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HOW RAPIDLY DID MARINE
REPTILES DIVERSIFY AFTER THE
PERMO-TRIASSIC MASS
EXTINCTION?
THAMASHA PERERA
The Permo-Triassic Mass
Extinction (PTME)
 252 MYA
 Earth’s most severe extinction
event
 A.K.A “The Great Dying”…
 Gradual environmental change
 Catastrophic event
 Devastated marine ecosystems
(80  96% marine spp. extinct)
 Resulted in evolutionary
opportunities
Source: Wikipedia
Origin of marine reptiles
Benton, 2015
Recovery dynamics – fast or
slow?
 1) Stepwise recovery of life – slow and gradual
 - delayed by biotic drivers
 - physical perturbations (high temps/CO2/acid rain) during Induan
to Anisian (first 6MY)
 - the addition of progressively higher trophic levels
 - spanning some 8-9 Myr well into Middle Triassic
 2) Rapid recovery
 - environmental inhibition not so significant
 - suggested burst of diversification in Anisian (Mid Triassic)
Measures of diversity/trends
of change
1) Counts of major clades
2) Feeding guilds
3) Morphology
1. Counts of major clades
Chinese marine reptile Lagerstatten
richest fauna
most continuous over long span of time
Key vertebrate biotas occur in 3
formations
International Union of Geological Sciences, 2009
Guanling formation (Luoping and Panxian biota)
Zhuganpo formation (Xingyi biota)
Xiowa formation (Guanling biota)
Early Triassic (fossiliferous)
 Chaohu area, famous reptile fossil assemblage
 Late Spathian
 Basal ichthyopterygian present
 Chaohusarus geishanensis, <1 or 2m length
 Sauropterygians
 Hupehsuchia
 9 valid taxa reported
Source: Wikipedia
Clades in Triassic
CLADE: Sauropterygians Ichthyopterygians Thalattosaurs Hupehsuchians
EARLY
TRIASSIC
(Chaohu)
NO YES NO YES
EARLY MID
TRIASSIC
(Guanling)
YES YES YES NO
LATE MID
TRIASSIC
(Zhuganpo)
YES NO YES NO
LATE TRIASSIC
(Xiowa)
NO YES YES
(dominant)
NO
 A recent summary of Chinese Triassic marine reptiles implies early
diversification throughout much of the early Triassic and even close to
the PT boundary
However…
 Sampling bias possible
 Artificial diversity boosts
 Low taxon numbers – truly represent success?
 NO fossil Lagerstatten from Early Triassic
2. Feeding guilds
 Unique feeding adaptations - Early Triassic
 1) Feeding habitat – pelagic or demersal
 2) Prey capture – ram with biting, lunge, suction
 3) Tooth shape – pointed, rounded, filter, edentulous
 Use these 3 basic features to capture any large scale
patterns in the evolution of feeding guilds
Feeding guilds
 Examples: looking at variation in feeding styles of -
 Hupesuchians - Palate and mandible suggest – rare
lunge feeder - first in history of life
 Spathian (Early Triassic)
 Ichthyosauria (Omphalosaurus)
 Rounded teeth, rostral jaw, edentulous
 Spathian (Olenekian, Early Triassic)
 Sauropterygian (Atopodentatus)
 Filter feeder
 Anisian (early middle Triassic)
Source: http://alphynix.tumblr.com/image/120459753490
Feeding guilds
 Highest variation in marine reptile feeding
strategy reached in Spathian (Early
Triassic), indicates a rapid diversification
of prey preference in less than 1 MY
 Burst in early Triassic
However….
 Minor lineages of marine reptile - poorly understood –
conceal true diversity of feeding styles
 Incompleteness of fossil record – inherently biased
3. Morphology
 Unusually large ichthyosaur humerus about 28cm length recovered
from Spathian (Early Triassic) –humerus length correlated to overall
body length
 C. geishanensis of Chaohu was max 2m length
 Humerus – shows that Early Triassic ichthyosaur record ALREADY quite
diverse and large ichthyosaurs were already widespread during the
Spathian
 NO significant size increase of predators observable from Early Triassic
to the Anisian
But…
 Accurate age dating of these Early Triassic fossils
remains difficult – depending on the definition of the
Olenekian-Anisian boundary (Late Early to Early Mid
Triassic)
Further evidence –
Thalattoarchon saurophagis
 Ichthyosaur – large - 8.6m or over
 12 cm tooth, large bicatinate cutting,
macropredatory
 Discovered Nevada - indicates early and
middle-Triassic ichthyosaur radiation 
this large macrophagous apex predator
as early as Anisian (mid Triassic)
 Modern level complexity reached within
8 MY after PTME and within 4 MY of
when reptiles first invaded the sea
 Top tier – indicates rest of ecosystem
had recovered by its appearance, no
gradual build up of forms to get to it Source: National Geographic
But…
 Analyses of rich fossil Lagerstatten in S. China ALSO
show evolution of diverse marine reptile faunas by
ANISIAN (early middle Triassic) - BUT large and
macrophagous apex predator is unknown there
 Could be peculiarity of record in China
Why not slow?
