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+The Project Gutenberg EBook of A New Order of Fishlike Amphibia From the
+Pennsylvanian of Kansas, by Theodore H. Eaton and Peggy Lou Stewart
+
+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: A New Order of Fishlike Amphibia From the Pennsylvanian of Kansas
+
+Author: Theodore H. Eaton
+        Peggy Lou Stewart
+
+Release Date: January 23, 2010 [EBook #31050]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK NEW ORDER OF FISHLIKE AMPHIBIA ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper, Diane Monico, and
+the Online Distributed Proofreading Team at
+https://www.pgdp.net
+
+
+
+
+
+
+
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS
+
+MUSEUM OF NATURAL HISTORY
+
+Volume 12, No. 4, pp. 217-240, 12 figs.
+May 2, 1960
+
+
+A New Order of Fishlike Amphibia
+From the Pennsylvanian of Kansas
+
+BY
+
+THEODORE H. EATON, JR., AND PEGGY LOU STEWART
+
+
+UNIVERSITY OF KANSAS
+LAWRENCE
+1960
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
+Robert W. Wilson
+
+
+Volume 12, No. 4, pp. 217-240, 12 figs.
+Published May 2, 1960
+
+
+UNIVERSITY OF KANSAS
+Lawrence, Kansas
+
+
+PRINTED IN
+THE STATE PRINTING PLANT
+TOPEKA, KANSAS
+1960
+
+28-2495
+
+
+
+
+A New Order of Fishlike Amphibia
+From the Pennsylvanian of Kansas
+
+BY
+
+THEODORE H. EATON, JR., AND PEGGY LOU STEWART
+
+INTRODUCTION
+
+
+A slab of shale obtained in 1955 by Mr. Russell R. Camp from a
+Pennsylvanian lagoon-deposit in Anderson County, Kansas, has yielded in
+the laboratory a skeleton of the small amphibian _Hesperoherpeton
+garnettense_ Peabody (1958). This skeleton provides new and surprising
+information not available from the holotype, No. 9976 K. U., which
+consisted only of a scapulocoracoid, neural arch, and rib fragment. The
+new specimen, No. 10295 K. U., is of the same size and stage of
+development as the holotype and it is thought that both individuals are
+adults.
+
+The quarry, University of Kansas Museum of Natural History Locality KAN
+1/D, is approximately six miles northwest of Garnett, Anderson County,
+Kansas, in Sec. 5, T. 19S, R. 19E, 200 yards southwest of the place
+where _Petrolacosaurus kansensis_ Lane was obtained (see Peabody,
+1952). The Rock Lake shale, deposited under alternately marine and
+freshwater lagoon conditions, is a thin member of the Stanton limestone
+formation, Lansing group, Missourian series, and thus is in the lower
+part of the Upper Pennsylvanian.
+
+Peabody (1958) placed _Hesperoherpeton_ in the order Anthracosauria,
+suborder Embolomeri, family Cricotidae. Study of the second and more
+complete specimen reveals that _Hesperoherpeton_ is unlike the known
+Embolomeri in many important features. The limbs and braincase are more
+primitive than those so far described in any amphibian. The vertebrae
+are comparable to those of Ichthyostegalia (Jarvik, 1952), as well as
+to those of Embolomeri. The forelimb is transitional between the
+pectoral fin of Rhipidistia and the limb of early Amphibia. The pattern
+of the bones of the forelimb closely resembles, but is simpler than,
+that of the hypothetical transitional type suggested by Eaton (1951).
+The foot seemingly had only four short digits. The hind limb is not
+known.
+
+The new skeleton of _Hesperoherpeton_ lies in an oblong block of limy
+shale measuring approximately 100 × 60 mm. After preparation of the
+entire lower surface, the exposed bones and matrix were embedded in
+Bioplastic, in a layer thin enough for visibility but giving firm
+support. Then the specimen was inverted and the matrix removed from the
+opposite side; this has not been covered with Bioplastic. The bones lie
+in great disorder, except that some parts of the roof of the skull are
+associated, and the middle section of the vertebral column is
+approximately in place. The bones of the left forelimb are close
+together but not in a natural position. The tail, pelvis, hind limbs
+and right forelimb are missing. Nearly all the bones present are
+broken, distorted by crushing, incomplete and scattered out of place,
+probably by the action of currents. The complete skeleton, in life,
+probably measured between 150 and 200 mm. in length.
+
+The specimen was studied at the Museum of Natural History, University
+of Kansas, with the help of a grant from the National Science
+Foundation, number NSF-G8624. The specimen was discovered in the slab
+by Miss Sharon K. Moriarty, and was further cleaned by the authors. Mr.
+Merton C. Bowman assisted with the illustrations. We are indebted to
+Dr. Robert W. Wilson for critical comments.
+
+
+SKULL
+
+_Dorsal Aspect_ (Figs. 1, 2)
+
+In reconstruction, the skull measures approximately 8.0 mm.
+dorsoventrally at the posterior end. The height diminishes anteriorly
+to about 1.5 mm. at the premaxillary. The length is about 15.5 mm. in
+the median line, or 24.0 mm. to the tip of the tabular, and the width
+about 16.0 mm. posteriorly. The snout is blunt, continuing about 1-2
+mm. anterior to the external nares. Each of the tabulars has a slender
+posterior process 5.0 mm. long, which probably met the supracleithrum;
+the intertabular space is about 8.5 mm. wide. The orbits are
+approximately 5.5 mm. in diameter and extend from the maxillary to
+within about 3.0 mm. of the midline dorsally. The pineal opening is 1.8
+mm. anterior to the occipital margin of the skull.
+
+Reduction of bones at the back of the skull seems to have eliminated
+any dermal elements posterior to the squamosal, while enlargement of
+the orbit has removed most of the postorbital series, leaving the
+squamosal as the only cheekbone. There is apparently no jugal or
+postfrontal.
+
+The squamosal of _Acanthostega_ (Jarvik, 1952) is articulated under the
+tabular and reaches forward and down, much as if it were an opercular
+in reversed position. Internally, it must lie against the otic capsule
+below the tabular, partially concealing the stapes. The bone that we
+suppose to be the squamosal of _H. garnettense_ is of similar shape, of
+about the same size and has internally an articular surface at one
+corner, bounded by a pair of ridges in the shape of a V. This articular
+surface probably fitted on a lateral process extending from the roof of
+the neurocranium, over the front of the otic capsule.
+
+The premaxillary extends posterolaterally to a distance 5.5 mm. from
+the midline and attains a width at its broadest point of about 1.5 mm.
+The posterior edge is slightly concave and in part forms the anterior
+border of the naris.
+
+[Illustration: FIG. 1. _Hesperoherpeton garnettense_ Peabody. Skull,
+dorsal view. Postorbital processes of the neurocranium are shown in
+dotted outline. KU 10295, × 4.]
+
+The nasal is triangular and, with the lacrimal, forms the medial border
+of the naris. The length of the medial side of the nasal bone is
+approximately 5.0 mm., the transverse width is 3.8 mm., and the extent
+of the posterolateral border is 5.5 mm.
+
+The maxillary meets the premaxillary lateral to the naris, borders the
+naris posteroventrally, and continues posteriorly beneath the orbit, of
+which it forms the external border. The maxillary is about 8.5 mm.
+long, and immediately anterior to the orbit has a maximum width of 1.3
+mm.
+
+The lacrimal fills the remaining rim of the narial opening between the
+nasal and maxillary, and extends to the anterior edge of the orbit. The
+length, from naris to orbit, is 4.2 mm.; the width ranges from 1.0 mm.
+anteriorly to 2.5 mm. posteriorly.
+
+[Illustration: FIG. 2. _Hesperoherpeton garnettense_ Peabody. Skull,
+lateral view, showing relatively large orbit and absence of smaller
+circumorbital bones. KU 10295, × 4.]
+
+The external naris is approximately 1.0 mm. in diameter. It is slightly
+anterodorsal to the internal naris and 4.0 mm. lateral to the midline.
+
+The dorsal margin of the orbit appears to be formed by the frontal. The
+anterior part of this margin, however, may be formed by a prefrontal,
+which is not clearly set off by a suture. The frontal extends 3.8 mm.
+in the midline, and anteriorly and laterally borders the nasal and
+lacrimal, respectively. A faint pattern of pitting radiates on the
+surface from the center of ossification of the frontal. There is also a
+pit indicating the presence of a supraorbital sensory pore.
+
+The parietal bones enclose the pineal opening, approximately 2.5 mm.
+posterior to the suture with the frontal. The foramen is about 0.5 mm.
+in diameter. Laterally the parietal meets the medial angle of the
+postorbital and the medial border of the supratemporal. No bone of this
+animal shows the deep pitting and heavy ornamentation characteristic of
+many primitive Amphibia.
+
+The postorbital meets the anterolateral corner of the parietal for a
+distance of 0.5 mm., the anterior edge bordering the frontal bone and
+the orbit for a combined distance of about 3.0 mm. The lateral margin
+is slightly convex, and is probably interrupted behind by the anterior
+point of the tabular. Medially, the concave margin of the postorbital
+meets the supratemporal for about 3.5 mm.
+
+The supratemporal is thus wedge-shaped and located between the parietal
+and the postorbital. The posterior edge of the supratemporal protrudes
+as a convex border slightly behind the end of the parietal, and
+measures 3.0 mm. around the curve to the parietal suture.
+
+[Illustration: FIG. 3. _Hesperoherpeton garnettense_ Peabody. A, left
+squamosal, internal surface. B, left squamosal, external surface. C,
+right tabular internal surface. D, right tabular, external surface. KU
+10295, all × 4.]
+
+The squamosal (Fig. 3 A, B) is a large, somewhat rectangular bone
+extending from the back of the orbit to the posterior extremity of the
+cheek. It outlines almost entirely the posterior border of the orbit,
+the ventrolateral portion of the cheek region, and the lateral border
+of the top of the skull behind the orbit. Dorsally, the squamosal meets
+the anterior half of the tabular and the lateral border of the
+supratemporal. Near the anteroventral edge of the squamosal there is a
+small pit, probably related to a postorbital sensory pore in the skin.
+
+The tabular (Fig. 3 C, D) is pointed anteriorly, where it probably fits
+against the lateroposterior edge of the postorbital. The dorsal part of
+the bone flares out and down, forming a small otic notch at a point
+halfway back. Posteriorly, the flange attains a dorsoventral width of
+2.0 mm. at the edge of the notch. The slender posterior process of the
+tabular which continues beyond the flange is approximately 0.5 mm. in
+diameter and 5.0 mm. long.
+
+_Ventral Aspect_ (Fig. 4)
+
+The palatal view of the skull shows the paired premaxillary, maxillary,
+palatine, pterygoid, and quadrate bones. The openings for the internal
+nares, the ventral orbital fenestrae, and the subtemporal fossae are
+readily recognized. The quadrate processes extend posteriorly leaving a
+large gap medially at the posterior end of the skull.
+
+[Illustration: FIG. 4. _Hesperoherpeton garnettense_ Peabody. Palate
+reconstructed; ventral aspect at left, showing teeth, dorsal aspect at
+right. KU 10295, × 4.]
+
+The left quadrate appears to be in place on the posterior prong of the
+pterygoid. The dorsal side of the quadrate is grooved between two
+anterolaterally directed ridges. The groove, which probably held the
+end of the stapes, extends about half the width of the quadrate itself.
+The width of the quadrate is 4.0 mm., the length is 4.5 mm. medially
+and about 2.0 mm. laterally. In ventral view the quadrate appears to
+project laterally, but is incomplete and its shape uncertain. The
+distance from the posterior end of the quadrate to the visible
+posterior edge of the orbital fenestra, which opens ventrally, is 10.0
+mm.
+
+This region between the quadrate and the orbit is occupied by a
+pterygoid with three projections. Anteriorly, the pterygoid outlines
+most of the posterior edge of the orbit (a distance of about 6.5 mm.).
+A lateral process separates the orbit from the subtemporal fossa. A
+posteriorly directed edge defines the fossa, which extends about 6.5
+mm. anteroposteriorly. The lateral process of the pterygoid terminates
+10.0 mm. from the midline. Both the lateral and posterior pterygoid
+processes are approximately 2.0 mm. wide. The greatest width of the
+subtemporal fossa is about 2.0 mm. The medial border of the orbital
+fenestra is missing, but apparently consisted of the pterygoid for at
+least the posterior half.
+
+Along the posterior edge of the orbital fenestra, there is a narrow,
+dorsally projecting flange of the pterygoid. The lateral opening of the
+orbit is approximately 7.5 mm. wide.
+
+The remaining border of the orbital fenestra on the anterior and medial
+sides is formed by a bone occupying the position of palatine and vomer;
+for convenience we designate this as palatine. When reconstructed in
+its probable position in relation to the pterygoid, the left palatine
+lacks a section, on its medial and posterior edges, measuring about 2.5
+mm. by 9.0 mm. The lateral margin of the palatine is convex; about 5.5
+mm. anterior to the orbit this margin curves into a strong anteriorly
+pointing projection, medial to which is seen the internal narial
+opening. The remaining anterior edge is slightly convex, smoothly
+rounded, and meets the midline about 9.0 mm. anterior to the pterygoid.
+
+The void area medial to the palatine and anterior to the pterygoid does
+not fit any bone which we can recognize as the parasphenoid. It is thus
+suspected that this area is covered in part by the missing edge of the
+palatine and partly by an anteromedial extension of the pterygoid. Of
+course a parasphenoid may also have been present.
+
+The position, length, and shape of the premaxillary shown in palatal
+view (Fig. 4) are primarily based upon the dorsal appearance since
+ventrally most of it cannot be seen. At the point where it forms the
+anterior border of the internal naris, the premaxillary is slightly
+wider than the maxillary and seems to become narrower as it approaches
+the midline.
+
+The ethmosphenoid, which we cannot identify, may have been exposed in a
+gap between the premaxillary and the palatine. The gap measures
+approximately 8.0 mm. wide and ranges up to 1.0 mm. anteroposteriorly.
+
+The maxillary begins at a suture with the premaxillary lateral to the
+naris and continues posteriorly, bordering the orbit with a width of
+about 1.2 mm. It then tapers to a point approximately 2.0 mm. anterior
+to the lateral projection of the pterygoid. The width of the maxillary
+at this point is 0.8 mm. and the posterior end is broken; probably when
+complete it approached the pterygoid, and either met the latter or had
+a ligamentous connection with it. As nearly as can be determined, the
+total length of the maxillary is approximately 12.0 mm.
+
+The teeth on the maxillary are small and seem to be in two longitudinal
+rows. The palatine bears two large, grooved teeth anteriorly; the first
+is approximately 1.0 mm. posteromedial to the naris and the second is
+about 3.0 mm. posterior and slightly lateral to the naris. The flat
+ventral surfaces of the palatine and pterygoid bear numerous small
+teeth distributed as shown in Fig. 4.
+
+_Braincase and Occipital Region_ (Fig. 5)
+
+The parts of the neurocranium are scattered, disconnected and
+incomplete, but it is possible to make out a number of features of the
+otico-occipital section with fair assurance. In posterior view the
+notochordal canal and foramen magnum are confluent with each other, and
+of great size relative to the skull as a whole. The notochordal canal
+measures 2.8 mm. in diameter, and the foramen magnum about 4.0 mm. The
+crescent-shaped supraoccipital rests on the upright ends of the
+exoccipitals, but between the latter and the basioccipital no sutures
+can be seen. Probably the whole posterior surface of the braincase
+slanted posteroventrally; consequently the rim of the notochordal canal
+was about 3.0 mm. behind the margin of the parietals.
+
+The U-shaped border of the notochordal canal is a thick, rounded bone,
+comparable in appearance to the U-shaped intercentra of the vertebrae.
+This bone apparently rested upon a thinner, troughlike piece (Fig. 5 B)
+forming the floor of the braincase. The latter is broad, shallow,
+concave, open midventrally and narrowing anteriorly to form a pair of
+articular processes. Since no sutures can be seen in this structure, it
+probably is the ventral, ossified portion of the basioccipital. Watson
+(1926, Fig. 4 B) illustrates the floor of the braincase in
+_Eusthenopteron_, with its more lateral, anterior portion labelled
+prootic, but in our specimen the corresponding part could scarcely have
+formed the anterior wall of the otic capsule, being entirely in the
+plane of the floor. The two articular surfaces anteriorly near the
+midline suggest that a movable joint existed between the
+otico-occipital part of the braincase and the ethmosphenoid part, as in
+Rhipidistia (Romer, 1937). We have found nothing in the specimen that
+could be referred to the ethmosphenoid; it may have been unossified.
+
+[Illustration: FIG. 5. _Hesperoherpeton garnettense_ Peabody, KU 10295,
+× 4. A, occipital view of skull; B, basioccipital bone in dorsal
+(internal) view.]
+
+The otic capsules appear to have rested against lateral projections of
+the basioccipital. The single otic capsule that can be seen (the
+right) is massively built, apparently ossified in one piece, with a
+shallow dorsomedial excavation, probably the vestige of a supratemporal
+fossa. On the lateral face is a broad, shallow depression dorsally, and
+a narrower, deeper one anteroventrally; these we suppose to have
+received the broader and narrower heads of the stapes, respectively.
+The posterior wall of the otic capsule we have designated opisthotic in
+the figure. Anterior to the otic capsule the lateral wall of the
+braincase cannot be seen, and may not have been ossified.
+
+The roof of the braincase is visible in its ventral aspect, extending
+from approximately the occipital margin to a broken edge in front of
+the parietal foramen, and laterally to paired processes which overlie
+the otic capsules directly behind the orbits (see dotted outlines in
+Fig. 1). Each of these postorbital processes, seen from beneath,
+appears to be the lateral extension of a shallow groove beginning near
+the midline. Presumably this section of the roof is an ossification of
+the synotic tectum. It should be noted that the roof of the braincase
+proper is perfectly distinct from the overlying series of dermal bones,
+and that the parietal foramen can be seen in both. The roof of the
+braincase in our specimen seems to have been detached from the
+underlying otic capsules and the occipital wall.
