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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/30620-8.txt b/30620-8.txt new file mode 100644 index 0000000..34fc59b --- /dev/null +++ b/30620-8.txt @@ -0,0 +1,856 @@ +The Project Gutenberg EBook of Two New Pelycosaurs from the Lower Permian +of Oklahoma, by Richard C. Fox + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Two New Pelycosaurs from the Lower Permian of Oklahoma + +Author: Richard C. Fox + +Release Date: December 7, 2009 [EBook #30620] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK PELYCOSAURS, L. PERMIAN, OKLAHOMA *** + + + + +Produced by Chris Curnow, Woodie4, Joseph Cooper and the +Online Distributed Proofreading Team at https://www.pgdp.net + + + + + + + + + + + UNIVERSITY OF KANSAS PUBLICATIONS + + MUSEUM OF NATURAL HISTORY + + Volume 12, No. 6, pp. 297-307, 6 figs. + May 21, 1962 + + + Two New Pelycosaurs from the Lower Permian + of Oklahoma + + BY + + RICHARD C. FOX + + + UNIVERSITY OF KANSAS + LAWRENCE + 1962 + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, Henry S. Fitch, + Theodore H. Eaton, Jr. + + + Volume 12, No. 6, pp. 297-307, 6 figs. + Published May 21, 1962 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED BY + JEAN M. NEIBARGER, STATE PRINTER + TOPEKA, KANSAS + 1962 + + 29-3001 + + + + +Two New Pelycosaurs from the Lower Permian of Oklahoma + +BY + +RICHARD C. FOX + + +In the course of examining material from fissure deposits of early +Permian age collected from a limestone quarry near Fort Sill, Oklahoma, +the author recovered several tooth-bearing fragments of small +pelycosaurs. The fragments were examined, compared with descriptions of +known kinds appearing in the literature, and determined to be new genera +within the Nitosauridae (Edaphosauria) and Sphenacodontidae +(Sphenacodontia). + +Appreciation is expressed to Prof. Theodore H. Eaton, Jr., for +permission to examine the collections of the University of Kansas from +Fort Sill, and for the financial assistance furnished by his National +Science Foundation grant (NSF-G8624). I am grateful both to Prof. Eaton +and Mr. Dale L. Hoyt for their suggestions regarding this manuscript. +The accompanying figures have been drawn by the author. + + + + +Family NITOSAURIDAE + +=Delorhynchus priscus= new genus and new species + + +(_delos_, Gr., evident; _rhynchos_, Gr., neuter, nostril; _priscus_, L., +ancient. _Delorhynchus_ is masculine because of the ending that it +acquires when transliterated into Latin.) + +_Type specimen._--Fragmentary left maxilla, bearing four teeth, KU +11117. + +_Referred specimens._--Fragmentary right maxilla having four teeth, KU +11118; fragmentary left maxilla having four teeth, the most posterior of +which has been broken, KU 11119. + +_Horizon and locality._--A fissure deposit in the Arbuckle limestone at +the Dolese Brothers Limestone Quarry, approximately six miles north of +Fort Sill, in sec. 31, T. 4 N, R. 11 W, Comanche County, Oklahoma. These +sediments are of early Permian age, possibly equivalent to the Arroyo +formation, Lower Clear Fork Group of Texas (Vaughn, 1958: 981). + +_Diagnosis._--Small; marginal teeth conical, slender and recurved at +tips; marginal tooth-row without caniniform enlargement; narial opening +enlarged and bordered dorsally, posteriorly and ventrally by maxilla; +maxilla with foramen opening laterally at posteroventral corner of +naris. + +_Description_ (based on 3 maxillary fragments, see Table 1).--Each of +the maxillary fragments bears four thecodont teeth. These are conical, +slender and sharply pointed; in their distal third they are slightly +recurved, laterally compressed, and have anterior and posterior +non-serrated cutting edges. In medial aspect at their bases, the teeth +are longitudinally striated. The bases of the teeth are circular in +cross-section and are slightly bulbous. There is no caniniform +enlargement of any of the teeth, the longest tooth of each fragment +being differently placed in the series of teeth and little longer than +the others. There is no swelling on either the internal or external +surfaces of the maxillae. The teeth are in a continuous series; no +diastema or maxillary step is evident. + +[Illustration: FIGURES 1-3. _Delorhynchus priscus_, lower Permian, 6 +miles north of Fort Sill, Comanche County, Oklahoma. All × 3. + +FIG. 1. KU 11117 (type specimen), lateral view of left maxilla. +FIG. 2. KU 11118, lateral view of right maxilla. +FIG. 3. KU 11119, lateral view of left maxilla.] + +The fragments have been broken along similar lines of fracture, and each +is approximately rhomboidal in shape. The maxilla encircles the +posterior border of the naris and extends dorsally above the naris to an +extent sufficient to indicate the probable exclusion of the lacrimal +bone from the narial border. At the posteroventral corner of the naris a +foramen opens onto the lateral surface of the maxilla. The opening is +the entrance to a canal that runs posteriorly above the tooth-row +throughout the length of each specimen. Beneath the naris the maxilla +extends as a broad tapering shelf, the ventral surface of which +articulates with the premaxilla. The narial rim is wide, but wider +ventrally than dorsally. The plane of the narial rim is oblique to the +lateral surface of the maxilla. The external surface of each fragment is +grooved and pitted. The ossification of each fragment appears to have +been complete. + +TABLE 1.--DIMENSIONS, IN MILLIMETERS, OF THREE MAXILLARY FRAGMENTS OF +DELORHYNCHUS PRISCUS + +Key: + +A. Anterior height of fragment +B. Posterior height of fragment +C. Length of fragment at tooth-row +D. Dorsal length of fragment +E. Mean length of teeth +F. Anterior width of naris + +============================================================ +CATALOGUE NUMBER | A. | B. | C. | D. | E. | F. +AND MEAN | | | | | | +-----------------+------+------+------+------+------+------ +KU 11117 | 6.0 | 8.0 | 6.0 | 8.0 | 3.0 | 3.0 +KU 11118 | 6.0 | 6.0 | 9.0 | 8.0 | 2.0 | 3.0 +KU 11119 | 6.6 | 8.0 | 10.0 | 11.0 | ? | 4.6 +-----------------+------+------+------+------+------+------ +Mean | 6.2 | 7.3 | 8.3 | 9.0 | 2.5 | 4.5 +-----------------+------+------+------+------+------+------ + +_Discussion._--The Nitosauridae are small primitive edaphosaurs with a +moderately elongate face, sharp subisodont teeth, little development of +canines and few specializations. The jaw is of a primitive type and +articulates on a level with the tooth-row. The palatal dentition is +primitive (Romer, 1956:280). The nitosaurids are thought to be related +to the later Caseidae, and the most obvious structural similarities are +found in the postcranial skeleton (Vaughn, 1958:989). Cranial +resemblances between the families are fewer, but nevertheless indicate +that a nitosaurid-caseid relationship exists. + +Vaughn (1958) described a small pelycosaur, _Colobomycter pholeter_ +(Eothyrididae, Ophiacodontia) that structurally resembles the Caseidae. +This individual also was obtained from the Fort Sill locality. In +Vaughn's opinion the features of _Colobomycter_ indicate a close +relationship between eothyridids and caseids and the possibility that +the caseids may well have been of eothyridid rather than nitosaurid +derivation. + +In view of this historical uncertainty of the relationships between the +Nitosauridae, the Eothyrididae and the Caseidae, it is well to consider +how the maxillary fragments described above differ from and resemble +representatives of each of these three families as reported in the +literature. + +_Delorhynchus_ resembles _Colobomycter_ in size. The mean +extra-maxillary length of the undamaged teeth of the three fragments is +2.5 mm., equal to that reported by Vaughn (1958:985) for teeth about +midway in the postcanine series of _Colobomycter_. None of the teeth of +_Delorhynchus_ extends beyond the maxillary rim as far as does the +canine of _Colobomycter_ (3.5 mm.). + +The teeth in both genera are conical and sharply pointed. The naris in +each is enlarged, and the lacrimal is excluded from the narial margin in +each (by inference in _Delorhynchus_.) + +The differences between the maxillae of _Colobomycter_ and +_Delorhynchus_ are most striking in the lack of canines in the latter +and the correlated absence of modifications of the maxillary for support +of canines. Additionally, _Delorhynchus_ bears an infraorbital canal in +contrast to the groove in similar position in _Colobomycter_. The +recurvature of the four teeth present in the