Cretaceous Period (135 - 65 m.y.a.

)
Extremely active geological period
Pangaea split into two segments by 125 m.y.a.

Human Evolution and Adaptation

Life Styles of the Merely Hominid

Northern land mass: Laurasia

Included North America, Europe, Most of Asia Rise of the Rocky Mountains

Southern land mass: Gondwana

Included South America, Africa, Australia, Antarctica, Indian subcontinent

Worldwide climate much warmer than today so tropical and sub-tropical fossils are found far from the equator

Cretaceous Period 2
Floral Shift from gymnosperms to angiosperms as dominant land plants
Gymnosperms are the vascular plants with seeds that are not enclosed in an ovary (naked seeds), mainly the cone-bearing trees (ferns, ginkos, cycads, and conifers) Predominate from the Carboniferous period (about 350 m.y.a.) when they began to displace the earliest spore-bearing land plants to the Cretaceous (about 125 m.y.a.)

Cretaceous Period 3
Angiosperms are the flowering plants, an advanced group of vascular plants with floral reproductive structures and encapsulated seeds including flowering herbs and trees, first appear near the end of the Mesozoic (135 m.y.a.) The flowering mechanism increased the potential for genetic diversity (decreasing self pollination)
Diversity of the angiosperms increased through coevolution with insect species, making for rapid adaptive radiation

Cretaceous Period 4
During the Cretaceous angiosperms spread to build forests of increasing complexity, and took over the dominant land plant role after the K/T (Cretaceous/Tertiary) extinction New econiches opened and old ones expanded
Frugivory: flowers and fruits are new food sources Gramnivory: encased seeds from the new plants Insectivory: bugs that co-evolved with flowering species multiply increasing bug eating opportunities

Cretaceous/Tertiary Event
Comet collision represented by the Chicxulub impact crater off the north west coast of the Yucatan Peninsula
Combined effects of terrestrial and marine impact Dust and debris cause cooling, break down of food webs

Cretaceous Continental Configurations

K/T
There is also a lot of volcanic activity on the Deccan Flats of the Indian subcontinent at this time, adding to cooling > 50% of extant genera disappear at this time
Terrestrial reptiles and marine invertebrates most heavily affected No land vertebrate larger than 50 pounds in body weight survived the K/T

Primate Adaptation
Our Cretaceous ancestors were small, fuzzy critters that were generalized enough in diet, morphology, and behavior to survive the K/T extinction Our primary adaptation at that time consisted of the Good Luck of being small, omnivorous, and flexible
Otherwise we would have gone the way of all large bodied terrestrial vertebrates This stochastic process appears to play a role in most mass extinctions including the Permian and K/T

Paleoclimates

Paleocene to Eocene Transition

Warming into the Eocene
Warmest epoch of the Tertiary

Eocene Adaptations
First true Prosimians (Euprimates) Primates spread with forested zones, appear to adapt to preying on small, quick moving prey in arboreal settings (visual predation)
Grasping hands and feet Nails instead of claws Eyes rotated forward, enhanced stereoscopic vision Elaboration of visual sensory pathways

Wide ranging evergreen rain forests throughout North America and Europe
Two cooling episodes in the Paleocene broke the tropicality From the mid-Eocene on there was a gradual cooling and drying of the higher latitudes

The primate rainforest habitat was very widespread during the peak of the Eocene

Cenozoic Climates

Eocene to Oligocene Transition

Continued cooling, lowered sea level
Extinction episode of many large bodied mammals at close of Eocene Mid to high latitude vegetation changed dramatically from broadleaf evergreen rain forest to deciduous forests
Remnant primates forced to cluster into smaller habitable forest areas near the equator (Fayum) Increased competition probably drove some changes in behavior and adaptive patterns

Habitats suitable for primates retreat into the current tropics where most Oligocene primates are found

Mid-Oligocene Extinction
There is a mass faunal extinction event spanning about 20,000 years at about 32 million years ago
Evident in the disappearance of archaic North American land mammals

Potential Causes:
Increased glaciation Worldwide cooling Retreat of the Oceans Floral changes related to Ocean circulation changes

Fayum

Early Oligocene, 35 m.y.a.