+
+The bone that we take to be the stapes is blunt, flattened (perhaps by
+crushing), 5.0 mm. in length, and has two unequal heads; its width
+across both of these is 4.0 mm. The length is appropriate to fit
+between the lateral face of the otic capsule and the dorsal edge of the
+quadrate; the wider head rests on a posterodorsal concavity on the otic
+capsule, and the smaller fits a lower, more anterior pit. Laterally the
+stapes carries a short, broad process that probably made contact with a
+dorsally placed tympanic membrane. Thus the bone was a hyomandibular in
+the sense that it articulated with the quadrate, but it may also have
+served as a stapes in sound-transmission. It contains no visible canal
+or foramen.
+
+_Mandible_ (Fig. 6)
+
+The crushed inner surface of the posterior part of the left mandible
+and most of the external surface of the right mandible are preserved in
+close proximity. Although the whole length of the tooth-bearing margins
+is missing, some parts of six elements of the right mandible can be
+seen. The pattern of sutures and the general contour closely resemble
+those of _Megalichthys_ (Watson, 1926, Figs. 37, 38) and other known
+Rhipidistia.
+
+The anteroposterior length of the mandible is about 23.8 mm., and the
+depth is 3.8 mm. The dentary extends approximately 17.6 mm. back from
+the symphysis, and its greatest width is probably 2.0 mm. Its lower
+edge meets all the other lateral bones of the jaw. The splenial and
+postsplenial form the curved anteroventral half of the jaw for a
+distance of about 9.0 mm. The fragmented articular, on the posterior
+end of the jaw, is 4.0 mm. long and 2.0 mm. deep, exhibiting a broken
+upper edge; presumably the surface for articulation with the quadrate
+was a shallow concavity, above the end of the articular.
+
+[Illustration: FIG. 6. _Hesperoherpeton garnettense_ Peabody. Right
+mandible, lateral view, KU 10295, × 4. External surfaces are pitted;
+broken surfaces are coarsely stippled.]
+
+
+VERTEBRAE (Fig. 7)
+
+The vertebrae that are visible from a lateral view are crushed and
+difficult to interpret. It is possible, nevertheless, to see that the
+trunk vertebrae resemble those of Ichthyostegalia (Jarvik, 1952, Fig.
+13 A, B), except that the pleurocentra are much larger. A few parts of
+additional vertebrae can be seen, but they are so scattered that it is
+impossible to be sure of their original location. Therefore comparisons
+between different regions cannot yet be made.
+
+The U-shaped intercentrum encloses the notochord and occupies an
+anteroventral position in the vertebra. Anteriorly, each intercentrum
+articulates with the pleurocentra of the next preceding vertebra by
+slightly concave surfaces. Dorsolaterally there is an articular surface
+for the capitulum of the rib.
+
+The two pleurocentra of each vertebra are separate ventrally as well as
+dorsally, but form thin, broad plates of about the same height as the
+notochord. The lateral surface appears to be depressed, allowing,
+perhaps, for movement of the rib. Above each pleurocentrum, on the
+lateral surface of the neural arch, there is a short diapophysis for
+articulation with the tuberculum of the rib.
+
+The margin of the neural spine is convex anteriorly and concave
+posteriorly, the tip reaching a point vertically above the
+postzygapophysis. The prezygapophysis of each vertebra articulates
+with the preceding postzygapophysis by a smooth dorsal surface. One
+nearly complete neural arch shows (Fig. 7 B) a pit above the neural
+canal, clearly corresponding to the canal for a dorsal ligament shown
+by Jarvik in _Ichthyostega_. Indeed this view of the neural arch and
+intercentrum together brings out the striking resemblance between the
+vertebrae of _Hesperoherpeton_ and those of the Ichthyostegids. The
+rounded intercentrum in both is an incomplete ring enclosing the
+notochordal canal.
+
+[Illustration: FIG. 7. _Hesperoherpeton garnettense_ Peabody. A, End
+view of incomplete vertebra, probably near anterior end of column. B,
+Neural arch and intercentrum in end view, showing probable association.
+C, Left lateral view of trunk vertebra. All figures: KU 10295, × 4.]
+
+TABLE 1.--AVERAGE MEASUREMENTS OF THE TRUNK VERTEBRAE (in mm.).
+NUMBERS IN PARENTHESES INDICATE THE NUMBER OF PIECES AVAILABLE FOR
+MEASURING
+
+----------------------------+------------+-------------+---------------
+            PARTS           | Ant.-post. | Dors.-vent. | Transv. width
+----------------------------+------------+-------------+---------------
+Neural spine                |  1.5 (3)   |   3.0  (3)  |     --
+----------------------------+------------+-------------+---------------
+Neural spine and arch       |  2.0 (4)   |   4.5? (4)  |     --
+----------------------------+------------+-------------+---------------
+Neural canal                |  2.0 (4)   |   2.0  (1)  |    1.0 (1)
+----------------------------+------------+-------------+---------------
+Intercentrum                |  1.5 (5)   |   3.5  (4)  |    3.0 (1)
+----------------------------+------------+-------------+---------------
+Pleurocentrum               |  1.5 (3)   |   3.0  (2)  |     --
+----------------------------+------------+-------------+---------------
+
+The shape, in end view, of a partly preserved neural arch (Fig. 7 A)
+seems to account for the incompleteness of the intercentrum just
+mentioned; the ventral edge of the arch is emarginate in such a way as
+to fit the dorsal surface of the notochord. The dorsal portion of this
+neural arch is not present (either broken or not yet ossified), but
+the opening of the neural canal is comparable in width to the foramen
+magnum. Hence this vertebra may be one of the most anterior in the
+column. In comparison with the trunk vertebrae seen farther posteriorly
+it appears that there may be a progressive ossification of neural
+arches toward their dorsal ends, and of intercentra around the
+notochord, with probable fusion of the intercentra and neural arches in
+the posterior part of the trunk. The notochord seems to have been
+slightly constricted by the intercentra, but not interrupted.
+
+
+RIBS
+
+The proximal ends of the ribs expand dorsoventrally to a width
+approximately four times that of their slender shafts. The tuberculum
+and capitulum on each of the trunk ribs are separated only by a shallow
+concavity. These two articular surfaces are so situated that the rib
+must tilt downward from the horizontal plane. The shaft flares
+terminally in some ribs, and the distal end is convex. Ribs in the
+trunk region differ little if any in size. Five that can be measured
+vary in length from 5.0 to 7.0 mm. One short, bent rib 3.5 mm. long
+perhaps is sacral or caudal.
+
+
+PECTORAL GIRDLE (Figs. 8, 9, 10)
+
+The right scapulocoracoid is almost complete, and the left one is
+present but partly broken into three pieces, somewhat pushed out of
+position. With the advantage of this new material, we may comment on
+the scapulocoracoid of _H. garnettense_ as described by Peabody (1958).
+In size and contour, the slight differences between the type (KU 9976)
+and the new skeleton (KU 10295) are considered to be no more than
+individual variation. We have redrawn the type (Fig. 8) in order to
+show the resemblances more clearly.
+
+The small sections that were missing from the type are present in KU
+10295. The jagged edge directly posterior to the area occupied by the
+neural arch in the type extends 0.5 mm. farther back in our specimen.
+The angle formed between the recurved dorsal ramus and the edge of the
+ventral flange is seen in our specimen to be less than 90°. The glenoid
+fossa, appearing as a concave articular surface for the cap of the
+humerus, was in part covered by cartilage and shows as "unfinished"
+bone (Peabody, 1958, p. 572); this area is more oval than triangular,
+as Peabody thought. The obstruction of a clear view of this part of the
+type is the result of the accidental position of a neural arch. The
+raised portion immediately dorsal to the glenoid fossa exhibits an
+unfinished surface, suggesting the presence of either cartilage or a
+ligament.
+
+[Illustration: FIG. 8. _Hesperoherpeton garnettense_ Peabody. Type
+specimen redrawn. Right scapulocoracoid in external view (at left), and
+internal view (at right). KU 9976, × 4.]
+
+
+[Illustration: FIG. 9. _Hesperoherpeton garnettense_ Peabody. Right
+scapulocoracoid in external view, showing part of interclavicle, and
+position occupied by clavicle. The specimen is flattened and lies
+entirely in one plane. KU 10295, × 4.]
+
+
+[Illustration: FIG. 10. _Hesperoherpeton garnettense_ Peabody. Right
+clavicle in external view. Anterior edge to right. KU 10295, × 4.]
+
+The right clavicle is complete, and resembles a spoon having a slender
+handle. The dorsal tip of the handle is L-shaped. The expanded ventral
+part is convex externally, and rested upon the anteroventral surface of
+the scapulocoracoid. The lateral edge next to the "stem" is distinctly
+concave, abruptly becoming similar in contour to the opposite edge, and
+giving the impression of an unsymmetrical spoon. The left clavicle is
+present in scattered fragments, its dorsal hooklike end being intact.
+
+The posterior end of the interclavicle lies in contact with the right
+scapulocoracoid. There are short lateral processes at the point where
+the interclavicle was overlapped by the clavicles, but we cannot be
+sure of the extent of this bone anteriorly or posteriorly.
+
+The presumed left cleithrum, a long rectangle, is approximately equal
+in length to the rodlike stem of the clavicle, and is about as wide as
+the dorsal L-shaped tip of the clavicle. The posterior end of the
+cleithrum presumably met the tip of the clavicle, while the rest of it
+was directed anteriorly and a little dorsally. There seems to be a
+small articular surface near the anterior extremity which suggests the
+presence of a supracleithrum. The upper border of the cleithrum is
+slightly convex and the lower concave.
+
+
+FORELIMB (Fig. 11)
+
+The left forelimb is the only one present and appears to be nearly
+complete, although the elements are scattered almost at random. The
+only parts of the forelimb known to be missing are two subterminal and
+two terminal phalanges, probably of the first and third digits, and the
+proximal end of the second metacarpal. The smooth and relatively flat
+surfaces suggest an aquatic rather than terrestrial limb; only the
+proximal half of the humerus bears any conspicuous ridges or
+depressions. As we restore the skeleton of the limb, several features
+are remarkable: The humerus, ulna, and ulnare align themselves as the
+major axis of the limb, each carrying on its posterior edge a process
+or flange comparable to those in the axial series of a rhipidistian
+fin. The remaining elements take positions comparable to the diagonally
+placed preaxial radials in such a fin. The digits appear to have been
+short, perhaps with no more than two phalanges. There is only one row
+of carpals present (the proximal row of other tetrapods). A second and
+third row would be expected in primitive Amphibia; if they existed in
+_Hesperoherpeton_ they must either have been wholly cartilaginous or
+washed away from the specimen. Neither of these alternatives seems at
+all likely to us in view of the well-ossified condition of the elements
+that are present, and the occurrence of both the proximal carpals and
+the metacarpals. The space available for metacarpals probably could not
+have contained more than the four that are recognized.
+
+[Illustration: FIG. 11. _Hesperoherpeton garnettense_ Peabody. Left
+forelimb, showing characters of both a crossopterygian fin and an
+amphibian foot. KU 10295, × 4.]
+
+The proximal end of the humerus is more rounded anteriorly than
+posteriorly, and has a thin articular border that bore a cartilaginous
+cap as the primary surface for articulation with the scapulocoracoid.
+Although the unfinished surface of the head extends down the anterior
+margin about a third the length of the humerus, the shaft has been
+broken and so twisted that the distal part is not in the same plane as
+the proximal. Immediately posterior to the cartilaginous cap is a
+round, deep notch bordered posteriorly by the dorsal process of the
+head.
+
+The shaft is longer and narrower than would be anticipated in a
+primitive amphibian limb (cf. Romer, 1947). The distal end bears two
+surfaces for articulation with the radius and ulna. The full extent of
+the former surface was not determined because the more anterior part of
+the expanded end is represented only by an impression. The surface
+nearest the ulna was partially rounded for articulation with that
+element, the remaining posterior edge being broadly concave. The most
+striking feature of the humerus is a slender hooklike process on the
+posterior edge near the distal end, probably homologous with (1) the
+posterior flange on the "humerus" in Rhipidistia, and (2) the
+entepicondyle of the humerus in _Archeria_ (Romer, 1957) and other
+tetrapods.
+
+The radius is about the same width proximally as distally. The
+curvature of the shaft is approximately alike on both sides. Distally
+the surface is rounded for articulation with the radiale and perhaps
+the intermedium.
+
+The proximal end of the ulna is similar to that of the radius but is
+slightly larger. Posteriorly, there is a short, broad expansion
+resembling the entepicondyle of the humerus, and even more nearly like
+the postaxial flanges in a crossopterygian fin.
+
+The ends of the radiale are expanded and rounded, the entire bone being
+approximately twice as long as wide. The three sides of the intermedium
+are similarly convex. The surface of this bone is unfinished, showing
+that it must have been embedded in cartilage. The ulnare is
+conspicuously similar to the ulna in bearing a posterior hooklike
+expansion, and is larger than the radiale.
+
+The four metacarpals are slightly expanded proximally and distally.
+Although measurements of length and width are tabulated below (Table
+2), we are not certain of the sequence of these bones in the row.
+
+The dimensions of the two proximal phalanges are alike. The shape of
+these elements is similar to that of the metacarpals. The two terminal
+phalanges are somewhat triangular in shape, the lateral edges being
+concave and the proximal convex.
+
+TABLE 2.--APPROXIMATE MEASUREMENTS OF THE FORELIMB (in mm.)
+
+--------------------------+-------------------------------------------
+                          |                 Dimensions
+                          +----------+--------------------------------
+ ELEMENT                  |          |              Width
+                          |  Length  +----------+----------+----------
+                          |          | Proximal |  Midway  |  Distal
+--------------------------+----------+----------+----------+----------
+Humerus                   |   16.0   |    5.0   |    2.0   |    7.5?
+Radius                    |    9.0   |    4.0   |    1.5   |    3.5
+Ulna                      |    8.5   |    4.5   |    1.5   |    3.5
+Radiale                   |    3.0   |    2.0   |    1.5   |    2.0
+Intermedium               |    1.5   |     --   |    2.0   |     --
+Ulnare                    |    3.5   |    2.0   |    2.0   |    2.5
+Metacarpal        A       |    4.5   |    2.5   |    1.0   |    2.0
+Metacarpal        B       |    4.5   |    3.0?  |    1.5   |    2.5
+Metacarpal        C       |    4.0   |    2.0   |    1.5   |    2.0
+Metacarpal        D       |    3.5   |    2.5   |    1.0   |    1.5
+Proximal Phalanx  A       |    2.0   |    1.5   |    1.0   |    1.5
+Proximal Phalanx  B       |    2.0   |    1.5   |    1.0   |    1.5
+Terminal Phalanx  A       |    1.5   |    1.5   |    1.0   |    1.0
+Terminal Phalanx  B       |    1.5   |    1.5   |    1.0   |    1.0
+--------------------------+----------+----------+----------+----------
+
+
+COMPARISONS AND DISCUSSION
+
+Apparently primitive rhipidistian characters in _Hesperoherpeton_ are:
+Braincase in two sections, posterior one containing an expanded
+notochordal canal; lateral series of mandibular bones closely
+resembling that of _Megalichthys_, as figured by Watson (1926); tabular
+having long process probably articulating with pectoral girdle; lack of
+movement between head and trunk correlated with absence of occipital
+condyle; sensory pits present on frontal and squamosal.
+
+Although we are unable to separate, by sutures, the vomers from the
+palatines, the palatal surface of these bones and of the pterygoids is
+studded by numerous small teeth, as in Rhipidistia (Jarvik, 1954) and
+some of the early Amphibia (Romer, 1947). The stapes apparently reaches
+the quadrate, and could therefore serve in hyostylic suspension of the
+upper jaw.
+
+The pectoral limb has an axial series of bones carrying hooklike
+flanges on their posterior edges. The other bones of the limb show
+little modification of form beyond the nearly flat, aquatic type seen
+in Rhipidistia. No distinct elbow or wrist joints are developed.
+
+Characters of _Hesperoherpeton_ common to most primitive Amphibia, in
+contrast with Crossopterygii, are: Nares separated from edge of jaw;
+stapes having external process that may have met a tympanic membrane,
+thus giving the bone a sound-transmitting function. Apparently none of
+the opercular series was present.
+
+There are two large palatal teeth, slightly labyrinthine in character,
+adjacent to each internal naris. The scapulocoracoid, as shown by
+Peabody (1958), is Anthracosaurian in structure, as are the
+long-stemmed clavicles. The limbs have digits rather than fin-lobes,
+although the digital number apparently is four and the number of bones
+in the manus is less than would be expected in a primitive amphibian.
+The vertebrae are similar to those of Ichthyostegids, as described by
+Jarvik (1952), except that the pleurocentra are much larger.
+
+In addition to this remarkable combination of crossopterygian and
+amphibian characters, _Hesperoherpeton_ is specialized in certain
+features of the skull. The orbits are much enlarged, probably in
+correlation with the diminutive size of the animal, and this has been
+accompanied by loss of several bones. The frontal and squamosal nearly
+meet each other, and both form part of the rim of the orbit. The bones
+of the posterior part of the dermal roof are greatly reduced, and there
+is none behind the squamosal except the projecting tabular; there is no
+indication of quadratojugal, jugal, intertemporal or postparietal. The
+foramen magnum is enormous. The external surfaces of the bones of the
+skull are nearly smooth.
+
+Is it possible that the "primitive" and "specialized" features of this
+animal are actually larval? Are they not just the kind of characters
+that would be expected in an immature, aquatic embolomere of
+Pennsylvanian time? For several reasons we do not think this is the
+case. Except for the anterior part of the braincase, there is no
+indication that the skeleton was not well ossified. The postaxial
+processes on the humerus, ulna and ulnare could scarcely have been
+larval features only, since they are so clearly homologous with those
+in adult Rhipidistia; a larval limb should indeed be simple, but its
+simplicity is unlikely to involve paleotelic adult characters. The
+scapulocoracoid of our specimen is of practically the same shape and
+size as that in the only other known individual, the type; this would
+be probable if both were adults, but somewhat less likely if they were
+larvae of a much larger animal. The form of the stapes, tabular and
+otic notch suggest a functional tympanic membrane, which could not have
+occurred in a gill-breathing larva. On the other hand, an adult animal
+of pigmy size might be expected to have large orbits, large otic
+capsules and a large foramen magnum.
