Edaphosaurus

{{Short description|Extinct genus of synapsids}}

{{Automatic taxobox

| fossil_range = Pennsylvanian to Cisuralian (Gzhelian to Kungurian), {{Fossil range|303.4|272.5}}

| image = Edaphosaurus boanerges AMNH.jpg

| image_caption = Restored specimen of E. boanerges, AMNH

| display_parents = 2

| taxon = Edaphosaurus

| authority = Cope, 1882

| type_species = {{extinct}}Edaphosaurus pogonias

| type_species_authority = Cope, 1882

| subdivision_ranks = Species

| subdivision = * {{extinct}}E. cruciger (Cope, 1878) (originally Dimetrodon)

  • {{extinct}}E. pogonias Cope, 1882
  • {{extinct}}E. novomexicanus
    Williston and Case, 1913
  • {{extinct}}E. boanerges
    Romer & Price, 1940
  • {{extinct}}E. colohistion Berman, 1979

| synonyms = * Brachycnemius (Williston, 1911)

  • Naosaurus (Cope, 1886)

}}

Edaphosaurus ({{IPAc-en|ˌ|ɛ|d|ə|f|oʊ-|ˈ|s|ɔːr|ə|s}}, meaning "pavement lizard" for dense clusters of its teeth) is a genus of extinct edaphosaurid synapsids that lived in what is now North America and Europe around 303.4 to 272.5 million years ago,{{cite web |url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=38911 |title=Paleobiology Database: Edaphosaurus Cope 1882 (synapsid) |website=Fossilworks |access-date=17 December 2021}} during the Late Carboniferous to Early Permian. The American paleontologist Edward Drinker Cope first described Edaphosaurus in 1882,{{cite journal |last= Cope|first= E. D.|year= 1882|title= Third contribution to the History of the Vertebrata of the Permian Formation of Texas|journal= Proceedings of the American Philosophical Society|volume= 20|pages= 447–474}} naming it for the "dental pavement" on both the upper and lower jaws, from the Greek {{lang|grc-Latn|edaphos}} {{lang|grc|έδαφος}} ("ground"; also "pavement")An Intermediate Greek-English Lexicon. Oxford. Clarendon Press. 1889. "ἔδαφος [edaphos]... 2. The ground-floor, pavement..." [https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0058%3Aentry%3De)%2Fdafos] and {{lang|grc|σαῦρος}} ({{lang|grc-Latn|sauros}}) ("lizard").

Edaphosaurus is important as one of the earliest-known, large, plant-eating (herbivorous), amniote tetrapods (four-legged land-living vertebrates). In addition to the large tooth plates in its jaws, the most characteristic feature of Edaphosaurus is a sail on its back. A number of other synapsids from the same time period also have tall dorsal sails, most famously the large apex predator Dimetrodon. However, the sail on Edaphosaurus is different in shape and morphology. The first fossils of Edaphosaurus came from the Texas Red Beds in North America, with later finds in New Mexico, Oklahoma, West Virginia, and Ohio. Fragmentary fossils attributed to Edaphosaurus have also been found in eastern Germany in Central Europe.

Etymology

File:Edaphosaurus NT small.jpg

The name Edaphosaurus, meant as "pavement lizard",Miller, S. A. (1889). North American Geology and Palaeontology for the Use of Amateurs, Students, and Scientists. Western Methodist Book Concern, Cincinnati. 718 pp. is often translated inaccurately as "earth lizard", "ground lizard", or "foundation lizard" on the basis of other meanings for the Greek {{lang|grc-Latn|edaphos}}, such as "soil, earth, ground, land, base" used in Neo-Latin scientific nomenclature (edaphology). However, older names in paleontology, such as Edaphodon Buckland, 1838 "pavement tooth" (a fossil fish), match Cope's clearly intended meaning "pavement" for Greek edaphos in reference to the animal's teeth.

Description and paleobiology

File:Edaphosaurus Scale.svg

Edaphosaurus species measured from {{convert|0.5|to|3.5|m|ft|1}} in length and weighed over 300 kg (660 lb).{{cite web|title=Edaphosaurus|url=http://palaeos.com/vertebrates/synapsida/edaphosauridae2.html#Edaphosaurus|website=Palaeos|access-date=9 September 2022|archive-url=https://web.archive.org/web/20220220162206/http://palaeos.com/vertebrates/synapsida/edaphosauridae2.html|archive-date=20 February 2022}} In keeping with its tiny head, the cervical vertebrae are reduced in length, while the dorsal vertebrae are massive, the tail is deep, the limbs are short and robust, and the ribs form a wide ribcage. Like most herbivores, Edaphosaurus would have had a capacious gut and symbiotic bacteria to aid in the breakdown of cellulose and other indigestible plant material. Like its more famous relative Dimetrodon, Edaphosaurus had a sail-like fin that was supported by bones of the vertebral column. Edaphosaurus differs from Dimetrodon in having cross-bars on the spines that supported its fin.{{cite web|title=Edaphosaurus pogonias|url=https://www.utexas.edu/tmm/sponsored_sites/dino_pit/edaphosaurus.html|website=The Dino Pit Fossils|access-date=22 May 2015}}

