How did fins evolve? How did limbs evolve? Fin Fold Hypothesis for ...

How did fins evolve? How did limbs evolve?
Fossil “Agnathan”
Petraspis
Gnathostomes: mobile, paired pectoral & pelvic fins
Fossil Placodermi:
Dunkleosteus
Kardong Fig. 9.5
Fossil Acanthodii: Climatius
Fossil Chondrichthyes: Cladoselache
Lecture 5, Exam 2, Biol. 453 - Comp. Vert. Anat.,
Autumn 2012; Read in Kardong Ch. 9. pg. 325-350.
“Archipterygium” Hypothesis for Paired Fin Evolution
Towers & Tickle, 2009
Apical ectodermal ridges (AER) define
distal edge of gill arches, fins & limbs:
Shared regulatory genes:
retinoic acid (RA), sonic hedgehog
(Shh) & fibroblast growth factor (Fgf8)
Model to convert gill arch into a fin
Skate Gill arch (I) Skate fin (J) Chick limb (K)
Gillis, Dahn, & Shubin, 2009
Fin Fold Hypothesis for Paired Fin Evolution
Hypothetical ancestral vertebrate with
paired, ventrolateral fin folds
Scapulocoracoid
Ischiopubic
Serial homology in fin design
Which HOX genes are expressed in the
pectoral & pelvic fins/limbs?
Chick
Pectoral
Chick
Pelvic
Compare the fins of the Chondrichthyes with other fishes
Compare with Kardong Fig. 9.10, 9.11, 9.13
Fossil Shark:
Hybodus
White-tip
Reef Shark
http://www.arkive.org/whitetip-reef-shark/triaenodon-obesus/video-11.html

How were fins modified in Actinopterygians?
Actinopterygians
Ischiopubic
http://www.arkive.org/threadfin-butterflyfish/chaetodon-auriga/video-00.html
When did the first tetrapods appear?
Niedz´wiedzki, et al. 2010;
Janvier & Clément. 2010.
Scale bars = 10 mm.
Tracks from marine
shoreline, Poland.
Holland, 2012
A recent phylogeny of the “tetrapodomorphs”
Tetrapodomorph fossil “fish”
phylogeny based on 81
morphological characters.
How did pterygiophores alter in the evolution of tetrapods?
Coelacanth Tetrapodomorphs Dipnoan
Asymmetric
radials
Uniserial Fins
Raff, 2007

How do pectoral girdles differ in fish & tetrapods?
Sarcopterygian Fossil Tetrapod Early Amniote
New post.
coracoid
Cleithrum
lost
Added
scapular
spine
Synapsida Mammalia: Prototheria Mammalia: Theria
What germ layers contribute to the appendicular skeleton?
Epimere/Paraxial: Dermatome
Dermal bones in pectoral girdle
Dorsal fins
Hypomere (Somatic Layer):
Pectoral girdle (part)
Pelvic girdle
Pectoral fins/limbs
Pelvic fins/limbs
How do pelvic girdles of fish & tetrapods differ?
Fossil Sarcopterygian: Ischiopubic plates
Early Synapsid
Therapsid
Early Tetrapod: ilium
attachment to sacral vert.
Mammals
How do endochondral & membranous (dermal) bone growth differ?
Cartilage Endochondral Bone Formation
Compact Bone
Membranous (Dermal) Bone Formation

What are the epiphyseal regions on long bones?
Endochondral Bone Development: Mammal
Selected References
Carollia:
Fruit Bat
Ahlberg PE, Clack JA & Blom H. 2005. The axial skeleton of the Devonian tetrapod Ichthyostega. Nature 437:137-140.
Ahn D, & Ho RK. 2008. Tri-phasic expression of posterior Hox genes during development of pectoral fins in zebrafish: Implications for the evolution of vertebrate paired appendages. Dev Biol. 322(1):220-233.
