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Fruitbat's Pdf Library - Pelycosaurs

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These are a few of the pdf files (and a few Microsoft Word documents) that I've accumulated in my web browsing. 

MOST of these are hyperlinked to their source.

If you want one that is not hyperlinked or if the link isn't working, e-mail me at joegallo1954@gmail.com and I'll be happy to send it to you.


Please note that this list will be updated continuously as I find more available resources. All of these files are freely available on the Internet so there should be no copyright issues.



Articles with author names in RED are new additions since July 29, 2018.


Class (Clade) Synapsida


Order(?) Pelycosauria


Family Caseidae

Brocklehurst, N. and J. Fröbisch (2017). A re-examination of the enigmatic Russian tetrapod Phreatophasma aenigmaticum and its evolutionary implications. Fossil Record, 20.

Brocklehurst, N., M. Romano and J. Fröbisch (2016). Principal Component Analysis as an Alternative Treatment for Morphometric Characters: Phylogeny of Caseids as a Case Study. Palaeontology, 59(6).

LeBlanc, A.R.H. and R.R. Reisz (2014). New Postcranial Material of the Early Caseid Casea broilii Williston, 1910 (Synapsida: Caseidae) with a Review of the Evolution of the Sacrum in Paleozoic Non-Mammalian Synapsids. PLoS ONE, 9(12).

Maddin, H.C., C.A. Sidor and R.R. Reisz (2008). Cranial Anatomy of Ennatosaurus tecton (Synapsida: Caseidae) from the Middle Permian of Russia and the Evolutionary Relationships of Caseidae. Journal of Vertebrate Paleontology, 28(1).

Olson, E.C. (1954). Fauna of the Vale and Choza: 7. Pelycosauria: Family Caseidae. Fieldiana Geology, Vol.10, Number 17.

Reisz, R.R. and J. Frӧbisch (2014). The Oldest Caseid Synapsid from the Late Pennsylvanian of Kansas, and the Evolution of Herbivory in Terrestrial Vertebrates. PLoS ONE, 9(4).

Reisz, R.R., S.J. Godfrey and D. Scott (2009). Eothyris and Oedalops: Do These Early Permian Synapsids from Texas and New Mexico Form a Clade? Journal of Vertebrate Paleontology, 29(1).

Reisz, R.R., et al. (2011). A new large caseid (Synapsida, Caseasauria) from the Permian of Rodez (France), including a reappraisal of "Casea" rutena Sigogneau-Russell & Russell, 1974. Geodiversitas, 33(2).

Romano, M. (2017). Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach. Lethaia.

Romano, M.  Phylogenetic analysis of the Family Caseidae (Synapsida, Caseasauria) with description and morphometric analysis of the new giant caseid Alierasaurus ronchii from the Permian of Sardinia, Italy. Ph.D. Thesis - Universita di Roma.

Romano, M. and U. Nicosia (2015). Cladistic Analysis of Caseidae (Caseasauria, Synapsida): Using the Gap-Weighting Method to Include Data Based on Incomplete Specimens. Palaeontology, Vol.58, Part 6.

Romano, M. and U. Nicosia (2014). Alierasaurus ronchii, gen.et sp.nov., A Caseid from the Permian of Sardinia, Italy. Journal of Vertebrate Paleontology, 34(4).

Romano, M., N. Brocklehurst and J. Fröbisch (2017). The postcranial skeleton of Ennatosaurus tecton (Synapsida, Caseidae). Journal of Systematic Palaeontology, 2017.

Romano, M., et al. (2017). New material of Alierasaurus ronchii (Synapsida, Caseidae) from the Permian of Sardinia (Italy), and its phylogenetic affinities. Palaeontologia Electronica, 20.2.26A.

Ronchi, A., et al. (2011). A huge caseid pelycosaur from north-western Sardinia and its bearing on European Permian stratigraphy and palaeobiogeography. Acta Palaeontologica Polonica, 56(4).

