CBO9780511793844.005 Fossil Primates - The World of The Past

Cambridge Books Online
http://ebooks.cambridge.org/
Fossil Primates
Susan Cachel
Book DOI: http://dx.doi.org/10.1017/CBO9780511793844
Online ISBN: 9780511793844
Hardback ISBN: 9781107005303
Paperback ISBN: 9780521183024
Chapter
4 - The world of the past pp. 59-65
Chapter DOI: http://dx.doi.org/10.1017/CBO9780511793844.005
Cambridge University Press
The world of the past
The origin of continents and oceans

The title of this section is a translation of a book rst written by the meteorologist Alfred Wegener in 1915, and revised and translated many times afterwards.
Wegener proposed that a giant supercontinent (Pangaea) had once existed.
Portions of this supercontinent split apart and moved away, creating ocean
basins and the modern geography of the earth. This hypothesis was called
continental drift. However, although Wegener amassed data on the similarity
of continental coastlines (especially South America and South Africa), geological strata, and animal and plant distributions to support his hypothesis, he
had no workable mechanism to explain the movements of the fragmented
supercontinent. Most geologists considered Wegener to be an eccentric crank,
although biologists appreciated the fact that continental drift eliminated the
necessity for invoking land bridges to explain animal and plant distributions;
and biologists further applauded the fact that continental drift rendered massive
parallel evolution unnecessary. Wegeners hypothesis was resurrected in the
1960s, which witnessed a revolution in geological thought. Using new evidence
about earthquake and volcanic belts, paleomagnetism, and sea-oor spreading,
geologists outlined a new mechanism of continental movement called plate
tectonics.
Movements (tectonics) affect plates of lithosphere. Convection currents deep
within the mantle of the earth are the ultimate cause of these movements. These
convection currents are driven by heat. Radioactive materials exist within the
mantle. Their natural decay processes emit heat, and thus a molten iron core lies
underneath the mantle. Movements within the mantle drag the overlying crust,
and upwelling mantle plumes create local hotspots of volcanic activity, as
observed in the Hawaiian Islands. These deep mantle plumes and hotspots can
be tracked by volcanic islands and submarine sea mounts as plates pass over the
plumes. These plumes can also affect the speed of plate movements and the
intensity of volcanic eruptions. For example, the Runion plume in the Indian
Ocean is responsible for the simultaneous rapid motion of the Indian plate and
slowing of the African plate beginning 67 mya (Cande & Stegman, 2011). The
enormous Indian Deccan ood basalts are adjacent to the Runion plume, and
begin erupting at the same time. These massive volcanic eruptions contribute to
the major extinctions that occur at the K/T boundary.
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60
Tectonic plates can separate from each other, slide past each other, or converge.
Plates separate from each other when new lava is extruded at their boundaries.
This can be observed on land today in the Afar Triangle of Ethiopia. When plates
separate at the central oceanic ridges, sea-oor spreading occurs, and ocean
basins are formed. The denser lava sinks, and the lighter continental crust sits
high on the surface of a plate. Most of the ocean oor is young. About 50 percent
of it is no older than 65 my, the length of primate evolution. We primates are
equal in age with much of the ocean oor! The rate of sea-oor spread can be
quick (6 cm per year at the San Andreas Fault, which is sliding part of California
north past the North American Plate), moderate (3 cm per year at the South
Atlantic Ridge), or catastrophic. The entire island of Sumatra was shifted by
10 meters during the colossal magnitude 9.3 earthquake of December 26, 2004.
The fault line created by this earthquake extended 1,500 km, the surface of the
entire planet was raised and lowered by at least 1 cm, and the earth rang for weeks
with free oscillations caused by the earthquakes seismic forces. Plate tectonic
movements can therefore sometimes be quick and violent. When plates converge,
a plate that carries oceanic crust subducts or descends into the earths mantle, and
creates a series of volcanic islandsan island arcas the descending plate deforms
the mantle. When plates carrying continental crust collide, the lithosphere is
driven upward, and mountains are formed. The Andes Mountains and the
Himalayas result from this type of plate movement.
The movement and dispersal of land animals is obviously affected by the
opening of water gaps and ocean basins. G. G. Simpson (1940a) rst examined
the nature of land mammal dispersal by showing the decreasing likelihood of
land mammals moving through land bridges or corridors, passing climatic or
geographic lters, and crossing substantial water gaps in a sweepstakes fashion
(Chapters 10 and 13). To this list of decreasingly less probable movement
(corridors, lters, and sweepstakes dispersal), McKenna (1973) added two new
elements that were made possible by plate tectonic theory: Noahs Arks and
Beached Viking Funeral Ships. In the Noahs Ark scenario, a lithospheric plate
moves and bears a cargo of living land animals. In the Beached Viking Funeral
Ships scenario, a moving plate bears fossils. If the plate eventually contacts
another plate, the fossils are now xed to another continent. However, these
fossils are immigrants, and have no biogeographic relationship to fossils or
living animals found in the new land. Because India was a Noahs Ark for much of
the Tertiary, and did not contact Asia until the Miocene, primate evolution and
dispersal in India and Asia is affected by plate tectonic theory.
Climates of the past

