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


[Pg 219]

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[Pg 220] 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.[Pg 221] 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.

Fig. 1. Hesperoherpeton garnettense Peabody. Skull, dorsal
view. Postorbital processes of the neurocranium are shown in
dotted outline. KU 10295, × 4. 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.[Pg 222]

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.

Fig. 2. Hesperoherpeton garnettense Peabody. Skull, lateral view,
showing relatively large orbit and absence of smaller circumorbital
bones. KU 10295, × 4. 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.[Pg 223]

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.

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. 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[Pg 224] 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.

Fig. 4. Hesperoherpeton garnettense Peabody. Palate reconstructed;
ventral aspect at left, showing teeth, dorsal aspect at
right. KU 10295, × 4. Fig. 4. Hesperoherpeton garnettense Peabody. Palate reconstructed; ventral aspect at left, showing teeth, dorsal aspect at right. KU 10295, × 4.

[Pg 225]

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.[Pg 226]

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[Pg 227] 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.

Fig. 5. Hesperoherpeton garnettense Peabody, KU
10295, × 4. A, occipital view of skull; B, basioccipital
bone in dorsal (internal) view. 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[Pg 228] 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.[Pg 229]

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.

Fig. 6. Hesperoherpeton garnettense Peabody. Right mandible, lateral
view, KU 10295, × 4. External surfaces are pitted; broken surfaces
are coarsely stippled. 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.[Pg 230] 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.

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. 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

PartsAnt.-post.Dors.-vent.Transv. width
Neural spine1.5 (3)3.0 (3)
Neural spine and arch2.0 (4)4.5? (4)
Neural canal2.0 (4)2.0 (1)1.0 (1)
Intercentrum1.5 (5)3.5 (4)3.0 (1)
Pleurocentrum1.5 (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[Pg 231] 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[Pg 232] immediately dorsal to the glenoid fossa exhibits an unfinished surface, suggesting the presence of either cartilage or a ligament.

Fig. 8. Hesperoherpeton garnettense Peabody. Type specimen redrawn.
Right scapulocoracoid in external view (at left), and internal view (at right).
KU 9976, × 4. Fig. 8. Hesperoherpeton garnettense Peabody. Type specimen redrawn. Right scapulocoracoid in external view (at left), and internal view (at right). KU 9976, × 4.

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. 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.

Fig. 10. Hesperoherpeton garnettense Peabody. Right clavicle in external
view. Anterior edge to right. KU 10295, × 4. Fig. 10. Hesperoherpeton garnettense Peabody. Right clavicle in external view. Anterior edge to right. KU 10295, × 4.

[Pg 233]

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[Pg 234] 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.

Fig. 11. Hesperoherpeton garnettense Peabody. Left forelimb,
showing characters of both a crossopterygian fin and an amphibian
foot. KU 10295, × 4. 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[Pg 235] 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.[Pg 236]

Table 2.—Approximate Measurements of the Forelimb (in mm.)

ElementDimensions
LengthWidth
ProximalMidwayDistal
Humerus16.05.02.07.5?
Radius9.04.01.53.5
Ulna8.54.51.53.5
Radiale3.02.01.52.0
Intermedium1.52.0
Ulnare3.52.02.02.5
Metacarpal           A4.52.51.02.0
Metacarpal           B4.53.0?1.52.5
Metacarpal           C4.02.01.52.0
Metacarpal           D3.52.51.01.5
Proximal Phalanx  A2.01.51.01.5
Proximal Phalanx  B2.01.51.01.5
Terminal Phalanx  A1.51.51.01.0
Terminal Phalanx  B1.51.51.01.0

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.[Pg 237]

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[Pg 238] 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.

Fig. 12. Hesperoherpeton garnettense Peabody. Probable appearance
in life. × 0.5. 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.[Pg 239]

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.[Pg 240]

LITERATURE CITED

Eaton, T. H., Jr.
1951. Origin of tetrapod limbs. Amer. Midl. Nat., 46: 245-251.

Jarvik, E.
1952. On the fish-like tail in the ichthyostegid stegocephalians. Meddel. om Grønland, 114: 1-90.
1954. On the visceral skeleton in Eusthenopteron with a discussion of the parasphenoid and palatoquadrate in fishes. Kgl. Svenska Vetenskapsakad. Handl., 5: 1-104.
1955. The oldest tetrapods and their forerunners. Sci. Monthly, 80: 141-154.

