Large-scale

evolutionary trends

Foraminiferal size, oxygen and

photosymbiosis
Outline
Copes rule
What are foraminifera?
Dramatic size increase in late Paleozoic
fusulinid foraminifera
Passive and driven evolutionary trends
Tests for analyzing the fusulinid size trend
Interpretation of results
Evolution of size

Copes rule:
o Tendency in animal groups to
evolve toward larger size
o First articulated in 1870s
o Size trends recognized in reptiles,
mammals, arthropods, mollusks
Copes rule: Traditional explanation
The largest size class is always unoccupied. Therefore,
over time the number of size classes will increase since the
one at the top is always open and available to be filled.

If extinction vacates organisms in a given

size class, others from adjacent size classes
absolute might increase or decrease in size in order to fill the void Theres
minimum always room
size at the top

Increasing size
What are foraminifera?
Living protists with fossil record dating back to
Cambrian Period (500 myr)
5,000 living species; >100,000 fossil species
Marine, brackish and freshwater
Benthic and planktonic (20% of total modern
carbonate production)
Most studied group of fossils
~ 3 cm
Foram sculpture park (China)
live foram assemblage
semelparous
reproduction
Fusulinid forams
Originated ~330 Ma; became extinct ~250 Ma
Very abundant & diverse; rock-building protists
Many lineages achieved gigantic size
 arrowhead made of silicified
fusulinid limestone

fusulinid limestones
Fusulinid forams

Large specimens can reach 16 mm in length and 8 mm in diameter

(volume = 500 mm3 surface area = 340 mm2)

Smallest specimen is 0.06 mm in length and 0.15 mm in diameter

(volume = 0.01 mm3 surface area = 0.04 mm2)
dramatic size evolution
in fusulinids
Passive vs. Driven trends

PASSIVE DRIVEN

No confining boundary;
increases more likely
Confining lower boundary; than decreases (implies
increases and decreases selection for large size)
equally likely
McShea 1994 Evolution
 Minimum test

PASSIVE DRIVEN

Minimum does not increase Minimum increases

McShea 1994 Evolution

Minimum test
suggests a
driven trend
Subclade test:
Size distribution of parent clade is nearly always right-skewed.

Subclade from the tail of the

parent clades distribution
is not skewed

Subclade from the tail of the

parent clades distribution
is right-skewed

McShea 1994 Evolution

Fusulinid size distribution

Volume (mm3)
Subclade test suggests a driven trend

parent clade
Quantifying passive and driven
components of large-scale trends

Wang (2001) recognized that large-scale trends are unlikely to

be entirely passive or entirely driven, but rather a combination
of both types
Analysis of skewness test determines the proportional
influence of passive and driven mechanisms: Sums of Cubes

SCtotal = SCbetween groups + SCwithin groups + SCheteroskedacticity

 indicates
passive trend

Each subclade exhibits a normal distribution, but

subclade means are not normally distributed
about the parent clade mean

Wang 2001 Evolution

 indicates
driven trend

Subclade means are normally distributed

about the parent clade mean, but each
subclade is right-skewed

Wang 2001 Evolution

 indicates
passive trend

Subclade means are normally distributed about the parent clade mean,
and each subclade is normally distributed, but standard deviation is
greater for subclades near right tail of parent clades distribution
Wang 2001 Evolution
Analysis of skewness
(fusulinid dataset)

Total skewness of fusulinoidean volume distribution as the sum of three components.

Category Value % Trend indicated

Skewness within subclades 204,892,468 67.01 driven

Skewness between subclades -295,643 -0.10 passive

Heteroskedasticity skewness 101,161,672 33.09 passive

Total skewness 305,758,497 100.00

Interpretation of results
Size trend in fusulinids is 2/3 driven and 1/3
passive
Driven component likely reflects selection for
large size
o Large size as a result of photosymbiosis
Passive component likely reflects relaxed
constraints on size
o Large size permitted by hyperoxia
Photosymbiosis in forams
Early suggestions of photosymbiosis in living
forams (1880s 1950s)
Lee et al. (1965) established first unequivocal
evidence for photosymbiosis in living forams
Photosymbionts now confirmed in 12 extant
families
o Symbionts include diatoms, dinoflagellates, unicellular
green algae, unicellular red algae and cyanobacteria
Photosymbionts in live
foram and coral

National Geographic

image courtesy of Pam Hallock

 foram with photosymbionts

image courtesy of Pam Hallock

Symbionts cultured from live foram

20 m

image courtesy of Scott Fay & Jere Lipps

Benefits of photosymbiosis
Energy
o Mixotrophic nutrition (feeding &
photosynthesis)
Calcification
o ATP energy for concentrating inorganic carbon;
removal of ions that inhibit calcification
Removal of host metabolites by symbionts
Characteristics of modern,
symbiont-bearing forams

Preference for tropical, oligotrophic habitats

o Stable environment; protected from continental and
seasonal influences
Unique life history strategy
o Large size (delayed reproductive maturity)
o Production of few, large embryons with low mortality
Giant embryons?

Fusulinid with
spherical adult shell
and elongate interior
Oxygen & size
Availability of oxygen constrains maximum
cell size

Surface
Volume 2/3
As the linear dimensions of an object increase by a factor of X, its
surface area increases by X2 while its volume increases by X3

2 Radius = 2
Surface = 50.3
Volume = 33.5

Radius = 1
Surface = 12.6 Four-fold increase in surface
Volume = 4.2 Eight-fold increase in volume
Late Paleozoic hyperoxia
Oxygen & size

size increase associated

with equally dramatic
increase in atmospheric
oxygen
Linear regression analysis

p = 0.0002
r2 = 0.41
Conclusions

Fusulinid size evolution was

mostly a driven response to
photosymbiosis (selection for large
size)

BUT, a significant part of the trend

was passive size increase in
response to increasing oxygen
availability (increase in the upper
bound to cell size)