fragments of _Delorhynchus_ +differs from that in the teeth of _Colobomycter_ in which only the +canine and precanine are recurved. Vaughn implies that anterior and +posterior cutting edges extend the length of the teeth in +_Colobomycter_; these are restricted to the distal third of the teeth in +_Delorhynchus_. The external surfaces of the maxillae of _Delorhynchus_ +are pitted and ridged; Vaughn was unable to discern sculpturing of the +corresponding surfaces in _Colobomycter_. + +_Delorhynchus_ resembles the nitosaurids in size, the shape and +sharpness of the teeth, their recurvature and the slight enlargement of +their bases, the exclusion of the lacrimal bone from the narial margin +(in _Mycterosaurus_) and the apparent lack of a special canine pair of +teeth. Resemblances to the caseids are to be noted in the enlargement of +the naris (4.5 mm. in height as opposed to 1.7 mm. in _Colobomycter_), +lack of development of canines, presence of an infraorbital canal (in +_Cotylorhynchus_) and absence of many replacement gaps in the marginal +row of teeth. + +The absence of caniniform enlargement and the extension of the maxilla +dorsad of the naris exclude _Delorhynchus_ from the Eothyrididae +(Ophiacodontia) but are no bar to its inclusion in the Nitosauridae +(Edaphosauria). The marginal teeth of _Delorhynchus_ are simple and +primitive, being much like those of the nitosaurids that are described +in the literature. + +The large narial opening and its posterior, dorsal and ventral enclosure +by the maxilla, the infraorbital canal, and the sculptured external +surfaces of the maxillary fragments indicate that _Delorhynchus_, in +these features at least, is close to achieving the caseid grade. + + + + +Family SPHENACODONTIDAE + +=Thrausmosaurus serratidens= new genus and new species + + +(_Thrausmosaurus_ is formed from the neuter Greek noun, _thrausma_, +meaning fragment, and the masculine Greek noun, _sauros_, meaning +reptile. The specific name, _serratidens_, is formed from the Latin +_serratus_, meaning serrate, and the masculine Latin noun, _dens_, +meaning tooth. The specific name is used as a substantive in apposition +with the generic name.) + +_Type specimen._--Fragmentary left dentary, bearing five teeth, the +most posterior of which is broken at the base, KU 11120. + +_Referred specimens._--Fragmentary ?left maxilla, having two teeth, KU +11121; fragmentary left dentary having two teeth, KU 11122. + +_Horizon and locality._--From the early Permian fissure deposits in the +Arbuckle limestone of the Dolese Brothers Limestone Quarry, +approximately 6 miles north of Fort Sill, in sec. 31, T. 4N, R. 11 W, +Comanche County, Oklahoma. + +_Diagnosis._--Small; teeth thecodont, compressed laterally, recurved +distally, and bearing anterior and posterior cutting edges; anterior +serrations limited to recurved portions of teeth, posterior serrations +extending nearly entire length of teeth; lateral compression of teeth +more pronounced medially than laterally; bases of teeth expanded. + +_Description._--The type specimen is 16 mm. long. It bears five teeth +that are implanted in a straight row. Empty sockets are present between +the first and second teeth, and the third and fourth teeth. The first +tooth is 3.0 mm. long, the middle two are each 2.5 mm. long, and the +fourth tooth is 2.0 mm. long. The fifth tooth is broken off at its base. + +The empty sockets are large. The mouth of each is circular and +approximately 2.0 mm. in diameter. Both sockets are 1.25 mm. deep. The +bases of the teeth are expanded to fill the sockets, although the blades +of the teeth arise from only the lateral portions of the bases. The edge +of the dentary rises above the bases of the teeth medially, thereby +producing a small depression at the junction of each base with the +dentary bone. + +The lateral compression of the teeth is pronounced but asymmetrical, in +that the lateral surface of each blade is more convex than the medial +surface. + +[Illustration: FIGURES 4-6. _Thrausmosaurus serratidens_, lower Permian, +6 miles north of Fort Sill, Comanche County, Oklahoma. All × 3. + +FIG. 4. KU 11120 (type specimen), lateral view of left dentary. +FIG. 5. KU 11121, lateral view of ?left maxilla. +FIG. 6. KU 11122, lateral view of left dentary.] + +The recurvature of the anterior cutting edges is much more severe than +that of the posterior edges, but the recurvature of both is limited to +the distal half of each tooth. + +The serrations of the cutting edges are not visible to the naked eye and +are limited on the anterior edges of the teeth to those portions of the +blades that are recurved. The posterior serrations extend nearly to the +junction of the blade of each tooth with its base. The serrations tend +to be more nearly crenulate than cuspidate. + +A portion of the lateral wall of the dentary surrounding the Meckelian +canal is present. The external surface of the wall is gently convex and +smooth, without sculpturing. The internal surfaces of the canal are +unmarked either by muscle scars or foramina. + +The fragment is a piece from the posterior portion of the dentary, since +the decrease in height from the first tooth to the fourth is pronounced. + +KU 11122, a fragment of the left dentary bearing two teeth, is 7.5 mm. +long. The anterior tooth is 3.0 mm. long; the posterior tooth is 3.5 mm. +long. The shape of the teeth and their implantation conform to the +description of the type specimen. The lateral surface of the fragment is +smooth and gently convex. What little is present of the surface +bordering the Meckelian canal is unmarked. + +The ?maxillary fragment bears two teeth which are 3.0 mm. long, and +which conform in their characters to the type. The lateral, medial and +ventral surfaces of the fragment have been sheared off, so that an exact +identification of the bone is impossible. Presumably the fragment is too +deep dorsoventrally to be a piece of the dentary, and no sign of the +Meckelian canal is present. + +_Discussion._--The implantation, lateral compression, recurvature and +cutting edges of the teeth borne by these fragments make clear their +sphenacodontid nature. The characters of the fragments are too few to +determine subfamilial affinities, however. That the fragments are the +remains of adult animals can be only surmised from the lack of bones or +teeth of large pelycosaurs in the extensive collections of the +University of Kansas from the Fort Sill locality. + +If _Thrausmosaurus_ is, in fact, adult, the genus is an unusually small +sphenacodontid, and of significance both on that account and because of +the resemblance of the teeth presently known to those of its far larger +relatives. + +_The Fort Sill Locality._--Peabody (1961) suggested that the fissures of +Fort Sill had been used as dens by predatory animals in the early +Permian, and that the unusually abundant bones in the fissures were the +remains of animals eaten there by these occupants. Evidence now known to +me affords an alternative explanation that is presented here as a +preliminary to a more complete study of the fauna and paleoecology of +these deposits currently being undertaken. + +The suggestion that the skeletal material found in the fissures is the +remnant of the prey of other animals is questionable because of: + + 1. The absence of tooth marks on the fossils. + + 2. The recovery from the matrix of skulls and portions of + articulated skeletons that are undamaged or damaged only by + pressure after burial. + + 3. The rarity in the deposits of animals of larger body size than + _Captorhinus_, the exceptions being a few limb fragments and skull + fragments of labyrinthodont or pelycosaurian nature. + + 4. The absence of coprolites in the matrix. + +If the fissures were the dens of predators, at least some and probably +many of the bones would show tooth marks. A predator feeding on other +animals would be expected to leave some evidence of its habits on the +bones of its prey. No such evidence is known to me, either from my own +examination of several thousand bones or from the reports in the +literature by others who have studied aspects of the early Permian fauna +of Fort Sill. + +If the predators were larger than _Captorhinus_ and occupied the +fissures for a long enough time to account for the accumulation of the +tremendous numbers of individuals that are represented, a considerable +amount of the skeletal material of the larger animals would be present +in the fissure deposits. Even if for some reason the