Oligocene Adaptations
First Anthropoid primates appear just after the Oligocene extinction
Body size larger than earlier primates
Larger brains, possibly slower development

Oligocene to Miocene Transition

Warmer than the late Oligocene with higher sea levels
Temperature peaks in mid-Miocene, about 15 m.y.a, then cools and climate becomes drier
Variability in temperature becomes extreme

Primarily fruit eating Sexual dimorphism suggests early anthropoids were living in large, complex polygynous groups

Continents were in nearly modern positions

Impact of India causes uplift in South Eurasia, building Himalayan Mountains African collision with Southwest Eurasia causes rift valley system and allowed faunal dispersal

Learning and social complexity may have served to buffer them from the extremes of the Oligocene climate, or the increasing variability of climate

Miocene Climates Miocene Habitats

The increase in climate variability would have been evident in many regions as increased seasonal variability in temperature and rainfall
One result was the increasing spread of grasslands Animals with large bodies to travel long distances and large guts to process the fibrous grasses sprang up on every continent

F o r e st

Gr ass

Gr

a ss

Cool/Dr y Phase Modified fr om Conr oy, 1990:192a)

Moder ate Phase Modified fr om Conr oy, 1990:192b)

Miocene Adaptations
About 18 million years ago we begin to see the expansion of the hominoid apes
F o r e st
Fossils occur in more open woodland and woodland to forest transition areas
Apes have larger bodies, larger brain to body size ratios
Thicker enamel enables a broader diet of soft and hard foods

Gr

a ss

Likely that developmental slow down is becoming pronounced in these animals

Longer developmental period enables more learning of complex behaviors to take place in larger brains Provides a behavioral buffer for the extreme environmental variability of the Miocene

War m/Wet Phase Modified fr om Conr oy, 1990:193c)

Cenozoic Climates and Primates

I: Height H: Radiation primate diversity hominoids, G: Post of episode appearance of Purgatorius F: DeclineofandPlesiadapiforms Europe E: NadirK/TprimateHominoids inOmomyids D:Evolutionand extinction ofandnorthern C: B: Appearance of of Northern A: Disappearance Adapids primates in of dispersal of in diversity humans extensive continents America Europe and North America and Asia ecological radiation of North cercopithecoids, and emergence of hominids

Late Miocene Climates

Relative climatic stability occurs from around 10 mya to about 6.4 mya
Cooler temperatures prevailed than earlier in the Miocene, and ice sheets were constant From 6.4 to 4.6 mya, large scale climatic fluctuations returned, larger than 16 -12 mya, with glaciations being large enough to isolate the Mediterranean from the Atlantic and to dry the Mediterranean up periodically (the Messinian crisis) A small, insignificant hominoid species began to walk upr ight during this time

Climate Change Last 6,000,000 Years Miocene Habitats

The increase in climate variability would have been evident in many regions as increased seasonal variability in temperature and rainfall
One result was the increasing spread of grasslands Animals with large bodies to travel long distances and large guts to process the fibrous grasses sprang up on every continent

Miocene African Forests

Early Hominid Patterns

The vertebral column articulates at almost a right angle to the base of the skull creating a sharply angled vocal tract Larger group sizes, facilitated by bipedalism, leads to increased opportunity for learning and communication
Dunbar (1996) posits this as the time for the transition from grooming to gossip as a means of group solidarity

Humans versus Apes

To understand the patterning of hominid adaptations, we need to be aware of the differences between humans and our closest relatives, the African anthropoid apes: Gorillas, Chimpanzees, and Bonobos The two key morphological differences include reduced canines and bipedalism
Bipedalism changed elements throughout the body and also must have had significant behavioral ramifications

Locomotor differences
By comparison to apes, humans have:
a foramen magnum that points down a curved lumbar spine a short, flared (versus long and thin) ilium (the upper most section of the hip bone or pelvis) a strong, robust talus (ankle bone) a strong, non-opposable big toe a complex two-way arch system in the foot

Hip Comparisons Posture Comparisons

Knee Comparisons

Foot Comparisons

3 - 4 mya

A few of the Many Just-So Stories about Hominid Origins

Tool manufacture and use Hunting Seed eating Prostitution Aquatic adaptations Walking efficiency Vertical Climbing Terrestrial Quadrupedalism

Tool manufacture and use

Bipedalism, small canines, and large brains are all linked to the manufacture and/or use of tools for early hominid adaptation to the environment
Larger Brains Tool Use Smaller Canines

Bipedalism

Tool Use According to Darwin

In The Descent of Man (1871), Darwin proposed an adaptive model that views encephalization, dental reduction, and bipedalism as systems among which feedback from the use of tools has occurred
Many have followed his proposals suggesting that all hominid characteristics are considered to be a complex of interrelated traits that evolved more or less simultaneously

Darwin proposed that bipedalism arose to free mans hands for tool use
Man alone has become a biped; and we can, I think, partly see how he has come to assume his erect attitude, which forms one of his most conspicuous characters. Man could not have attained his present dominant position in the world without the use of his hands, which are so admirably adapted to act in obedience to his will. But the hands and arms could hardly have become perfect enough to have manufactured weapons, or to have hurled stones and spears with a true aim, as long as they were habitually used for locomotion and for supporting the whole weight of the body, or, as before remarked, so long as they were especially fitted for climbing trees. Such rough treatment would also have blunted the sense of touch, on which their delicate use largely depends. (Darwin, 1871)

