to nests (slow moving), young are carried on fur, baby parking when they feed, female ranges are relatively small and do overlap while males have larger home ranges that overlap multiple female ranges v. Female offspring , as well as males, develop their own home ranges after weaning, no matriarchies vi. Depend more on scent marking rather than calls because they do not want to attract attention of predators b. Arctocebus calabarensis (Calabar angwantibo) 200 grams i. Different set of smaller predators, rather than facing the predator, will move into dense vegetation and will face away, tail almost acts as a target, will bite the nose of the predator, when it is shaken off, it will let go of its grip and fall to the ground ii. Stayed in the undergrowth, primarily between 0-5 meters, may move upward if disturbed, spent time in bushes and on lianas between bushes, 40% of their supports were less than 1 cm, 52% were 1-10 cm, found 20% on horizontal, 30% on oblique, and 50% of the time on vertical supports iii. 85% of the diet was animal, 14% was fruit, and 0% was gum (insectivore) iv. 70% of their insects are caterpillars, most of which are also toxic, large quantity available because other animals do not predate upon them often, slowmoving, smelly or visible c. Protection against predation - have developed specialized anti-predation techniques, very cryptic, slow-moving and try not to disturb vegetation 5) Lorisinae (Asian slow lorises) a. Protection against predation - smell bad, glands secrete toxins, can kill a human fairly easily, will lick babies to cover them with these toxins

The Adaptive Array of the Lemurs of Madagascar 1) About 500 miles long and 250 miles wide, third largest island in the world, has almost all of the same ecosystems as can be found on mainland Africa a. The central plateau is high and wet, where most of the people live, deforested a great deal of the land area though was probably densely forested before humans settled b. Eastern portion has mountains and rainforest areas c. The north has deciduous forest, gets drier along the western coast until desert in the southern portion d. Major groups are divided into east (adapted to rainforest) and north and west (adapted to drier deciduous forest), another split is between nocturnal and diurnal (approximately 50% each) i. Most of the new species being discovered are in the east where one would expect higher diversity, very few species are found in both habitat types ii. Most Lemuridae are diurnal, a few are cathemeral (active during the day and night), none are completely nocturnal, Lepilemuridae are all nocturnal, Indriidae are primarily diurnal iii. At least 16 species of lemur have gone extinct within the past 2000 years (since humans came to the island), all of which were larger than our modern species, all filled different niches e. Southwestern and Western Forests i. 3 diurnal species (only 2 as you move further south) and 5 nocturnal species f. Northwestern Forests i. 5 diurnal species, though Lemur catta has fallen out, replaced by arboreal species, 6 nocturnal species g. Eastern Forests i. 7 diurnal species (more arboreal and bamboo-eating species), 6 nocturnal species (different from those found in NW Forests) ii. Diversified the lemurs (belong to more families) h. Western Lemurs from October to January, have many fruits and flowers, between May and September, conditions become much harsher as fruits and flowers disappear during the dry season i. Lemur catta (Diurnal) Ring-tailed Lemur 1. Only living lemur that spends the majority of its time on the ground (33%), and varying amounts of time in all other stratum, with the exception of the emergent layer, feeds mostly on the

ii.

iii.

iv.

v.