Spindler, F., J. Falconnet and J. Fröbisch (2016). Callibrachion and Datheosaurus, two historical and previously mistaken basal caseasaurian synapsids from Europe. Acta Palaeontologica Polonica, 61(3).

Stovall, J.W., L.I. Price, and A.S. Romer (1966). The Postcranial Skeleton of the Giant Permian Pelycosaur Cotylorhynchus romeri. Bulletin Museum of Comparative Zoology, Vol.135, Number 1.


Family Edaphosauridae

Case, E.C. (1918). A Mounted Skeleton of Edaphosaurus cruciger Cope, in the Geological Collection of the University of Michigan. Occasional Papers of the Museum of Zoology - University of Michigan, Number 62.

Case, E.C. (1906). On the Skull of Edaphosaurus pogonias Cope. Bulletin American Museum of Natural History, Vol.XXII, Article II.

Huttenlocker, A.K., D. Mazierski and R.R. Reisz (2011). Comparative Osteohistology of Hyperelongate Neural Spines in the Edaphosauridae (Amniota: Synapsida). Palaeontology, Vol.54, Part 3.

Mazierski, D.M. and R.R. Reisz (2010). Description of a new specimen of Ianthasaurus hardestiorum (Eupelycosauria: Edaphosauridae) and a re-evaluation of edaphosaurid phylogeny. Can.J. Earth Sci., 47.

Modesto, S.P. (1995). The Skull of the Herbivorous Synapsid Edaphosaurus boanerges from the Lower Permian of Texas. Palaeontology, Vol.38, Part 1.

Modesto, S.P. (1994). The Lower Permian Synapsid Glaucosaurus from Texas. Palaeontology, Vol.37, Part 1.

Modesto, S.P. and R.R. Reisz (2011). A new skeleton of Ianthasaurus hardestii, a primitive edaphosaur (Synapsida: Pelycosauria) from the Upper Pennsylvanian of Kansas. Can.J. Earth Sci., 27(6).

Modesto, S.P. and R.R. Reisz (1992). Restudy of Permo-Carboniferous synapsid Edaphosaurus novomexicanus Williston and Case, the oldest known herbivorous amniote. Can.J. Earth Sci., 29.

Modesto, S.P. and R.R. Reisz (1990). Taxonomic Status of Edaphosaurus raymondi Case. J.Paleont., 64(6).

Osborn, H.F. (1907). A Mounted Skeleton of Naosaurus , A Pelycosaur from the Permian of Texas. Bulletin American Museum of Natural History, Vol.XXIII, Article XIV.

Reisz, R.R. and D.S. Berman (2011). Ianthasaurus hardestii n.sp., a primitive edaphosaur (Reptilia, Pelycosauria) from the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas. Can.J. Earth Sci., 23(1).

Sumida, S. (1989). New information on the pectoral girdle and vertebral column in Lupeosaurus (Reptilia, Pelycosauria). Can.J. Earth Sci., Vol.26.


Family Eothyrididae


Brocklehurst, N., et al. (2016). A Re-Description of 'Mycterosaurus' smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids. PLoS ONE, 11(6).

Langston, W. (1965). Oedaleops campi (Reptilia: Pelycosauria) New Genus and Species from the Lower Permian of New Mexico, and the Family Eothyrididae. Bulletin of the Texas Memorial Museum, Number 9.

Reisz, R.R., S.J. Godfrey and D.Scott (2009). Eothyris and Oedaleops: Do These Early Permian Synapsids from Texas and New Mexico Form a Clade? Journal of Vertebrate Paleontology, 29(1).


Family Ophiacodontidae


Berman, D.S., A.C. Henrici and S.G. Lucas (2013). Ophiacodon (Synapsida, Ophiacodontidae) from the Lower Permian Sangre de Cristo Formation of New Mexico. In: The Carboniferous-Permian Transition. Lucas, S.G., et al. (eds). New Mexico Museum of Natural History and Science, Bulletin 60.