Detailed climatic records for over 150 years illustrate how mammals react to
climatic change. Different species respond in different ways (Blois & Hadly,
2009). At the local level, seasonal changes might inuence diet, mating, home
range size, migration, or hibernation. Increasing seasonality might lead to shifts in
Climates of the past
diet, habitat, and geographic range. At the regional level, differences between
populations could lead to divergence and speciation. Prolonged climatic change is
likely to be associated with immigration, speciation, or extinction of species.
Up until about 34 mya, the climate of the earth was distinctly warmer than
at present. Mean global temperatures were much higher, and there was no ice,
or virtually no ice, at the poles. The atmosphere of the earth had higher
concentrations of greenhouse gasses. Volcanic eruptions were frequent from
about 60 to 40 mya, which partly accounts for this higher concentration.
Methane emissions from shallow ocean sediments bubble up to the atmosphere
and also affect greenhouse gas concentrations (Chapter 8). On the other hand,
beginning at 34 mya, Antarctic ice becomes permanent. Ephemeral ice sheets
appear in the northern hemisphere at 10 mya, during the Late Miocene.
Northern ice sheets become permanent, and reach a continental scale at about
3 mya. The growth and recession of the great continental ice sheets has been a
focus of intense study. Geologists have been debating the cause of the mysteriously increasing and diminishing northern ice sheets for nearly 170 years
(Ramo & Huybers, 2008). Major plate tectonic events clearly set the stage. Land
surfaces sitting over the poles, or polar ocean surrounded by land, allow ice to
build up in high latitudes. Land surfaces that restrict oceanic circulation are
also important. With the rise of the Isthmus of Panama at 3.5 mya, warm
equatorial water could no longer ow around the entire globe. This appears to
have been the nal trigger for the advent of permanent ice sheets in the
northern hemisphere.
It has been difcult to ne-tune our understanding of the activity of these
northern ice sheets. Besides the difculties of dating their presence and extent,
models that explain their waxing and waning are proving difcult to understand.
From about 3 to 1 mya, glacial cycles were regular, and lasted about 41,000 yrs
(Ramo & Huybers, 2008). This matches the hypothesis of Milutin Milankovi, who
argued in the 1930s that variations in the earths orbit are responsible for glaciation. When the northern hemisphere receives less solar radiation during the
summer, snow lasts throughout the year, and gradually builds up into continental
ice sheets. Known astronomical variations led Milankovi to predict a 41,000 yr
glacial cycle. Beginning at about 1 mya, and continuing to the present, Ice Age
climate appears to show 100,000 yr cycles (Ramo & Huybers, 2008:Fig. 1).
Questions about solar radiation, the coupling or decoupling of northern and
southern hemisphere glaciation, the geographic extent of early ice sheets, and
the thickness and ablation of ice sheets are currently being debated. In particular,
climatologists are still uncertain about how changes in solar radiation that affect
the earths upper atmosphere are translated into drastic variations in ice volume at
ground level. Hypotheses about the relative length of ice growth and melt seasons
are now being tested.
For the Cenozoic, at least, the paleoclimatic record is detailed enough to show
that the carbon cycle is intimately linked to climate. At the end of the nineteenth
century, the American geologist Thomas C. Chamberlain hypothesized that
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uctuations in the carbon cycle were the trigger for global ice ages. He argued that
the weathering of high mountain ranges and the reduction of oceanic calcium
carbonate would reduce atmospheric CO2 levels, causing lower global temperatures. Low rock weathering rates and high oceanic carbonate production would
increase atmospheric CO2, causing higher global temperatures. But carbon dioxide
levels in the atmosphere uctuate in a more complex way. These levels are
affected both by inuxes from volcanic out-gassing and by the sequestering or
release of organic carbon. Changes in continental ice volume are strongly correlated to the amount of CO2 in the atmosphere. As the partial pressure of atmospheric CO2 changes, paleobotanical data also change to reect either tropical or
temperate shifts. Atmospheric carbon dioxide thus affects climate, but other