Darwin also proposed that the free, manipulative hands led to the reduction of dentition found in humans relative to our pongid relatives
The free use of the arms and hands, partly the cause and partly the result of mans erect position, appears to have led in an indirect manner to other modifications of structure. The early male forefathers of man were, as previously stated, probably furnished with great canine teeth; but as they gradually acquired the habit of using stones, clubs, or other weapons, for fighting with their enemies or rivals, they would use their jaws and teeth less and less. In this case, the jaws, together with the teeth, would become reduced in size, as we may feel almost sure from innumerable analogous cases. (Darwin, 1871)

Finally, Darwin argues that the extreme encephalization seen in humans would have been driven by the demands of more and more taxing intellectual challenges associated with tool manufacture and use
As the various mental faculties gradually developed themselves the brain would almost certainly become larger. No one, I presume, doubts that the large proportion which the size of mans brain bears to his body, compared to the same proportion in the gorilla or orang, is closely connected with his higher mental powers. (Darwin, 1871)
He is explicitly refuting Wallaces position in this argument, I cannot, therefore, understand how it is that Mr. Wallace (1869) maintains, that natural selection could only have endowed the savage with a brain a little superior to that of an ape.

Problems with the Tool Use Just-So Story

The brain, teeth, and bipedalism evolved at very different rates and different times Many small brained, non-bipedal species use tools; chimps manufacture and carry even stone tools Bipedalism shows up at least 2 million years before stone tools appear

Seed Eating
Jolly (1970) looked to gelada baboons to explain human dentition and posture, based on the assumption that early hominid diets would have been grass seed based in the drier environment
Smaller canines and heavy occlusal wear are found for processing seeds Seated, upright posture accompanies the squatting processing of grasses

But bipedalism clearly evolved in heavily forested, not savanna setting.

Hunting
Washburn focused on the role of men in hunting as an explanation for bipedalism
He argued that bipedalism evolved as an enabling mechanism for men to carry food from the hunt to provision their mates and offspring Those most successful would leave the healthiest offspring most likely to survive

The Prostitute Model

Lovejoy proposed a model of hominid origins based on increased reproductive efficiency
In this scenario, males who were mobile and successful at hunting and carrying nutritious food back to females were more successful at reproducing than were less mobile males (i.e., those less bipedal)

Serious hunting appears millions of years after bipedalism evolves

Chimps do it, Baboons do it.

The Prostitute Model--2

Females stayed close to a home base and invested their time in rearing offspring
Food that was readily available to females and children near the home base was not usually of high quality Females who were provisioned by males would have better success at raising young

The Prostitute Model--3

Provisioning also would have shortened the interval necessary between births
Provisioned females would have to expend less of their own energy to sustain infants, both in terms of carrying and in terms of breast feeding Provisioned females would return to an ovulatory state sooner, and be able to produce more offspring on average than nonprovisioned females

The Prostitute Model--4

Sexual selection operated to reinforce the provisioning of females by males
Fatty storage especially in breasts and buttocks marked fertile, desirable females Cryptic estrus allows females to fake it by copulating throughout the menstrual cycle without external evidence of peak fertility periods to restrict sexual activity Both of these facilitate long-term pair bonding

The Aquatic Ape

Partly in response to Man the Hunter evolutionary scenarios, Elaine Morgan proposed that hominid evolution was primarily a female phenomenon
She says that several female characteristics such as higher body fat levels and less body hair came along with standing bipedally in the surf off East Africa
In this model, males splash along behind

Thermoregulation
Upright, bipedal stance means that a hominoid absorbs 60% less heat at noon by reducing body surface area exposed to sun
Loss of body hair also facilitates cooling by sweating
77 pound bipedal hominid requires 3.3 pints of water a day for cooling compared to 5.5 pints for a hairy quadruped of the same weight

Posture selection
Jablonski: Typically hominoid threat display is bipedal stancetaller = more impressive
Intrasexual selection would favor long, bipedal legs for males and females would be included because of the necessary changes

How much were the early hominids exposed to direct sunlight for foraging, if they were forest dwellers
Mad dogs and Englishmen (Wheeler from Liverpool)

Hunt: 80% of bipedal behavior among chimps was found to be related to stationary feeding on fruits from bushes and low branches in small trees (the deer do this in my front yard)
Bipedal morphology would be selected for by enhancing resource competition

Walking Efficiency
Why would our ancestors have invested in this awkward way of getting around on the ground? Human bipedal running is both slower and less energetically efficient than is chimpanzee knuckle-walking or quadrupedalism But at a normal walking pace, for instance the way Australopithecus might have gotten from one food patch to another, human bipedal walking is more energetically efficient than is chimpanzee knuckle-walking Walking speed 2.9 km/hr 4.5 km/hr

Walking Efficiency, 2
Energetic Efficiency Energy cost Energy cost relative to ml 02/kg/hr quadruped 0.522 149% 0.193 0.426 0.170 86% 148% 94%