Berman, D.S., et al. (1995). The Cranial Anatomy and Relationships of the Synapsid Varanosaurus (Eupelycosauria: Ophiacodontidae) from the Early Permian of Texas and Oklahoma. Annals of Carnegie Museum, Vol.64, Number 2.

Harris, S.K., S.G. Lucas and J.A. Spielmann (2010). A Partial Skeleton of Ophiacodon navajovicus (Eupelycosauria: Ophiacodontidae) from the Upper Pennsylvanian of Cañon del Cobre, New Mexico. In: Carb-Permian transition in Cañon del Cobre, New Mexico. Lucas, S.G., et al. (eds.), New Mexico Museum of Natural History and Science, Bulletin 49.

Romer, A.S. (1961). A large ophiacodont pelycosaur from the Pennsylvanian of the Pittsburgh region. Breviora, 144.

Shelton, C. and P.M. Sander (2015). Ophiacodon long bone histology: the earliest occurrence of FLB in the mammalian stem lineage. PeerJ Preprints.

Spielmann, J.A. and S.G. Lucas (2010). Re-Evaluation of Ruthiromia elcobriensis (Eupelycosauria: Ophiacodontidae?) from the Lower Permian (Seymouran?) of Cañon del Cobre, Northern New Mexico. In: Carb-Permian transition in Cañon del Cobre, New Mexico. Lucas, S.G., et al. (eds.), New Mexico Museum of Natural History and Science, Bulletin 49.

Stephens, J.J. (1964). Ophiacodon from Ohio. The Ohio Journal of Science, 64(3).


Family Sphenacodontidae


Falconett, J. (2015). A sphenacodontid synapsid Neosaurus cynodus, and related material, from the Permo-Carboniferous of France. Acta Palaeontologica Polonica, 60(1).

Reisz, R.R., D.S. Berman and D. Scott (1992). The cranial anatomy and relationships of Secodontosaurus, an unusual mammal-like reptile (Synapsida: Sphenacodontidae) from the early Permian of Texas. Zoological Journal of the Linnean Society, 104.

Spindler, F., D. Scott and R.R. Reisz (2015). New information on the cranial and postcranial anatomy of the early synapsid Ianthodon schultzei (Sphenacomorpha: Sphenacodontia), and its evolutionary significance.  Fossil Record, 18.


Subfamily Sphenacodontinae


Berman, D.S., et al. (2001). A new species of Dimetrodon (Synapsida: Sphenacodontidae) from the Lower Permian of Germany records first occurrence of genus outside of North America. Can.J. Earth Sci., 38.

Brink, K.S. (2015). Case 3695. Dimetrodon Cope, 1878 (Synapsida, SPHENACODONTIDAE): proposed conservation by reversal of precedence with Bathygnathus Leidy, 1853. Bulletin of Zoological Nomenclature, 72(4).

Brink, K.S. and R.R. Reisz (2014). Hidden dental diversity in the oldest terrestrial apex predator Dimetrodon. Nature Communications, 5:3269.

Brink, K.S. and R.R. Reisz (2012). Morphology of the palate and braincase of Dimetrodon milleri. In: 12th International Symposium on Early Vertebrates/Lower Vertebrates Special Issue, Historical Biology, Vol.24, Number 4.

Brink, K.S., et al. (2015). Re-evaluation of the historic Canadian fossil Bathygnathus borealis from the Early Permian of Prince Edward Island. Can.J. Earth Sci., Vol.52.

Case, E.C. (1924). A Possible Explanation of Fenestration in the Primitive Reptilian Skull, With Notes on the Temporal Region of the Genus Dimetrodon. Contributions from the Museum of Paleontology - The University of Michigan, Vol.II, Number 1.

Case, E.C. (1910). Description of a Skeleton of Dimetrodon incisivus Cope. Bulletin American Musem of Natural History, Vol.XXVIII, Article XIX.