+
+We conclude that _Hesperoherpeton_ lived and sought food in the weedy
+shallows at the margin of a pond or lagoon, and that for much of the
+time its head was partly out of water (Fig. 12). The animal could
+either steady itself or crawl around by means of the paddlelike limbs,
+but these probably could not be used in effective locomotion on land.
+Like the Ichthyostegids, it probably swam by means of a fishlike tail.
+
+[Illustration: FIG. 12. _Hesperoherpeton garnettense_ Peabody. Probable
+appearance in life. × 0.5.]
+
+
+TAXONOMY
+
+Evidently _Hesperoherpeton_ is a small, lagoon-dwelling survivor of
+the Devonian forms that initiated the change from Crossopterygii to
+Amphibia (Jarvik, 1955). It shows, however, that this transition did
+not affect all structures at the same time, for some, as the braincase
+with its notochordal canal, the mandibular bones and axial limb bones,
+are unchanged from the condition normal for the Rhipidistia, but most
+other characters are of amphibian grade. To express these facts
+taxonomically requires that _Hesperoherpeton_ be removed from the
+family Cricotidae, suborder Embolomeri, order Anthracosauria, and
+placed in a new order and family of labyrinthodont Amphibia.
+
+     Order PLESIOPODA
+
+     (_plesios_, Gr., near, almost; _podos_, Gr., foot)
+
+     Labyrinthodontia having limbs provided with digits, but
+     retaining posterior flanges on axial bones as in
+     Rhipidistia, without joint-structure at elbow and wrist
+     essential for terrestrial locomotion; neurocranium having
+     separate otico-occipital section, large notochordal canal,
+     no occipital condyle, as in Rhipidistia; nares separate from
+     rim of mouth; pectoral girdle anthracosaurian; vertebrae
+     having U-shaped intercentrum and paired, but large,
+     pleurocentra.
+
+Probably associated with the characters of the order, as given above,
+are the connection of pectoral girdle with skull, and the presence of a
+tympanic membrane, the stapes functioning in both sound-transmission
+and palatoquadrate suspension.
+
+     Family HESPEROHERPETONIDAE
+
+     Orbits and foramen magnum unusually large in correlation
+     with reduced size of animal; squamosal forming posterior
+     margin of orbit; circumorbital series absent (except for
+     postorbital); sensory pits on squamosal and frontal.
+
+Characters defining the family are evidently the more specialized
+cranial features, which probably evolved during Mississippian and early
+Pennsylvanian times.
+
+The definition of the genus and species may be left to rest upon
+Peabody's (1958) original description and the present account, until
+the discovery of other members of the family gives reason for making
+further distinctions.
+
+
+SUMMARY
+
+_Hesperoherpeton garnettense_ Peabody (1958), based on a
+scapulocoracoid and part of a vertebra, was originally placed in the
+order Anthracosauria, suborder Embolomeri, family Cricotidae. A new
+skeleton from the type locality near Garnett, Kansas (Rock Lake shale,
+Stanton formation, Upper Pennsylvanian), shows that the animal has the
+following rhipidistian characters: Large notochordal canal below
+foramen magnum, otico-occipital block separate from ethmosphenoid,
+postaxial processes on three axial bones of forelimb, pectoral girdle
+(probably) articulated with tabular. Nevertheless, _Hesperoherpeton_
+has short digits, an anthracosaurian type of pectoral girdle, an otic
+rather than spiracular notch, nostrils separate from the mouth, and
+vertebrae in which the intercentrum is U-shaped and the pleurocentra
+large but paired. The stapes reaches the quadrate.
+
+_Hesperoherpeton_ is placed in a new order, PLESIOPODA, on the basis of
+the characters stated above, and a new family, HESPEROHERPETONIDAE.
+Specialized characters of the family include: Reduction of
+circumorbital bones, bringing the squamosal to the edge of the orbit,
+loss of certain bones of the temporal region, and relative enlargement
+of the orbits and foramen magnum, in correlation with the diminutive
+size of the animal. The structural characters of _Hesperoherpeton_
+suggest to us that it lived in the shallow, weedy margins of lagoons,
+rested with its head partly out of water, and normally did not walk on
+land.
+
+
+LITERATURE CITED
+
+EATON, T. H., JR.
+    1951. Origin of tetrapod limbs. Amer. Midl. Nat., 46: 245-251.
+
+JARVIK, E.
+    1952. On the fish-like tail in the ichthyostegid stegocephalians.
+          Meddel. om Grønland, 114: 1-90.
+    1954. On the visceral skeleton in _Eusthenopteron_ with a discussion
+          of the parasphenoid and palatoquadrate in fishes. Kgl. Svenska
+          Vetenskapsakad. Handl., 5: 1-104.
+    1955. The oldest tetrapods and their forerunners. Sci. Monthly,
+          80: 141-154.
+
+MOORE, R. C., FRYE, J. C., and JEWETT, J. M.
+    1944. Tabular description of outcropping rocks in Kansas. Kansas
+          State Geol. Surv. Bull., 52: 137-212.
+
+PEABODY, F. E.
+    1952. _Petrolacosaurus kansensis_ Lane, a Pennsylvanian reptile
+          from Kansas. Univ. Kansas Paleont. Contrib., Vertebrata,
+          Art. 1: 1-41.
+    1958. An embolomerous amphibian in the Garnett fauna (Pennsylvanian)
+          of Kansas. Jour. Paleont., 32: 571-573.
+
+ROMER, A. S.
+    1937. The braincase of the Carboniferous crossopterygian
+          _Megalichthys nitidus_. Mus. Comp. Zool. Bull., 82: 1-73.
+    1947. Review of the Labyrinthodontia. Mus. Comp. Zool. Bull.,
+          99: 1-368.
+    1957. The appendicular skeleton of the Permian embolomerous
+          amphibian _Archeria_. Univ. Michigan Contrib. Mus. Paleont.,
+          13: 103-159.
+
+WATSON, D. M. S.
+    1926. The evolution and origin of the Amphibia. Phil. Trans. Roy.
+          Soc. London, (B) 214: 189-257.
+
+_Transmitted January 13, 1960._
+
+
+28-2495
+
+
+
+
+
+End of the Project Gutenberg EBook of A New Order of Fishlike Amphibia From
+the Pennsylvanian of Kansas, by Theodore H. Eaton and Peggy Lou Stewart
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+  <head>
+    <meta http-equiv="Content-Type" content="text/html;charset=iso-8859-1" />
+    <meta http-equiv="Content-Style-Type" content="text/css" />
+    <title>
+      The Project Gutenberg eBook of A New Order of Fishlike Amphibia
+From the Pennsylvanian of Kansas, by Theodore H. Eaton, Jr., and Peggy Lou Stewart.
+    </title>
+    <style type="text/css">
+
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+<pre>
+
+The Project Gutenberg EBook of A New Order of Fishlike Amphibia From the
+Pennsylvanian of Kansas, by Theodore H. Eaton and Peggy Lou Stewart
+
+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: A New Order of Fishlike Amphibia From the Pennsylvanian of Kansas
+
+Author: Theodore H. Eaton
+        Peggy Lou Stewart
+
+Release Date: January 23, 2010 [EBook #31050]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK NEW ORDER OF FISHLIKE AMPHIBIA ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper, Diane Monico, and
+the Online Distributed Proofreading Team at
+https://www.pgdp.net
+
+
+
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+</pre>
+
+
+
+
+
+
+<p class="title">
+<span class="smcap">University of Kansas Publications</span><br />
+
+<span class="smcap">Museum of Natural History</span><br /><br />
+
+Volume 12, No. 4, pp. 217-240, 12 figs.<br />
+May 2, 1960</p>
+<hr style="width: 45%;" />
+
+
+<h1>A New Order of Fishlike Amphibia<br />
+From the Pennsylvanian of Kansas</h1>
+
+<p class="title"><small>BY</small><br /><br />
+
+THEODORE H. EATON, JR., AND PEGGY LOU STEWART<br /><br /><br />
+
+
+<span class="smcap">University of Kansas</span><br />
+<span class="smcap">Lawrence</span><br />
+1960
+</p>
+<hr style="width: 45%;" />
+
+
+
+<p class="title">
+<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br />
+<br />
+Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br />
+Robert W. Wilson<br />
+<br />
+<br />
+Volume 12, No. 4, pp. 217-240, 12 figs.<br />
+Published May 2, 1960<br />
+<br />
+<br />
+<span class="smcap">University of Kansas</span><br />
+Lawrence, Kansas<br />
+<br />
+<br />
+<small>PRINTED IN</small><br />
+<small>THE STATE PRINTING PLANT</small><br />
+<small>TOPEKA, KANSAS</small><br />
+<small>1960</small><br />
+<br />
+<small>28-2495</small><br />
+</p>
+<hr style="width: 45%;" />
+
+
+<p><span class='pagenum'><a name="Page_219" id="Page_219">[Pg 219]</a></span></p>
+
+<h2><big>A New Order of Fishlike Amphibia<br />
+From the Pennsylvanian of Kansas<br /></big></h2>
+
+<p class="center"><small>BY</small><br /><br />
+
+THEODORE H. EATON, JR., AND PEGGY LOU STEWART</p>
+
+<h2>INTRODUCTION</h2>
+
+
+<p>A slab of shale obtained in 1955 by Mr. Russell R. Camp from a
+Pennsylvanian lagoon-deposit in Anderson County, Kansas, has
+yielded in the laboratory a skeleton of the small amphibian <i>Hesperoherpeton
+garnettense</i> Peabody (1958). This skeleton provides
+new and surprising information not available from the holotype,
+No. 9976 K. U., which consisted only of a scapulocoracoid,
+neural arch, and rib fragment. The new specimen, No. 10295 K. U.,
+is of the same size and stage of development as the holotype and it
+is thought that both individuals are adults.</p>
+
+<p>The quarry, University of Kansas Museum of Natural History
+Locality KAN 1/D, is approximately six miles northwest of Garnett,
+Anderson County, Kansas, in Sec. 5, T. 19S, R. 19E, 200 yards southwest
+of the place where <i>Petrolacosaurus kansensis</i> Lane was obtained
+(see Peabody, 1952). The Rock Lake shale, deposited under
+alternately marine and freshwater lagoon conditions, is a thin member
+of the Stanton limestone formation, Lansing group, Missourian
+series, and thus is in the lower part of the Upper Pennsylvanian.</p>
+
+<p>Peabody (1958) placed <i>Hesperoherpeton</i> in the order Anthracosauria,
+suborder Embolomeri, family Cricotidae. Study of the
+second and more complete specimen reveals that <i>Hesperoherpeton</i>
+is unlike the known Embolomeri in many important features. The
+limbs and braincase are more primitive than those so far described
+in any amphibian. The vertebrae are comparable to those of Ichthyostegalia
+(Jarvik, 1952), as well as to those of Embolomeri. The
+forelimb is transitional between the pectoral fin of Rhipidistia and
+the limb of early Amphibia. The pattern of the bones of the forelimb
+closely resembles, but is simpler than, that of the hypothetical
+transitional type suggested by Eaton (1951). The foot seemingly
+had only four short digits. The hind limb is not known.</p>
+
+<p>The new skeleton of <i>Hesperoherpeton</i> lies in an oblong block of
+limy shale measuring approximately 100&nbsp;&times;&nbsp;60 mm. After preparation
+of the entire lower surface, the exposed bones and matrix
+were embedded in Bioplastic, in a layer thin enough for visibility<span class='pagenum'><a name="Page_220" id="Page_220">[Pg 220]</a></span>
+but giving firm support. Then the specimen was inverted and the
+matrix removed from the opposite side; this has not been covered
+with Bioplastic. The bones lie in great disorder, except that some
+parts of the roof of the skull are associated, and the middle section
+of the vertebral column is approximately in place. The bones of the
+left forelimb are close together but not in a natural position. The
+tail, pelvis, hind limbs and right forelimb are missing. Nearly all
+the bones present are broken, distorted by crushing, incomplete and
+scattered out of place, probably by the action of currents. The
+complete skeleton, in life, probably measured between 150 and 200
+mm. in length.</p>
+
+<p>The specimen was studied at the Museum of Natural History,
+University of Kansas, with the help of a grant from the National
+Science Foundation, number NSF-G8624. The specimen was discovered
+in the slab by Miss Sharon K. Moriarty, and was further
+cleaned by the authors. Mr. Merton C. Bowman assisted with the
+illustrations. We are indebted to Dr. Robert W. Wilson for critical
+comments.</p>
+
+
+<h2>SKULL</h2>
+
+<h3><i>Dorsal Aspect</i> (Figs. <a href="#fig1">1</a>, <a href="#fig2">2</a>)</h3>
+
+<p>In reconstruction, the skull measures approximately 8.0 mm. dorsoventrally
+at the posterior end. The height diminishes anteriorly
+to about 1.5 mm. at the premaxillary. The length is about 15.5 mm.
+in the median line, or 24.0 mm. to the tip of the tabular, and the
+width about 16.0 mm. posteriorly. The snout is blunt, continuing
+about 1-2 mm. anterior to the external nares. Each of the tabulars
+has a slender posterior process 5.0 mm. long, which probably met
+the supracleithrum; the intertabular space is about 8.5 mm. wide.
+The orbits are approximately 5.5 mm. in diameter and extend from
+the maxillary to within about 3.0 mm. of the midline dorsally. The
+pineal opening is 1.8 mm. anterior to the occipital margin of the
+skull.</p>
+
+<p>Reduction of bones at the back of the skull seems to have eliminated
+any dermal elements posterior to the squamosal, while enlargement
+of the orbit has removed most of the postorbital series,
+leaving the squamosal as the only cheekbone. There is apparently
+no jugal or postfrontal.</p>
+
+<p>The squamosal of <i>Acanthostega</i> (Jarvik, 1952) is articulated
+under the tabular and reaches forward and down, much as if it were
+an opercular in reversed position. Internally, it must lie against
+the otic capsule below the tabular, partially concealing the stapes.<span class='pagenum'><a name="Page_221" id="Page_221">[Pg 221]</a></span>
+The bone that we suppose to be the squamosal of <i>H. garnettense</i>
+is of similar shape, of about the same size and has internally an
+articular surface at one corner, bounded by a pair of ridges in the
+shape of a V. This articular surface probably fitted on a lateral
+process extending from the roof of the neurocranium, over the front
+of the otic capsule.</p>
+
+<p>The premaxillary extends posterolaterally to a distance 5.5 mm.
+from the midline and attains a width at its broadest point of about
+1.5 mm. The posterior edge is slightly concave and in part forms the
+anterior border of the naris.</p>
+
+<p class="figcenter" style="width: 538px;">
+<a name="fig1" id="fig1"></a>
+<img src="images/image001.png" width="538" height="600" alt="Fig. 1. Hesperoherpeton garnettense Peabody. Skull, dorsal
+view. Postorbital processes of the neurocranium are shown in
+dotted outline. KU&nbsp;10295, &times;&nbsp;4." title="Fig. 1. Hesperoherpeton garnettense Peabody. Skull, dorsal
+view. Postorbital processes of the neurocranium are shown in
+dotted outline. KU&nbsp;10295, &times;&nbsp;4." />
+<span class="caption">Fig. 1.<i> Hesperoherpeton garnettense</i> Peabody. Skull, dorsal
+view. Postorbital processes of the neurocranium are shown in
+dotted outline. KU&nbsp;10295, &times;&nbsp;4.</span>
+</p>
+
+<p>The nasal is triangular and, with the lacrimal, forms the medial
+border of the naris. The length of the medial side of the nasal bone
+is approximately 5.0 mm., the transverse width is 3.8 mm., and the
+extent of the posterolateral border is 5.5 mm.<span class='pagenum'><a name="Page_222" id="Page_222">[Pg 222]</a></span></p>
+
+<p>The maxillary meets the premaxillary lateral to the naris, borders
+the naris posteroventrally, and continues posteriorly beneath the
+orbit, of which it forms the external border. The maxillary is
+about 8.5 mm. long, and immediately anterior to the orbit has a
+maximum width of 1.3 mm.</p>
+
+<p>The lacrimal fills the remaining rim of the narial opening between
+the nasal and maxillary, and extends to the anterior edge of the
+orbit. The length, from naris to orbit, is 4.2 mm.; the width ranges
+from 1.0 mm. anteriorly to 2.5 mm. posteriorly.</p>
+
+<p class="figcenter" style="width: 600px;">
+<a name="fig2" id="fig2"></a>
+<img src="images/image002.png" width="600" height="300" alt="Fig. 2. Hesperoherpeton garnettense Peabody. Skull, lateral view,
+showing relatively large orbit and absence of smaller circumorbital
+bones. KU&nbsp;10295, &times;&nbsp;4." title="Fig. 2. Hesperoherpeton garnettense Peabody. Skull, lateral view,
+showing relatively large orbit and absence of smaller circumorbital
+bones. KU&nbsp;10295, &times;&nbsp;4." />
+<span class="caption">Fig. 2. <i>Hesperoherpeton garnettense</i> Peabody. Skull, lateral view,
+showing relatively large orbit and absence of smaller circumorbital
+bones. KU&nbsp;10295, &times;&nbsp;4.</span>
+</p>
+
+<p>The external naris is approximately 1.0 mm. in diameter. It is
+slightly anterodorsal to the internal naris and 4.0 mm. lateral to
+the midline.</p>
+
+<p>The dorsal margin of the orbit appears to be formed by the
+frontal. The anterior part of this margin, however, may be formed
+by a prefrontal, which is not clearly set off by a suture. The frontal
+extends 3.8 mm. in the midline, and anteriorly and laterally borders
+the nasal and lacrimal, respectively. A faint pattern of pitting
+radiates on the surface from the center of ossification of the frontal.