predators died in +areas other than within the fissures, thereby accounting for the absence +of large bones, coprolites should appear in the deposits if, in fact, +the fissures were feeding places. In view of the nearly undamaged +condition of many of the bones recovered from the fissures, it is +reasonable to expect that fecal material would be preserved. + +The character of the matrix of the deposits varies from a homogeneous +clay to clay interrupted by layers of soft, limey, conglomeratic rock, +to a hard, well-cemented, calcareous conglomerate. In general the bone +in each kind of matrix is colored characteristically and exhibits a +characteristic degree of wear. The bones entrapped in the homogeneous +clay are relatively few, black, usually disarticulated, little worn and +not unduly fragmented; consequently the discovery of undamaged limb +bones, for example, from this kind of matrix is not unusual. The bones +found in the stratified portion of the matrix are more numerous within +the layers of conglomerate than between. The bones are black, brown or +white, highly fragmented and waterworn to a variable degree. The +fragments recovered from the hard, calcareous matrix are numerous, range +in color from white through various shades of brown, to black, are +highly fragmented, and are usually worn by water. + +These categories for bone and matrix, however, are not mutually +exclusive, since bones of any of these colors and exhibiting any degree +of wear and fragmentation are found in any of the kinds of matrix +described above. That water was the agent of wear is suggested by the +highly polished appearance of the worn bones and pebbles that are found +in the matrix. + +The variability of the matrix and of the color and condition of the +bones indicates that the agencies of burial and fossilization differed +from time to time and that the agency of transportation of the bones +from the site of burial to the fissures was running water. One can +easily visualize a stream coursing the early Permian landscape that was +subject to periodic flooding and droughts. Along the banks of the stream +and in its pools lived a variety of microsaurs, captorhinids, small +labyrinthodonts and small pelycosaurs. Some of the animals, after they +died, were either buried near the site of their death or were swept +along and buried in sediments further downstream. Burial was for a +length of time sufficient to impart a color to the bones characteristic +of the site in which they were buried. Later floods reexposed the sites +of burial, picked up the bones and carried them to the openings into the +fissures. Presumably, too, a proportion of the bones was carried to the +fissures without previous burial. + +The differences in wear exhibited by different bones within the same +block of matrix is attributable to differences in distance that the +bones were transported before final deposition. The final sites of +deposition, the fissures, were inundated occasionally by floods alone, +or because of changes in location of the channel of the stream at the +time of flooding. The periodicity of deposition of the sediments within +portions of the fissures is indicated by the stratification of the bone +conglomerate mentioned earlier. + +In summary, it seems that there is little or no evidence beyond the +numbers of bones involved to support the hypothesis that the +concentration of bones in the fissures of Fort Sill represents the +remains of food of predators, and that the fissures were used as dens by +their predatory occupants. On the contrary, the evidence indicates that +the deposition of the bones in the fissures was secondary and that the +agency of transportation, deposition and accumulation of the bones was +an early Permian stream characterized by periodic flooding. + + + + +LITERATURE CITED + + +PEABODY, F. E. + 1961. Annual growth zones in living and fossil + vertebrates. Jour. Morph. 108 (1): 11-62, 69 figs., January. + +ROMER, A. S. + 1956. Osteology of the reptiles. The University of Chicago + Press, Chicago, xxi + 772 pp., 248 figs. + +ROMER, A. S., and PRICE, L. I. + 1940. Review of the Pelycosauria. Geol. + Soc. America, Spec. Pap., 28: x + 538 pp., 71 figs., 46 pls., + 8 tables, December 6. + +VAUGHN, P. P. + 1958. On a new pelycosaur from the lower Permian of + Oklahoma, and the origin of the family Caseidae. Jour. Paleont., + 32:981-991, 1 fig., September. + +_Transmitted March 15, 1962._ + + + + + +End of the Project Gutenberg EBook of Two New Pelycosaurs from the Lower +Permian of Oklahoma, by Richard C. Fox + +*** END OF THIS PROJECT GUTENBERG EBOOK PELYCOSAURS, L. 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Fox. + </title> + <style type="text/css"> + +body { + margin-left: 10%; + margin-right: 10%; +} + + h1,h2,h3,h4,h5 { + text-align: center; /* all headings centered */ + clear: both; +} + +p { + margin-top: .75em; + text-align: justify; + margin-bottom: .75em; +} + +hr { + width: 33%; + margin-top: 2em; + margin-bottom: 2em; + margin-left: auto; + margin-right: auto; + clear: both; +} + +table { + margin-left: auto; + margin-right: auto; +} + +.pagenum { /* uncomment the next line for invisible page numbers */ + visibility: hidden; + position: absolute; + left: 92%; + font-size: smaller; + text-align: right; +} /* page numbers */ + + +.center {text-align: center;} + +.smcap {font-variant: small-caps;} + +/* Images */ +.figcenter { + margin: auto; + text-align: center; +} + + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of Two New Pelycosaurs from the Lower Permian +of Oklahoma, by Richard C. Fox + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Two New Pelycosaurs from the Lower Permian of Oklahoma + +Author: Richard C. Fox + +Release Date: December 7, 2009 [EBook #30620] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK PELYCOSAURS, L. PERMIAN, OKLAHOMA *** + + + + +Produced by Chris Curnow, Woodie4, Joseph Cooper and the +Online Distributed Proofreading Team at https://www.pgdp.net + + + + + + +</pre> + + +<p><span class="pagenum"><a name="Page_297" id="Page_297">[Pg 297]</a></span></p> + +<hr style="width: 90%;" /> + +<h3><span class="smcap">University of Kansas Publications</span><br /> + +<span class="smcap">Museum of Natural History</span></h3> + +<hr style="width: 25%;" /> + +<h4>Volume 12, No. 6, pp. 297-307, 6 figs.</h4> +<h4>May 21, 1962</h4> + +<hr style="width: 90%;" /> + + +<h1>Two New Pelycosaurs from the Lower Permian +of Oklahoma</h1> + +<h3>BY</h3> + +<h2>RICHARD C. FOX</h2> + + +<h3><span class="smcap">University of Kansas</span></h3> +<h3><span class="smcap">Lawrence</span></h3> +<h3>1962<br /><br /><br /><br /></h3> + +<p><span class="pagenum"><a name="Page_298" id="Page_298">[Pg 298]</a></span></p> + + +<h4><span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Theodore H. Eaton, Jr.<br /><br /><br /></h4> + +<h4>Volume 12, No. 6, pp. 297-307, 6 figs.<br /> +Published May 21, 1962<br /><br /><br /></h4> + +<h4><span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /><br /><br /></h4> + +<h5>PRINTED BY<br /> +JEAN M. NEIBARGER, STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1962<br /></h5> + +<h5>29-3001<br /></h5> +<p><span class="pagenum"><a name="Page_299" id="Page_299">[Pg 299]</a></span></p> + + + +<hr style="width: 65%;" /> +<h2>Two New Pelycosaurs from the Lower Permian of Oklahoma</h2> + +<h5>BY</h5> + +<h4><span class="smcap">Richard C. Fox</span></h4> + + +<p>In the course of examining material from fissure deposits of early +Permian age collected from a limestone quarry near Fort Sill, Oklahoma, +the author recovered several tooth-bearing fragments of small +pelycosaurs. The fragments were examined, compared with descriptions of +known kinds appearing in the literature, and determined to be new genera +within the Nitosauridae (Edaphosauria) and Sphenacodontidae +(Sphenacodontia).</p> + +<p>Appreciation is expressed to Prof. Theodore H. Eaton, Jr., for +permission to examine the collections of the University of Kansas from +Fort Sill, and for the financial assistance furnished by his National +Science Foundation grant (NSF-G8624). I am grateful both to Prof. Eaton +and Mr. Dale L. Hoyt for their suggestions regarding this manuscript. +The accompanying figures have been drawn by the author.</p> + + + +<hr style="width: 65%;" /> +<h2>Family NITOSAURIDAE</h2> + +<h3><b>Delorhynchus priscus</b> new genus and new species</h3> + + +<p>(<i>delos</i>, Gr., evident; <i>rhynchos</i>, Gr., neuter, nostril; <i>priscus</i>, L., +ancient. <i>Delorhynchus</i> is masculine because of the ending that it +acquires when transliterated into Latin.)