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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/31050-8.txt b/31050-8.txt new file mode 100644 index 0000000..c52bc8b --- /dev/null +++ b/31050-8.txt @@ -0,0 +1,1263 @@ +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. 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Thus, we do not necessarily +keep eBooks in compliance with any particular paper edition. + + +Most people start at our Web site which has the main PG search facility: + + https://www.gutenberg.org + +This Web site includes information about Project Gutenberg-tm, +including how to make donations to the Project Gutenberg Literary +Archive Foundation, how to help produce our new eBooks, and how to +subscribe to our email newsletter to hear about new eBooks. diff --git a/31050-8.zip b/31050-8.zip Binary files differnew file mode 100644 index 0000000..c55870f --- /dev/null +++ b/31050-8.zip diff --git a/31050-h.zip b/31050-h.zip Binary files differnew file mode 100644 index 0000000..9d0886e --- /dev/null +++ b/31050-h.zip diff --git a/31050-h/31050-h.htm b/31050-h/31050-h.htm new file mode 100644 index 0000000..c5a128a --- /dev/null +++ b/31050-h/31050-h.htm @@ -0,0 +1,1473 @@ +<!DOCTYPE html PUBLIC "-//W3C//DTD XHTML 1.0 Strict//EN" + "http://www.w3.org/TR/xhtml1/DTD/xhtml1-strict.dtd"> + +<html xmlns="http://www.w3.org/1999/xhtml" xml:lang="en" lang="en"> + <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"> + + + p { margin-top: .75em; + text-align: justify; + margin-bottom: .75em; + } + + h1 { text-align: center; line-height: 1.5; clear: both; } + + p.title { text-align: center; text-indent: 0; + font-weight: bold; + line-height: 1.4; margin-bottom: 3em; } + + h2,h3 { text-align: center; clear: both; } + + hr { width: 33%; + margin-top: 2em; + margin-bottom: 2em; + margin-left: auto; + margin-right: auto; + clear: both; + } + + table {margin-left: auto; margin-right: auto;} + + body{margin-left: 10%; + margin-right: 10%; + } + + .pagenum { /* uncomment the next line for invisible page numbers +*/ + /* visibility: hidden; */ + position: absolute; + left: 92%; + font-size: smaller; + text-align: right; + } /* page numbers */ + + .blockquot{margin-left: 5%; margin-right: 10%;} + .i4 {display: block; margin-left: 2.5em; + padding-left: 2.5em; text-indent: -2.5em;} + + .center {text-align: center;} + .smcap {font-variant: small-caps;} + .u {text-decoration: underline;} + + .caption {font-weight: bold;} + + .figcenter {margin: auto; text-align: center;} + + + </style> + </head> +<body> + + +<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 + + + + + + +</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 × 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 10295, × 4." title="Fig. 1. Hesperoherpeton garnettense Peabody. Skull, dorsal +view. Postorbital processes of the neurocranium are shown in +dotted outline. KU 10295, × 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 10295, × 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 10295, × 4." title="Fig. 2. Hesperoherpeton garnettense Peabody. Skull, lateral view, +showing relatively large orbit and absence of smaller circumorbital +bones. KU 10295, × 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 10295, × 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 10295, all × 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 10295, all × 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 10295, all × 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 10295, × 4." title="Fig. 4. Hesperoherpeton garnettense Peabody. Palate reconstructed; +ventral aspect at left, showing teeth, dorsal aspect at +right. KU 10295, × 4." /> +<span class="caption">Fig. 4. <i>Hesperoherpeton garnettense</i> Peabody. Palate reconstructed; +ventral aspect at left, showing teeth, dorsal aspect at +right. KU 10295, × 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, × 4. A, occipital view of skull; B, basioccipital +bone in dorsal (internal) view." title="Fig. 5. Hesperoherpeton garnettense Peabody, KU +10295, × 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, × 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 10295, × 4. External surfaces are pitted; broken surfaces +are coarsely stippled." title="Fig. 6. Hesperoherpeton garnettense Peabody. Right mandible, lateral +view, KU 10295, × 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 10295, × 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 10295, × 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 10295, × 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 10295, × 4.</span><br /><br /> +</p> + +<p class="center"><b> +<span class="smcap">Table 1.—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'>—</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'>—</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'>—</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 9976) and the new skeleton (KU 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 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<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 9976, × 4." title="Fig. 8. Hesperoherpeton garnettense Peabody. Type specimen redrawn. +Right scapulocoracoid in external view (at left), and internal view (at right). +KU 9976, × 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 9976, × 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 10295, × 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 10295, × 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 10295, × 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 10295, × 4." title="Fig. 10. Hesperoherpeton garnettense Peabody. Right clavicle in external +view. Anterior edge to right. KU 10295, × 4." /> +<span class="caption">Fig. 10. <i>Hesperoherpeton garnettense</i> Peabody. Right clavicle in external +view. Anterior edge to right. KU 10295, × 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 10295, × 4." title="Fig. 11. Hesperoherpeton garnettense Peabody. Left forelimb, +showing characters of both a crossopterygian fin and an amphibian +foot. KU 10295, × 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 10295, × 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.—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'>—</td><td align='center'>2.0</td><td align='center'>—</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"> 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"> 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"> 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"> 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"> 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"> 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"> 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"> 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. × 0.5." title="Fig. 12. Hesperoherpeton garnettense Peabody. Probable appearance +in life. × 0.5." /> +<span class="caption">Fig. 12. <i>Hesperoherpeton garnettense</i> Peabody. Probable appearance +in life. × 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 Grønland, 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. 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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. 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