+There is also a pit indicating the presence of a supraorbital sensory
+pore.</p>
+
+<p>The parietal bones enclose the pineal opening, approximately 2.5
+mm. posterior to the suture with the frontal. The foramen is about
+0.5 mm. in diameter. Laterally the parietal meets the medial angle
+of the postorbital and the medial border of the supratemporal. No
+bone of this animal shows the deep pitting and heavy ornamentation
+characteristic of many primitive Amphibia.<span class='pagenum'><a name="Page_223" id="Page_223">[Pg 223]</a></span></p>
+
+<p>The postorbital meets the anterolateral corner of the parietal for
+a distance of 0.5 mm., the anterior edge bordering the frontal bone
+and the orbit for a combined distance of about 3.0 mm. The lateral
+margin is slightly convex, and is probably interrupted behind by the
+anterior point of the tabular. Medially, the concave margin of the
+postorbital meets the supratemporal for about 3.5 mm.</p>
+
+<p>The supratemporal is thus wedge-shaped and located between
+the parietal and the postorbital. The posterior edge of the supratemporal
+protrudes as a convex border slightly behind the end of
+the parietal, and measures 3.0 mm. around the curve to the parietal
+suture.</p>
+
+<p class="figcenter" style="width: 600px;">
+<a name="fig3" id="fig3"></a>
+<img src="images/image003.png" width="600" height="292" alt="Fig. 3. Hesperoherpeton garnettense Peabody. A, left squamosal, internal
+surface. B, left squamosal, external surface. C, right tabular
+internal surface. D, right tabular, external surface. KU&nbsp;10295, all &times;&nbsp;4." title="Fig. 3. Hesperoherpeton garnettense Peabody. A, left squamosal, internal
+surface. B, left squamosal, external surface. C, right tabular
+internal surface. D, right tabular, external surface. KU&nbsp;10295, all &times;&nbsp;4." />
+<span class="caption">Fig. 3. <i>Hesperoherpeton garnettense</i> Peabody. A, left squamosal, internal
+surface. B, left squamosal, external surface. C, right tabular
+internal surface. D, right tabular, external surface. KU&nbsp;10295, all &times;&nbsp;4.</span>
+</p>
+
+<p>The squamosal (Fig. <a href="#fig3">3 A, B</a>) is a large, somewhat rectangular
+bone extending from the back of the orbit to the posterior extremity
+of the cheek. It outlines almost entirely the posterior border of the
+orbit, the ventrolateral portion of the cheek region, and the lateral
+border of the top of the skull behind the orbit. Dorsally, the
+squamosal meets the anterior half of the tabular and the lateral
+border of the supratemporal. Near the anteroventral edge of the
+squamosal there is a small pit, probably related to a postorbital
+sensory pore in the skin.</p>
+
+<p>The tabular (Fig. <a href="#fig3">3 C, D</a>) is pointed anteriorly, where it probably
+fits against the lateroposterior edge of the postorbital. The dorsal
+part of the bone flares out and down, forming a small otic notch at
+a point halfway back. Posteriorly, the flange attains a dorsoventral
+width of 2.0 mm. at the edge of the notch. The slender posterior<span class='pagenum'><a name="Page_224" id="Page_224">[Pg 224]</a></span>
+process of the tabular which continues beyond the flange is approximately
+0.5 mm. in diameter and 5.0 mm. long.</p>
+
+<h3><i>Ventral Aspect</i> (Fig. <a href="#fig4">4</a>)</h3>
+
+<p>The palatal view of the skull shows the paired premaxillary,
+maxillary, palatine, pterygoid, and quadrate bones. The openings
+for the internal nares, the ventral orbital fenestrae, and the subtemporal
+fossae are readily recognized. The quadrate processes extend
+posteriorly leaving a large gap medially at the posterior end of
+the skull.</p>
+
+<p class="figcenter" style="width: 415px;">
+<a name="fig4" id="fig4"></a>
+<img src="images/image004.png" width="415" height="580" alt="Fig. 4. Hesperoherpeton garnettense Peabody. Palate reconstructed;
+ventral aspect at left, showing teeth, dorsal aspect at
+right. KU&nbsp;10295, &times;&nbsp;4." title="Fig. 4. Hesperoherpeton garnettense Peabody. Palate reconstructed;
+ventral aspect at left, showing teeth, dorsal aspect at
+right. KU&nbsp;10295, &times;&nbsp;4." />
+<span class="caption">Fig. 4. <i>Hesperoherpeton garnettense</i> Peabody. Palate reconstructed;
+ventral aspect at left, showing teeth, dorsal aspect at
+right. KU&nbsp;10295, &times;&nbsp;4.</span>
+</p><p><span class='pagenum'><a name="Page_225" id="Page_225">[Pg 225]</a></span></p>
+
+<p>The left quadrate appears to be in place on the posterior prong
+of the pterygoid. The dorsal side of the quadrate is grooved between
+two anterolaterally directed ridges. The groove, which probably
+held the end of the stapes, extends about half the width of the
+quadrate itself. The width of the quadrate is 4.0 mm., the length is
+4.5 mm. medially and about 2.0 mm. laterally. In ventral view the
+quadrate appears to project laterally, but is incomplete and its shape
+uncertain. The distance from the posterior end of the quadrate to
+the visible posterior edge of the orbital fenestra, which opens
+ventrally, is 10.0 mm.</p>
+
+<p>This region between the quadrate and the orbit is occupied by a
+pterygoid with three projections. Anteriorly, the pterygoid outlines
+most of the posterior edge of the orbit (a distance of about 6.5 mm.).
+A lateral process separates the orbit from the subtemporal fossa. A
+posteriorly directed edge defines the fossa, which extends about 6.5
+mm. anteroposteriorly. The lateral process of the pterygoid terminates
+10.0 mm. from the midline. Both the lateral and posterior pterygoid
+processes are approximately 2.0 mm. wide. The greatest width
+of the subtemporal fossa is about 2.0 mm. The medial border of
+the orbital fenestra is missing, but apparently consisted of the pterygoid
+for at least the posterior half.</p>
+
+<p>Along the posterior edge of the orbital fenestra, there is a narrow,
+dorsally projecting flange of the pterygoid. The lateral opening of
+the orbit is approximately 7.5 mm. wide.</p>
+
+<p>The remaining border of the orbital fenestra on the anterior and
+medial sides is formed by a bone occupying the position of palatine
+and vomer; for convenience we designate this as palatine. When
+reconstructed in its probable position in relation to the pterygoid,
+the left palatine lacks a section, on its medial and posterior edges,
+measuring about 2.5 mm. by 9.0 mm. The lateral margin of the
+palatine is convex; about 5.5 mm. anterior to the orbit this margin
+curves into a strong anteriorly pointing projection, medial to
+which is seen the internal narial opening. The remaining anterior
+edge is slightly convex, smoothly rounded, and meets the midline
+about 9.0 mm. anterior to the pterygoid.</p>
+
+<p>The void area medial to the palatine and anterior to the pterygoid
+does not fit any bone which we can recognize as the parasphenoid.
+It is thus suspected that this area is covered in part by the
+missing edge of the palatine and partly by an anteromedial extension
+of the pterygoid. Of course a parasphenoid may also have
+been present.<span class='pagenum'><a name="Page_226" id="Page_226">[Pg 226]</a></span></p>
+
+<p>The position, length, and shape of the premaxillary shown in
+palatal view (Fig. <a href="#fig4">4</a>) are primarily based upon the dorsal appearance
+since ventrally most of it cannot be seen. At the point where
+it forms the anterior border of the internal naris, the premaxillary is
+slightly wider than the maxillary and seems to become narrower as
+it approaches the midline.</p>
+
+<p>The ethmosphenoid, which we cannot identify, may have been
+exposed in a gap between the premaxillary and the palatine. The
+gap measures approximately 8.0 mm. wide and ranges up to 1.0 mm.
+anteroposteriorly.</p>
+
+<p>The maxillary begins at a suture with the premaxillary lateral
+to the naris and continues posteriorly, bordering the orbit with a
+width of about 1.2 mm. It then tapers to a point approximately 2.0
+mm. anterior to the lateral projection of the pterygoid. The width
+of the maxillary at this point is 0.8 mm. and the posterior end is
+broken; probably when complete it approached the pterygoid, and
+either met the latter or had a ligamentous connection with it. As
+nearly as can be determined, the total length of the maxillary is
+approximately 12.0 mm.</p>
+
+<p>The teeth on the maxillary are small and seem to be in two
+longitudinal rows. The palatine bears two large, grooved teeth
+anteriorly; the first is approximately 1.0 mm. posteromedial to the
+naris and the second is about 3.0 mm. posterior and slightly lateral
+to the naris. The flat ventral surfaces of the palatine and pterygoid
+bear numerous small teeth distributed as shown in Fig. <a href="#fig4">4</a>.</p>
+
+<h3><i>Braincase and Occipital Region</i> (Fig. <a href="#fig5">5</a>)</h3>
+
+<p>The parts of the neurocranium are scattered, disconnected and
+incomplete, but it is possible to make out a number of features of
+the otico-occipital section with fair assurance. In posterior view
+the notochordal canal and foramen magnum are confluent with
+each other, and of great size relative to the skull as a whole. The
+notochordal canal measures 2.8 mm. in diameter, and the foramen
+magnum about 4.0 mm. The crescent-shaped supraoccipital rests
+on the upright ends of the exoccipitals, but between the latter and
+the basioccipital no sutures can be seen. Probably the whole posterior
+surface of the braincase slanted posteroventrally; consequently
+the rim of the notochordal canal was about 3.0 mm. behind
+the margin of the parietals.</p>
+
+<p>The U-shaped border of the notochordal canal is a thick, rounded
+bone, comparable in appearance to the U-shaped intercentra of the<span class='pagenum'><a name="Page_227" id="Page_227">[Pg 227]</a></span>
+vertebrae. This bone apparently rested upon a thinner, troughlike
+piece (Fig. <a href="#fig5">5 B</a>) forming the floor of the braincase. The latter is
+broad, shallow, concave, open midventrally and narrowing anteriorly
+to form a pair of articular processes. Since no sutures can
+be seen in this structure, it probably is the ventral, ossified portion
+of the basioccipital. Watson (1926, Fig. 4 B) illustrates the floor
+of the braincase in <i>Eusthenopteron</i>, with its more lateral, anterior
+portion labelled prootic, but in our specimen the corresponding part
+could scarcely have formed the anterior wall of the otic capsule,
+being entirely in the plane of the floor. The two articular surfaces
+anteriorly near the midline suggest that a movable joint existed between
+the otico-occipital part of the braincase and the ethmosphenoid
+part, as in Rhipidistia (Romer, 1937). We have found
+nothing in the specimen that could be referred to the ethmosphenoid;
+it may have been unossified.</p>
+
+<p class="figcenter" style="width: 600px;">
+<a name="fig5" id="fig5"></a>
+<img src="images/image005.png" width="600" height="586" alt="Fig. 5. Hesperoherpeton garnettense Peabody, KU
+10295, &times;&nbsp;4. A, occipital view of skull; B, basioccipital
+bone in dorsal (internal) view." title="Fig. 5. Hesperoherpeton garnettense Peabody, KU
+10295, &times;&nbsp;4. A, occipital view of skull; B, basioccipital
+bone in dorsal (internal) view." />
+<span class="caption">Fig. 5. <i>Hesperoherpeton garnettense</i> Peabody, KU
+10295, &times;&nbsp;4. A, occipital view of skull; B, basioccipital
+bone in dorsal (internal) view.</span>
+</p>
+
+<p>The otic capsules appear to have rested against lateral projections
+of the basioccipital. The single otic capsule that can be seen (the<span class='pagenum'><a name="Page_228" id="Page_228">[Pg 228]</a></span>
+right) is massively built, apparently ossified in one piece, with a
+shallow dorsomedial excavation, probably the vestige of a supratemporal
+fossa. On the lateral face is a broad, shallow depression
+dorsally, and a narrower, deeper one anteroventrally; these we
+suppose to have received the broader and narrower heads of the
+stapes, respectively. The posterior wall of the otic capsule we
+have designated opisthotic in the figure. Anterior to the otic capsule
+the lateral wall of the braincase cannot be seen, and may not
+have been ossified.</p>
+
+<p>The roof of the braincase is visible in its ventral aspect, extending
+from approximately the occipital margin to a broken edge in front
+of the parietal foramen, and laterally to paired processes which
+overlie the otic capsules directly behind the orbits (see dotted outlines
+in Fig. <a href="#fig1">1</a>). Each of these postorbital processes, seen from
+beneath, appears to be the lateral extension of a shallow groove
+beginning near the midline. Presumably this section of the roof
+is an ossification of the synotic tectum. It should be noted that the
+roof of the braincase proper is perfectly distinct from the overlying
+series of dermal bones, and that the parietal foramen can be seen
+in both. The roof of the braincase in our specimen seems to have
+been detached from the underlying otic capsules and the occipital
+wall.</p>
+
+<p>The bone that we take to be the stapes is blunt, flattened (perhaps
+by crushing), 5.0 mm. in length, and has two unequal heads; its
+width across both of these is 4.0 mm. The length is appropriate to
+fit between the lateral face of the otic capsule and the dorsal edge of
+the quadrate; the wider head rests on a posterodorsal concavity
+on the otic capsule, and the smaller fits a lower, more anterior
+pit. Laterally the stapes carries a short, broad process that
+probably made contact with a dorsally placed tympanic membrane.
+Thus the bone was a hyomandibular in the sense that it articulated
+with the quadrate, but it may also have served as a stapes in sound-transmission.
+It contains no visible canal or foramen.</p>
+
+<h3><i>Mandible</i> (Fig. <a href="#fig6">6</a>)</h3>
+
+<p>The crushed inner surface of the posterior part of the left mandible
+and most of the external surface of the right mandible are preserved
+in close proximity. Although the whole length of the tooth-bearing
+margins is missing, some parts of six elements of the right
+mandible can be seen. The pattern of sutures and the general contour
+closely resemble those of <i>Megalichthys</i> (Watson, 1926, Figs.
+37, 38) and other known Rhipidistia.<span class='pagenum'><a name="Page_229" id="Page_229">[Pg 229]</a></span></p>
+
+<p>The anteroposterior length of the mandible is about 23.8 mm.,
+and the depth is 3.8 mm. The dentary extends approximately 17.6
+mm. back from the symphysis, and its greatest width is probably
+2.0 mm. Its lower edge meets all the other lateral bones of the
+jaw. The splenial and postsplenial form the curved anteroventral
+half of the jaw for a distance of about 9.0 mm. The fragmented
+articular, on the posterior end of the jaw, is 4.0 mm. long and 2.0
+mm. deep, exhibiting a broken upper edge; presumably the surface
+for articulation with the quadrate was a shallow concavity, above
+the end of the articular.</p>
+
+<p class="figcenter" style="width: 600px;">
+<a name="fig6" id="fig6"></a>
+<img src="images/image006.png" width="600" height="170" alt="Fig. 6. Hesperoherpeton garnettense Peabody. Right mandible, lateral
+view, KU&nbsp;10295, &times;&nbsp;4. External surfaces are pitted; broken surfaces
+are coarsely stippled." title="Fig. 6. Hesperoherpeton garnettense Peabody. Right mandible, lateral
+view, KU&nbsp;10295, &times;&nbsp;4. External surfaces are pitted; broken surfaces
+are coarsely stippled." />
+<span class="caption">Fig. 6. <i>Hesperoherpeton garnettense</i> Peabody. Right mandible, lateral
+view, KU&nbsp;10295, &times;&nbsp;4. External surfaces are pitted; broken surfaces
+are coarsely stippled.</span>
+</p>
+
+
+<h2>VERTEBRAE (Fig. <a href="#fig7">7</a>)</h2>
+
+<p>The vertebrae that are visible from a lateral view are crushed
+and difficult to interpret. It is possible, nevertheless, to see that
+the trunk vertebrae resemble those of Ichthyostegalia (Jarvik, 1952,
+Fig. 13 A, B), except that the pleurocentra are much larger. A few
+parts of additional vertebrae can be seen, but they are so scattered
+that it is impossible to be sure of their original location. Therefore
+comparisons between different regions cannot yet be made.</p>
+
+<p>The U-shaped intercentrum encloses the notochord and occupies
+an anteroventral position in the vertebra. Anteriorly, each intercentrum
+articulates with the pleurocentra of the next preceding
+vertebra by slightly concave surfaces. Dorsolaterally there is an
+articular surface for the capitulum of the rib.</p>
+
+<p>The two pleurocentra of each vertebra are separate ventrally as
+well as dorsally, but form thin, broad plates of about the same
+height as the notochord. The lateral surface appears to be depressed,
+allowing, perhaps, for movement of the rib. Above each
+pleurocentrum, on the lateral surface of the neural arch, there is a
+short diapophysis for articulation with the tuberculum of the rib.</p>
+
+<p>The margin of the neural spine is convex anteriorly and concave
+posteriorly, the tip reaching a point vertically above the postzygapophysis.<span class='pagenum'><a name="Page_230" id="Page_230">[Pg 230]</a></span>
+The prezygapophysis of each vertebra articulates with
+the preceding postzygapophysis by a smooth dorsal surface. One
+nearly complete neural arch shows (Fig. <a href="#fig7">7 B</a>) a pit above the neural
+canal, clearly corresponding to the canal for a dorsal ligament shown
+by Jarvik in <i>Ichthyostega</i>. Indeed this view of the neural arch and
+intercentrum together brings out the striking resemblance between
+the vertebrae of <i>Hesperoherpeton</i> and those of the Ichthyostegids.