</p> + +<p><i>Type specimen.</i>—Fragmentary left maxilla, bearing four teeth, KU +11117.</p> + +<p><i>Referred specimens.</i>—Fragmentary right maxilla having four teeth, KU +11118; fragmentary left maxilla having four teeth, the most posterior of +which has been broken, KU 11119.</p> + +<p><i>Horizon and locality.</i>—A fissure deposit in the Arbuckle limestone at +the Dolese Brothers Limestone Quarry, approximately six miles north of +Fort Sill, in sec. 31, T. 4 N, R. 11 W, Comanche County, Oklahoma. These +sediments are of early Permian age, possibly equivalent to the Arroyo +formation, Lower Clear Fork Group of Texas (Vaughn, 1958: 981).</p> + +<p><i>Diagnosis.</i>—Small; marginal teeth conical, slender and recurved at +tips; marginal tooth-row without caniniform enlargement; narial opening +enlarged and bordered dorsally, posteriorly and ventrally by maxilla; +maxilla with foramen opening laterally at posteroventral corner of +naris.</p> + +<p><i>Description</i> (based on 3 maxillary fragments, see Table 1).—Each of +the maxillary fragments bears four thecodont teeth. These are conical, +slender and sharply pointed; in their distal third they are slightly +recurved, laterally compressed, and have anterior and posterior +non-serrated cutting edges. In medial aspect at their bases, the teeth +are longitudinally striated. The bases of the teeth<span class="pagenum"><a name="Page_300" id="Page_300">[Pg 300]</a></span> are circular in +cross-section and are slightly bulbous. There is no caniniform +enlargement of any of the teeth, the longest tooth of each fragment +being differently placed in the series of teeth and little longer than +the others. There is no swelling on either the internal or external +surfaces of the maxillae. The teeth are in a continuous series; no +diastema or maxillary step is evident.</p> + +<div class="figcenter" style="width: 600px;"> +<img src="images/i004.png" width="600" height="315" alt="Figures 1-3." title="" /> +</div> + +<p>The fragments have been broken along similar lines of fracture, and each +is approximately rhomboidal in shape. The maxilla encircles the +posterior border of the naris and extends dorsally above the naris to an +extent sufficient to indicate the probable exclusion of the lacrimal +bone from the narial border. At the posteroventral corner of the naris a +foramen opens onto the lateral surface of the maxilla. The opening is +the entrance to a canal that runs posteriorly<span class="pagenum"><a name="Page_301" id="Page_301">[Pg 301]</a></span> above the tooth-row +throughout the length of each specimen. Beneath the naris the maxilla +extends as a broad tapering shelf, the ventral surface of which +articulates with the premaxilla. The narial rim is wide, but wider +ventrally than dorsally. The plane of the narial rim is oblique to the +lateral surface of the maxilla. The external surface of each fragment is +grooved and pitted. The ossification of each fragment appears to have +been complete.<br /><br /></p> + +<p><span class="smcap">Table 1.—Dimensions, in Millimeters, of Three Maxillary Fragments of +Delorhynchus priscus</span></p> + +<p style="margin-left: 35%;"> +Key:<br /> + +A. Anterior height of fragment<br /> +B. Posterior height of fragment<br /> +C. Length of fragment at tooth-row<br /> +D. Dorsal length of fragment<br /> +E. Mean length of teeth<br /> +F. Anterior width of naris<br /><br /></p> + + <div class="center"> +<table border="2" cellpadding="8" cellspacing="0" summary="maxillar dimensions"> +<tr><td align="left"><span class="smcap">Catalogue Number<br /> +and Mean</span></td><td align="center">A.</td><td align="center">B.</td><td align="center">C.</td><td align="center">D.</td><td align="center">E.</td><td align="center">F.</td></tr> +<tr><td align="left">KU 11117</td><td align="center">6.0</td><td align="center">8.0</td><td align="center">6.0</td><td align="center">8.0</td><td align="center">3.0</td><td align="center">3.0</td></tr> +<tr><td align="left">KU 11118</td><td align="center">6.0</td><td align="center">6.0</td><td align="center">9.0</td><td align="center">8.0</td><td align="center">2.0</td><td align="center">3.0</td></tr> +<tr><td align="left">KU 11119</td><td align="center">6.6</td><td align="center">8.0</td><td align="center">10.0</td><td align="center">11.0</td><td align="center">?</td><td align="center">4.6</td></tr> +<tr><td align="left">Mean</td><td align="center">6.2</td><td align="center">7.3</td><td align="left">8.3</td><td align="center">9.0</td><td align="center">2.5</td><td align="center">4.5</td></tr> +</table><br /></div> + + +<p><i>Discussion.</i>—The Nitosauridae are small primitive edaphosaurs with a +moderately elongate face, sharp subisodont teeth, little development of +canines and few specializations. The jaw is of a primitive type and +articulates on a level with the tooth-row. The palatal dentition is +primitive (Romer, 1956:280). The nitosaurids are thought to be related +to the later Caseidae, and the most obvious structural similarities are +found in the postcranial skeleton (Vaughn, 1958:989). Cranial +resemblances between the families are fewer, but nevertheless indicate +that a nitosaurid-caseid relationship exists.</p> + +<p>Vaughn (1958) described a small pelycosaur, <i>Colobomycter pholeter</i> +(Eothyrididae, Ophiacodontia) that structurally resembles the Caseidae. +This individual also was obtained from the Fort Sill locality. In +Vaughn's opinion the features of <i>Colobomycter</i> indicate a close +relationship between eothyridids and caseids and the possibility that +the caseids may well have been of eothyridid rather than nitosaurid +derivation.</p> + +<p>In view of this historical uncertainty of the relationships between the +Nitosauridae, the Eothyrididae and the Caseidae, it is well to consider +how the maxillary fragments described above differ from and resemble +representatives of each of these three families as reported in the +literature.</p> + +<p><i>Delorhynchus</i> resembles <i>Colobomycter</i> in size. The mean +extra-maxillary length of the undamaged teeth of the three fragments is +2.5 mm., equal to that reported by Vaughn (1958:985) for teeth about +midway in the postcanine series of <i>Colobomycter</i>. None of the teeth of +<i>Delorhynchus</i> extends beyond the maxillary rim as far as does the +canine of <i>Colobomycter</i> (3.5 mm.).</p> + +<p>The teeth in both genera are conical and sharply pointed. The naris in +each is enlarged, and the lacrimal is excluded from the narial margin in +each (by inference in <i>Delorhynchus</i>.)</p> + +<p>The differences between the maxillae of <i>Colobomycter</i> and +<i>Delorhynchus</i> are most striking in the lack of canines in the latter +and the correlated absence of modifications of the maxillary for support +of canines. Additionally, <i>Delorhynchus</i> bears an infraorbital canal<span class="pagenum"><a name="Page_302" id="Page_302">[Pg 302]</a></span> in +contrast to the groove in similar position in <i>Colobomycter</i>. The +recurvature of the four teeth present in the fragments of <i>Delorhynchus</i> +differs from that in the teeth of <i>Colobomycter</i> in which only the +canine and precanine are recurved. Vaughn implies that anterior and +posterior cutting edges extend the length of the teeth in +<i>Colobomycter</i>; these are restricted to the distal third of the teeth in +<i>Delorhynchus</i>. The external surfaces of the maxillae of <i>Delorhynchus</i> +are pitted and ridged; Vaughn was unable to discern sculpturing of the +corresponding surfaces in <i>Colobomycter</i>.</p> + +<p><i>Delorhynchus</i> resembles the nitosaurids in size, the shape and +sharpness of the teeth, their recurvature and the slight enlargement of +their bases, the exclusion of the lacrimal bone from the narial margin +(in <i>Mycterosaurus</i>) and the apparent lack of a special canine pair of +teeth. Resemblances to the caseids are to be noted in the enlargement of +the naris (4.5 mm. in height as opposed to 1.7 mm. in <i>Colobomycter</i>), +lack of development of canines, presence of an infraorbital canal (in +<i>Cotylorhynchus</i>) and absence of many replacement gaps in the marginal +row of teeth.</p> + +<p>The absence of caniniform enlargement and the extension of the maxilla +dorsad of the naris exclude <i>Delorhynchus</i> from the Eothyrididae +(Ophiacodontia) but are no bar to its inclusion in the Nitosauridae +(Edaphosauria). The marginal teeth of <i>Delorhynchus</i> are simple and +primitive, being much like those of the nitosaurids that are described +in the literature.