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Allometry: the study of the relationship between size and shape
Galileo, 1636: Discorsi
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Allometry of Stegosaur plates
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Allometry in Tyrannosaurus forelegs
Adaptive explanations Allometry in combination with Cope’s Rule
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Surface/volume considerations and Allometry
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Assume a spherical cow . . . Surface area of sphere: 4p r 2 Volume of sphere: 4/3 p r 3
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Julian Huxley and Georges Tessier
Exponential form y=bx a Linear form: log(y) = a log(x)+log(b)
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Surface/volume and respiration in arthropods
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Problems of scaling to larger size
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Problems of scale at smaller size
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Surface/volume and locomotion
Surface area of wings vs mass Cross sectional surface area of legs vs mass
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The “Irish Elk” Actually a deer Lived across N. Europe and Asia
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But a very large deer . . .
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Antler span up to 12 feet
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Prospecting for fossil Megaloceras in Irish peat bogs
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Allometry in the white-tailed deer
Allometry can be measured within a species, usually by comparing juveniles to adults
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Allometry in Megaloceras
Allometry of antler size comparing adults of different species of deer “M” indicates the Irish Elk, Megaloceras
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Negative allometry of metabolic heat production on body weight
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Do we really have big brains?
Allometric curve has slope of 0.667, indicating the real relationship is between surface area of brain and size of animal Reptiles have the same slope, lower intercept on the y axis
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Primate brain weights plotted on the same field
The real measure of brain size is neither simple weight nor a ratio of brain to body weight, but deflection from the allometric curve
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Heterochrony: the evolution of allometry (rate) and timing in organismal development
Ernst Haeckel Met Darwin in 1866 and became defender of Natural Selection “Ontogeny recapitulates Phylogeny”
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Ontogeny and Phylogeny defined
Ontogeny: the developmental sequence of an individual organism Phylogeny: the evolutionary history of a species or lineage (phylogeny is to a species as genealogy is to an individual)
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Evolution in fish tails
Diphycercal tail in lungfish Protopterus
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Evolution in fish tails
Heterocercal tail in the sturgeon and other intermediate fish
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Evolution in fish tails
Heterocercal tail in modern teleostean fish (e.g. perch, bass) Note vestigial internal assymmetry of axial skeleton
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Ontogeny and phylogeny in fish tail evolution
A good example of recapitulation, consistent with Haeckel’s theory
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However . . . The Axolotl salamander: an inconvenient counter-example
Descendant (bottom) resembles juvenile of the ancestor (top)
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Synthesizing the literature on heterochrony
Paedomorphosis (juvenilization) Neoteny (K, slope, decreases) Progenesis (b, offset, moves left) Post-displacement (a , onset, moves right) Peramorphosis (“adultization”) Acceleration (K, slope, increases) Hypermorphosis (b, offset, moves right) Pre-displacement (a , onset, moves left)
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Neoteny in Mountain Sheep
Study by Geist compares ontogenies of male to female sheep
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Synthesizing the literature on heterochrony
Paedomorphosis (juvenilization) Neoteny (K, slope, decreases) Progenesis (b, offset, moves left) Post-displacement (a , onset, moves right) Peramorphosis (“adultization”) Acceleration (K, slope, increases) Hypermorphosis (b, offset, moves right) Pre-displacement (a , onset, moves left)
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Progenesis in Micromalthus, the gall midge
Wingless females reproduce and feed on mushrooms and other fungi
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r selection vs K selection
No=ert Small High juvenile mortality Unstable environments Large # offspring No parental care No=ert (K-N)/K Large Low juvenile mortality Stable environments Small # offspring Parental care
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Synthesizing the literature on heterochrony
Paedomorphosis (juvenilization) Neoteny (K, slope, decreases) Progenesis (b, offset, moves left) Post-displacement (a , onset, moves right) Peramorphosis (“adultization”) Acceleration (K, slope, increases) Hypermorphosis (b, offset, moves right) Pre-displacement (a , onset, moves left)
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Post-displacement in the Dachshund
Legs begin developing late and never catch up
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Astogeny (colonial development) recapitulates phylogeny?
Earliest colonial forms (analogous to juveniles) carried forward into later astogeny
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Synthesizing the literature on heterochrony
Paedomorphosis (juvenilization) Neoteny (K, slope, decreases) Progenesis (b, offset, moves left) Post-displacement (a , onset, moves right) Peramorphosis (“adultization”) Acceleration (K, slope, increases) Hypermorphosis (b, offset, moves right) Pre-displacement (a , onset, moves left)
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Hypermorphosis in the Irish Elk
Postponing the offset of development leads to a larger adult – one way of achieving Cope’s Rule of phyletic size increase
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Acceleration in ammonite phylogeny
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Heterochrony in Paleozoic zaphrentid corals
Note adult shape compressed back into early ontogeny, resulting in recapitulation in descendant Acceleration of development
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Human allometry
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Primate allometric patterns
Positive allometry of jaw, brow ridge on skull Negative allometry of cranial vault on skull
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Human skull allometry Follows same allometric pattern as primate ancestors, but slowed down (neotenic)
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Allometry in Pan troglodites
Note: upright posture Vaulted cranium Flattened alveolar Region Less pronounced Brow ridge Of juvenile
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Primate allometry of birth weight
Humans retain rapid fetal growth rates longer, hence a relatively larger fetus
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Effect of neoteny on brain weight
Humans retain rapid fetal growth rate of brain longer, shifting flexure in brain-body curve to right, resulting in larger brains
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Allometry in toy dogs: King Charles Spaniel (top)
Irish Wolfhound (bottom) Note negative allometry of cranial cavity and positive allometry of prognathous jaw – both standard mammalian ontogenetic patterns
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Paedomorphosis in foraminifera: Paleogene genus Morozovella
Juvenile form angulose (visible in lower cross section), carried forward to adult morphology in descendant
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Ecological heterochrony?
Note steepening curve showing increasing angularity (left) Increasing isotopic difference between juvenile and adult in descendant (right)
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Adult vs juvenile morphology in Mickey
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Paedomorphosis in Mickey
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Comparative embryology and heterochrony
Early embryonic stages are comparable Differentiation is increasingly expressed later in ontogeny
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Allometry in birth weight
Birth weight compared to gestation time in mammals
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