Moore, R. C., Frye, J. C., and Jewett, J. M.
1944. Tabular description of outcropping rocks in Kansas. Kansas State Geol. Surv. Bull., 52: 137-212.

Peabody, F. E.
1952. Petrolacosaurus kansensis Lane, a Pennsylvanian reptile from Kansas. Univ. Kansas Paleont. Contrib., Vertebrata, Art. 1: 1-41.
1958. An embolomerous amphibian in the Garnett fauna (Pennsylvanian) of Kansas. Jour. Paleont., 32: 571-573.

Romer, A. S.
1937. The braincase of the Carboniferous crossopterygian Megalichthys nitidus. Mus. Comp. Zool. Bull., 82: 1-73.
1947. Review of the Labyrinthodontia. Mus. Comp. Zool. Bull., 99: 1-368.
1957. The appendicular skeleton of the Permian embolomerous amphibian Archeria. Univ. Michigan Contrib. Mus. Paleont., 13: 103-159.

Watson, D. M. S.
1926. The evolution and origin of the Amphibia. Phil. Trans. Roy. Soc. London, (B) 214: 189-257.

Transmitted January 13, 1960.

28-2495






End of the Project Gutenberg EBook of A New Order of Fishlike Amphibia From
the Pennsylvanian of Kansas, by Theodore H. Eaton and Peggy Lou Stewart

*** END OF THIS PROJECT GUTENBERG EBOOK NEW ORDER OF FISHLIKE AMPHIBIA ***

***** This file should be named 31050-h.htm or 31050-h.zip *****
This and all associated files of various formats will be found in:
        https://www.gutenberg.org/3/1/0/5/31050/

Produced by Chris Curnow, Joseph Cooper, Diane Monico, and
the Online Distributed Proofreading Team at
https://www.pgdp.net


Updated editions will replace the previous one--the old editions
will be renamed.

Creating the works from public domain print editions means that no
one owns a United States copyright in these works, so the Foundation
(and you!) can copy and distribute it in the United States without
permission and without paying copyright royalties.  Special rules,
set forth in the General Terms of Use part of this license, apply to
copying and distributing Project Gutenberg-tm electronic works to
protect the PROJECT GUTENBERG-tm concept and trademark.  Project
Gutenberg is a registered trademark, and may not be used if you
charge for the eBooks, unless you receive specific permission.  If you
do not charge anything for copies of this eBook, complying with the
rules is very easy.  You may use this eBook for nearly any purpose
such as creation of derivative works, reports, performances and
research.  They may be modified and printed and given away--you may do
practically ANYTHING with public domain eBooks.  Redistribution is
subject to the trademark license, especially commercial
redistribution.



*** START: FULL LICENSE ***

THE FULL PROJECT GUTENBERG LICENSE
PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK

To protect the Project Gutenberg-tm mission of promoting the free
distribution of electronic works, by using or distributing this work
(or any other work associated in any way with the phrase "Project
Gutenberg"), you agree to comply with all the terms of the Full Project
Gutenberg-tm License (available with this file or online at
https://gutenberg.org/license).


Section 1.  General Terms of Use and Redistributing Project Gutenberg-tm
electronic works

1.A.  By reading or using any part of this Project Gutenberg-tm
electronic work, you indicate that you have read, understand, agree to
and accept all the terms of this license and intellectual property
(trademark/copyright) agreement.  If you do not agree to abide by all
the terms of this agreement, you must cease using and return or destroy
all copies of Project Gutenberg-tm electronic works in your possession.
If you paid a fee for obtaining a copy of or access to a Project
Gutenberg-tm electronic work and you do not agree to be bound by the
terms of this agreement, you may obtain a refund from the person or
entity to whom you paid the fee as set forth in paragraph 1.E.8.

1.B.  "Project Gutenberg" is a registered trademark.  It may only be
used on or associated in any way with an electronic work by people who
agree to be bound by the terms of this agreement.  There are a few
things that you can do with most Project Gutenberg-tm electronic works
even without complying with the full terms of this agreement.  See
paragraph 1.C below.  There are a lot of things you can do with Project
Gutenberg-tm electronic works if you follow the terms of this agreement
and help preserve free future access to Project Gutenberg-tm electronic
works.  See paragraph 1.E below.