</p> + +<p>The large narial opening and its posterior, dorsal and ventral enclosure +by the maxilla, the infraorbital canal, and the sculptured external +surfaces of the maxillary fragments indicate that <i>Delorhynchus</i>, in +these features at least, is close to achieving the caseid grade.</p> + + + +<hr style="width: 65%;" /> +<h2>Family SPHENACODONTIDAE</h2> + +<h3><b>Thrausmosaurus serratidens</b> new genus and new species</h3> + + +<p>(<i>Thrausmosaurus</i> is formed from the neuter Greek noun, <i>thrausma</i>, +meaning fragment, and the masculine Greek noun, <i>sauros</i>, meaning +reptile. The specific name, <i>serratidens</i>, is formed from the Latin +<i>serratus</i>, meaning serrate, and the masculine Latin noun, <i>dens</i>, +meaning tooth. The specific name is used as a substantive in apposition +with the generic name.)</p> + +<p><i>Type specimen.</i>—Fragmentary left dentary, bearing five teeth, the +most posterior of which is broken at the base, KU 11120.</p> + +<p><i>Referred specimens.</i>—Fragmentary ?left maxilla, having two teeth, KU +11121; fragmentary left dentary having two teeth, KU 11122.</p> + +<p><i>Horizon and locality.</i>—From the early Permian fissure deposits in the +Arbuckle limestone of the Dolese Brothers Limestone Quarry, +approximately 6 miles north of Fort Sill, in sec. 31, T. 4N, R. 11 W, +Comanche County, Oklahoma.<span class="pagenum"><a name="Page_303" id="Page_303">[Pg 303]</a></span></p> + +<p><i>Diagnosis.</i>—Small; teeth thecodont, compressed laterally, recurved +distally, and bearing anterior and posterior cutting edges; anterior +serrations limited to recurved portions of teeth, posterior serrations +extending nearly entire length of teeth; lateral compression of teeth +more pronounced medially than laterally; bases of teeth expanded.</p> + +<p><i>Description.</i>—The type specimen is 16 mm. long. It bears five teeth +that are implanted in a straight row. Empty sockets are present between +the first and second teeth, and the third and fourth teeth. The first +tooth is 3.0 mm. long, the middle two are each 2.5 mm. long, and the +fourth tooth is 2.0 mm. long. The fifth tooth is broken off at its base.</p> + +<p>The empty sockets are large. The mouth of each is circular and +approximately 2.0 mm. in diameter. Both sockets are 1.25 mm. deep. The +bases of the teeth are expanded to fill the sockets, although the blades +of the teeth arise from only the lateral portions of the bases. The edge +of the dentary rises above the bases of the teeth medially, thereby +producing a small depression at the junction of each base with the +dentary bone.</p> + +<p>The lateral compression of the teeth is pronounced but asymmetrical, in +that the lateral surface of each blade is more convex than the medial +surface.</p> + +<div class="figcenter" style="width: 600px;"> +<img src="images/i007.png" width="600" height="253" alt="Figures 4-6." title="" /> +</div> + +<p>The recurvature of the anterior cutting edges is much more severe than +that of the posterior edges, but the recurvature of both is limited to +the distal half of each tooth.</p> + +<p>The serrations of the cutting edges are not visible to the naked eye and +are limited on the anterior edges of the teeth to those portions of the +blades that are recurved. The posterior serrations extend nearly to the +junction of the blade of each tooth with its base. The serrations tend +to be more nearly crenulate than cuspidate.<span class="pagenum"><a name="Page_304" id="Page_304">[Pg 304]</a></span></p> + +<p>A portion of the lateral wall of the dentary surrounding the Meckelian +canal is present. The external surface of the wall is gently convex and +smooth, without sculpturing. The internal surfaces of the canal are +unmarked either by muscle scars or foramina.</p> + +<p>The fragment is a piece from the posterior portion of the dentary, since +the decrease in height from the first tooth to the fourth is pronounced.</p> + +<p>KU 11122, a fragment of the left dentary bearing two teeth, is 7.5 mm. +long. The anterior tooth is 3.0 mm. long; the posterior tooth is 3.5 mm. +long. The shape of the teeth and their implantation conform to the +description of the type specimen. The lateral surface of the fragment is +smooth and gently convex. What little is present of the surface +bordering the Meckelian canal is unmarked.</p> + +<p>The ?maxillary fragment bears two teeth which are 3.0 mm. long, and +which conform in their characters to the type. The lateral, medial and +ventral surfaces of the fragment have been sheared off, so that an exact +identification of the bone is impossible. Presumably the fragment is too +deep dorsoventrally to be a piece of the dentary, and no sign of the +Meckelian canal is present.</p> + +<p><i>Discussion.</i>—The implantation, lateral compression, recurvature and +cutting edges of the teeth borne by these fragments make clear their +sphenacodontid nature. The characters of the fragments are too few to +determine subfamilial affinities, however. That the fragments are the +remains of adult animals can be only surmised from the lack of bones or +teeth of large pelycosaurs in the extensive collections of the +University of Kansas from the Fort Sill locality.</p> + +<p>If <i>Thrausmosaurus</i> is, in fact, adult, the genus is an unusually small +sphenacodontid, and of significance both on that account and because of +the resemblance of the teeth presently known to those of its far larger +relatives.</p> + +<p><i>The Fort Sill Locality.</i>—Peabody (1961) suggested that the fissures of +Fort Sill had been used as dens by predatory animals in the early +Permian, and that the unusually abundant bones in the fissures were the +remains of animals eaten there by these occupants. Evidence now known to +me affords an alternative explanation that is presented here as a +preliminary to a more complete study of the fauna and paleoecology of +these deposits currently being undertaken.</p> + +<p>The suggestion that the skeletal material found in the fissures is the +remnant of the prey of other animals is questionable because of:<span class="pagenum"><a name="Page_305" id="Page_305">[Pg 305]</a></span></p> + + +<blockquote><p>1. The absence of tooth marks on the fossils.</p> + +<p>2. The recovery from the matrix of skulls and portions of +articulated skeletons that are undamaged or damaged only by +pressure after burial.</p> + +<p>3. The rarity in the deposits of animals of larger body size than +<i>Captorhinus</i>, the exceptions being a few limb fragments and skull +fragments of labyrinthodont or pelycosaurian nature.</p> + +<p>4. The absence of coprolites in the matrix.</p></blockquote> + +<p>If the fissures were the dens of predators, at least some and probably +many of the bones would show tooth marks. A predator feeding on other +animals would be expected to leave some evidence of its habits on the +bones of its prey. No such evidence is known to me, either from my own +examination of several thousand bones or from the reports in the +literature by others who have studied aspects of the early Permian fauna +of Fort Sill.</p> + +<p>If the predators were larger than <i>Captorhinus</i> and occupied the +fissures for a long enough time to account for the accumulation of the +tremendous numbers of individuals that are represented, a considerable +amount of the skeletal material of the larger animals would be present +in the fissure deposits. Even if for some reason the predators died in +areas other than within the fissures, thereby accounting for the absence +of large bones, coprolites should appear in the deposits if, in fact, +the fissures were feeding places. In view of the nearly undamaged +condition of many of the bones recovered from the fissures, it is +reasonable to expect that fecal material would be preserved.</p> + +<p>The character of the matrix of the deposits varies from a homogeneous +clay to clay interrupted by layers of soft, limey, conglomeratic rock, +to a hard, well-cemented, calcareous conglomerate. In general the bone +in each kind of matrix is colored characteristically and exhibits a +characteristic degree of wear. The bones entrapped in the homogeneous +clay are relatively few, black, usually disarticulated, little worn and +not unduly fragmented; consequently the discovery of undamaged limb +bones, for example, from this kind of matrix is not unusual. The bones +found in the stratified portion of the matrix are more numerous within +the layers of conglomerate than between. The bones are black, brown or +white, highly fragmented and waterworn to a variable degree. The +fragments recovered from the hard, calcareous matrix are numerous, range +in color from white through various shades of brown, to black, are +highly fragmented, and are usually worn by water.