Case, E.C. (1904).  The Osteology of the Skull of the Pelycosaurian Genus, Dimetrodon. The Journal of Geology, Vol.12, Number 4.

Eberth, D.A. (1985). The skull of Sphenacodon ferocior, and comparisons with other sphenacodontines (Reptilia: Pelycosauria). New Mexico Bureau of Mines & Mineral Resources, Circular 190.

Fröbisch, J., et al. (2011). A new basal sphenacodontid synapsid from the Late Carboniferous of the Saar-Nahe Basin, Germany. Acta Palaeontologica Polonica, 56(1).

Henrici, A.C., et al. (2005). The Carpus and Tarsus of the Early Permian Synapsid Sphenacodon ferox (Eupelycosauria: Sphenacodontidae). In: The Nonmarine Permian. Lucas, S.G. and K.E. Ziegler (eds.), New Mexico Museum of Natural History and Science Bulletin Number 30.

Lucas, S.G., et al. (2009). Dimetrodon (Amniota: Synapsida: Sphenacodontidae) from the Lower Permian Abo Formation, Socorro County, New Mexico. New Mexico Geological Society Guidebook, 60th Field Conference, Geology of the Chupadera Mesa Region.

Shelton, C.D., et al. (2013). Long bone histology indicates sympatric species of Dimetrodon (Lower Permian, Sphenacodontidae). Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 103.

Spielmann, J.A., et al. (2010). Redescription of the Cranial Anatomy of Sphenacodon ferox Marsh (Eupelycosauria: Sphenacodontidae) from the Late Pennsylvanian - Early Permian of New Mexico. In: The Carb-Permian transition in Cancun del Cobre. Lucas, et al. (eds.), New Mexico Museum of Natural History and Science, Bulletin 49.

Williston, S.W. (1916). Sphenacodon Marsh, A Permocarboniferous Theromorph Reptile from New Mexico.


General Sphenacodontidae


Brink, K.S., A.R.H. LeBlanc and R.R. Reisz (2014). First record of plicidentine in Synapsida and patterns of tooth root shape change in Early Permian sphenacodontians. Naturwissenschaften, 10:1.

Laurin, M. (1993). Anatomy and Relationships of Haptodus garnettensis, A Pennsylvanian Synapsid from Kansas. Journal of Vertebrate Paleontology, 13(2).

Spindler, F. (2015). The basal Sphenacodontia - systematic revision and evolutionary implications. Ph.D. Dissertation - Technischen Universitat Bergakademie Freiberg.


Family Varanopidae


Anderson, J.S. and R.R. Reisz (2004). Pyozia mesenensis, a New Small Varanopid (Synapsida, Eupelycosauria) from Russia: "Pelycosaur" Diversity in the Middle Permian. Journal of Vertebrate Paleontology, 24(1).

Reisz, R.R. and D.W. Dilkes (2003). Archaeovenator hamiltonensis, a new varanopid (Synapsida: Eupelycosauria) from the Upper Carboniferous of Kansas. Can.J.Earth Sci., 40.

Reisz, R.R., M. Laurin and D. Marjanović (2010). Apsisaurus witteri from the Lower Permian of Texas: Yet Another Small Varanopid Synapsid, Not a Diapsid. Journal of Vertebrate Paleontology, 30(5).

Spindler, F., et al. (2018). First arboreal 'pelycosaurs' (Synaspsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte , SE Germany,  with a review of varanopid phylogeny. Pal.Z., (2018), 5.


Subfamily Mycterosaurinae


Broom, R. (1930). On a New Primitive Theromorph (Eumatthevia bolli). American Museum Novitates, Number 446.

Modesto, S.P., et al. (2011). The last "pelycosaur": a varanopid synapsid from the Pristerognathus Assemblage Zone, Middle Permian of South Africa. Naturwissenschaften, 98.

Reisz, R.R. and D.S. Berman (2001). The Skull of Mesenosaurus romeri, A Small Varanopseid (Synapsida: Eupelycosauria) from the Upper Permian of the Mezen River Basin, Northern Russia. Annals of Carnegie Museum, Vol.70, Number 2.