1.C.  The Project Gutenberg Literary Archive Foundation ("the Foundation"
or PGLAF), owns a compilation copyright in the collection of Project
Gutenberg-tm electronic works.  Nearly all the individual works in the
collection are in the public domain in the United States.  If an
individual work is in the public domain in the United States and you are
located in the United States, we do not claim a right to prevent you from
copying, distributing, performing, displaying or creating derivative
works based on the work as long as all references to Project Gutenberg
are removed.  Of course, we hope that you will support the Project
Gutenberg-tm mission of promoting free access to electronic works by
freely sharing Project Gutenberg-tm works in compliance with the terms of
this agreement for keeping the Project Gutenberg-tm name associated with
the work.  You can easily comply with the terms of this agreement by
keeping this work in the same format with its attached full Project
Gutenberg-tm License when you share it without charge with others.

1.D.  The copyright laws of the place where you are located also govern
what you can do with this work.  Copyright laws in most countries are in
a constant state of change.  If you are outside the United States, check
the laws of your country in addition to the terms of this agreement
before downloading, copying, displaying, performing, distributing or
creating derivative works based on this work or any other Project
Gutenberg-tm work.  The Foundation makes no representations concerning
the copyright status of any work in any country outside the United
States.

1.E.  Unless you have removed all references to Project Gutenberg:

1.E.1.  The following sentence, with active links to, or other immediate
access to, the full Project Gutenberg-tm License must appear prominently
whenever any copy of a Project Gutenberg-tm work (any work on which the
phrase "Project Gutenberg" appears, or with which the phrase "Project
Gutenberg" is associated) is accessed, displayed, performed, viewed,
copied or distributed:

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

1.E.2.  If an individual Project Gutenberg-tm electronic work is derived
from the public domain (does not contain a notice indicating that it is
posted with permission of the copyright holder), the work can be copied
and distributed to anyone in the United States without paying any fees
or charges.  If you are redistributing or providing access to a work
with the phrase "Project Gutenberg" associated with or appearing on the
work, you must comply either with the requirements of paragraphs 1.E.1
through 1.E.7 or obtain permission for the use of the work and the
Project Gutenberg-tm trademark as set forth in paragraphs 1.E.8 or
1.E.9.

1.E.3.  If an individual Project Gutenberg-tm electronic work is posted
with the permission of the copyright holder, your use and distribution
must comply with both paragraphs 1.E.1 through 1.E.7 and any additional
terms imposed by the copyright holder.  Additional terms will be linked
to the Project Gutenberg-tm License for all works posted with the
permission of the copyright holder found at the beginning of this work.

1.E.4.  Do not unlink or detach or remove the full Project Gutenberg-tm
License terms from this work, or any files containing a part of this
work or any other work associated with Project Gutenberg-tm.

1.E.5.  Do not copy, display, perform, distribute or redistribute this
electronic work, or any part of this electronic work, without
prominently displaying the sentence set forth in paragraph 1.E.1 with
active links or immediate access to the full terms of the Project
Gutenberg-tm License.

1.E.6.  You may convert to and distribute this work in any binary,
compressed, marked up, nonproprietary or proprietary form, including any
word processing or hypertext form.  However, if you provide access to or
distribute copies of a Project Gutenberg-tm work in a format other than
"Plain Vanilla ASCII" or other format used in the official version
posted on the official Project Gutenberg-tm web site (www.gutenberg.org),
you must, at no additional cost, fee or expense to the user, provide a
copy, a means of exporting a copy, or a means of obtaining a copy upon
request, of the work in its original "Plain Vanilla ASCII" or other
form.  Any alternate format must include the full Project Gutenberg-tm
License as specified in paragraph 1.E.1.

1.E.7.  Do not charge a fee for access to, viewing, displaying,
performing, copying or distributing any Project Gutenberg-tm works
unless you comply with paragraph 1.E.8 or 1.E.9.