 Pattern - feeding guilds - does not support slow stepwise
 Delayed recovery could reflect poor sampling in the Early
Triassic – not true absence
 Already by the Induan had several trophic levels for global
marine ecosystem present - Marine trophic pyramid was
NOT truncated in early Triassic
 Prolonged stepwise recovery pattern of marine ecosystems
after PTME needs reconsideration
Summary – how rapid was marine
reptile diversification?
 Clade diversity –peak already reached in Anisian (Early Mid Triassic)
 Feeding guild diversification – RAPID – highest variation- Spathian –
EARLY TRIASSIC
 morphological disparity - suggests this 1st radiation already diversified
broadly to a variety of trophic guilds (like feeding guilds)
 Discovery of Thalattoarchon shows full ecosystem recovery early on in
Triassic
 High degree of adaptation to aquatic environment of marine reptiles,
infer that they evolved during EARLIER stages of Early Triassic e.g.
Induan
 Likely follows rapid recovery model for diversification
References
Benton MJ, Zhang Q, Hu S, Chen Z-Q, Wen W, et al. (2013) Exceptional
vertebrate biotas from the Triassic of China, and the expansion of marine
ecosystems after the Permo-Triassic mass extinction. Earth Sci Rev 125: 199–
243
Chen, Z-Q and Benton, M.J. (2012) The timing and pattern of biotic recovery
folllowing the end-Permian mass extinction. Nature Geoscience. 5:375-383
Fröbisch, N. B., Fröbisch, J., Sander, P. M., Schmitz, L., & Rieppel, O. 2013.
Macropredatory ichthyosaur from the Middle Triassic and the origin of
modern trophic networks. Proceedings of the National Academy of
Sciences, 110: 1393-1397
Motani, R., Chen, X. H., Jiang, D. Y., Cheng, L., Tintori, A., & Rieppel, O.
(2015). Lunge feeding in early marine reptiles and fast evolution of marine
tetrapod feeding guilds. Scientific reports, 5.
Scheyer, T. M., Romano, C., Jenks, J., & Bucher, H. 2014. Early Triassic Marine
Biotic Recovery: The Predators' Perspective. PloS one. 9: 3.

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FINAL-Current-controv-Marine-reptiles-presentation-DONE

  • 1. HOW RAPIDLY DID MARINE REPTILES DIVERSIFY AFTER THE PERMO-TRIASSIC MASS EXTINCTION? THAMASHA PERERA
  • 2. The Permo-Triassic Mass Extinction (PTME)  252 MYA  Earth’s most severe extinction event  A.K.A “The Great Dying”…  Gradual environmental change  Catastrophic event  Devastated marine ecosystems (80  96% marine spp. extinct)  Resulted in evolutionary opportunities Source: Wikipedia
  • 3. Origin of marine reptiles Benton, 2015
  • 4. Recovery dynamics – fast or slow?  1) Stepwise recovery of life – slow and gradual  - delayed by biotic drivers  - physical perturbations (high temps/CO2/acid rain) during Induan to Anisian (first 6MY)  - the addition of progressively higher trophic levels  - spanning some 8-9 Myr well into Middle Triassic  2) Rapid recovery  - environmental inhibition not so significant  - suggested burst of diversification in Anisian (Mid Triassic)
  • 5. Measures of diversity/trends of change 1) Counts of major clades 2) Feeding guilds 3) Morphology
  • 6. 1. Counts of major clades Chinese marine reptile Lagerstatten richest fauna most continuous over long span of time Key vertebrate biotas occur in 3 formations
  • 7. International Union of Geological Sciences, 2009 Guanling formation (Luoping and Panxian biota) Zhuganpo formation (Xingyi biota) Xiowa formation (Guanling biota)
  • 8. Early Triassic (fossiliferous)  Chaohu area, famous reptile fossil assemblage  Late Spathian  Basal ichthyopterygian present  Chaohusarus geishanensis, <1 or 2m length  Sauropterygians  Hupehsuchia  9 valid taxa reported Source: Wikipedia
  • 9. Clades in Triassic CLADE: Sauropterygians Ichthyopterygians Thalattosaurs Hupehsuchians EARLY TRIASSIC (Chaohu) NO YES NO YES EARLY MID TRIASSIC (Guanling) YES YES YES NO LATE MID TRIASSIC (Zhuganpo) YES NO YES NO LATE TRIASSIC (Xiowa) NO YES YES (dominant) NO
  • 10.  A recent summary of Chinese Triassic marine reptiles implies early diversification throughout much of the early Triassic and even close to the PT boundary
  • 11. However…  Sampling bias possible  Artificial diversity boosts  Low taxon numbers – truly represent success?  NO fossil Lagerstatten from Early Triassic
  • 12. 2. Feeding guilds  Unique feeding adaptations - Early Triassic  1) Feeding habitat – pelagic or demersal  2) Prey capture – ram with biting, lunge, suction  3) Tooth shape – pointed, rounded, filter, edentulous  Use these 3 basic features to capture any large scale patterns in the evolution of feeding guilds