=Skull=

File:The_Osteology_of_the_Reptiles_p50.png

The head of Edaphosaurus was short, relatively broad, triangular in outline, and remarkably small compared to its body size. The deep lower jaw likely had powerful muscles and the marginal teeth along the front and sides of its jaws had serrated tips, helping Edaphosaurus to crop bite-sized pieces from tough terrestrial plants. Back parts of the roof of the mouth and the inside of the lower jaw held dense batteries of peglike teeth, forming a broad crushing and grinding surface on each side above and below. Its jaw movements were propalinal (front to back). Early descriptions suggested that Edaphosaurus fed on invertebrates such as mollusks, which it would have crushed with its tooth plates. However, paleontologists now think that Edaphosaurus ate plants, although tooth-on-tooth wear between its upper and lower tooth plates indicates only "limited processing of food"{{cite journal |last= Reisz|first= R. R.|year= 2006|title= Origin of dental occlusion in tetrapods: signal for terrestrial vertebrate evolution? |journal= Journal of Experimental Zoology Part B |volume= 306B |issue= 3|pages= 261–277 |doi=10.1002/jez.b.21115|pmid= 16683226|bibcode= 2006JEZB..306..261R}} compared to other early plant-eaters such as Diadectes, a large nonamniote reptiliomorph (Diadectidae) that lived at the same time.

The recently described Melanedaphodon from the Middle Pennsylvanian subperiod of the Carboniferous Period in North America is currently the earliest known edaphosaurid and represents a transitional stage from a diet of hard-shelled invertebrates such as insects and mollusks to fibrous plants. Melanedaphodon had large and bulbous teeth along its upper and lower jaws, but also had "a moderately-developed tooth battery" on its palate, "which appears intermediary towards the condition seen in Edaphosaurus" and would have helped process tough plant material.{{cite journal|last1=Mann|first1=A.|last2=Henrici|first2=A. C.|last3=Sues|first3=H.-D.|last4=Pierce|first4=S. E.|date=2023|title=A new Carboniferous edaphosaurid and the origin of herbivory in mammal forerunners|journal=Scientific Reports|volume=13|number=4459|page=4459 |doi=10.1038/s41598-023-30626-8|pmid=37019927 |pmc=10076360 |bibcode=2023NatSR..13.4459M |doi-access=free}} Melanedaphodon was found to be a sister taxon to Edaphosaurus and lived earlier than the edaphosaurid Ianthasaurus, which lacked tooth plates and ate insects.

=Sail=

File:Boston Edaphosaurus.jpg

The sail along the back of Edaphosaurus was supported by hugely elongated neural spines from neck to lumbar region, connected by tissue in life. When compared with the sail of Dimetrodon, the vertebral spines are shorter and heavier, and bear numerous small crossbars. Edaphosaurus and other members of the Edaphosauridae evolved tall dorsal sails independently of sail-back members of the Sphenacodontidae such as Dimetrodon and Secodontosaurus that lived at the same time, an unusual example of parallel evolution. The {{Not a typo|function(s)}} of the sail in both groups is still debated. Researchers have suggested that such sails could have provided camouflage, wind-powered sailing over water, anchoring for extra muscle support and rigidity for the backbone, protection against predator attacks, fat-storage areas, body-temperature control surfaces, or sexual display and species recognition. The height of the sail, curvature of the spines, and shape of the crossbars are distinct in each of the described species of Edaphosaurus and show a trend for larger and more elaborate (but fewer) projecting processes over time. The possible function (or functions) of the bony tubercles on the spines remains uncertain. Romer and Price suggested that the projections on the spines of Edaphosaurus might have been embedded in tissue under the skin and might have supported food-storage or fat similar to the hump of a camel.{{cite journal |last=Romer |first=A.S. |author2=Price, L.I. |year=1940 |title=Review of the Pelycosauria |journal=Geological Society of America Special Paper |volume=28 |pages=1–538 |doi=10.1130/spe28-p1|series=Geological Society of America Special Papers }} Bennett argued that the bony projections on Edaphosaurus spines were exposed and could create air turbulence for more efficient cooling over the surface of the sail to regulate body temperature.{{cite journal |last= Bennett|first= S. C.|year= 1996|title= Aerodynamics and thermoregulatory function of the dorsal sail of Edaphosaurus |journal= Paleobiology |volume= 22 |issue= 4|pages= 496–506|doi= 10.1017/S0094837300016481|bibcode= 1996Pbio...22..496B|s2cid= 89276555}} Recent research that examined the microscopic bone structure of the tall neural spines in edaphosaurids has raised doubts about a thermoregulatory role for the sail and suggests that a display function is more plausible.{{cite journal |last= Huttenlocker |first= A. K. |author2=Mazierski, D. |author3=Reisz, R. R. |year=2011 |title= Comparative osteohistology of hyperelongate neural spines in the Edaphosauridae (Amniota: Synapsida) |journal= Palaeontology |volume= 54 |issue= 3 |pages= 573–590 |doi=10.1111/j.1475-4983.2011.01047.x|bibcode= 2011Palgy..54..573H |doi-access= free }}