greenhouse gasses also force climate change. Atmospheric methane has a greater
effect than CO2 does. Eruptions of methane into the atmosphere from sea-oor
sediments were apparently responsible for the terric rapid upward spike of global
temperature at the Paleocene/Eocene boundary.
Marine sediments are currently emitting oil and gasses, including methane. The
rate of this emission is not constant. A number of factors affect the rate of
hydrocarbon ooze. During periods of Pleistocene deglaciation, tar deposits occur
in sediments off the California coast, indicating greater seepage of hydrocarbons
as the climate ameliorated (Hill et al., 2006). Incorporation of methane into the
atmosphere would further intensify the natural climatic warming.
Much of the ne-grained analysis of climate change in the Late Cenozoic is
conducted through geochemistry. Measurements are made of the ratio of heavy to
light oxygen extracted from sea-oor sediments and terrestrial ice cores. Reconstructing uctuations between cold and warm global climate is based on these
oxygen isotope ratios.
Habitat reconstruction
It is not enough to outline the broad details of geography or climate. When one
considers the life of the past, one desires knowledge of vegetation, precipitation,
and seasonality before limning in the fossil animal species of an ancient
community. Pollen has an outer surface that is not only extremely durable, but
also has a characteristic morphology for genera. Pollen can sometimes be
retrieved from sites. Impressions of leaves, fruits, owers, stems, and sometimes
whole tree trunks of fossil woodmacrobotanical remainscan be recovered from
sites. Thus, one can gain some understanding of plant species that were present at
fossil sites. By measuring the topography of leaf edges that occur in different
climates today, paleobotanists have established a link between leaf edge structure
and mean annual temperature. The simpler the leaf margin (e.g. palm leaves), the
higher the temperature. Complicated leaf margins (e.g. oak leaves) occur in more
temperate conditions. Empirical regression lines between the degree of leaf evagination (i.e. the total length of leaf margin) and mean annual temperature are
generated from living plant species (Wolfe, 1978, 1993). The margins of fossil leaf
assemblages retrieved from ancient sediments are then measured, and ancient
temperatures are inferred from the results.
Many Cenozoic sites have plant genera and species that are alive today, and this
allows botanists to reconstruct an ancient habitat in some detail. Often, the plant
taxa are out of place. For example, tropical trees occur in what are now desolate
badlands in the American West during the Paleocene and Eocene. These ancient
trees occur in what is obviously a very different habitat from the modern landscape. Sometimes taxa occur in suites that are not observed in modern times. This
situation is more difcult to interpret, because it implies that community structure
and species interactions were different in the past. These comments also apply to
animal genera and species. A thorny complication in both cases takes place when
an ancient taxon has no living representatives, which occurs with increasing
frequency as sites become more ancient. The solution for enigmatic plant and
animal taxa is to investigate their anatomy in detail in an attempt to infer their
lifeways. For example, does the plant have woody tissue? What is the bone density
and cross-sectional area of an animal limb? Chapter 5 will highlight ways in
which the lifeways of extinct animals can be inferred.
Ancient soils (paleosols) can sometimes be recovered from sites. Detailed knowledge of the temperature and rainfall regimes that create similar soils in the
modern world allows specialists to reconstruct ancient habitats based on the depth
and nature of ancient soil horizons. Miocene ape habitats in Africa and southern
Asia have been reconstructed using paleosol data (Retallack, 1991), and habitats
dating back to the Mesozoic are routinely studied using paleosols. Nevertheless,
one would like even ner detailan impression of the degree of open country or
wooded area at a site. Is this possible? Yes. Correlating the carbon isotopes from
thousands of samples of modern soil associated with satellite photographs of
vegetation in the same area allows one to discover the relationship between
modern soil temperature and ground cover. Carbon isotopes retrieved from
ancient soils (paleosols) can then be used to reconstruct ancient soil temperatures