1.E.8.  You may charge a reasonable fee for copies of or providing
access to or distributing Project Gutenberg-tm electronic works provided
that

- You pay a royalty fee of 20% of the gross profits you derive from
     the use of Project Gutenberg-tm works calculated using the method
     you already use to calculate your applicable taxes.  The fee is
     owed to the owner of the Project Gutenberg-tm trademark, but he
     has agreed to donate royalties under this paragraph to the
     Project Gutenberg Literary Archive Foundation.  Royalty payments
     must be paid within 60 days following each date on which you
     prepare (or are legally required to prepare) your periodic tax
     returns.  Royalty payments should be clearly marked as such and
     sent to the Project Gutenberg Literary Archive Foundation at the
     address specified in Section 4, "Information about donations to
     the Project Gutenberg Literary Archive Foundation."

- You provide a full refund of any money paid by a user who notifies
     you in writing (or by e-mail) within 30 days of receipt that s/he
     does not agree to the terms of the full Project Gutenberg-tm
     License.  You must require such a user to return or
     destroy all copies of the works possessed in a physical medium
     and discontinue all use of and all access to other copies of
     Project Gutenberg-tm works.

- You provide, in accordance with paragraph 1.F.3, a full refund of any
     money paid for a work or a replacement copy, if a defect in the
     electronic work is discovered and reported to you within 90 days
     of receipt of the work.

- You comply with all other terms of this agreement for free
     distribution of Project Gutenberg-tm works.

1.E.9.  If you wish to charge a fee or distribute a Project Gutenberg-tm
electronic work or group of works on different terms than are set
forth in this agreement, you must obtain permission in writing from
both the Project Gutenberg Literary Archive Foundation and Michael
Hart, the owner of the Project Gutenberg-tm trademark.  Contact the
Foundation as set forth in Section 3 below.

1.F.

1.F.1.  Project Gutenberg volunteers and employees expend considerable
effort to identify, do copyright research on, transcribe and proofread
public domain works in creating the Project Gutenberg-tm
collection.  Despite these efforts, Project Gutenberg-tm electronic
works, and the medium on which they may be stored, may contain
"Defects," such as, but not limited to, incomplete, inaccurate or
corrupt data, transcription errors, a copyright or other intellectual
property infringement, a defective or damaged disk or other medium, a
computer virus, or computer codes that damage or cannot be read by
your equipment.

1.F.2.  LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for the "Right
of Replacement or Refund" described in paragraph 1.F.3, the Project
Gutenberg Literary Archive Foundation, the owner of the Project
Gutenberg-tm trademark, and any other party distributing a Project
Gutenberg-tm electronic work under this agreement, disclaim all
liability to you for damages, costs and expenses, including legal
fees.  YOU AGREE THAT YOU HAVE NO REMEDIES FOR NEGLIGENCE, STRICT
LIABILITY, BREACH OF WARRANTY OR BREACH OF CONTRACT EXCEPT THOSE
PROVIDED IN PARAGRAPH F3.  YOU AGREE THAT THE FOUNDATION, THE
TRADEMARK OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL NOT BE
LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT, CONSEQUENTIAL, PUNITIVE OR
INCIDENTAL DAMAGES EVEN IF YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH
DAMAGE.

1.F.3.  LIMITED RIGHT OF REPLACEMENT OR REFUND - If you discover a
defect in this electronic work within 90 days of receiving it, you can
receive a refund of the money (if any) you paid for it by sending a
written explanation to the person you received the work from.  If you
received the work on a physical medium, you must return the medium with
your written explanation.  The person or entity that provided you with
the defective work may elect to provide a replacement copy in lieu of a
refund.  If you received the work electronically, the person or entity
providing it to you may choose to give you a second opportunity to
receive the work electronically in lieu of a refund.  If the second copy
is also defective, you may demand a refund in writing without further
opportunities to fix the problem.

1.F.4.  Except for the limited right of replacement or refund set forth
in paragraph 1.F.3, this work is provided to you 'AS-IS' WITH NO OTHER
WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED, INCLUDING BUT NOT LIMITED TO
WARRANTIES OF MERCHANTIBILITY OR FITNESS FOR ANY PURPOSE.

1.F.5.  Some states do not allow disclaimers of certain implied
warranties or the exclusion or limitation of certain types of damages.
If any disclaimer or limitation set forth in this agreement violates the
law of the state applicable to this agreement, the agreement shall be
interpreted to make the maximum disclaimer or limitation permitted by
the applicable state law.  The invalidity or unenforceability of any
provision of this agreement shall not void the remaining provisions.