ground and below the canopy, travels primarily on the ground 2. Mainly frugivorous, continues eating fruits and herbs off the ground during the dry season, but will eat a slightly generalized diet of leaves Eulemur fulvus rufus (Diurnal) Brown Lemur 1. Almost exactly the same as ring-tailed lemur, very subtle and specific differences, spend most time in the canopy stratum of the forest (71%), most of the feeding time is in the region as well, travel through the canopy 2. Specialized folivore, feeds a great deal on a few species (11), 3 species make up 80%, has adapted to the toxins of these plants, prefers fruits during the wet season, but will eat leaves during dry season Propithecus verreauxi (Diurnal) 1. Much larger, about the size of a gibbon rather than a cat, spend most of their time on the vertical trunks beneath the canopy, and some time in the canopy and the emergents where it does most of its feeding, travel beneath the canopy, but off the ground 2. Generalized folivore, lots of different species of plants (85), limit the amount of any one toxin, 24 species make up 80% of their diet Cheiriogaleus medius (Nocturnal) Fat-tailed Dwarf Lemur 1. Unlike any other primate, actually hibernates during the dry season when fruits and flowers disappear, stay in hollows in tree trunks, emerges in November, quite active during the rainy season, opportunistic frugivore, increase amount of food with some insects, prior to hibernation, will accumulate fats from fruits in their tails, may gain up to 50% of their body weight in tail 2. Scurry like mice, not particularly agile, fat tail is not conducive to very acrobatic locomotion, come to the ground for insects, but focus primarily on fruits Microcebus murinus (Nocturnal) Mouse Lemur 1. Go into torpor (state of partial hibernation), females are less active, come out of torpor less often than males, people believed the populations were strongly weighted towards

males, actually just during the dry season, stay in hollows for a number of consecutive days 2. Live mainly in the canopy and eat primarily fruits and insects (approximately 50% of diet is mistletoe-like fruit) vi. Microcebus (Mirza) coquereli (Nocturnal) 1. Lives like Mouse lemur in dense foliage of canopy, eats insects and fruits, during the dry season, focuses on a specific insect, eating its secretion rather than the insect itself, high carbohydrate, so can make it through dry season without fruit 2. All three species eat the same things during the wet season when resources are abundant, all have very special adaptations to get them through the dry season vii. Phaner furcifer (Nocturnal) 1. Active throughout the year, specialized to focus on gums throughout the year, especially during the dry season, moves through the canopy both vertically and horizontally, will eat fruits and floral nectar during the wet season 2. Adaptations for eating gums: has claws (secondarily adapted from nails), procumbent incisors for gnawing into bark, long, pointed tongue for reaching gums viii. Lepilemur mustelinus (Nocturnal) 1. Specialized folivore, can eat mature leaves (91% of diet), which many primates have trouble digesting, large caecum and wide colon adapted for eating leaves, shearing crests on molars adapted for grinding leaves, can detoxify certain toxins, so can eat as many as 24 species, or as few as 2 i. Northern Lemurs i. Eulemur fulvus fulvus brown lemur 1. Different from the E. f. rufus found in the west 2. With E. mongoz: niches overlap heavily during wet season, during resource-poor season, fulvus remains pretty much the same, but mongoz becomes nocturnal, will eat primarily flowers and nectaries (81% of diet), some fruits, and very few leaves 3. With E. macaco: nobody has done a good study on their interactions, assume they divide up the landscape

4. With E. coronatus: found primarily in the under story while the brown lemur is found primarily in the middle story, this means they are using different trees, 18% of crowned lemur diet is shared with brown lemur, about 75% is different (similar species, but different parts of the trees due to their choice in strata) ii. Eulemur mongoz, Eulemur macaco, Eulemur coronatus 1. Ecological equivalents with one another, replaced as you move north, ranges do not overlap, though all overlap with E. fulvus iii. Propithecus spp. 1. A number of different species are ecological equivalents, all fill the same niches, so will never be found in the same area, changes as you move around the island iv. Hapalemur griseus v. Avahi langier (Nocturnal folivore) vi. Lepilemur mustelinus (Nocturnal folivore) vii. All other nocturnal species also present j. Eastern Lemurs i. Eulemur fulvus rufous (Frugivore) 1. With E. rubriventer: only 1 species in their top 5 most commonly eaten plant species overlaps between the two, different species, but similar parts, though E. rubriventer eats more fruits and less leaves than E. fulvus, not as well adapted to toxic parts of the plant 2. With V. variegata: most of their time is spent up higher in the forest (15-25 meters) while E. fulvus tends to be lower, V. variegata is a specialized frugivore in these taller trees ii. Eulemur rubriventer (Frugivore) 1. Live in pair bonded groups rather than large social collections iii. Varecia variegata (Frugivore) iv. Propithecus diadema (Folivore) 1. Multi-female groups 2. With I. indri: eat similar things, but at different rates, Indri is much more specialized towards leaves, while P. diadema eats almost as much fruit (actually seeds within) as leaves v. Indri indri (Folivore) 1. Family groups (mother, father, and offspring), probably related to resource distribution