Baier DB, Gatesy SM, & Jenkins Jr, FA. 2007. A critical ligamentous mechanism in the evolution of avian flight. Nature 445:307-310.
Boisvert CA, Mark-Kurik E, Ahlberg PE. 2008. The pectoral fin of Panderichthys and the origin of digits. Nature. 456(7222):636-638.
Botha-Brink J, & Angielczyk KD. 2010. Do extraordinarily high growth rates in Permo-Triassic dicynodonts (Therapsida, Anomodontia) explain their success before and after the end-Permian extinction? Zool. J Linn.
Soc. 160:341–365.
Canoville A, & Laurin M. 2010. Evolution of humeral microanatomy and lifestyle in amniotes, and some comments on palaeobiological inferences. Biol J Linnean Soc 100:384–406.
Davis MC, Dahn RD, & Shubin NH. 2007. An autopodial-like pattern of Hox expression in the fins of a basal actinopterygian fish. Nature 447:473-477.
Daeschle, EB, Shubin NH, & Jenkins FA, Jr. 2006. A Devonian tetrapod-like fish and the evolution of the tetrapod body plan. Nature 440:757-763.
Dean MN., & Summers AP. 2006. Mineralized cartilage in the skeleton of Chondrichthyan fishes. Zoology 109:164-168.
Dean MN, Mull CG, Gorb SN, & Summers AP. 2009. Ontogeny of the tessellated skeleton: insight from the skeletal growth of the round stingray Urobatis halleri. J Anat. 215(3):227–239.
Erickson GM 2005. Assessing dinosaur growth patterns: a microscopic revolution. Trends in Ecology and Evolution 20(12):677-684.
Erickson GM 2005. Dinosaur growth. AccessScience, ©McGraw-Hill Companies http://www.accessscience.com/popup.aspx?id=YB050600&name=print
Freitas R, Zhang GJ, & Cohn MJ. 2007. Biphasic Hoxd gene expression in shark paired fins reveals an ancient origin of the distal limb domain, PLoS ONE 2(8): e754.
Gillis JA, Dahn RD, & Shubin NH. 2009 . Shared developmental mechanisms pattern the vertebrate gill arch and paired fin skeletons. PNAS 106(14):5720–5724.
Holland T. 2012. Pectoral girdle and fin anatomy of Gogonasus andrewsae long, 1985: Implications for tetrapodomorph limb evolution. J Morph. Early View (published before inclusion in an issue)
http://onlinelibrary.wiley.com/doi/10.1002/jmor.20078/full
Hutchinson JR & Allen V. 2009. The evolutionary continuum of limb function from early theropods to birds. Naturwissenschaften 96:423–448.
Janvier P & Clément G. 2010. Muddy tetrapod origins. Nature 463(7):40-41.
Johansonn Z, et. al. 2007. Fish Fingers: Digit Homologues in Sarcopterygian Fish Fins. J Exp Zoolog B Mol Dev Evol. 308(6):757-768.
Kriloff A, Germain D, Vincent P, et al. 2008. Evolution of bone microanatomy of the tetrapod tibia and its use in palaeobiological inference. J Evol. BIol. 21(3):807-826.
Kronenberg HM. 2003. Developmental regulation of the growth plate. Nature 423:332-336.
Laroche M. 2008. Treatment of osteoporosis: all the questions we still cannot answer. Am J Medicine. 121(9):744-747.
Martin RA. 2010. Skeleton in the corset. Biology of Sharks & Rays. ReefQuest Centre for Shark Research. http://www.elasmo-research.org/education/white_shark/skeleton.htm
Mukherjee D, Ray S, & Sengupta DP. 2010. Preliminary observations on the bone microstructure, growth patterns, and life habits of some Triassic temnospondyls from India. J Vert Paleontology 30(1):78-93.
Murphy DC. 2006. Tetrapods Answer. The Devonian Times. http://www.devoniantimes.org/opportunity/tetrapodsAnswer.html
Niedz´wiedzki G. et al. 2010. Tetrapod trackways from the early Middle Devonian period of Poland. Nature 463(7):43-48.