and infer the degree of shade or tree cover that existed at an ancient site (Cerling
et al., 2011; Feibel, 2011). Quantication of shade allows modern biologists to
categorize habitats as grasslands, open woodlands, or dense forests. These paleosol researchers use a worldwide database of modern sites to dene grasslands as
having less than 40 percent tree cover; woodlands as having greater than 40 percent tree cover; and forests as having greater than 80 percent tree cover. Examining sites in East Africa back to 7 my reveals that most of the sites with fossil
humans have less than 40 percent canopy cover (Cerling et al., 2011). They are
thus dened as grasslands, which has important implications both for human
evolution and for the evolution of Old World monkeys that are sympatric with
humans at these sites (Chapter 15).
When coexisting or sympatric species are considered together, paleontologists
explore the way that ecosystems function. In the modern world, a number of
factors are known to affect the structure and function of ecosystems (Figure 4.1).
Some of these operate at a local level; some of these operate at a continental level.
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Figure 4.1. The relative ranking of factors that affect ecosystem structure and processes.
Figure 4.2. Ecology versus geography as a determinant of community assembly. Evidence
indicates that a true equilibrium state (stasis) does not exist. That is, if perturbations occur,
the community does not return to its initial state. At any given time, a community is an
assemblage of species that are accreting or being removed in a partly random fashion.
Many researchers study living communities, and lament species loss within these
communities. Ecological and geographical factors are known to affect species
composition within communities (Figure 4.2). Comparison between communities
can yield insights into the way that natural selection or random events affect
evolution. Similarity between communities in the percentage of species present
can be caused by a resemblance in available niches or by the dispersal abilities of
taxa. However, community composition in the modern world appears to be very
plastic or labile. Native species go extinct in a local area, and are easily dislodged
or replaced by invasive species. If perturbations occur, the community does not
return to its initial state. Evidence indicates that a true equilibrium state (stasis)
does not exist. At any given time, a community is an assemblage of species that
are accreting or being removed in a partly random fashion. An ecological niche
does not need to be occupied by any given, single species, but by similar species
with similar nichesmembers of a guild. The same processes occur in the fossil
record. Paleontologists have falsied the idea of coordinated stasis. Coordinated
stasis is the idea that a network of species interactions is so strong that it resists
change. New or invasive species therefore cannot penetrate an existing ecosystem.
No evidence for this has been found in the fossil record.
What affects primate presence in a community? This is partly determined by
how primates disperse. They cannot y and they cannot swim. Another factor
limiting primates is their nearly universal reliance on tropical or subtropical forest
habitats. Tropical or low-latitude animals are much more sensitive to changes in
the physical environment than temperate animals are. This is known as Rapaports
rule (Stevens, 1989; Cachel, 2006). Primate sensitivity to climatic or habitat
change quickly removes them from an ecosystem. When living primates are
studied, there is no evidence that they experience competition from sympatric
primates. This may be caused by recent extinction events since the Late Miocene
that removed many primate species. Primate survivors have a wealth of food and
space that fossil primatesshoehorned into species-rich ancient communitiesdid
not have. If living primates experience competition, it may come from frugivorous
birds and bats or other arboreal mammals, such as squirrels or the small arboreal
marsupials of South America.
65

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Cambridge Books Online

The world of the past

The origin of continents and oceans

The world of the past

Climates of the past

Climates of the past

The world of the past

The world of the past

Figure 4.2. Ecology versus geography as a determinant of community assembly. Evidence

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