</p> + +<p>These categories for bone and matrix, however, are not mutually<span class="pagenum"><a name="Page_306" id="Page_306">[Pg 306]</a></span> +exclusive, since bones of any of these colors and exhibiting any degree +of wear and fragmentation are found in any of the kinds of matrix +described above. That water was the agent of wear is suggested by the +highly polished appearance of the worn bones and pebbles that are found +in the matrix.</p> + +<p>The variability of the matrix and of the color and condition of the +bones indicates that the agencies of burial and fossilization differed +from time to time and that the agency of transportation of the bones +from the site of burial to the fissures was running water. One can +easily visualize a stream coursing the early Permian landscape that was +subject to periodic flooding and droughts. Along the banks of the stream +and in its pools lived a variety of microsaurs, captorhinids, small +labyrinthodonts and small pelycosaurs. Some of the animals, after they +died, were either buried near the site of their death or were swept +along and buried in sediments further downstream. Burial was for a +length of time sufficient to impart a color to the bones characteristic +of the site in which they were buried. Later floods reexposed the sites +of burial, picked up the bones and carried them to the openings into the +fissures. Presumably, too, a proportion of the bones was carried to the +fissures without previous burial.</p> + +<p>The differences in wear exhibited by different bones within the same +block of matrix is attributable to differences in distance that the +bones were transported before final deposition. The final sites of +deposition, the fissures, were inundated occasionally by floods alone, +or because of changes in location of the channel of the stream at the +time of flooding. The periodicity of deposition of the sediments within +portions of the fissures is indicated by the stratification of the bone +conglomerate mentioned earlier.</p> + +<p>In summary, it seems that there is little or no evidence beyond the +numbers of bones involved to support the hypothesis that the +concentration of bones in the fissures of Fort Sill represents the +remains of food of predators, and that the fissures were used as dens by +their predatory occupants. On the contrary, the evidence indicates that +the deposition of the bones in the fissures was secondary and that the +agency of transportation, deposition and accumulation of the bones was +an early Permian stream characterized by periodic flooding.<span class="pagenum"><a name="Page_307" id="Page_307">[Pg 307]</a></span></p> + + + +<hr style="width: 65%;" /> +<h2>LITERATURE CITED</h2> + + +<p><span class="smcap">Peabody, F. E.</span><br /> 1961. Annual growth zones in living and fossil +vertebrates. Jour. Morph. 108 (1): 11-62, 69 figs., January.</p> + +<p><span class="smcap">Romer, A. S.</span><br /> 1956. Osteology of the reptiles. The University of Chicago +Press, Chicago, xxi + 772 pp., 248 figs.</p> + +<p><span class="smcap">Romer, A. S.</span>, and <span class="smcap">Price, L. I.</span><br /> 1940. Review of the Pelycosauria. Geol. +Soc. America, Spec. Pap., 28: x + 538 pp., 71 figs., 46 pls., 8 tables, +December 6.</p> + +<p><span class="smcap">Vaughn, P. P.</span><br /> 1958. On a new pelycosaur from the lower Permian of +Oklahoma, and the origin of the family Caseidae. Jour. Paleont., +32:981-991, 1 fig., September.</p> + +<p><i>Transmitted March 15, 1962.</i></p> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Two New Pelycosaurs from the Lower +Permian of Oklahoma, by Richard C. Fox + +*** END OF THIS PROJECT GUTENBERG EBOOK PELYCOSAURS, L. 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Fox + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Two New Pelycosaurs from the Lower Permian of Oklahoma + +Author: Richard C. Fox + +Release Date: December 7, 2009 [EBook #30620] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK PELYCOSAURS, L. PERMIAN, OKLAHOMA *** + + + + +Produced by Chris Curnow, Woodie4, Joseph Cooper and the +Online Distributed Proofreading Team at https://www.pgdp.net + + + + + + + + + + + UNIVERSITY OF KANSAS PUBLICATIONS + + MUSEUM OF NATURAL HISTORY + + Volume 12, No. 6, pp. 297-307, 6 figs. + May 21, 1962 + + + Two New Pelycosaurs from the Lower Permian + of Oklahoma + + BY + + RICHARD C. FOX + + + UNIVERSITY OF KANSAS + LAWRENCE + 1962 + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, Henry S. Fitch, + Theodore H. Eaton, Jr. + + + Volume 12, No. 6, pp. 297-307, 6 figs. + Published May 21, 1962 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED BY + JEAN M. NEIBARGER, STATE PRINTER + TOPEKA, KANSAS + 1962 + + 29-3001 + + + + +Two New Pelycosaurs from the Lower Permian of Oklahoma + +BY + +RICHARD C. FOX + + +In the course of examining material from fissure deposits of early +Permian age collected from a limestone quarry near Fort Sill, Oklahoma, +the author recovered several tooth-bearing fragments of small +pelycosaurs. The fragments were examined, compared with descriptions of +known kinds appearing in the literature, and determined to be new genera +within the Nitosauridae (Edaphosauria) and Sphenacodontidae +(Sphenacodontia). + +Appreciation is expressed to Prof. Theodore H. Eaton, Jr., for +permission to examine the collections of the University of Kansas from +Fort Sill, and for the financial assistance furnished by his National +Science Foundation grant (NSF-G8624). I am grateful both to Prof. Eaton +and Mr. Dale L. Hoyt for their suggestions regarding this manuscript. +The accompanying figures have been drawn by the author. + + + + +Family NITOSAURIDAE + +=Delorhynchus priscus= new genus and new species + + +(_delos_, Gr., evident; _rhynchos_, Gr., neuter, nostril; _priscus_, L., +ancient. _Delorhynchus_ is masculine because of the ending that it +acquires when transliterated into Latin.) + +_Type specimen._--Fragmentary left maxilla, bearing four teeth, KU +11117. + +_Referred specimens._--Fragmentary right maxilla having four teeth, KU +11118; fragmentary left maxilla having four teeth, the most posterior of +which has been broken, KU 11119. + +_Horizon and locality._--A fissure deposit in the Arbuckle limestone at +the Dolese Brothers Limestone Quarry, approximately six miles north of +Fort Sill, in sec. 31, T. 4 N, R. 11 W, Comanche County, Oklahoma. These +sediments are of early Permian age, possibly equivalent to the Arroyo +formation, Lower Clear Fork Group of Texas (Vaughn, 1958: 981). + +_Diagnosis._--Small; marginal teeth conical, slender and recurved at +tips; marginal tooth-row without caniniform enlargement; narial opening +enlarged and bordered dorsally, posteriorly and ventrally by maxilla; +maxilla with foramen opening laterally at posteroventral corner of +naris. + +_Description_ (based on 3 maxillary fragments, see Table 1).--Each of +the maxillary fragments bears four thecodont teeth. These are conical, +slender and sharply pointed; in their distal third they are slightly +recurved, laterally compressed, and have anterior and posterior +non-serrated cutting edges. In medial aspect at their bases, the teeth +are longitudinally striated. The bases of the teeth are circular in +cross-section and are slightly bulbous. There is no caniniform +enlargement of any of the teeth, the longest tooth of each fragment +being differently placed in the series of teeth and little longer than +the others. There is no swelling on either the internal or external +surfaces of the maxillae. The teeth are in a continuous series; no +diastema or maxillary step is evident. + +[Illustration: FIGURES 1-3. _Delorhynchus priscus_, lower Permian, 6 +miles north of Fort Sill, Comanche County, Oklahoma. All x 3. + +FIG. 1. KU 11117 (type specimen), lateral view of left maxilla. +FIG. 2. KU 11118, lateral view of right maxilla. +FIG. 3. KU 11119, lateral view of left maxilla.] + +The fragments have been broken along similar lines of fracture, and each +is approximately rhomboidal in shape. The maxilla encircles the +posterior border of the naris and extends dorsally above the naris to an +extent sufficient to indicate the probable exclusion of the lacrimal +bone from the narial border. At the posteroventral corner of the naris a +foramen opens onto the lateral surface of the maxilla. The opening is +the entrance to a canal that runs posteriorly above the tooth-row +throughout the length of each specimen. Beneath the naris the maxilla +extends as a broad tapering shelf, the ventral surface of which +articulates with the premaxilla. The narial rim is wide, but wider +ventrally than dorsally. The plane of the narial rim is oblique to the +lateral surface of the maxilla. The external surface of each fragment is +grooved and pitted. The ossification of each fragment appears to have +been complete. + +TABLE 1.