Pourlis AF, & Antonopoulos,J. 2011. The ossification of the pectoral girdle and wing skeleton of the Quail (Coturnix coturnix japonica). Anat. Histol. Embryol. 40:219–225.
Qiang, J, Zhexi L, & Shu-a J. 1999. A Chinese triconodont mammal and mosaic evolution of the mammalian skeleton. Nature 398:326-330.
Raff RA. 2007. Written in stone: fossils, genes and evo–devo. Nature Reviews: Genetics 8:911-920.
Ray S, Mukherjee D, & Bandyopadhyay S. 2009. Growth patterns of fossil vertebrates as deduced from bone microstructure: case studies from India. J. Biosci. 34:661–672.
Sanchez S, et al. 2010. Limb-bone histology of temnospondyls: implications for understanding the diversification of palaeoecologies and patterns of locomotion of Permo-Triassic tetrapods. J . Evol. Biol.
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Scheye, TM, Klein N, & Sander PM. 2010. Developmental palaeontology of Reptilia as revealed by histological studies. Seminars in Cell & Developmental Biology 21:462–470.
Shubin , NH, Daeschler EB, & Jenkins FA, Jr. 2006. The pectoral fin of Tiktaalik roseae and the origin of the tetrapod limb. Nature 440:764-771.
Shubin N, Tabin C, & Carroll S. 2000. Fossils, Genes and the Evolution of Animal Limbs. In Gee, H., ed. 2000. Shaking the Tree: Readings from Nature in the History of Life.
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Science Museum of Minnesota. 2003. Dinosaur Bone Histology. http://ltc.smm.org/histology/
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Wisenden B. 2005. Chapter 7: The Skeletal System: Appendicular Division. Evolution Atlas. Pearson Education. http://cwx.prenhall.com/bookbind/pubbooks/martini10/chapter7/custom2/deluxe-content.html
How can bones be used to age a fossil & study growth?
Scheyer, Klein, & Sander, 2010.
Sketch of prosauropod, Plateosaurus engelhardti
http://palaeo-electronica.org/2009_2/185/fig_14.htm
Sampling is done by complete cross-sectioning or core-drilling of the bone at mid-diaphysis.
(B) Polished section of humerus. (C) Close-up of the outer region of the cortex of that specimen.
(D) Interpretative drawing of C. Note lines of arrested growth (LAGs).
Review Questions
1. Define appendicular skeleton. Describe the “Archipterygium & fin-fold hypotheses for the
evolution of vertebrate fins/limbs. What evidence supports each hypothesis? Describe the
HOX gene pattern within the appendicular skeleton.
2. Compare the position & arrangement of girdle cartilages/bones in the Chondrichthyes,
Actinopterygians, Sarcopterygians & early tetrapods. What is the advantage of the derived
Actinopterygian relocation of the pelvic fins? What design features are shared by the fossil
Sarcopterygian fin bones & the tetrapod limbs?
3. Diagram the major elements in an early tetrapod pectoral girdle. Why did it become free of
the skull? Describe the 2 major trends amongst these bones with evolution of upright
posture (which elements were enlarged, which were reduced?
4. Diagram & compare the bones that form the pelvic girdle of a fossil sarcopterygian & of an
early fossil tetrapod. In the tetrapod, label the acetabulum (socket for the femur). What new
connection is made in the tetrapod pelvic girdle? In general how did the relative size of
ilium, ischium & pubis change with the evolution of upright posture in synapsids to
mammals?
5. Compare the growth of replacement (endochondral) bones & membranous (dermal) bones.
Diagram a long bone in early development & label the diaphysis, epiphyses & epiphyseal
disks. How can growth rates be studied in fossils like the dinosaurs? Diagram a transverse
section of an embryo & label the regions of mesoderm. Describe & name the germ layer
origin of the appendicular skeleton.