=Growth and metabolism=

A study comparing the microscopic bone histology of the vertebral centra of Edaphosaurus and Dimetrodon found that the plant-eating Edaphosaurus "grew distinctly more slowly" than the predator Dimetrodon, which had a higher growth rate, reflecting an "elevated metabolism".{{Cite journal|author1=Amin Agliano |author2=P. Martin Sander |author3=Tanja Wintrich |year=2020 |title=Bone histology and microanatomy of Edaphosaurus and Dimetrodon (Amniota, Synapsida) vertebrae from the Lower Permian of Texas |journal=The Anatomical Record |volume=304 |issue=3 |pages=570–583 |doi=10.1002/ar.24468 |pmid=32484294 |s2cid=219172923 |doi-access=free }} Earlier studies of Edaphosaurus limb bones had also indicated slower growth and a lower metabolism, reflecting an ectothermic (cold-blooded) animal, although the plant-eating early synapsid caseids had a lower growth rate than Edaphosaurus.Christen Don Shelton (2015) Origins of endothermy in the mammalian lineage the evolutionary beginning of fibrolamellar bone in the "mammal-like" reptiles.(Ph.D. Dissertation) https://bonndoc.ulb.uni-bonn.de/xmlui/handle/20.500.11811/6495 Evidence of growth rates include the number of blood vessels in the bones (with more vascularization in the rapidly growing Dimetrodon) and the presence of lamellar bone in the cancellous part. In contrast to slow growth in overall body size and in most bones, the histology of the tall dorsal spines on Edaphosaurus suggests that the projecting bony tubercles developed "by sudden, rapid growth over a few seasons", unlike the incremental growth of the tubercles in the earlier edaphosaurid Ianthasaurus.{{cite journal|author1= Frederik Spindler |author2= Sebastian Voigt |author3= Jan Fischer |year=2020 |title= Edaphosauridae (Synapsida, Eupelycosauria) from Europe and their relationship to North American representatives |journal= PalZ |volume=94 |issue= 1 |pages=125–153 |doi=10.1007/s12542-019-00453-2 |bibcode= 2020PalZ...94..125S |s2cid= 198140317 }}

Species

class="wikitable sortable"
SpeciesAuthorityLocationStatusSynonymsclass="unsortable" | Images
Edaphosaurus boanergesRomer & Price, 1940TexasValid125px
Edaphosaurus colohistionBerman, 1979West VirginiaValid
Edaphosaurus crucigerCope, 1878Texas and OklahomaValidEdaphosaurus microdus125px
Edaphosaurus novomexicanusWilliston & Case, 1913New MexicoValid125px
Edaphosaurus pogoniasCope, 1882TexasValidEdaphosaurus claviger125px

Discovery and classification

File:Edaphosaurus-Field Museum.jpg]]

Edward Drinker Cope named and described Edaphosaurus ("pavement lizard") in 1882, based on a crushed skull and a left lower jaw from the Texas Red Beds. He noted in particular the "dense body of teeth" on both the upper and lower jaws, and used the term "dental pavement" in a table in his description. The type species name pogonias means "bearded" in Greek, referring to the enlarged inward sloping chin on the lower jaw. Cope classified Edaphosaurus as a member of his Pelycosauria and created the new family Edaphosauridae. The type material did not include any of the post-cranial skeleton apart from an axis vertebra and Cope was unaware of the animal's large sail, a feature then known only for Dimetrodon.