vi. Hapalemur grieus (Bamboo and Grass Eaters) 1. Most generalized, smaller, different types of grasses and bamboo, sometimes fruits as well, detoxifies bamboo vii. Hapalemur aureus (Bamboo and Grass Eaters) 1. Very localized, slightly more generalized than H. simus, fairly small, detoxifies bamboo viii. Hapelemur simus (Bamboo and Grass Eaters) 1. Highly specialized for one type of bamboo, bigger, all three detoxify bamboo differently k. Lemur Conservation i. 85% correlation between human population density increase and deforestation of forested regions ii. After economic surge that brought people into the city in the 1970s, people began returning to rural areas and deforesting 1. Cutting forests for corn plantations, firewood, and slash and burn agriculture Quantitative Study of Primates 1) A study was performed in which a Brown Lemur (Eulemur fulvus fulvus) was released in an area where Ring-tailed Lemurs (Lemur catta) were found, but where Brown Lemurs were not normally found, the Brown actually adapted completely to the Ring-tails lifestyle a. This was presumed to mean that the two species would compete directly with one another b. Hypothesis was that the habitat and diet was actually divided between the two species when they were found together i. Found three different types of forests (catta alone, fulvus alone, both together) to determine behavioral differences 2) Needed to collect quantitative data a. Scan-Sampling Method every five minutes, determine the number of individuals (single or pair) participating in each type of task (feeding, resting, grooming, individual moving, group travelling, other), also noted which level of the forest strata they were performing these tasks, ad-lib notes as well for other, may take note of plant species they are eating b. Ring-tailed Lemur are found throughout the south of the island, Brown Lemur are found everywhere else (wetter forests), despite differences in coat color, actually very similar, both species ranges are sympatric with Propithecus verreauxi i. In most cases, when there is an edge habitat, it is more diverse than the main forest and nearby

savannah or grassland ii. Ring-tails tend to be found in edge forest and brush as well as gallery forest, while Browns were found in only gallery iii. Structure of forest at three different sites was similar, though the species differed c. Utilization of Forest Strata i. Regardless of Activity 1. Brown Lemur spent the great majority of their time in levels 3, 4, and 5 (essentially remains in the canopy), Ring-tail spend a lot of time on the group and significant time in 3 and 4 (move vertically a great deal), unlike Brown lemur, the Ring-tail actually spent 60% of their time at the edge or in the brush areas 2. Apparently activity did not change when the two species were together or separate 3. Unlike the Brown Lemur, and all other lemurs, heel of the Ring-tail is not covered in hair, more like monkeys an apes, different means of locomotion due to movement on ground rather than in trees a. Amount of rotation at joints in Brown lemur is much more like arboreal primates, while Ring-tails were more like terrestrial primates (more stability), densities differ this way d. Utilization of Horizontal Space i. Ring-tails covered a large portion of the forest, in and out, one group of 19 animals covered about 20 square hectares, 9 groups of 5-7 Brown Lemurs utilized the same 20 hectare space, remained in small home ranges of 1-1.5 hectares ii. Ring-tails would move approximately 1000 meters in a single day, cover their entire range every 7-10 days, covering a number of different habitat types in a single day iii. Brown Lemurs moved about 250 meters within much smaller day range, but tended to move around almost their entire range in a single day e. Diet i. Ring-tails ate a much larger variety (24 species) of plants, very similar between sites, only about 25% of their diet is Tamarindus indica no matter what the season, even during the dry season, a significant portion of the diet came from fruits and herbs (leaves