+The rounded intercentrum in both is an incomplete ring enclosing
+the notochordal canal.</p>
+
+<p class="figcenter" style="width: 600px;">
+<a name="fig7" id="fig7"></a>
+<img src="images/image007.png" width="600" height="246" alt="Fig. 7. Hesperoherpeton garnettense Peabody. A, End view of incomplete
+vertebra, probably near anterior end of column. B, Neural arch and intercentrum
+in end view, showing probable association. C, Left lateral view of
+trunk vertebra. All figures: KU&nbsp;10295, &times;&nbsp;4." title="Fig. 7. Hesperoherpeton garnettense Peabody. A, End view of incomplete
+vertebra, probably near anterior end of column. B, Neural arch and intercentrum
+in end view, showing probable association. C, Left lateral view of
+trunk vertebra. All figures: KU&nbsp;10295, &times;&nbsp;4." />
+<span class="caption">Fig. 7. <i>Hesperoherpeton garnettense</i> Peabody. A, End view of incomplete
+vertebra, probably near anterior end of column. B, Neural arch and intercentrum
+in end view, showing probable association. C, Left lateral view of
+trunk vertebra. All figures: KU&nbsp;10295, &times;&nbsp;4.</span><br /><br />
+</p>
+
+<p class="center"><b>
+<span class="smcap">Table 1.&mdash;Average Measurements of the Trunk Vertebrae</span> (in mm.).<br />
+<span class="smcap">Numbers in Parentheses Indicate the Number of Pieces Available for Measuring</span><br />
+</b></p>
+
+
+<div class='center'>
+<table border="1" cellpadding="4" cellspacing="0" summary="avgtrnk">
+<tr><td align='center'>Parts</td><td align='center'>Ant.-post.</td><td align='center'>Dors.-vent.</td><td align='center'>Transv. width</td></tr>
+<tr><td align='left'>Neural spine</td><td align='center'>1.5 (3)</td><td align='center'>3.0 (3)</td><td align='center'>&mdash;</td></tr>
+<tr><td align='left'>Neural spine and arch</td><td align='center'>2.0 (4)</td><td align='center'>4.5? (4)</td><td align='center'>&mdash;</td></tr>
+<tr><td align='left'>Neural canal</td><td align='center'>2.0 (4)</td><td align='center'>2.0 (1)</td><td align='center'>1.0 (1)</td></tr>
+<tr><td align='left'>Intercentrum</td><td align='center'>1.5 (5)</td><td align='center'>3.5 (4)</td><td align='center'>3.0 (1)</td></tr>
+<tr><td align='left'>Pleurocentrum</td><td align='center'>1.5 (3)</td><td align='center'>3.0 (2)</td><td align='center'>&mdash;</td></tr>
+</table></div>
+
+
+<p>The shape, in end view, of a partly preserved neural arch (Fig.
+<a href="#fig7">7 A</a>) seems to account for the incompleteness of the intercentrum
+just mentioned; the ventral edge of the arch is emarginate in such
+a way as to fit the dorsal surface of the notochord. The dorsal
+portion of this neural arch is not present (either broken or not yet<span class='pagenum'><a name="Page_231" id="Page_231">[Pg 231]</a></span>
+ossified), but the opening of the neural canal is comparable in
+width to the foramen magnum. Hence this vertebra may be one
+of the most anterior in the column. In comparison with the trunk
+vertebrae seen farther posteriorly it appears that there may be a
+progressive ossification of neural arches toward their dorsal ends,
+and of intercentra around the notochord, with probable fusion of
+the intercentra and neural arches in the posterior part of the trunk.
+The notochord seems to have been slightly constricted by the intercentra,
+but not interrupted.</p>
+
+
+<h2>RIBS</h2>
+
+<p>The proximal ends of the ribs expand dorsoventrally to a width
+approximately four times that of their slender shafts. The tuberculum
+and capitulum on each of the trunk ribs are separated only
+by a shallow concavity. These two articular surfaces are so situated
+that the rib must tilt downward from the horizontal plane.
+The shaft flares terminally in some ribs, and the distal end is convex.
+Ribs in the trunk region differ little if any in size. Five that can
+be measured vary in length from 5.0 to 7.0 mm. One short, bent
+rib 3.5 mm. long perhaps is sacral or caudal.</p>
+
+
+<h2>PECTORAL GIRDLE (Figs. <a href="#fig8">8</a>, <a href="#fig9">9</a>, <a href="#fig10">10</a>)</h2>
+
+<p>The right scapulocoracoid is almost complete, and the left one
+is present but partly broken into three pieces, somewhat pushed out
+of position. With the advantage of this new material, we may comment
+on the scapulocoracoid of <i>H. garnettense</i> as described by Peabody
+(1958). In size and contour, the slight differences between
+the type (KU&nbsp;9976) and the new skeleton (KU&nbsp;10295) are considered
+to be no more than individual variation. We have redrawn
+the type (Fig. <a href="#fig8">8</a>) in order to show the resemblances more clearly.</p>
+
+<p>The small sections that were missing from the type are present
+in KU&nbsp;10295. The jagged edge directly posterior to the area occupied
+by the neural arch in the type extends 0.5 mm. farther back
+in our specimen. The angle formed between the recurved dorsal
+ramus and the edge of the ventral flange is seen in our specimen
+to be less than 90&deg;. The glenoid fossa, appearing as a concave
+articular surface for the cap of the humerus, was in part covered
+by cartilage and shows as "unfinished" bone (Peabody, 1958, p. 572);
+this area is more oval than triangular, as Peabody thought. The
+obstruction of a clear view of this part of the type is the result
+of the accidental position of a neural arch. The raised portion<span class='pagenum'><a name="Page_232" id="Page_232">[Pg 232]</a></span>
+immediately dorsal to the glenoid fossa exhibits an unfinished surface,
+suggesting the presence of either cartilage or a ligament.<br /><br /></p>
+
+<p class="figcenter" style="width: 600px;">
+<a name="fig8" id="fig8"></a>
+<img src="images/image008.png" width="600" height="338" alt="Fig. 8. Hesperoherpeton garnettense Peabody. Type specimen redrawn.
+Right scapulocoracoid in external view (at left), and internal view (at right).
+KU&nbsp;9976, &times;&nbsp;4." title="Fig. 8. Hesperoherpeton garnettense Peabody. Type specimen redrawn.
+Right scapulocoracoid in external view (at left), and internal view (at right).
+KU&nbsp;9976, &times;&nbsp;4." />
+<span class="caption">Fig. 8. <i>Hesperoherpeton garnettense</i> Peabody. Type specimen redrawn.
+Right scapulocoracoid in external view (at left), and internal view (at right).
+KU&nbsp;9976, &times;&nbsp;4.</span>
+<br /><br /></p>
+
+
+<p class="figcenter" style="width: 446px;">
+<a name="fig9" id="fig9"></a>
+<img src="images/image009.png" width="446" height="500" alt="Fig. 9. Hesperoherpeton garnettense Peabody. Right scapulocoracoid in external
+view, showing part of interclavicle, and position occupied by clavicle.
+The specimen is flattened and lies entirely in one plane. KU&nbsp;10295, &times;&nbsp;4." title="Fig. 9. Hesperoherpeton garnettense Peabody. Right scapulocoracoid in external
+view, showing part of interclavicle, and position occupied by clavicle.
+The specimen is flattened and lies entirely in one plane. KU&nbsp;10295, &times;&nbsp;4." />
+<span class="caption">Fig. 9. <i>Hesperoherpeton garnettense</i> Peabody. Right scapulocoracoid in external
+view, showing part of interclavicle, and position occupied by clavicle.
+The specimen is flattened and lies entirely in one plane. KU&nbsp;10295, &times;&nbsp;4.</span>
+<br /></p>
+
+<p class="figcenter" style="width: 209px;">
+<a name="fig10" id="fig10"></a>
+<img src="images/image010.png" width="209" height="500" alt="Fig. 10. Hesperoherpeton garnettense Peabody. Right clavicle in external
+view. Anterior edge to right. KU&nbsp;10295, &times;&nbsp;4." title="Fig. 10. Hesperoherpeton garnettense Peabody. Right clavicle in external
+view. Anterior edge to right. KU&nbsp;10295, &times;&nbsp;4." />
+<span class="caption">Fig. 10. <i>Hesperoherpeton garnettense</i> Peabody. Right clavicle in external
+view. Anterior edge to right. KU&nbsp;10295, &times;&nbsp;4.</span>
+</p>
+
+<p><span class='pagenum'><a name="Page_233" id="Page_233">[Pg 233]</a></span></p>
+
+<p>The right clavicle is complete, and resembles a spoon having a
+slender handle. The dorsal tip of the handle is L-shaped. The
+expanded ventral part is convex externally, and rested upon the
+anteroventral surface of the scapulocoracoid. The lateral edge next
+to the "stem" is distinctly concave, abruptly becoming similar in
+contour to the opposite edge, and giving the impression of an unsymmetrical
+spoon. The left clavicle is present in scattered fragments,
+its dorsal hooklike end being intact.</p>
+
+<p>The posterior end of the interclavicle lies in contact with the right
+scapulocoracoid. There are short lateral processes at the point
+where the interclavicle was overlapped by the clavicles, but we cannot
+be sure of the extent of this bone anteriorly or posteriorly.</p>
+
+<p>The presumed left cleithrum, a long rectangle, is approximately
+equal in length to the rodlike stem of the clavicle, and is about
+as wide as the dorsal L-shaped tip of the clavicle. The posterior
+end of the cleithrum presumably met the tip of the clavicle, while
+the rest of it was directed anteriorly and a little dorsally. There
+seems to be a small articular surface near the anterior extremity
+which suggests the presence of a supracleithrum. The upper border
+of the cleithrum is slightly convex and the lower concave.</p>
+
+
+<h2>FORELIMB (Fig. <a href="#fig11">11</a>)</h2>
+
+<p>The left forelimb is the only one present and appears to be nearly
+complete, although the elements are scattered almost at random.
+The only parts of the forelimb known to be missing are two subterminal
+and two terminal phalanges, probably of the first and
+third digits, and the proximal end of the second metacarpal. The
+smooth and relatively flat surfaces suggest an aquatic rather than
+terrestrial limb; only the proximal half of the humerus bears any
+conspicuous ridges or depressions. As we restore the skeleton of
+the limb, several features are remarkable: The humerus, ulna, and
+ulnare align themselves as the major axis of the limb, each carrying
+on its posterior edge a process or flange comparable to those in the
+axial series of a rhipidistian fin. The remaining elements take positions
+comparable to the diagonally placed preaxial radials in such
+a fin. The digits appear to have been short, perhaps with no more
+than two phalanges. There is only one row of carpals present (the
+proximal row of other tetrapods). A second and third row would
+be expected in primitive Amphibia; if they existed in <i>Hesperoherpeton</i>
+they must either have been wholly cartilaginous or washed<span class='pagenum'><a name="Page_234" id="Page_234">[Pg 234]</a></span>
+away from the specimen. Neither of these alternatives seems at all
+likely to us in view of the well-ossified condition of the elements that
+are present, and the occurrence of both the proximal carpals and
+the metacarpals. The space available for metacarpals probably
+could not have contained more than the four that are recognized.</p>
+
+<p class="figcenter" style="width: 438px;">
+<a name="fig11" id="fig11"></a>
+<img src="images/image011.png" width="438" height="600" alt="Fig. 11. Hesperoherpeton garnettense Peabody. Left forelimb,
+showing characters of both a crossopterygian fin and an amphibian
+foot. KU&nbsp;10295, &times;&nbsp;4." title="Fig. 11. Hesperoherpeton garnettense Peabody. Left forelimb,
+showing characters of both a crossopterygian fin and an amphibian
+foot. KU&nbsp;10295, &times;&nbsp;4." />
+<span class="caption">Fig. 11. <i>Hesperoherpeton garnettense</i> Peabody. Left forelimb,
+showing characters of both a crossopterygian fin and an amphibian
+foot. KU&nbsp;10295, &times;&nbsp;4.</span>
+</p>
+
+<p>The proximal end of the humerus is more rounded anteriorly
+than posteriorly, and has a thin articular border that bore a cartilaginous<span class='pagenum'><a name="Page_235" id="Page_235">[Pg 235]</a></span>
+cap as the primary surface for articulation with the
+scapulocoracoid. Although the unfinished surface of the head extends
+down the anterior margin about a third the length of the
+humerus, the shaft has been broken and so twisted that the distal
+part is not in the same plane as the proximal. Immediately posterior
+to the cartilaginous cap is a round, deep notch bordered posteriorly
+by the dorsal process of the head.</p>
+
+<p>The shaft is longer and narrower than would be anticipated in a
+primitive amphibian limb (cf. Romer, 1947). The distal end bears
+two surfaces for articulation with the radius and ulna. The full extent
+of the former surface was not determined because the more anterior
+part of the expanded end is represented only by an impression.
+The surface nearest the ulna was partially rounded for articulation
+with that element, the remaining posterior edge being broadly concave.
+The most striking feature of the humerus is a slender hooklike
+process on the posterior edge near the distal end, probably
+homologous with (1) the posterior flange on the "humerus" in Rhipidistia,
+and (2) the entepicondyle of the humerus in <i>Archeria</i> (Romer,
+1957) and other tetrapods.</p>
+
+<p>The radius is about the same width proximally as distally. The
+curvature of the shaft is approximately alike on both sides. Distally
+the surface is rounded for articulation with the radiale and perhaps
+the intermedium.</p>
+
+<p>The proximal end of the ulna is similar to that of the radius but is
+slightly larger. Posteriorly, there is a short, broad expansion resembling
+the entepicondyle of the humerus, and even more nearly
+like the postaxial flanges in a crossopterygian fin.</p>
+
+<p>The ends of the radiale are expanded and rounded, the entire bone
+being approximately twice as long as wide. The three sides of the
+intermedium are similarly convex. The surface of this bone is unfinished,
+showing that it must have been embedded in cartilage.
+The ulnare is conspicuously similar to the ulna in bearing a posterior
+hooklike expansion, and is larger than the radiale.</p>
+
+<p>The four metacarpals are slightly expanded proximally and distally.
+Although measurements of length and width are tabulated
+below (Table 2), we are not certain of the sequence of these bones
+in the row.</p>
+
+<p>The dimensions of the two proximal phalanges are alike. The
+shape of these elements is similar to that of the metacarpals. The
+two terminal phalanges are somewhat triangular in shape, the lateral
+edges being concave and the proximal convex.<span class='pagenum'><a name="Page_236" id="Page_236">[Pg 236]</a></span></p>
+
+<p class="center">
+<b><span class="smcap">Table 2.&mdash;Approximate Measurements of the Forelimb</span> (in mm.)<br />
+</b></p>
+
+
+<div class='center'>
+<table border="1" cellpadding="4" cellspacing="0" summary="forelimb">
+<tr><th rowspan="3">Element</th><th colspan="4">Dimensions</th></tr>
+<tr><th rowspan="2">Length</th><th colspan="3">Width</th></tr>
+<tr><th>Proximal</th><th>Midway</th><th>Distal</th></tr>
+<tr><td align='left'>Humerus</td><td align='center'>16.0</td><td align='center'>5.0</td><td align='center'>2.0</td><td align='center'>7.5?</td></tr>
+<tr><td align='left'>Radius</td><td align='center'>9.0</td><td align='center'>4.0</td><td align='center'>1.5</td><td align='center'>3.5</td></tr>
+<tr><td align='left'>Ulna</td><td align='center'>8.5</td><td align='center'>4.5</td><td align='center'>1.5</td><td align='center'>3.5</td></tr>
+<tr><td align='left'>Radiale</td><td align='center'>3.0</td><td align='center'>2.0</td><td align='center'>1.5</td><td align='center'>2.0</td></tr>
+<tr><td align='left'>Intermedium</td><td align='center'>1.5</td><td align='center'>&mdash;</td><td align='center'>2.0</td><td align='center'>&mdash;</td></tr>
+<tr><td align='left'>Ulnare</td><td align='center'>3.5</td><td align='center'>2.0</td><td align='center'>2.0</td><td align='center'>2.5</td></tr>
+<tr><td align='left'>Metacarpal<span style="text-align:right">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;A</span></td><td align='center'>4.5</td><td align='center'>2.5</td><td align='center'>1.0</td><td align='center'>2.0</td></tr>
+<tr><td align='left'>Metacarpal<span style="text-align:right">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;B</span></td><td align='center'>4.5</td><td align='center'>3.0?</td><td align='center'>1.5</td><td align='center'>2.5</td></tr>
+<tr><td align='left'>Metacarpal<span style="text-align:right">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;C</span></td><td align='center'>4.0</td><td align='center'>2.0</td><td align='center'>1.5</td><td align='center'>2.0</td></tr>
+<tr><td align='left'>Metacarpal<span style="text-align:right">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;D</span></td><td align='center'>3.5</td><td align='center'>2.5</td><td align='center'>1.0</td><td align='center'>1.5</td></tr>
+<tr><td align='left'>Proximal Phalanx<span style="text-align:right">&nbsp;&nbsp;A</span></td><td align='center'>2.0</td><td align='center'>1.5</td><td align='center'>1.0</td><td align='center'>1.5</td></tr>
+<tr><td align='left'>Proximal Phalanx<span style="text-align:right">&nbsp;&nbsp;B</span></td><td align='center'>2.0</td><td align='center'>1.5</td><td align='center'>1.0</td><td align='center'>1.5</td></tr>
+<tr><td align='left'>Terminal Phalanx<span style="text-align:right">&nbsp;&nbsp;A</span></td><td align='center'>1.5</td><td align='center'>1.5</td><td align='center'>1.0</td><td align='center'>1.0</td></tr>
+<tr><td align='left'>Terminal Phalanx<span style="text-align:right">&nbsp;&nbsp;B</span></td><td align='center'>1.5</td><td align='center'>1.5</td><td align='center'>1.0</td><td align='center'>1.0</td></tr>
+</table></div>
+
+
+
+<h2>COMPARISONS AND DISCUSSION</h2>
+
+<p>Apparently primitive rhipidistian characters in <i>Hesperoherpeton</i>
+are: Braincase in two sections, posterior one containing an expanded
+notochordal canal; lateral series of mandibular bones closely resembling
+that of <i>Megalichthys</i>, as figured by Watson (1926); tabular
+having long process probably articulating with pectoral girdle; lack
+of movement between head and trunk correlated with absence of
+occipital condyle; sensory pits present on frontal and squamosal.</p>
+
+<p>Although we are unable to separate, by sutures, the vomers from
+the palatines, the palatal surface of these bones and of the pterygoids
+is studded by numerous small teeth, as in Rhipidistia (Jarvik,
+1954) and some of the early Amphibia (Romer, 1947). The stapes
+apparently reaches the quadrate, and could therefore serve in hyostylic
+suspension of the upper jaw.</p>
+
+<p>The pectoral limb has an axial series of bones carrying hooklike
+flanges on their posterior edges. The other bones of the limb show
+little modification of form beyond the nearly flat, aquatic type seen
+in Rhipidistia. No distinct elbow or wrist joints are developed.</p>
+
+<p>Characters of <i>Hesperoherpeton</i> common to most primitive Amphibia,
+in contrast with Crossopterygii, are: Nares separated from
+edge of jaw; stapes having external process that may have met a
+tympanic membrane, thus giving the bone a sound-transmitting
+function. Apparently none of the opercular series was present.<span class='pagenum'><a name="Page_237" id="Page_237">[Pg 237]</a></span></p>
+
+<p>There are two large palatal teeth, slightly labyrinthine in character,
+adjacent to each internal naris. The scapulocoracoid, as
+shown by Peabody (1958), is Anthracosaurian in structure, as are the
+long-stemmed clavicles. The limbs have digits rather than fin-lobes,
+although the digital number apparently is four and the number of
+bones in the manus is less than would be expected in a primitive amphibian.
