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Only a theory: Why birds fly, why mammals stayed small, and why dinosaurs became big and died out Marcus Clauss Clinic for Zoo Animals, Exotic Pets and Wildlife, Vetsuisse Faculty, University of Zurich, Switzerland Science and Barbecue Zürich August 2012

Only a theory: Why birds fly, why mammals stayed small ...ffffffff-f77d-1dab... · The reproductive mode of dinosaurs (ovipary), with its intrisic limited juvenile start size linked

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Only a theory: Why birds fly, why mammals stayed small, and why dinosaurs became

big and died out

Marcus Clauss Clinic for Zoo Animals, Exotic Pets and Wildlife, Vetsuisse Faculty, University of

Zurich, Switzerland Science and Barbecue Zürich August 2012

... or how listening to interdisciplinary gossip, riding trains and drawing

pictures can lead to high attention-generating and completely non-

useful hypotheses.

DFG Research Group 533 Sauropod Biology - the Evolution of Gigantism

DFG Research Group 533 Sauropod Biology - the Evolution of Gigantism

Digestive physiology supervisors

Marcus Clauss Jürgen Hummel

DFG Research Group 533 Sauropod Biology - the Evolution of Gigantism

Digestive physiology supervisors

Digestive physiology PhD students & associates

Marcus Clauss Jürgen Hummel

Julia Fritz Ragna Franz Patrick Steuer

DFG Research Group 533 Sauropod Biology - the Evolution of Gigantism

(Digestive physiology) supervisors

Digestive physiology PhD students & associates

Marcus Clauss Jürgen Hummel

Julia Fritz Ragna Franz Patrick Steuer

Chris Carbone

DFG Research Group 533 Sauropod Biology - the Evolution of Gigantism

(Digestive physiology) supervisors

Digestive physiology PhD students & associates

Marcus Clauss Jürgen Hummel

Julia Fritz Ragna Franz Patrick Steuer Dennis Müller Daryl Codron Irina

Nurutdinova

Chris Carbone

DFG Research Group 533 Sauropod Biology - the Evolution of Gigantism

(Digestive physiology) supervisors

Digestive physiology PhD students & associates

Marcus Clauss Jürgen Hummel

Julia Fritz Ragna Franz Patrick Steuer Dennis Müller Daryl Codron Irina

Nurutdinova

Chris Carbone

Assumptions

- Niche stratification according to body size (=body mass) - Eggs cannot increase endlessly in size because of

physical constraints on egg shell thickness (stronger shells needed for larger eggs) and diffusion (thicker shells prevent diffusion of oxygen)

- The K-T extinction event affected all animals above a certain body size threshold

Niches of parent and offspring niche space

A

A - 1

A - 2

A - 3

adult

Niches of parent and offspring niche space

A

A - 1

A - 2

A - 3

adult

egg

Niches of parent and offspring niche space

A

A - 1

A - 2

A - 3

adult

egg

Niches of parent and offspring niche space

A

A - 1

A - 2

A - 3

adult

egg neonate

Niches of parent and offspring niche space

A

A - 1

A - 2

A - 3

adult

juvenile

egg neonate

Niches of parent and offspring niche space

A

A - 1

A - 2

A - 3

adult

juvenile

juvenile

egg neonate

Niches of parent and offspring niche space

adult A

A - 1

A - 2

A - 3

Niches of parent and offspring niche space

adult

foetus A

A - 1

A - 2

A - 3

Niches of parent and offspring niche space

adult

neonate

A

A - 1

A - 2

A - 3

Niches of parent and offspring niche space

adult

juvenile

A

A - 1

A - 2

A - 3

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

niche space Niches of parent and offspring

Resulting hypothesis I

The reproductive mode of dinosaurs (ovipary), with its intrisic limited juvenile start size linked to ontogenetic niche stratification, may have led to a parallel decrease of small dinosaur species with the increase of larger species, because offspring of large species fill the niche space of the smaller species.

A catastrophic event, truncating adult body size, could therefore, in theory, wipe out the whole reproductive dinosaur population.

In contrast, the reproductive mode of mammals (and also that of many birds due to parental care) does not lead to an ontogenetic niche stratification, but offspring use the parents’ niche.

Thus, with increasing body mass of larger species, niches of smaller species are not usurped by the larger species’ offspring to the same extent.

Die-offs of larger species would not leave niches of smaller life forms empty.

Resulting hypothesis II

Check species assemblages for distribution patterns (is there a ‘gap’)?

Design deterministic model to test effect of interspecific competition on body mass distribution of mammals and dinosaurs before and after a simulated K-T-event

Work plan

Check species assemblages for distribution patterns (is there a ‘gap’)?