make up 43% and 24% of dry and wet season diet, respectively) 1. Adjust their home range to find what trees were flowering at any given point during the year, flock to that for an number of days (frugivore) 2. Due to their exploitation of the ground and ability to move over large areas ii. Brown Lemurs ate a small variety (8 species), Tamarindus indica could make up nearly 80% of the diet during the dry season (eating primarily leaves), though only about 50% in the wet season (eating much more fruits and flowers) 1. What they ate really depended on their home range, some group could get fruit while others had no flowering trees in their home range (folivore) f. Activity Budgets i. Ring tails would eat a bit in the morning, but primarily rest, feed throughout most of the day, rest for a short period during the hottest time of the day, continue eating into the afternoon, rest towards the night ii. Brown Lemur would eat primarily in the morning and afternoon (sometimes going into the night), rest for the majority of the middle of the day g. Group size and composition i. Both species live in multi-male multi-female groups (3.4 male 4.3 female in Brown Lemur, 6 male 6.3 female in Ring-tailed Lemur), also have some juveniles (1.2 in fulvus, 3 in catta) and infants (1 and 3, respectively) ii. Ring-tailed lemurs tend to have larger groups (10-30 members) with a distinct hierarchy with sub-groups, central males in center with females, sub-adult males are peripheral, females also have hierarchy based on matrilines (alpha female family is dominant to beta female) iii. Brown lemurs tend to have smaller groups (4-17 members) in an egalitarian manner with no real hierarchy h. Summary i. Social structure and behavior are directly related to their differential use of time, food, and space ii. Behave the same whether they are alone or together, no character displacement, behavior based on long evolutionary history utilizing different niches (Brown adapted to wet forests while ring-tails are adapted to

drier deciduous forests where resources are more uneven) iii. Parallels between behavior and ecology in these species and similarly-adapted monkeys in Africa (higher density of folivores, smaller home ranges in arboreal) iv. Can relate morpho-physiology, social behavior and structure to ecological differences Groups and Social Structure 1) Groups are a number of individuals of the same species who interact socially more frequently among themselves than other conspecifics, and these group members exhibit different behavior to conspecifics not in their group, and occupy the same home range, groups also tend to be fairly stable 2) Social Structure is divided into group structure and populations structure a. Group structure is the size and composition of the group b. Population Structure is the distribution of groups within the population 3) Social Organization is the pattern of interactions between individuals both within and between groups 4) Home Range is the area over which group members travel in pursuit of their everyday tasks over their lifetime a. Core Area is the portion of the home range used with greater regularity than other areas (may define as 50% or 95%), tend to be most resource-rich 5) Day Range is where the group travels in any given day 6) Territoriality is the behavioral mechanisms that control space between two animals such as ritualized behavioral displays, must also have very little overlap between groups to be considered territorial 7) Types of Group Structures we find in primates a. Solitary but Social sleep together, but do not move together, orangutans and most nocturnal prosimians b. Multi-male/Multi-female group more than one male and more than one female, found mostly in diurnal lemurs, many New World Monkeys, most terrestrial African Monkeys, some colobines, and gorillas c. One-male/Multi-female group harem with one male and many females which whom he may mate, Old World Monkeys d. Fission-Fusion number of animals interact in a community, but come together and move apart from time to time i. (New World) Spider-monkeys different individuals on different days, but always the same animals in the long term

ii. Chimpanzees live in large groups iii. Baboons not the individuals that come together and break up, but subgroups (Gelata baboons have onemale reproductive groups, Hamadryas have one-male groups together in bands and even larger troops) e. Pair-bonding Groups male and female with their offspring, found in Indri, rubriventer spp., colocebus, and gibbons, tend to be territorial f. Polyandrous groups one reproducing female with more than one male in the group, may also have non-reproductive females, found only in callitrichids