+The vertebrae are similar to those of Ichthyostegids, as
+described by Jarvik (1952), except that the pleurocentra are much
+larger.</p>
+
+<p>In addition to this remarkable combination of crossopterygian
+and amphibian characters, <i>Hesperoherpeton</i> is specialized in certain
+features of the skull. The orbits are much enlarged, probably in correlation
+with the diminutive size of the animal, and this has been
+accompanied by loss of several bones. The frontal and squamosal
+nearly meet each other, and both form part of the rim of the orbit.
+The bones of the posterior part of the dermal roof are greatly reduced,
+and there is none behind the squamosal except the projecting
+tabular; there is no indication of quadratojugal, jugal, intertemporal
+or postparietal. The foramen magnum is enormous. The external
+surfaces of the bones of the skull are nearly smooth.</p>
+
+<p>Is it possible that the "primitive" and "specialized" features of
+this animal are actually larval? Are they not just the kind of characters
+that would be expected in an immature, aquatic embolomere
+of Pennsylvanian time? For several reasons we do not think this is
+the case. Except for the anterior part of the braincase, there is no
+indication that the skeleton was not well ossified. The postaxial
+processes on the humerus, ulna and ulnare could scarcely have been
+larval features only, since they are so clearly homologous with those
+in adult Rhipidistia; a larval limb should indeed be simple, but its
+simplicity is unlikely to involve paleotelic adult characters. The
+scapulocoracoid of our specimen is of practically the same shape and
+size as that in the only other known individual, the type; this would
+be probable if both were adults, but somewhat less likely if they
+were larvae of a much larger animal. The form of the stapes, tabular
+and otic notch suggest a functional tympanic membrane, which
+could not have occurred in a gill-breathing larva. On the other
+hand, an adult animal of pigmy size might be expected to have
+large orbits, large otic capsules and a large foramen magnum.</p>
+
+<p>We conclude that <i>Hesperoherpeton</i> lived and sought food in the
+weedy shallows at the margin of a pond or lagoon, and that for much
+of the time its head was partly out of water (Fig. <a href="#fig12">12</a>). The animal
+could either steady itself or crawl around by means of the paddlelike<span class='pagenum'><a name="Page_238" id="Page_238">[Pg 238]</a></span>
+limbs, but these probably could not be used in effective locomotion
+on land. Like the Ichthyostegids, it probably swam by means
+of a fishlike tail.</p>
+
+<p class="figcenter" style="width: 600px;">
+<a name="fig12" id="fig12"></a>
+<img src="images/image012.png" width="600" height="203" alt="Fig. 12. Hesperoherpeton garnettense Peabody. Probable appearance
+in life. &times;&nbsp;0.5." title="Fig. 12. Hesperoherpeton garnettense Peabody. Probable appearance
+in life. &times;&nbsp;0.5." />
+<span class="caption">Fig. 12. <i>Hesperoherpeton garnettense</i> Peabody. Probable appearance
+in life. &times;&nbsp;0.5.</span>
+</p>
+
+
+<h2>TAXONOMY</h2>
+
+<p>Evidently <i>Hesperoherpeton</i> is a small, lagoon-dwelling survivor
+of the Devonian forms that initiated the change from Crossopterygii
+to Amphibia (Jarvik, 1955). It shows, however, that this transition
+did not affect all structures at the same time, for some, as the braincase
+with its notochordal canal, the mandibular bones and axial limb
+bones, are unchanged from the condition normal for the Rhipidistia,
+but most other characters are of amphibian grade. To express these
+facts taxonomically requires that <i>Hesperoherpeton</i> be removed from
+the family Cricotidae, suborder Embolomeri, order Anthracosauria,
+and placed in a new order and family of labyrinthodont Amphibia.</p>
+
+<h3><big>Order PLESIOPODA</big></h3>
+
+<p class="center">(<i>plesios</i>, Gr., near, almost; <i>podos</i>, Gr., foot)</p>
+
+<div class="blockquot">
+<p>Labyrinthodontia having limbs provided with digits, but retaining
+posterior flanges on axial bones as in Rhipidistia, without
+joint-structure at elbow and wrist essential for terrestrial
+locomotion; neurocranium having separate otico-occipital section,
+large notochordal canal, no occipital condyle, as in Rhipidistia;
+nares separate from rim of mouth; pectoral girdle anthracosaurian;
+vertebrae having U-shaped intercentrum and
+paired, but large, pleurocentra.</p></div>
+
+<p>Probably associated with the characters of the order, as given
+above, are the connection of pectoral girdle with skull, and the presence
+of a tympanic membrane, the stapes functioning in both sound-transmission
+and palatoquadrate suspension.<span class='pagenum'><a name="Page_239" id="Page_239">[Pg 239]</a></span></p>
+
+<h3><big>Family HESPEROHERPETONIDAE</big></h3>
+
+<div class="blockquot">
+<p>Orbits and foramen magnum unusually large in correlation
+with reduced size of animal; squamosal forming posterior margin
+of orbit; circumorbital series absent (except for postorbital);
+sensory pits on squamosal and frontal.</p></div>
+
+<p>Characters defining the family are evidently the more specialized
+cranial features, which probably evolved during Mississippian and
+early Pennsylvanian times.</p>
+
+<p>The definition of the genus and species may be left to rest upon
+Peabody's (1958) original description and the present account,
+until the discovery of other members of the family gives reason for
+making further distinctions.</p>
+
+
+<h2>SUMMARY</h2>
+
+<p><i>Hesperoherpeton garnettense</i> Peabody (1958), based on a scapulocoracoid
+and part of a vertebra, was originally placed in the order
+Anthracosauria, suborder Embolomeri, family Cricotidae. A new
+skeleton from the type locality near Garnett, Kansas (Rock Lake
+shale, Stanton formation, Upper Pennsylvanian), shows that the animal
+has the following rhipidistian characters: Large notochordal
+canal below foramen magnum, otico-occipital block separate from
+ethmosphenoid, postaxial processes on three axial bones of forelimb,
+pectoral girdle (probably) articulated with tabular. Nevertheless,
+<i>Hesperoherpeton</i> has short digits, an anthracosaurian type of pectoral
+girdle, an otic rather than spiracular notch, nostrils separate
+from the mouth, and vertebrae in which the intercentrum is U-shaped
+and the pleurocentra large but paired. The stapes reaches the quadrate.</p>
+
+<p><i>Hesperoherpeton</i> is placed in a new order, PLESIOPODA, on the
+basis of the characters stated above, and a new family, HESPEROHERPETONIDAE.
+Specialized characters of the family include:
+Reduction of circumorbital bones, bringing the squamosal to the
+edge of the orbit, loss of certain bones of the temporal region, and
+relative enlargement of the orbits and foramen magnum, in correlation
+with the diminutive size of the animal. The structural
+characters of <i>Hesperoherpeton</i> suggest to us that it lived in the
+shallow, weedy margins of lagoons, rested with its head partly out
+of water, and normally did not walk on land.<span class='pagenum'><a name="Page_240" id="Page_240">[Pg 240]</a></span></p>
+
+
+<h2>LITERATURE CITED</h2>
+
+<p>
+<span class="smcap">Eaton, T. H., Jr.</span><br />
+<span class="i4">1951. Origin of tetrapod limbs. Amer. Midl. Nat., 46: 245-251.</span><br />
+<br />
+<span class="smcap">Jarvik, E.</span><br />
+<span class="i4">1952. On the fish-like tail in the ichthyostegid stegocephalians. Meddel.
+om Gr&oslash;nland, 114: 1-90.</span><br />
+<span class="i4">1954. On the visceral skeleton in <i>Eusthenopteron</i> with a discussion of the
+parasphenoid and palatoquadrate in fishes. Kgl. Svenska Vetenskapsakad.
+Handl., 5: 1-104.</span><br />
+<span class="i4">1955. The oldest tetrapods and their forerunners. Sci. Monthly, 80: 141-154.</span><br />
+<br />
+<span class="smcap">Moore, R. C.</span>, <span class="smcap">Frye, J. C.</span>, and <span class="smcap">Jewett, J. M.</span><br />
+<span class="i4">1944. Tabular description of outcropping rocks in Kansas. Kansas State
+Geol. Surv. Bull., 52: 137-212.</span><br />
+<br />
+<span class="smcap">Peabody, F. E.</span><br />
+<span class="i4">1952. <i>Petrolacosaurus kansensis</i> Lane, a Pennsylvanian reptile from Kansas.
+Univ. Kansas Paleont. Contrib., Vertebrata, Art. 1: 1-41.</span><br />
+<span class="i4">1958. An embolomerous amphibian in the Garnett fauna (Pennsylvanian)
+of Kansas. Jour. Paleont., 32: 571-573.</span><br />
+<br />
+<span class="smcap">Romer, A. S.</span><br />
+<span class="i4">1937. The braincase of the Carboniferous crossopterygian <i>Megalichthys</i>
+<i>nitidus</i>. Mus. Comp. Zool. Bull., 82: 1-73.</span><br />
+<span class="i4">1947. Review of the Labyrinthodontia. Mus. Comp. Zool. Bull., 99: 1-368.</span><br />
+<span class="i4">1957. The appendicular skeleton of the Permian embolomerous amphibian
+<i>Archeria</i>. Univ. Michigan Contrib. Mus. Paleont., 13: 103-159.</span><br />
+<br />
+<span class="smcap">Watson, D. M. S.</span><br />
+<span class="i4">1926. The evolution and origin of the Amphibia. Phil. Trans. Roy. Soc.
+London, (B) 214: 189-257.</span><br />
+</p>
+
+<p><i>Transmitted January 13, 1960.</i></p>
+
+
+<p class="center"><small>28-2495</small></p>
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
+End of the Project Gutenberg EBook of A New Order of Fishlike Amphibia From
+the Pennsylvanian of Kansas, by Theodore H. Eaton and Peggy Lou Stewart
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+The Project Gutenberg EBook of A New Order of Fishlike Amphibia From the
+Pennsylvanian of Kansas, by Theodore H. Eaton and Peggy Lou Stewart
+
+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: A New Order of Fishlike Amphibia From the Pennsylvanian of Kansas
+
+Author: Theodore H. Eaton
+        Peggy Lou Stewart
+
+Release Date: January 23, 2010 [EBook #31050]
+
+Language: English
+
+Character set encoding: ASCII
+
+*** START OF THIS PROJECT GUTENBERG EBOOK NEW ORDER OF FISHLIKE AMPHIBIA ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper, Diane Monico, and
+the Online Distributed Proofreading Team at
+https://www.pgdp.net
+
+
+
+
+
+
+
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS
+
+MUSEUM OF NATURAL HISTORY
+
+Volume 12, No. 4, pp. 217-240, 12 figs.
+May 2, 1960
+
+
+A New Order of Fishlike Amphibia
+From the Pennsylvanian of Kansas
+
+BY
+
+THEODORE H. EATON, JR., AND PEGGY LOU STEWART
+
+
+UNIVERSITY OF KANSAS
+LAWRENCE
+1960
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
+Robert W. Wilson
+
+
+Volume 12, No. 4, pp. 217-240, 12 figs.
+Published May 2, 1960
+
+
+UNIVERSITY OF KANSAS
+Lawrence, Kansas
+
+
+PRINTED IN
+THE STATE PRINTING PLANT
+TOPEKA, KANSAS
+1960
+
+28-2495
+
+
+
+
+A New Order of Fishlike Amphibia
+From the Pennsylvanian of Kansas
+
+BY
+
+THEODORE H. EATON, JR., AND PEGGY LOU STEWART
+
+INTRODUCTION
+
+
+A slab of shale obtained in 1955 by Mr. Russell R. Camp from a
+Pennsylvanian lagoon-deposit in Anderson County, Kansas, has yielded in
+the laboratory a skeleton of the small amphibian _Hesperoherpeton
+garnettense_ Peabody (1958). This skeleton provides new and surprising
+information not available from the holotype, No. 9976 K. U., which
+consisted only of a scapulocoracoid, neural arch, and rib fragment. The
+new specimen, No. 10295 K. U., is of the same size and stage of
+development as the holotype and it is thought that both individuals are
+adults.
+
+The quarry, University of Kansas Museum of Natural History Locality KAN
+1/D, is approximately six miles northwest of Garnett, Anderson County,
+Kansas, in Sec. 5, T. 19S, R. 19E, 200 yards southwest of the place
+where _Petrolacosaurus kansensis_ Lane was obtained (see Peabody,
+1952). The Rock Lake shale, deposited under alternately marine and
+freshwater lagoon conditions, is a thin member of the Stanton limestone
+formation, Lansing group, Missourian series, and thus is in the lower
+part of the Upper Pennsylvanian.
+
+Peabody (1958) placed _Hesperoherpeton_ in the order Anthracosauria,
+suborder Embolomeri, family Cricotidae. Study of the second and more
+complete specimen reveals that _Hesperoherpeton_ is unlike the known
+Embolomeri in many important features. The limbs and braincase are more
+primitive than those so far described in any amphibian. The vertebrae
+are comparable to those of Ichthyostegalia (Jarvik, 1952), as well as
+to those of Embolomeri. The forelimb is transitional between the
+pectoral fin of Rhipidistia and the limb of early Amphibia. The pattern
+of the bones of the forelimb closely resembles, but is simpler than,
+that of the hypothetical transitional type suggested by Eaton (1951).
+The foot seemingly had only four short digits. The hind limb is not
+known.
+
+The new skeleton of _Hesperoherpeton_ lies in an oblong block of limy
+shale measuring approximately 100 x 60 mm. After preparation of the
+entire lower surface, the exposed bones and matrix were embedded in
+Bioplastic, in a layer thin enough for visibility but giving firm
+support. Then the specimen was inverted and the matrix removed from the
+opposite side; this has not been covered with Bioplastic. The bones lie
+in great disorder, except that some parts of the roof of the skull are
+associated, and the middle section of the vertebral column is
+approximately in place. The bones of the left forelimb are close
+together but not in a natural position. The tail, pelvis, hind limbs
+and right forelimb are missing. Nearly all the bones present are
+broken, distorted by crushing, incomplete and scattered out of place,
+probably by the action of currents. The complete skeleton, in life,
+probably measured between 150 and 200 mm. in length.
+
+The specimen was studied at the Museum of Natural History, University
+of Kansas, with the help of a grant from the National Science
+Foundation, number NSF-G8624. The specimen was discovered in the slab
+by Miss Sharon K. Moriarty, and was further cleaned by the authors. Mr.
+Merton C. Bowman assisted with the illustrations. We are indebted to
+Dr. Robert W. Wilson for critical comments.
+
+
+SKULL
+
+_Dorsal Aspect_ (Figs. 1, 2)
+
+In reconstruction, the skull measures approximately 8.0 mm.
+dorsoventrally at the posterior end. The height diminishes anteriorly
+to about 1.5 mm. at the premaxillary. The length is about 15.5 mm. in
+the median line, or 24.0 mm. to the tip of the tabular, and the width
+about 16.0 mm. posteriorly. The snout is blunt, continuing about 1-2
+mm. anterior to the external nares. Each of the tabulars has a slender
+posterior process 5.0 mm. long, which probably met the supracleithrum;
+the intertabular space is about 8.5 mm. wide. The orbits are
+approximately 5.5 mm. in diameter and extend from the maxillary to
+within about 3.0 mm. of the midline dorsally. The pineal opening is 1.8
+mm. anterior to the occipital margin of the skull.
+
+Reduction of bones at the back of the skull seems to have eliminated
+any dermal elements posterior to the squamosal, while enlargement of
+the orbit has removed most of the postorbital series, leaving the
+squamosal as the only cheekbone. There is apparently no jugal or
+postfrontal.
+
+The squamosal of _Acanthostega_ (Jarvik, 1952) is articulated under the
+tabular and reaches forward and down, much as if it were an opercular
+in reversed position. Internally, it must lie against the otic capsule
+below the tabular, partially concealing the stapes. The bone that we
+suppose to be the squamosal of _H. garnettense_ is of similar shape, of
+about the same size and has internally an articular surface at one
+corner, bounded by a pair of ridges in the shape of a V. This articular
+surface probably fitted on a lateral process extending from the roof of
+the neurocranium, over the front of the otic capsule.
+
+The premaxillary extends posterolaterally to a distance 5.5 mm. from
+the midline and attains a width at its broadest point of about 1.5 mm.
+The posterior edge is slightly concave and in part forms the anterior
+border of the naris.
+
+[Illustration: FIG. 1. _Hesperoherpeton garnettense_ Peabody. Skull,
+dorsal view. Postorbital processes of the neurocranium are shown in
+dotted outline. KU 10295, x 4.]