Design deterministic model to test effect of interspecific competition on body mass distribution of mammals and dinosaurs before and after a simulated K-T-event

Work plan

Daryl Codron

Species distributions

Deterministic model

• Populations of different sized species of: Dinosaurs: i1,i2…ik = {2 g to ~131 tons} Mammals: i1,i2…ik = {2 g to ~16 tons}

 Each structured by size (log2M increments, j) from neonate-adult

Mneonate = aMadultb bdinosaurs ~0.6

bmammals ~0.9

 For each Mi,j, estimate: 1. initial abundance n (allometry) 2. survivorship p (r-selecting or K-selecting) 3. fecundity f (allometry; add fini,k to ni,1)

• Assume: each log2M step = ontogenetic niche shift

Deterministic model

• Density of each i after competition:

Ni = pi,j ni,j – α(Σ[pi,j nj] – pi,jni,j)

  Independent α for D on M and M on D:

Dinosaurs: NDi = Σ[nDi,j – αDD(ΣnDj–nDi,j) - αMD(ΣnMj)]

Mammals: NMi = Σ[nMi,j – αMM(ΣnMj – nMi,j) - αDM(ΣnDj)]

 Simulate K-T: kill all individuals >25 kg

Competition-induced mortality (α) occurs amongst all similarly-sized individuals (symmetric)

Model results - no competition

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.00

0.02

0.04

0.06 Dinosaurs

(non-avian) Aves Mammalia

αDD = 0.00αMM = 0.00αDM = 0.00αMD = 0.00

Model results - mild competition-induced mortality

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.00

0.02

0.04

0.06 Dinosaurs

(non-avian) Aves Mammalia

αDD = 0.1αMM = 0.1αDM = 0.00αMD = 0.00

Model results - stronger competition

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.00

0.02

0.04

0.06

0.08

0.10 Dinosaurs (non-avian)

Aves Mammalia

αDD = 0.15αMM = 0.15αDM = 0.00αMD = 0.00

Model results - including competition from dinosaurs on mammals

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.00

0.02

0.04

0.06

0.08

0.10

0.12

Dinosaurs (non-avian)

Aves Mammalia

αDD = 0.15αMM = 0.15αDM = 0.15αMD = 0.00

Model results - ... and from mammals on dinosaurs

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.00

0.02

0.04

0.06

0.08

0.10

0.12

0.14

0.16

0.18

0.20 Dinosaurs

(non-avian) Aves Mammalia

αDD = 0.15αMM = 0.15αDM = 0.15αMD = 0.15

Model results - ... and from mammals on dinosaurs

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.00

0.02

0.04

0.06

0.08

0.10

0.12

0.14

0.16

0.18

0.20 Dinosaurs

(non-avian) Aves Mammalia

αDD = 0.15αMM = 0.15αDM = 0.15αMD = 0.15

Model results - after K/T, with competition

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.00

0.02

0.04

0.06

0.08

0.10

0.12 Dinosaurs (non-avian)

Aves Mammalia

αDD = 0.15αMM = 0.15

Deterministic model

• Robustness and sensitivity  Changes and variability in M effects on:

1. life history strategy 2. reproductive output 3. Madult:Mneonate ratio 4. competition co-efficient (α) 5. symmetry of α 6. abundance

 Stochastic parameter estimates (bounded within empirical limits for birds/herpetiles or mammals), 103 iterations

Model results - variation in neonate scaling, demography, and ecology

pre K-T post K-T

log2Madult (kg)

relative abundance

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.050.100.150.200.25

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.020.040.060.080.100.120.14

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.040.080.120.160.20

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.020.040.060.080.100.12

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.020.040.060.080.10

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.020.040.060.080.100.12

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.050.100.150.200.25

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.020.040.060.080.100.12

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.100.200.300.400.50

-9 -7 -5 -3 -1 1 3 5 7 9 11 13 15 170.000.040.080.120.160.20

Dinosaurs (non-avian); Aves; Mammalia

variable adult-neonate mass scaling exponent variable adult-neonate mass scaling exponent

variable initial abundance variable initial abundance

variable abundance-mass scaling exponentvariable abundance-mass scaling exponent

variable fertility (birth rate) variable fertility (birth rate)

variable α variable α

Conclusions

Size-specific competition influences species diversity - Effects largest amongst species with high intraspecific

niche breadth (e.g. large, oviparous)…

- … favouring large-bodied taxa

Explains paucity of small-bodied dinosaurs - Dominance of large dinosaurs then limited max BM of

other vertebrate groups in Mesozoic…

Extinction event above body mass threshold = remaining species diversity too low for recovery

- … resultant dominance of small taxa amongst mammals further limited small dinosaurs (=aerial niche?)