  • 13. Feeding guilds  Examples: looking at variation in feeding styles of -  Hupesuchians - Palate and mandible suggest – rare lunge feeder - first in history of life  Spathian (Early Triassic)  Ichthyosauria (Omphalosaurus)  Rounded teeth, rostral jaw, edentulous  Spathian (Olenekian, Early Triassic)  Sauropterygian (Atopodentatus)  Filter feeder  Anisian (early middle Triassic) Source: http://alphynix.tumblr.com/image/120459753490
  • 14. Feeding guilds  Highest variation in marine reptile feeding strategy reached in Spathian (Early Triassic), indicates a rapid diversification of prey preference in less than 1 MY  Burst in early Triassic
  • 15. However….  Minor lineages of marine reptile - poorly understood – conceal true diversity of feeding styles  Incompleteness of fossil record – inherently biased
  • 16. 3. Morphology  Unusually large ichthyosaur humerus about 28cm length recovered from Spathian (Early Triassic) –humerus length correlated to overall body length  C. geishanensis of Chaohu was max 2m length  Humerus – shows that Early Triassic ichthyosaur record ALREADY quite diverse and large ichthyosaurs were already widespread during the Spathian  NO significant size increase of predators observable from Early Triassic to the Anisian
  • 17. But…  Accurate age dating of these Early Triassic fossils remains difficult – depending on the definition of the Olenekian-Anisian boundary (Late Early to Early Mid Triassic)
  • 18. Further evidence – Thalattoarchon saurophagis  Ichthyosaur – large - 8.6m or over  12 cm tooth, large bicatinate cutting, macropredatory  Discovered Nevada - indicates early and middle-Triassic ichthyosaur radiation  this large macrophagous apex predator as early as Anisian (mid Triassic)  Modern level complexity reached within 8 MY after PTME and within 4 MY of when reptiles first invaded the sea  Top tier – indicates rest of ecosystem had recovered by its appearance, no gradual build up of forms to get to it Source: National Geographic
  • 19. But…  Analyses of rich fossil Lagerstatten in S. China ALSO show evolution of diverse marine reptile faunas by ANISIAN (early middle Triassic) - BUT large and macrophagous apex predator is unknown there  Could be peculiarity of record in China
  • 20. Why not slow?  Pattern - feeding guilds - does not support slow stepwise  Delayed recovery could reflect poor sampling in the Early Triassic – not true absence  Already by the Induan had several trophic levels for global marine ecosystem present - Marine trophic pyramid was NOT truncated in early Triassic  Prolonged stepwise recovery pattern of marine ecosystems after PTME needs reconsideration
  • 21. Summary – how rapid was marine reptile diversification?  Clade diversity –peak already reached in Anisian (Early Mid Triassic)  Feeding guild diversification – RAPID – highest variation- Spathian – EARLY TRIASSIC  morphological disparity - suggests this 1st radiation already diversified broadly to a variety of trophic guilds (like feeding guilds)  Discovery of Thalattoarchon shows full ecosystem recovery early on in Triassic  High degree of adaptation to aquatic environment of marine reptiles, infer that they evolved during EARLIER stages of Early Triassic e.g. Induan  Likely follows rapid recovery model for diversification
  • 22. References Benton MJ, Zhang Q, Hu S, Chen Z-Q, Wen W, et al. (2013) Exceptional vertebrate biotas from the Triassic of China, and the expansion of marine ecosystems after the Permo-Triassic mass extinction. Earth Sci Rev 125: 199– 243 Chen, Z-Q and Benton, M.J. (2012) The timing and pattern of biotic recovery folllowing the end-Permian mass extinction. Nature Geoscience. 5:375-383 Fröbisch, N. B., Fröbisch, J., Sander, P. M., Schmitz, L., & Rieppel, O. 2013. Macropredatory ichthyosaur from the Middle Triassic and the origin of modern trophic networks. Proceedings of the National Academy of Sciences, 110: 1393-1397 Motani, R., Chen, X. H., Jiang, D. Y., Cheng, L., Tintori, A., & Rieppel, O. (2015). Lunge feeding in early marine reptiles and fast evolution of marine tetrapod feeding guilds. Scientific reports, 5. Scheyer, T. M., Romano, C., Jenks, J., & Bucher, H. 2014. Early Triassic Marine Biotic Recovery: The Predators' Perspective. PloS one. 9: 3.