Williston, S.W. (1915). A New Genus and Species of American Theromorpha - Mycterosaurus longiceps. The Journal of Geology, Vol.23, Number 6.


Subfamily Varanopinae


Langston, W. and R.R. Reisz (1981). Aerosaurus wellesi, New Species, A Varanopseid Mammal-Like Reptile (Synapsida: Pelycosauria) from the Lower Permian of New Mexico. Journal of Vertebrate Paleontology, 1(1).

Maddin, H.C., D.C. Evans, and R.R. Reisz (2006). An Early Permian Varanodontine Varanopid (Synapsida: Eupelycosauria) from the Richards Spur Locality, Oklahoma. Journal of Vertebrate Paleontology, 26(4).

Modesto, S., et al. (2001). A second varanopseid skull from the Upper Permian of South Africa: implications for Late Permian 'pelycosaur' evolution. Lethaia, Vol.34.

Reisz, R.R. and L.A. Tsuji (2006). An Articulated Skeleton of Varanops With Bite Marks: The Oldest Known Evidence of Scavenging Among Terrestrial Vertebrates. Journal of Vertebrate Paleontology, 26(4).

Reisz, R.R. and M. Laurin (2004). A reevaluation of the enigmatic Permian synapsid Watongia and of its stratigraphic significance. Can.Jour.Earth Sci., 41.

Reisz, R.R., D.W. Dilkes and D.S. Berman (1998). Anatomy and Relationships of Elliotsmithia longiceps Broom, A Small Synapsid (Eupelycosauria: Varanopseidae) from the Late Permian of South Africa. Journal of Vertebrate Paleontology, 18(3).


General Varanopidae


Botha-Brink, J. and S.P. Modesto (2007). A mixed-age classed 'pelycosaur' aggregation from South Africa: earliest evidence of parental care in amniotes? Proc.R.Soc. B, 274.

Campione, N.E. and R.R. Reisz (2011). Morphology and evolutionary significance of the atlas-axis complex in varanopid synapsids. Acta Palaeontologica Polonica, 56(4).

Piñeiro, G., et al. (2003). First Basal Synapsids ("Pelycosaurs") from the Upper Permian-?Lower Triassic of Uruguay, South America. J.Paleont., 77(2).


General Pelycosauria

Brocklehurst, N. and J. Fröbisch (2014). Current and historical perspectives on the completeness of the fossil record of pelycosaurian-grade synapsids. Palaeogeography, Palaeoclimatology, Palaeoecology, 399.

Case, E.C. (1907). Revision of the Pelycosauria of North America. Carnegie Institute of Washington. (285 pages)

Paton, R.L. (1974). Lower Permian Pelycosaurs from the English Midlands. Palaeontology, Vol.17, Part 3.

Pineiro, J., et al. (2003). First Basal Synapsids ("Pelycosaurs") from the Upper Permian - ?Lower Triassic of Uruguay, South America. J.Paleont., 77(2).

Reisz, R.R. Pelycosaurian Reptiles from the Middle Pennsylvanian of North America. Masters Thesis - McGill University.

Reisz, R.R., M.J. Heaton and B.R. Pynn  (1982). Vertebrate Fauna of Late Pennsylvanian Rock Lake Shale Near Garnett, Kansas: Pelycosauria. Journal of Paleontology, Vol.56, Number 3.

Van Valkenberg, B. and I. Jenkins (2002). Evolutionary Patterns in the History of Permo-Triassic and Cenozoic Synapsid Predators. Paleontological Society Papers, Vol.8.

Watson, D.M.S.  (1916). Reconstructions of the Skulls of Three Pelycosaurs in the American Museum of Natural History. Bulletin American Museum of Natural History, Vol.XXXV, Article XXXII.

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Created and updated - October 3, 2017.



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Updated July 29, 2018.



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