Species Chimp Human Chimp Human

2.9 km/hr is normal knuckle-walking speed of chimps, 4.5 km/hr is normal bipedal walking speed of humans (Rodman and McHenry 1980)

Bipedal Efficiency Walking efficiency, 3

Bipedalism would confer an adaptive advantage especially in times of low food availability and sparse distribution of food resources
Bipedal hominids would expend less energy to move from patch to patch of food, and would get there faster on average than knucklewalking early pongids

Walking efficiency, 4
In a highly variable and changing environment such as that found at the close of the Miocene, several patterns would emerge:
Forested areas with nesting and food resources would become more sparsely distributed
In the wet/dry cycling, forest will not quickly replace savanna once the grassland is established

Walking efficiency, 5
In this late Miocene setting, the Chimphuman common ancestor could have adapted in two different ways:
Remain quadrupedal/knuckle-walking and reduce group size so that less travel was necessary for foraging Increase the energetic efficiency of travel from one patch to another by becoming bipedal, allowing expansion of group size

There would be less certainty about the location of food resources from generation to generation

Early Australopithecines became bipedal while retaining climbing limb proportions and abilities

Social Group Size

Pliocene
This bipedal/climbing adaptation of early australopithecines remained stable through much of the Pliocene, despite the high levels of climatic variability
Adaptation includes retention of the Miocene trait of thick enamel in the cheek teeth so that both hard and soft foods were consumable

Elaboration of the pattern results of substantial regional and temporal variation and speciation

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Late Pliocene
Around 3 mya the Isthmus of Panama emerged as a permanent separation of the ocean circulation between the Pacific and the Atlantic/Caribbean
Several other planetary and continental forces combine so that by around 2 mya there is a major expansion of the Antarctic ice sheet and major ice rafts are recorded in the Northern Hemisphere There is a critical drop in temperature about this time, along with a nearly doubling of the level of climatic variability

Climate Change Last 6,000,000 Years

Late Pliocene, 2
Savanna habitats spread into previous woodland settings, along with increased climate variability
Some hominids appear to have adapted to this change by increasing dental surface for grinding tough, drought resistant plant foods (e.g., Australopithecus aethiopicus, A. boisei, A. robustus) Other hominids adopted toolmaking, transport of resources, and dietary change including increased use of animal resources (e.g., Homo habilis)

Plio-Pleistocene Setting
There is little average change in temperature between 2.5 mya and today
Dramatic changes in amplitude of changes Breakpoint about 600,000 years ago
The Ice Ages with periods of extensive glaciation in high latitudes alternating with brief warm periods

The survivor of this transition appears by 1.8 1.6 mya, (Homo erectus) with substantially enlarged brains and large erect bodies

The beginning of extreme fluctuation coincides with advanced Homo erectus

Continuing fluctuation produces Archaic Homo sapiens

Plio-Pleistocene Climates
Homo neanderthalensis Homo heidelbergensis Homo erectus Homo habilis/rudolfensis Less Ice

Pliocene Lifecycles
Australopithecus afarensis and earlier forms appear to have developed much like modern chimps
Leigh would add brief adolescent spurt to Bogin and Smiths model

Australopithecus africanus may have had slightly extended infancy and childhood
Increased adolescence in Leighs model
Mor e Ice

Homo habilis adds a childhood phase, and probably increased adolescence

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Pleistocene Lifecycles, 2
Early Homo erectus increases the childhood phase, but according to Bogin and Smith still had no adolescent spurt
Based on Leighs analysis, this group, represented by the oversized adolescent KNM-WT 15000 would be uniquely large bodied among primates to not have an adolescent spurt

Lifecycles

Later Homo erectus would have increased childhood and exhibits an adolescent spurt in both models
Possibility that the extended childhood is beginning to play a role in transmitting communication skills

Alternate Lifecycles

Pleistocene Lifecycles, 3
Early Homo erectus may have lived long enough for post-reproductive females to experience menopause--a first in mammalian evolutionary history A small fraction of later Homo erectus populations appears to have lived substantially past menopause for females and into old age for males
If relatively sophisticated communication systems existed at this point, the later life could have served a valuable adaptive purpose in passing on information about rare events (the once in a century drought or freeze) in the face of unprecedented climatic variability

Menopause

Late Pleistocene Happenings

The warm peak of the last interglacial (ca. 120,000 - 125,000 years ago) suspiciously coincides with the consensus date estimate of several genetic studies for the origin of modern Homo sapiens
The last refinements on language and improvements in technology may have occurred and been a substantial cultural advantage in this time of extreme climatic change The final expansion of childhood to allow the perfecting of language skills may have accompanied this change

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Consensus date of genetic models for appear ance of Moder n Homo sapiens

Recent Climates

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