In 1886, Cope erected the new genus Naosaurus "ship lizard" (from Greek naos "ship") for skeletal remains similar to those of the long-spined Dimetrodon, but with distinctive "transverse processes or branches, which resemble the yardarms of a ship's mast".{{cite journal |last= Cope|first= E. D.|year= 1886|title= The long-spined Theromorpha of the Permian epoch |journal= American Naturalist|volume= 20|pages= 544–545|doi=10.1086/274275 |doi-access= free}} He speculated that "the yardarms were connected by membranes with the neural spine or mast, thus serving the animal as a sail with which he navigated the waters of the Permian lakes". He recognized three species: Naosaurus claviger "club-bearer" (for the projections on its spines; now considered a synonym of Edaphosaurus pogonias); Naosaurus cruciger "cross-bearer" (for the projections on its spines; first described by Cope as Dimetrodon cruciger in 1878; now Edaphosaurus cruciger, the largest species in size); and Naosaurus microdus "small tooth" (first described as Edaphosaurus microdus in 1884). Cope noted some incomplete skull material found associated with the specimens of N. claviger and N. microdus, but thought Naosaurus was distinct from Edaphosaurus. He later decided that Naosaurus must have had a large carnivorous skull similar to Dimetrodon, although he had no direct fossil proof. In 1910, German paleontologist Otto Jaekel reported remains near Dresden in Saxony, which he called Naosaurus credneri.{{cite journal |last= Jaekel|first= O. M. J.|year= 1910|title= Naosaurus credneri im Rotliegenden von Sachsen |journal= Zeitschrift der Deutschen Geologischen Gesellschaft|volume= 62|pages= 526–535 }}

In 1907, American paleontologist Ermine Cowles Case suggested that the skull of Edaphosaurus might belong with skeletons called Naosaurus, based on a specimen found in 1906 that appeared to associate elements of both.{{cite book |last=Case |first=E.C. |year=1907 |title=Revision of the Pelycosauria of North America |publisher=Carnegie Institution of Washington |location=Washington, D.C.|pages=[https://archive.org/details/bub_gb_7QMDAAAAIAAJ/page/n6 1]–176 |url=https://archive.org/details/bub_gb_7QMDAAAAIAAJ}} In 1913, Samuel Wendell Williston and Case described the new species Edaphosaurus novomexicanus from a fairly complete specimen unearthed in New Mexico in 1910, in which a sailbacked Naosaurus-type skeleton was found with a small Edaphosaurus-type skull.{{cite journal |last= Williston |first=S.W. |author2=Case, E.C. |year=1913 |title= A Description of Edaphosaurus Cope |journal=Permo-Carboniferous Vertebrates from New Mexico. Carnegie Institution of Washington Geological Society of America Special Paper |volume=181 |pages=71–81}} The older generic name Edaphosaurus Cope, 1882 became the valid one.

In 1940, paleontologists Alfred Sherwood Romer and Llewellyn Ivor Price named the new species Edaphosaurus boanerges ("thunderous orator")Webster's Revised Unabridged Dictionary. Springfield, MA: C. & G. Merriam Co., 1913. [http://www.websters1913.com/words/Boanerges Boanerges. ] – an ironic reference to the remarkably small size of the holotype lower jaw on a composite skeleton originally mounted in the Museum of Comparative Zoology (Harvard University) with the head restored based on the larger species Edaphosaurus cruciger.

In 1979, paleontologist David Berman erected Edaphosaurus colohistion ("stunted sail") for an early species with a relatively small sail, based on fossils from West Virginia.{{cite journal |last= Berman|first= D. S.|year= 1979|title= Edaphosaurus (Reptilia, Pelycosauria) from the Lower Permian of Northeastern United States, with description of a new species |journal= Annals of the Carnegie Museum |volume= 48 | issue = 11 |pages= 185–202|doi= 10.5962/p.215833|s2cid= 191159505|doi-access= free}}

Reassigned species

Other proposed species of Edaphosaurus have been based on more fragmentary material that cannot be rigorously diagnosed to a genus/species level, but which may nonetheless represent edaphosaurids. The nominal species Naosaurus raymondi was assigned to Edaphosaurus by Romer and Price (1940), but Modesto and Reisz (1990) designated it a nomen vanum,Modesto, S.P. & Reisz, R.R., 1990. Taxonomic status of Edaphosaurus raymondi Case. Journal of Paleontology 64 (6): 1049‐1051. and Spindler (2015) considered it probably referable to Ianthasaurus due to its age and stratigraphy.http://tubaf.qucosa.de/api/qucosa%3A22988/attachment/ATT-0/ {{Dead link|date=February 2022}} The taxon Naosaurus mirabilis Fritsch, 1895 from the Czech Republic was given its own genus Bohemiclavulus by Spindler et al. (2019).

See also

{{Portal|Paleontology}}

References

;Notes

{{Reflist}}

;Bibliography

  • Carroll, R. L. (1988), Vertebrate Paleontology and Evolution, WH Freeman & Co.
  • Colbert, E. H., (1969), Evolution of the Vertebrates, John Wiley & Sons Inc (2nd ed.)
  • Romer, A. S., (1947, revised ed. 1966) Vertebrate Paleontology, University of Chicago Press, Chicago
  • Romer, A. S. and Price, L. I., (1940), Review of the Pelycosauria, Geological Society of America Special Papers, No 28