+
+The nasal is triangular and, with the lacrimal, forms the medial border
+of the naris. The length of the medial side of the nasal bone is
+approximately 5.0 mm., the transverse width is 3.8 mm., and the extent
+of the posterolateral border is 5.5 mm.
+
+The maxillary meets the premaxillary lateral to the naris, borders the
+naris posteroventrally, and continues posteriorly beneath the orbit, of
+which it forms the external border. The maxillary is about 8.5 mm.
+long, and immediately anterior to the orbit has a maximum width of 1.3
+mm.
+
+The lacrimal fills the remaining rim of the narial opening between the
+nasal and maxillary, and extends to the anterior edge of the orbit. The
+length, from naris to orbit, is 4.2 mm.; the width ranges from 1.0 mm.
+anteriorly to 2.5 mm. posteriorly.
+
+[Illustration: FIG. 2. _Hesperoherpeton garnettense_ Peabody. Skull,
+lateral view, showing relatively large orbit and absence of smaller
+circumorbital bones. KU 10295, x 4.]
+
+The external naris is approximately 1.0 mm. in diameter. It is slightly
+anterodorsal to the internal naris and 4.0 mm. lateral to the midline.
+
+The dorsal margin of the orbit appears to be formed by the frontal. The
+anterior part of this margin, however, may be formed by a prefrontal,
+which is not clearly set off by a suture. The frontal extends 3.8 mm.
+in the midline, and anteriorly and laterally borders the nasal and
+lacrimal, respectively. A faint pattern of pitting radiates on the
+surface from the center of ossification of the frontal. There is also a
+pit indicating the presence of a supraorbital sensory pore.
+
+The parietal bones enclose the pineal opening, approximately 2.5 mm.
+posterior to the suture with the frontal. The foramen is about 0.5 mm.
+in diameter. Laterally the parietal meets the medial angle of the
+postorbital and the medial border of the supratemporal. No bone of this
+animal shows the deep pitting and heavy ornamentation characteristic of
+many primitive Amphibia.
+
+The postorbital meets the anterolateral corner of the parietal for a
+distance of 0.5 mm., the anterior edge bordering the frontal bone and
+the orbit for a combined distance of about 3.0 mm. The lateral margin
+is slightly convex, and is probably interrupted behind by the anterior
+point of the tabular. Medially, the concave margin of the postorbital
+meets the supratemporal for about 3.5 mm.
+
+The supratemporal is thus wedge-shaped and located between the parietal
+and the postorbital. The posterior edge of the supratemporal protrudes
+as a convex border slightly behind the end of the parietal, and
+measures 3.0 mm. around the curve to the parietal suture.
+
+[Illustration: FIG. 3. _Hesperoherpeton garnettense_ Peabody. A, left
+squamosal, internal surface. B, left squamosal, external surface. C,
+right tabular internal surface. D, right tabular, external surface. KU
+10295, all x 4.]
+
+The squamosal (Fig. 3 A, B) is a large, somewhat rectangular bone
+extending from the back of the orbit to the posterior extremity of the
+cheek. It outlines almost entirely the posterior border of the orbit,
+the ventrolateral portion of the cheek region, and the lateral border
+of the top of the skull behind the orbit. Dorsally, the squamosal meets
+the anterior half of the tabular and the lateral border of the
+supratemporal. Near the anteroventral edge of the squamosal there is a
+small pit, probably related to a postorbital sensory pore in the skin.
+
+The tabular (Fig. 3 C, D) is pointed anteriorly, where it probably fits
+against the lateroposterior edge of the postorbital. The dorsal part of
+the bone flares out and down, forming a small otic notch at a point
+halfway back. Posteriorly, the flange attains a dorsoventral width of
+2.0 mm. at the edge of the notch. The slender posterior process of the
+tabular which continues beyond the flange is approximately 0.5 mm. in
+diameter and 5.0 mm. long.
+
+_Ventral Aspect_ (Fig. 4)
+
+The palatal view of the skull shows the paired premaxillary, maxillary,
+palatine, pterygoid, and quadrate bones. The openings for the internal
+nares, the ventral orbital fenestrae, and the subtemporal fossae are
+readily recognized. The quadrate processes extend posteriorly leaving a
+large gap medially at the posterior end of the skull.
+
+[Illustration: FIG. 4. _Hesperoherpeton garnettense_ Peabody. Palate
+reconstructed; ventral aspect at left, showing teeth, dorsal aspect at
+right. KU 10295, x 4.]
+
+The left quadrate appears to be in place on the posterior prong of the
+pterygoid. The dorsal side of the quadrate is grooved between two
+anterolaterally directed ridges. The groove, which probably held the
+end of the stapes, extends about half the width of the quadrate itself.
+The width of the quadrate is 4.0 mm., the length is 4.5 mm. medially
+and about 2.0 mm. laterally. In ventral view the quadrate appears to
+project laterally, but is incomplete and its shape uncertain. The
+distance from the posterior end of the quadrate to the visible
+posterior edge of the orbital fenestra, which opens ventrally, is 10.0
+mm.
+
+This region between the quadrate and the orbit is occupied by a
+pterygoid with three projections. Anteriorly, the pterygoid outlines
+most of the posterior edge of the orbit (a distance of about 6.5 mm.).
+A lateral process separates the orbit from the subtemporal fossa. A
+posteriorly directed edge defines the fossa, which extends about 6.5
+mm. anteroposteriorly. The lateral process of the pterygoid terminates
+10.0 mm. from the midline. Both the lateral and posterior pterygoid
+processes are approximately 2.0 mm. wide. The greatest width of the
+subtemporal fossa is about 2.0 mm. The medial border of the orbital
+fenestra is missing, but apparently consisted of the pterygoid for at
+least the posterior half.
+
+Along the posterior edge of the orbital fenestra, there is a narrow,
+dorsally projecting flange of the pterygoid. The lateral opening of the
+orbit is approximately 7.5 mm. wide.
+
+The remaining border of the orbital fenestra on the anterior and medial
+sides is formed by a bone occupying the position of palatine and vomer;
+for convenience we designate this as palatine. When reconstructed in
+its probable position in relation to the pterygoid, the left palatine
+lacks a section, on its medial and posterior edges, measuring about 2.5
+mm. by 9.0 mm. The lateral margin of the palatine is convex; about 5.5
+mm. anterior to the orbit this margin curves into a strong anteriorly
+pointing projection, medial to which is seen the internal narial
+opening. The remaining anterior edge is slightly convex, smoothly
+rounded, and meets the midline about 9.0 mm. anterior to the pterygoid.
+
+The void area medial to the palatine and anterior to the pterygoid does
+not fit any bone which we can recognize as the parasphenoid. It is thus
+suspected that this area is covered in part by the missing edge of the
+palatine and partly by an anteromedial extension of the pterygoid. Of
+course a parasphenoid may also have been present.
+
+The position, length, and shape of the premaxillary shown in palatal
+view (Fig. 4) are primarily based upon the dorsal appearance since
+ventrally most of it cannot be seen. At the point where it forms the
+anterior border of the internal naris, the premaxillary is slightly
+wider than the maxillary and seems to become narrower as it approaches
+the midline.
+
+The ethmosphenoid, which we cannot identify, may have been exposed in a
+gap between the premaxillary and the palatine. The gap measures
+approximately 8.0 mm. wide and ranges up to 1.0 mm. anteroposteriorly.
+
+The maxillary begins at a suture with the premaxillary lateral to the
+naris and continues posteriorly, bordering the orbit with a width of
+about 1.2 mm. It then tapers to a point approximately 2.0 mm. anterior
+to the lateral projection of the pterygoid. The width of the maxillary
+at this point is 0.8 mm. and the posterior end is broken; probably when
+complete it approached the pterygoid, and either met the latter or had
+a ligamentous connection with it. As nearly as can be determined, the
+total length of the maxillary is approximately 12.0 mm.
+
+The teeth on the maxillary are small and seem to be in two longitudinal
+rows. The palatine bears two large, grooved teeth anteriorly; the first
+is approximately 1.0 mm. posteromedial to the naris and the second is
+about 3.0 mm. posterior and slightly lateral to the naris. The flat
+ventral surfaces of the palatine and pterygoid bear numerous small
+teeth distributed as shown in Fig. 4.
+
+_Braincase and Occipital Region_ (Fig. 5)
+
+The parts of the neurocranium are scattered, disconnected and
+incomplete, but it is possible to make out a number of features of the
+otico-occipital section with fair assurance. In posterior view the
+notochordal canal and foramen magnum are confluent with each other, and
+of great size relative to the skull as a whole. The notochordal canal
+measures 2.8 mm. in diameter, and the foramen magnum about 4.0 mm. The
+crescent-shaped supraoccipital rests on the upright ends of the
+exoccipitals, but between the latter and the basioccipital no sutures
+can be seen. Probably the whole posterior surface of the braincase
+slanted posteroventrally; consequently the rim of the notochordal canal
+was about 3.0 mm. behind the margin of the parietals.
+
+The U-shaped border of the notochordal canal is a thick, rounded bone,
+comparable in appearance to the U-shaped intercentra of the vertebrae.
+This bone apparently rested upon a thinner, troughlike piece (Fig. 5 B)
+forming the floor of the braincase. The latter is broad, shallow,
+concave, open midventrally and narrowing anteriorly to form a pair of
+articular processes. Since no sutures can be seen in this structure, it
+probably is the ventral, ossified portion of the basioccipital. Watson
+(1926, Fig. 4 B) illustrates the floor of the braincase in
+_Eusthenopteron_, with its more lateral, anterior portion labelled
+prootic, but in our specimen the corresponding part could scarcely have
+formed the anterior wall of the otic capsule, being entirely in the
+plane of the floor. The two articular surfaces anteriorly near the
+midline suggest that a movable joint existed between the
+otico-occipital part of the braincase and the ethmosphenoid part, as in
+Rhipidistia (Romer, 1937). We have found nothing in the specimen that
+could be referred to the ethmosphenoid; it may have been unossified.
+
+[Illustration: FIG. 5. _Hesperoherpeton garnettense_ Peabody, KU 10295,
+x 4. A, occipital view of skull; B, basioccipital bone in dorsal
+(internal) view.]
+
+The otic capsules appear to have rested against lateral projections of
+the basioccipital. The single otic capsule that can be seen (the
+right) is massively built, apparently ossified in one piece, with a
+shallow dorsomedial excavation, probably the vestige of a supratemporal
+fossa. On the lateral face is a broad, shallow depression dorsally, and
+a narrower, deeper one anteroventrally; these we suppose to have
+received the broader and narrower heads of the stapes, respectively.
+The posterior wall of the otic capsule we have designated opisthotic in
+the figure. Anterior to the otic capsule the lateral wall of the
+braincase cannot be seen, and may not have been ossified.
+
+The roof of the braincase is visible in its ventral aspect, extending
+from approximately the occipital margin to a broken edge in front of
+the parietal foramen, and laterally to paired processes which overlie
+the otic capsules directly behind the orbits (see dotted outlines in
+Fig. 1). Each of these postorbital processes, seen from beneath,
+appears to be the lateral extension of a shallow groove beginning near
+the midline. Presumably this section of the roof is an ossification of
+the synotic tectum. It should be noted that the roof of the braincase
+proper is perfectly distinct from the overlying series of dermal bones,
+and that the parietal foramen can be seen in both. The roof of the
+braincase in our specimen seems to have been detached from the
+underlying otic capsules and the occipital wall.
+
+The bone that we take to be the stapes is blunt, flattened (perhaps by
+crushing), 5.0 mm. in length, and has two unequal heads; its width
+across both of these is 4.0 mm. The length is appropriate to fit
+between the lateral face of the otic capsule and the dorsal edge of the
+quadrate; the wider head rests on a posterodorsal concavity on the otic
+capsule, and the smaller fits a lower, more anterior pit. Laterally the
+stapes carries a short, broad process that probably made contact with a
+dorsally placed tympanic membrane. Thus the bone was a hyomandibular in
+the sense that it articulated with the quadrate, but it may also have
+served as a stapes in sound-transmission. It contains no visible canal
+or foramen.
+
+_Mandible_ (Fig. 6)
+
+The crushed inner surface of the posterior part of the left mandible
+and most of the external surface of the right mandible are preserved in
+close proximity. Although the whole length of the tooth-bearing margins
+is missing, some parts of six elements of the right mandible can be
+seen. The pattern of sutures and the general contour closely resemble
+those of _Megalichthys_ (Watson, 1926, Figs. 37, 38) and other known
+Rhipidistia.
+
+The anteroposterior length of the mandible is about 23.8 mm., and the
+depth is 3.8 mm. The dentary extends approximately 17.6 mm. back from
+the symphysis, and its greatest width is probably 2.0 mm. Its lower
+edge meets all the other lateral bones of the jaw. The splenial and
+postsplenial form the curved anteroventral half of the jaw for a
+distance of about 9.0 mm. The fragmented articular, on the posterior
+end of the jaw, is 4.0 mm. long and 2.0 mm. deep, exhibiting a broken
+upper edge; presumably the surface for articulation with the quadrate
+was a shallow concavity, above the end of the articular.
+
+[Illustration: FIG. 6. _Hesperoherpeton garnettense_ Peabody. Right
+mandible, lateral view, KU 10295, x 4. External surfaces are pitted;
+broken surfaces are coarsely stippled.]
+
+
+VERTEBRAE (Fig. 7)
+
+The vertebrae that are visible from a lateral view are crushed and
+difficult to interpret. It is possible, nevertheless, to see that the
+trunk vertebrae resemble those of Ichthyostegalia (Jarvik, 1952, Fig.
+13 A, B), except that the pleurocentra are much larger. A few parts of
+additional vertebrae can be seen, but they are so scattered that it is
+impossible to be sure of their original location. Therefore comparisons
+between different regions cannot yet be made.
+
+The U-shaped intercentrum encloses the notochord and occupies an
+anteroventral position in the vertebra. Anteriorly, each intercentrum
+articulates with the pleurocentra of the next preceding vertebra by
+slightly concave surfaces. Dorsolaterally there is an articular surface
+for the capitulum of the rib.
+
+The two pleurocentra of each vertebra are separate ventrally as well as
+dorsally, but form thin, broad plates of about the same height as the
+notochord. The lateral surface appears to be depressed, allowing,
+perhaps, for movement of the rib. Above each pleurocentrum, on the
+lateral surface of the neural arch, there is a short diapophysis for
+articulation with the tuberculum of the rib.
+
+The margin of the neural spine is convex anteriorly and concave
+posteriorly, the tip reaching a point vertically above the
+postzygapophysis. The prezygapophysis of each vertebra articulates
+with the preceding postzygapophysis by a smooth dorsal surface. One
+nearly complete neural arch shows (Fig. 7 B) a pit above the neural
+canal, clearly corresponding to the canal for a dorsal ligament shown
+by Jarvik in _Ichthyostega_. Indeed this view of the neural arch and
+intercentrum together brings out the striking resemblance between the
+vertebrae of _Hesperoherpeton_ and those of the Ichthyostegids. The
+rounded intercentrum in both is an incomplete ring enclosing the
+notochordal canal.
+
+[Illustration: FIG. 7. _Hesperoherpeton garnettense_ Peabody. A, End
+view of incomplete vertebra, probably near anterior end of column. B,
+Neural arch and intercentrum in end view, showing probable association.
+C, Left lateral view of trunk vertebra. All figures: KU 10295, x 4.]
+
+TABLE 1.--AVERAGE MEASUREMENTS OF THE TRUNK VERTEBRAE (in mm.).
+NUMBERS IN PARENTHESES INDICATE THE NUMBER OF PIECES AVAILABLE FOR
+MEASURING
+
+----------------------------+------------+-------------+---------------
+            PARTS           | Ant.-post. | Dors.-vent. | Transv. width
+----------------------------+------------+-------------+---------------
+Neural spine                |  1.5 (3)   |   3.0  (3)  |     --
+----------------------------+------------+-------------+---------------
+Neural spine and arch       |  2.0 (4)   |   4.5? (4)  |     --
+----------------------------+------------+-------------+---------------
+Neural canal                |  2.0 (4)   |   2.0  (1)  |    1.0 (1)
+----------------------------+------------+-------------+---------------
+Intercentrum                |  1.5 (5)   |   3.5  (4)  |    3.0 (1)
+----------------------------+------------+-------------+---------------
+Pleurocentrum               |  1.5 (3)   |   3.0  (2)  |     --
+----------------------------+------------+-------------+---------------
+
+The shape, in end view, of a partly preserved neural arch (Fig. 7 A)
+seems to account for the incompleteness of the intercentrum just
+mentioned; the ventral edge of the arch is emarginate in such a way as
+to fit the dorsal surface of the notochord. The dorsal portion of this
+neural arch is not present (either broken or not yet ossified), but
+the opening of the neural canal is comparable in width to the foramen
+magnum. Hence this vertebra may be one of the most anterior in the
+column. In comparison with the trunk vertebrae seen farther posteriorly
+it appears that there may be a progressive ossification of neural
+arches toward their dorsal ends, and of intercentra around the
+notochord, with probable fusion of the intercentra and neural arches in
+the posterior part of the trunk. The notochord seems to have been
+slightly constricted by the intercentra, but not interrupted.
+
+
+RIBS
+
+The proximal ends of the ribs expand dorsoventrally to a width
+approximately four times that of their slender shafts. The tuberculum
+and capitulum on each of the trunk ribs are separated only by a shallow
+concavity. These two articular surfaces are so situated that the rib
+must tilt downward from the horizontal plane. The shaft flares
+terminally in some ribs, and the distal end is convex. Ribs in the
+trunk region differ little if any in size. Five that can be measured
+vary in length from 5.0 to 7.0 mm. One short, bent rib 3.5 mm. long
+perhaps is sacral or caudal.
+
+
+PECTORAL GIRDLE (Figs. 8, 9, 10)
+
+The right scapulocoracoid is almost complete, and the left one is
+present but partly broken into three pieces, somewhat pushed out of
+position. With the advantage of this new material, we may comment on
+the scapulocoracoid of _H. garnettense_ as described by Peabody (1958).