--DIMENSIONS, IN MILLIMETERS, OF THREE MAXILLARY FRAGMENTS OF +DELORHYNCHUS PRISCUS + +Key: + +A. Anterior height of fragment +B. Posterior height of fragment +C. Length of fragment at tooth-row +D. Dorsal length of fragment +E. Mean length of teeth +F. Anterior width of naris + +============================================================ +CATALOGUE NUMBER | A. | B. | C. | D. | E. | F. +AND MEAN | | | | | | +-----------------+------+------+------+------+------+------ +KU 11117 | 6.0 | 8.0 | 6.0 | 8.0 | 3.0 | 3.0 +KU 11118 | 6.0 | 6.0 | 9.0 | 8.0 | 2.0 | 3.0 +KU 11119 | 6.6 | 8.0 | 10.0 | 11.0 | ? | 4.6 +-----------------+------+------+------+------+------+------ +Mean | 6.2 | 7.3 | 8.3 | 9.0 | 2.5 | 4.5 +-----------------+------+------+------+------+------+------ + +_Discussion._--The Nitosauridae are small primitive edaphosaurs with a +moderately elongate face, sharp subisodont teeth, little development of +canines and few specializations. The jaw is of a primitive type and +articulates on a level with the tooth-row. The palatal dentition is +primitive (Romer, 1956:280). The nitosaurids are thought to be related +to the later Caseidae, and the most obvious structural similarities are +found in the postcranial skeleton (Vaughn, 1958:989). Cranial +resemblances between the families are fewer, but nevertheless indicate +that a nitosaurid-caseid relationship exists. + +Vaughn (1958) described a small pelycosaur, _Colobomycter pholeter_ +(Eothyrididae, Ophiacodontia) that structurally resembles the Caseidae. +This individual also was obtained from the Fort Sill locality. In +Vaughn's opinion the features of _Colobomycter_ indicate a close +relationship between eothyridids and caseids and the possibility that +the caseids may well have been of eothyridid rather than nitosaurid +derivation. + +In view of this historical uncertainty of the relationships between the +Nitosauridae, the Eothyrididae and the Caseidae, it is well to consider +how the maxillary fragments described above differ from and resemble +representatives of each of these three families as reported in the +literature. + +_Delorhynchus_ resembles _Colobomycter_ in size. The mean +extra-maxillary length of the undamaged teeth of the three fragments is +2.5 mm., equal to that reported by Vaughn (1958:985) for teeth about +midway in the postcanine series of _Colobomycter_. None of the teeth of +_Delorhynchus_ extends beyond the maxillary rim as far as does the +canine of _Colobomycter_ (3.5 mm.). + +The teeth in both genera are conical and sharply pointed. The naris in +each is enlarged, and the lacrimal is excluded from the narial margin in +each (by inference in _Delorhynchus_.) + +The differences between the maxillae of _Colobomycter_ and +_Delorhynchus_ are most striking in the lack of canines in the latter +and the correlated absence of modifications of the maxillary for support +of canines. Additionally, _Delorhynchus_ bears an infraorbital canal in +contrast to the groove in similar position in _Colobomycter_. The +recurvature of the four teeth present in the fragments of _Delorhynchus_ +differs from that in the teeth of _Colobomycter_ in which only the +canine and precanine are recurved. Vaughn implies that anterior and +posterior cutting edges extend the length of the teeth in +_Colobomycter_; these are restricted to the distal third of the teeth in +_Delorhynchus_. The external surfaces of the maxillae of _Delorhynchus_ +are pitted and ridged; Vaughn was unable to discern sculpturing of the +corresponding surfaces in _Colobomycter_. + +_Delorhynchus_ resembles the nitosaurids in size, the shape and +sharpness of the teeth, their recurvature and the slight enlargement of +their bases, the exclusion of the lacrimal bone from the narial margin +(in _Mycterosaurus_) and the apparent lack of a special canine pair of +teeth. Resemblances to the caseids are to be noted in the enlargement of +the naris (4.5 mm. in height as opposed to 1.7 mm. in _Colobomycter_), +lack of development of canines, presence of an infraorbital canal (in +_Cotylorhynchus_) and absence of many replacement gaps in the marginal +row of teeth. + +The absence of caniniform enlargement and the extension of the maxilla +dorsad of the naris exclude _Delorhynchus_ from the Eothyrididae +(Ophiacodontia) but are no bar to its inclusion in the Nitosauridae +(Edaphosauria). The marginal teeth of _Delorhynchus_ are simple and +primitive, being much like those of the nitosaurids that are described +in the literature. + +The large narial opening and its posterior, dorsal and ventral enclosure +by the maxilla, the infraorbital canal, and the sculptured external +surfaces of the maxillary fragments indicate that _Delorhynchus_, in +these features at least, is close to achieving the caseid grade. + + + + +Family SPHENACODONTIDAE + +=Thrausmosaurus serratidens= new genus and new species + + +(_Thrausmosaurus_ is formed from the neuter Greek noun, _thrausma_, +meaning fragment, and the masculine Greek noun, _sauros_, meaning +reptile. The specific name, _serratidens_, is formed from the Latin +_serratus_, meaning serrate, and the masculine Latin noun, _dens_, +meaning tooth. The specific name is used as a substantive in apposition +with the generic name.) + +_Type specimen._--Fragmentary left dentary, bearing five teeth, the +most posterior of which is broken at the base, KU 11120. + +_Referred specimens._--Fragmentary ?left maxilla, having two teeth, KU +11121; fragmentary left dentary having two teeth, KU 11122. + +_Horizon and locality._--From the early Permian fissure deposits in the +Arbuckle limestone of the Dolese Brothers Limestone Quarry, +approximately 6 miles north of Fort Sill, in sec. 31, T. 4N, R. 11 W, +Comanche County, Oklahoma. + +_Diagnosis._--Small; teeth thecodont, compressed laterally, recurved +distally, and bearing anterior and posterior cutting edges; anterior +serrations limited to recurved portions of teeth, posterior serrations +extending nearly entire length of teeth; lateral compression of teeth +more pronounced medially than laterally; bases of teeth expanded. + +_Description._--The type specimen is 16 mm. long. It bears five teeth +that are implanted in a straight row. Empty sockets are present between +the first and second teeth, and the third and fourth teeth. The first +tooth is 3.0 mm. long, the middle two are each 2.5 mm. long, and the +fourth tooth is 2.0 mm. long. The fifth tooth is broken off at its base. + +The empty sockets are large. The mouth of each is circular and +approximately 2.0 mm. in diameter. Both sockets are 1.25 mm. deep. The +bases of the teeth are expanded to fill the sockets, although the blades +of the teeth arise from only the lateral portions of the bases. The edge +of the dentary rises above the bases of the teeth medially, thereby +producing a small depression at the junction of each base with the +dentary bone. + +The lateral compression of the teeth is pronounced but asymmetrical, in +that the lateral surface of each blade is more convex than the medial +surface. + +[Illustration: FIGURES 4-6. _Thrausmosaurus serratidens_, lower Permian, +6 miles north of Fort Sill, Comanche County, Oklahoma. All x 3. + +FIG. 4. KU 11120 (type specimen), lateral view of left dentary. +FIG. 5. KU 11121, lateral view of ?left maxilla. +FIG. 6. KU 11122, lateral view of left dentary.] + +The recurvature of the anterior cutting edges is much more severe than +that of the posterior edges, but the recurvature of both is limited to +the distal half of each tooth. + +The serrations of the cutting edges are not visible to the naked eye and +are limited on the anterior edges of the teeth to those portions of the +blades that are recurved. The posterior serrations extend nearly to the +junction of the blade of each tooth with its base. The serrations tend +to be more nearly crenulate than cuspidate. + +A portion of the lateral wall of the dentary surrounding the Meckelian +canal is present. The external surface of the wall is gently convex and +smooth, without sculpturing. The internal surfaces of the canal are +unmarked either by muscle scars or foramina. + +The fragment is a piece from the posterior portion of the dentary, since +the decrease in height from the first tooth to the fourth is pronounced. + +KU 11122, a fragment of the left dentary bearing two teeth, is 7.5 mm. +long. The anterior tooth is 3.0 mm. long; the posterior tooth is 3.5 mm. +long. The shape of the teeth and their implantation conform to the +description of the type specimen. The lateral surface of the fragment is +smooth and gently convex. What little is present of the surface +bordering the Meckelian canal is unmarked. + +The ?maxillary fragment bears two teeth which are 3.0 mm. long, and +which conform in their characters to the type. The lateral, medial and +ventral surfaces of the fragment have been sheared off, so that an exact +identification of the bone is impossible. Presumably the fragment is too +deep dorsoventrally to be a piece of the dentary, and no sign of the +Meckelian canal is present. + +_Discussion._--The implantation, lateral compression, recurvature and +cutting edges of the teeth borne by these fragments make clear their +sphenacodontid nature. The characters of the fragments are too few to +determine subfamilial affinities, however. That the fragments are the +remains of adult animals can be only surmised from the lack of bones or +teeth of large pelycosaurs in the extensive collections of the +University of Kansas from the Fort Sill locality. + +If _Thrausmosaurus_ is, in fact, adult, the genus is an unusually small +sphenacodontid, and of significance both on that account and because of +the resemblance of the teeth presently known to those of its far larger +relatives. + +_The Fort Sill Locality._--Peabody (1961) suggested that the fissures of +Fort Sill had been used as dens by predatory animals in the early +Permian, and that the unusually abundant bones in the fissures were the +remains of animals eaten there by these occupants. Evidence now known to +me affords an alternative explanation that is presented here as a +preliminary to a more complete study of the fauna and paleoecology of +these deposits currently being undertaken. + +The suggestion that the skeletal material found in the fissures is the +remnant of the prey of other animals is questionable because of: + + 1. The absence of tooth marks on the fossils. + + 2. The recovery from the matrix of skulls and portions of + articulated skeletons that are undamaged or damaged only by + pressure after burial. + + 3. The rarity in the deposits of animals of larger body size than + _Captorhinus_, the exceptions being a few limb fragments and skull + fragments of labyrinthodont or pelycosaurian nature. + + 4. The absence of coprolites in the matrix. + +If the fissures were the dens of predators, at least some and probably +many of the bones would show tooth marks. A predator feeding on other +animals would be expected to leave some evidence of its habits on the +bones of its prey. No such evidence is known to me, either from my own +examination of several thousand bones or from the reports in the +literature by others who have studied aspects of the early Permian fauna +of Fort Sill. + +If the predators were larger than _Captorhinus_ and occupied the +fissures for a long enough time to account for the accumulation of the +tremendous numbers of individuals that are represented, a considerable +amount of the skeletal material of the larger animals would be present +in the fissure deposits. Even if for some reason the predators died in +areas other than within the fissures, thereby accounting for the absence +of large bones, coprolites should appear in the deposits if, in fact, +the fissures were feeding places. In view of the nearly undamaged +condition of many of the bones recovered from the fissures, it is +reasonable to expect that fecal material would be preserved. + +The character of the matrix of the deposits varies from a homogeneous +clay to clay interrupted by layers of soft, limey, conglomeratic rock, +to a hard, well-cemented, calcareous conglomerate. In general the bone +in each kind of matrix is colored characteristically and exhibits a +characteristic degree of wear. The bones entrapped in the homogeneous +clay are relatively few, black, usually disarticulated, little worn and +not unduly fragmented; consequently the discovery of undamaged limb +bones, for example, from this kind of matrix is not unusual. The bones +found in the stratified portion of the matrix are more numerous within +the layers of conglomerate than between. The bones are black, brown or +white, highly fragmented and waterworn to a variable degree. The +fragments recovered from the hard, calcareous matrix are numerous, range +in color from white through various shades of brown, to black, are +highly fragmented, and are usually worn by water. + +These categories for bone and matrix, however, are not mutually +exclusive, since bones of any of these colors and exhibiting any degree +of wear and fragmentation are found in any of the kinds of matrix +described above. That water was the agent of wear is suggested by the +highly polished appearance of the worn bones and pebbles that are found +in the matrix. + +The variability of the matrix and of the color and condition of the +bones indicates that the agencies of burial and fossilization differed +from time to time and that the agency of transportation of the bones +from the site of burial to the fissures was running water. One can +easily visualize a stream coursing the early Permian landscape that was +subject to periodic flooding and droughts. Along the banks of the stream +and in its pools lived a variety of microsaurs, captorhinids, small +labyrinthodonts and small pelycosaurs. Some of the animals, after they +died, were either buried near the site of their death or were swept +along and buried in sediments further downstream. Burial was for a +length of time sufficient to impart a color to the bones characteristic +of the site in which they were buried. Later floods reexposed the sites +of burial, picked up the bones and carried them to the openings into the +fissures. Presumably, too, a proportion of the bones was carried to the +fissures without previous burial. + +The differences in wear exhibited by different bones within the same +block of matrix is attributable to differences in distance that the +bones were transported before final deposition. The final sites of +deposition, the fissures, were inundated occasionally by floods alone, +or because of changes in location of the channel of the stream at the +time of flooding. The periodicity of deposition of the sediments within +portions of the fissures is indicated by the stratification of the bone +conglomerate mentioned earlier. + +In summary, it seems that there is little or no evidence beyond the +numbers of bones involved to support the hypothesis that the +concentration of bones in the fissures of Fort Sill represents the +remains of food of predators, and that the fissures were used as dens by +their predatory occupants. On the contrary, the evidence indicates that +the deposition of the bones in the fissures was secondary and that the +agency of transportation, deposition and accumulation of the bones was +an early Permian stream characterized by periodic flooding. + + + + +LITERATURE CITED + + +PEABODY, F. E. + 1961. Annual growth zones in living and fossil + vertebrates. Jour. Morph. 108 (1): 11-62, 69 figs., January. + +ROMER, A. S. + 1956. Osteology of the reptiles. The University of Chicago + Press, Chicago, xxi + 772 pp., 248 figs. + +ROMER, A. S., and PRICE, L. I. + 1940. Review of the Pelycosauria. Geol. + Soc. America, Spec. Pap., 28: x + 538 pp., 71 figs., 46 pls., + 8 tables, December 6. + +VAUGHN, P. P. + 1958. On a new pelycosaur from the lower Permian of + Oklahoma, and the origin of the family Caseidae. Jour. Paleont., + 32:981-991, 1 fig., September. + +_Transmitted March 15, 1962._ + + + + + +End of the Project Gutenberg EBook of Two New Pelycosaurs from the Lower +Permian of Oklahoma, by Richard C. Fox + +*** END OF THIS PROJECT GUTENBERG EBOOK PELYCOSAURS, L. 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