+In size and contour, the slight differences between the type (KU 9976)
+and the new skeleton (KU 10295) are considered to be no more than
+individual variation. We have redrawn the type (Fig. 8) in order to
+show the resemblances more clearly.
+
+The small sections that were missing from the type are present in KU
+10295. The jagged edge directly posterior to the area occupied by the
+neural arch in the type extends 0.5 mm. farther back in our specimen.
+The angle formed between the recurved dorsal ramus and the edge of the
+ventral flange is seen in our specimen to be less than 90 deg.. The glenoid
+fossa, appearing as a concave articular surface for the cap of the
+humerus, was in part covered by cartilage and shows as "unfinished"
+bone (Peabody, 1958, p. 572); this area is more oval than triangular,
+as Peabody thought. The obstruction of a clear view of this part of the
+type is the result of the accidental position of a neural arch. The
+raised portion immediately dorsal to the glenoid fossa exhibits an
+unfinished surface, suggesting the presence of either cartilage or a
+ligament.
+
+[Illustration: FIG. 8. _Hesperoherpeton garnettense_ Peabody. Type
+specimen redrawn. Right scapulocoracoid in external view (at left), and
+internal view (at right). KU 9976, x 4.]
+
+
+[Illustration: FIG. 9. _Hesperoherpeton garnettense_ Peabody. Right
+scapulocoracoid in external view, showing part of interclavicle, and
+position occupied by clavicle. The specimen is flattened and lies
+entirely in one plane. KU 10295, x 4.]
+
+
+[Illustration: FIG. 10. _Hesperoherpeton garnettense_ Peabody. Right
+clavicle in external view. Anterior edge to right. KU 10295, x 4.]
+
+The right clavicle is complete, and resembles a spoon having a slender
+handle. The dorsal tip of the handle is L-shaped. The expanded ventral
+part is convex externally, and rested upon the anteroventral surface of
+the scapulocoracoid. The lateral edge next to the "stem" is distinctly
+concave, abruptly becoming similar in contour to the opposite edge, and
+giving the impression of an unsymmetrical spoon. The left clavicle is
+present in scattered fragments, its dorsal hooklike end being intact.
+
+The posterior end of the interclavicle lies in contact with the right
+scapulocoracoid. There are short lateral processes at the point where
+the interclavicle was overlapped by the clavicles, but we cannot be
+sure of the extent of this bone anteriorly or posteriorly.
+
+The presumed left cleithrum, a long rectangle, is approximately equal
+in length to the rodlike stem of the clavicle, and is about as wide as
+the dorsal L-shaped tip of the clavicle. The posterior end of the
+cleithrum presumably met the tip of the clavicle, while the rest of it
+was directed anteriorly and a little dorsally. There seems to be a
+small articular surface near the anterior extremity which suggests the
+presence of a supracleithrum. The upper border of the cleithrum is
+slightly convex and the lower concave.
+
+
+FORELIMB (Fig. 11)
+
+The left forelimb is the only one present and appears to be nearly
+complete, although the elements are scattered almost at random. The
+only parts of the forelimb known to be missing are two subterminal and
+two terminal phalanges, probably of the first and third digits, and the
+proximal end of the second metacarpal. The smooth and relatively flat
+surfaces suggest an aquatic rather than terrestrial limb; only the
+proximal half of the humerus bears any conspicuous ridges or
+depressions. As we restore the skeleton of the limb, several features
+are remarkable: The humerus, ulna, and ulnare align themselves as the
+major axis of the limb, each carrying on its posterior edge a process
+or flange comparable to those in the axial series of a rhipidistian
+fin. The remaining elements take positions comparable to the diagonally
+placed preaxial radials in such a fin. The digits appear to have been
+short, perhaps with no more than two phalanges. There is only one row
+of carpals present (the proximal row of other tetrapods). A second and
+third row would be expected in primitive Amphibia; if they existed in
+_Hesperoherpeton_ they must either have been wholly cartilaginous or
+washed away from the specimen. Neither of these alternatives seems at
+all likely to us in view of the well-ossified condition of the elements
+that are present, and the occurrence of both the proximal carpals and
+the metacarpals. The space available for metacarpals probably could not
+have contained more than the four that are recognized.
+
+[Illustration: FIG. 11. _Hesperoherpeton garnettense_ Peabody. Left
+forelimb, showing characters of both a crossopterygian fin and an
+amphibian foot. KU 10295, x 4.]
+
+The proximal end of the humerus is more rounded anteriorly than
+posteriorly, and has a thin articular border that bore a cartilaginous
+cap as the primary surface for articulation with the scapulocoracoid.
+Although the unfinished surface of the head extends down the anterior
+margin about a third the length of the humerus, the shaft has been
+broken and so twisted that the distal part is not in the same plane as
+the proximal. Immediately posterior to the cartilaginous cap is a
+round, deep notch bordered posteriorly by the dorsal process of the
+head.
+
+The shaft is longer and narrower than would be anticipated in a
+primitive amphibian limb (cf. Romer, 1947). The distal end bears two
+surfaces for articulation with the radius and ulna. The full extent of
+the former surface was not determined because the more anterior part of
+the expanded end is represented only by an impression. The surface
+nearest the ulna was partially rounded for articulation with that
+element, the remaining posterior edge being broadly concave. The most
+striking feature of the humerus is a slender hooklike process on the
+posterior edge near the distal end, probably homologous with (1) the
+posterior flange on the "humerus" in Rhipidistia, and (2) the
+entepicondyle of the humerus in _Archeria_ (Romer, 1957) and other
+tetrapods.
+
+The radius is about the same width proximally as distally. The
+curvature of the shaft is approximately alike on both sides. Distally
+the surface is rounded for articulation with the radiale and perhaps
+the intermedium.
+
+The proximal end of the ulna is similar to that of the radius but is
+slightly larger. Posteriorly, there is a short, broad expansion
+resembling the entepicondyle of the humerus, and even more nearly like
+the postaxial flanges in a crossopterygian fin.
+
+The ends of the radiale are expanded and rounded, the entire bone being
+approximately twice as long as wide. The three sides of the intermedium
+are similarly convex. The surface of this bone is unfinished, showing
+that it must have been embedded in cartilage. The ulnare is
+conspicuously similar to the ulna in bearing a posterior hooklike
+expansion, and is larger than the radiale.
+
+The four metacarpals are slightly expanded proximally and distally.
+Although measurements of length and width are tabulated below (Table
+2), we are not certain of the sequence of these bones in the row.
+
+The dimensions of the two proximal phalanges are alike. The shape of
+these elements is similar to that of the metacarpals. The two terminal
+phalanges are somewhat triangular in shape, the lateral edges being
+concave and the proximal convex.
+
+TABLE 2.--APPROXIMATE MEASUREMENTS OF THE FORELIMB (in mm.)
+
+--------------------------+-------------------------------------------
+                          |                 Dimensions
+                          +----------+--------------------------------
+ ELEMENT                  |          |              Width
+                          |  Length  +----------+----------+----------
+                          |          | Proximal |  Midway  |  Distal
+--------------------------+----------+----------+----------+----------
+Humerus                   |   16.0   |    5.0   |    2.0   |    7.5?
+Radius                    |    9.0   |    4.0   |    1.5   |    3.5
+Ulna                      |    8.5   |    4.5   |    1.5   |    3.5
+Radiale                   |    3.0   |    2.0   |    1.5   |    2.0
+Intermedium               |    1.5   |     --   |    2.0   |     --
+Ulnare                    |    3.5   |    2.0   |    2.0   |    2.5
+Metacarpal        A       |    4.5   |    2.5   |    1.0   |    2.0
+Metacarpal        B       |    4.5   |    3.0?  |    1.5   |    2.5
+Metacarpal        C       |    4.0   |    2.0   |    1.5   |    2.0
+Metacarpal        D       |    3.5   |    2.5   |    1.0   |    1.5
+Proximal Phalanx  A       |    2.0   |    1.5   |    1.0   |    1.5
+Proximal Phalanx  B       |    2.0   |    1.5   |    1.0   |    1.5
+Terminal Phalanx  A       |    1.5   |    1.5   |    1.0   |    1.0
+Terminal Phalanx  B       |    1.5   |    1.5   |    1.0   |    1.0
+--------------------------+----------+----------+----------+----------
+
+
+COMPARISONS AND DISCUSSION
+
+Apparently primitive rhipidistian characters in _Hesperoherpeton_ are:
+Braincase in two sections, posterior one containing an expanded
+notochordal canal; lateral series of mandibular bones closely
+resembling that of _Megalichthys_, as figured by Watson (1926); tabular
+having long process probably articulating with pectoral girdle; lack of
+movement between head and trunk correlated with absence of occipital
+condyle; sensory pits present on frontal and squamosal.
+
+Although we are unable to separate, by sutures, the vomers from the
+palatines, the palatal surface of these bones and of the pterygoids is
+studded by numerous small teeth, as in Rhipidistia (Jarvik, 1954) and
+some of the early Amphibia (Romer, 1947). The stapes apparently reaches
+the quadrate, and could therefore serve in hyostylic suspension of the
+upper jaw.
+
+The pectoral limb has an axial series of bones carrying hooklike
+flanges on their posterior edges. The other bones of the limb show
+little modification of form beyond the nearly flat, aquatic type seen
+in Rhipidistia. No distinct elbow or wrist joints are developed.
+
+Characters of _Hesperoherpeton_ common to most primitive Amphibia, in
+contrast with Crossopterygii, are: Nares separated from edge of jaw;
+stapes having external process that may have met a tympanic membrane,
+thus giving the bone a sound-transmitting function. Apparently none of
+the opercular series was present.
+
+There are two large palatal teeth, slightly labyrinthine in character,
+adjacent to each internal naris. The scapulocoracoid, as shown by
+Peabody (1958), is Anthracosaurian in structure, as are the
+long-stemmed clavicles. The limbs have digits rather than fin-lobes,
+although the digital number apparently is four and the number of bones
+in the manus is less than would be expected in a primitive amphibian.
+The vertebrae are similar to those of Ichthyostegids, as described by
+Jarvik (1952), except that the pleurocentra are much larger.
+
+In addition to this remarkable combination of crossopterygian and
+amphibian characters, _Hesperoherpeton_ is specialized in certain
+features of the skull. The orbits are much enlarged, probably in
+correlation with the diminutive size of the animal, and this has been
+accompanied by loss of several bones. The frontal and squamosal nearly
+meet each other, and both form part of the rim of the orbit. The bones
+of the posterior part of the dermal roof are greatly reduced, and there
+is none behind the squamosal except the projecting tabular; there is no
+indication of quadratojugal, jugal, intertemporal or postparietal. The
+foramen magnum is enormous. The external surfaces of the bones of the
+skull are nearly smooth.
+
+Is it possible that the "primitive" and "specialized" features of this
+animal are actually larval? Are they not just the kind of characters
+that would be expected in an immature, aquatic embolomere of
+Pennsylvanian time? For several reasons we do not think this is the
+case. Except for the anterior part of the braincase, there is no
+indication that the skeleton was not well ossified. The postaxial
+processes on the humerus, ulna and ulnare could scarcely have been
+larval features only, since they are so clearly homologous with those
+in adult Rhipidistia; a larval limb should indeed be simple, but its
+simplicity is unlikely to involve paleotelic adult characters. The
+scapulocoracoid of our specimen is of practically the same shape and
+size as that in the only other known individual, the type; this would
+be probable if both were adults, but somewhat less likely if they were
+larvae of a much larger animal. The form of the stapes, tabular and
+otic notch suggest a functional tympanic membrane, which could not have
+occurred in a gill-breathing larva. On the other hand, an adult animal
+of pigmy size might be expected to have large orbits, large otic
+capsules and a large foramen magnum.
+
+We conclude that _Hesperoherpeton_ lived and sought food in the weedy
+shallows at the margin of a pond or lagoon, and that for much of the
+time its head was partly out of water (Fig. 12). The animal could
+either steady itself or crawl around by means of the paddlelike limbs,
+but these probably could not be used in effective locomotion on land.
+Like the Ichthyostegids, it probably swam by means of a fishlike tail.
+
+[Illustration: FIG. 12. _Hesperoherpeton garnettense_ Peabody. Probable
+appearance in life. x 0.5.]
+
+
+TAXONOMY
+
+Evidently _Hesperoherpeton_ is a small, lagoon-dwelling survivor of
+the Devonian forms that initiated the change from Crossopterygii to
+Amphibia (Jarvik, 1955). It shows, however, that this transition did
+not affect all structures at the same time, for some, as the braincase
+with its notochordal canal, the mandibular bones and axial limb bones,
+are unchanged from the condition normal for the Rhipidistia, but most
+other characters are of amphibian grade. To express these facts
+taxonomically requires that _Hesperoherpeton_ be removed from the
+family Cricotidae, suborder Embolomeri, order Anthracosauria, and
+placed in a new order and family of labyrinthodont Amphibia.
+
+     Order PLESIOPODA
+
+     (_plesios_, Gr., near, almost; _podos_, Gr., foot)
+
+     Labyrinthodontia having limbs provided with digits, but
+     retaining posterior flanges on axial bones as in
+     Rhipidistia, without joint-structure at elbow and wrist
+     essential for terrestrial locomotion; neurocranium having
+     separate otico-occipital section, large notochordal canal,
+     no occipital condyle, as in Rhipidistia; nares separate from
+     rim of mouth; pectoral girdle anthracosaurian; vertebrae
+     having U-shaped intercentrum and paired, but large,
+     pleurocentra.
+
+Probably associated with the characters of the order, as given above,
+are the connection of pectoral girdle with skull, and the presence of a
+tympanic membrane, the stapes functioning in both sound-transmission
+and palatoquadrate suspension.
+
+     Family HESPEROHERPETONIDAE
+
+     Orbits and foramen magnum unusually large in correlation
+     with reduced size of animal; squamosal forming posterior
+     margin of orbit; circumorbital series absent (except for
+     postorbital); sensory pits on squamosal and frontal.
+
+Characters defining the family are evidently the more specialized
+cranial features, which probably evolved during Mississippian and early
+Pennsylvanian times.
+
+The definition of the genus and species may be left to rest upon
+Peabody's (1958) original description and the present account, until
+the discovery of other members of the family gives reason for making
+further distinctions.
+
+
+SUMMARY
+
+_Hesperoherpeton garnettense_ Peabody (1958), based on a
+scapulocoracoid and part of a vertebra, was originally placed in the
+order Anthracosauria, suborder Embolomeri, family Cricotidae. A new
+skeleton from the type locality near Garnett, Kansas (Rock Lake shale,
+Stanton formation, Upper Pennsylvanian), shows that the animal has the
+following rhipidistian characters: Large notochordal canal below
+foramen magnum, otico-occipital block separate from ethmosphenoid,
+postaxial processes on three axial bones of forelimb, pectoral girdle
+(probably) articulated with tabular. Nevertheless, _Hesperoherpeton_
+has short digits, an anthracosaurian type of pectoral girdle, an otic
+rather than spiracular notch, nostrils separate from the mouth, and
+vertebrae in which the intercentrum is U-shaped and the pleurocentra
+large but paired. The stapes reaches the quadrate.
+
+_Hesperoherpeton_ is placed in a new order, PLESIOPODA, on the basis of
+the characters stated above, and a new family, HESPEROHERPETONIDAE.
+Specialized characters of the family include: Reduction of
+circumorbital bones, bringing the squamosal to the edge of the orbit,
+loss of certain bones of the temporal region, and relative enlargement
+of the orbits and foramen magnum, in correlation with the diminutive
+size of the animal. The structural characters of _Hesperoherpeton_
+suggest to us that it lived in the shallow, weedy margins of lagoons,
+rested with its head partly out of water, and normally did not walk on
+land.
+
+
+LITERATURE CITED
+
+EATON, T. H., JR.
+    1951. Origin of tetrapod limbs. Amer. Midl. Nat., 46: 245-251.
+
+JARVIK, E.
+    1952. On the fish-like tail in the ichthyostegid stegocephalians.
+          Meddel. om Gronland, 114: 1-90.
+    1954. On the visceral skeleton in _Eusthenopteron_ with a discussion
+          of the parasphenoid and palatoquadrate in fishes. Kgl. Svenska
+          Vetenskapsakad. Handl., 5: 1-104.
+    1955. The oldest tetrapods and their forerunners. Sci. Monthly,
+          80: 141-154.
+
+MOORE, R. C., FRYE, J. C., and JEWETT, J. M.
+    1944. Tabular description of outcropping rocks in Kansas. Kansas
+          State Geol. Surv. Bull., 52: 137-212.
+
+PEABODY, F. E.
+    1952. _Petrolacosaurus kansensis_ Lane, a Pennsylvanian reptile
+          from Kansas. Univ. Kansas Paleont. Contrib., Vertebrata,
+          Art. 1: 1-41.
+    1958. An embolomerous amphibian in the Garnett fauna (Pennsylvanian)
+          of Kansas. Jour. Paleont., 32: 571-573.
+
+ROMER, A. S.
+    1937. The braincase of the Carboniferous crossopterygian
+          _Megalichthys nitidus_. Mus. Comp. Zool. Bull., 82: 1-73.
+    1947. Review of the Labyrinthodontia. Mus. Comp. Zool. Bull.,
+          99: 1-368.
+    1957. The appendicular skeleton of the Permian embolomerous
+          amphibian _Archeria_. Univ. Michigan Contrib. Mus. Paleont.,
+          13: 103-159.
+
+WATSON, D. M. S.
+    1926. The evolution and origin of the Amphibia. Phil. Trans. Roy.
+          Soc. London, (B) 214: 189-257.
+
+_Transmitted January 13, 1960._
+
+
+28-2495
+
+
+
+
+
+End of the Project Gutenberg EBook of A New Order of Fishlike Amphibia From
+the Pennsylvanian of Kansas, by Theodore H. Eaton and Peggy Lou Stewart
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