1.F.6.  INDEMNITY - You agree to indemnify and hold the Foundation, the
trademark owner, any agent or employee of the Foundation, anyone
providing copies of Project Gutenberg-tm electronic works in accordance
with this agreement, and any volunteers associated with the production,
promotion and distribution of Project Gutenberg-tm electronic works,
harmless from all liability, costs and expenses, including legal fees,
that arise directly or indirectly from any of the following which you do
or cause to occur: (a) distribution of this or any Project Gutenberg-tm
work, (b) alteration, modification, or additions or deletions to any
Project Gutenberg-tm work, and (c) any Defect you cause.


Section  2.  Information about the Mission of Project Gutenberg-tm

Project Gutenberg-tm is synonymous with the free distribution of
electronic works in formats readable by the widest variety of computers
including obsolete, old, middle-aged and new computers.  It exists
because of the efforts of hundreds of volunteers and donations from
people in all walks of life.

Volunteers and financial support to provide volunteers with the
assistance they need are critical to reaching Project Gutenberg-tm's
goals and ensuring that the Project Gutenberg-tm collection will
remain freely available for generations to come.  In 2001, the Project
Gutenberg Literary Archive Foundation was created to provide a secure
and permanent future for Project Gutenberg-tm and future generations.
To learn more about the Project Gutenberg Literary Archive Foundation
and how your efforts and donations can help, see Sections 3 and 4
and the Foundation web page at https://www.pglaf.org.


Section 3.  Information about the Project Gutenberg Literary Archive
Foundation

The Project Gutenberg Literary Archive Foundation is a non profit
501(c)(3) educational corporation organized under the laws of the
state of Mississippi and granted tax exempt status by the Internal
Revenue Service.  The Foundation's EIN or federal tax identification
number is 64-6221541.  Its 501(c)(3) letter is posted at
https://pglaf.org/fundraising.  Contributions to the Project Gutenberg
Literary Archive Foundation are tax deductible to the full extent
permitted by U.S. federal laws and your state's laws.

The Foundation's principal office is located at 4557 Melan Dr. S.
Fairbanks, AK, 99712., but its volunteers and employees are scattered
throughout numerous locations.  Its business office is located at
809 North 1500 West, Salt Lake City, UT 84116, (801) 596-1887, email
business@pglaf.org.  Email contact links and up to date contact
information can be found at the Foundation's web site and official
page at https://pglaf.org

For additional contact information:
     Dr. Gregory B. Newby
     Chief Executive and Director
     gbnewby@pglaf.org


Section 4.  Information about Donations to the Project Gutenberg
Literary Archive Foundation

Project Gutenberg-tm depends upon and cannot survive without wide
spread public support and donations to carry out its mission of
increasing the number of public domain and licensed works that can be
freely distributed in machine readable form accessible by the widest
array of equipment including outdated equipment.  Many small donations
($1 to $5,000) are particularly important to maintaining tax exempt
status with the IRS.

The Foundation is committed to complying with the laws regulating
charities and charitable donations in all 50 states of the United
States.  Compliance requirements are not uniform and it takes a
considerable effort, much paperwork and many fees to meet and keep up
with these requirements.  We do not solicit donations in locations
where we have not received written confirmation of compliance.  To
SEND DONATIONS or determine the status of compliance for any
particular state visit https://pglaf.org

While we cannot and do not solicit contributions from states where we
have not met the solicitation requirements, we know of no prohibition
against accepting unsolicited donations from donors in such states who
approach us with offers to donate.

International donations are gratefully accepted, but we cannot make
any statements concerning tax treatment of donations received from
outside the United States.  U.S. laws alone swamp our small staff.

Please check the Project Gutenberg Web pages for current donation
methods and addresses.  Donations are accepted in a number of other
ways including including checks, online payments and credit card
donations.  To donate, please visit: https://pglaf.org/donate


Section 5.  General Information About Project Gutenberg-tm electronic
works.

Professor Michael S. Hart was the originator of the Project Gutenberg-tm
concept of a library of electronic works that could be freely shared
with anyone.  For thirty years, he produced and distributed Project
Gutenberg-tm eBooks with only a loose network of volunteer support.


Project Gutenberg-tm eBooks are often created from several printed
editions, all of which are confirmed as Public Domain in the U.S.
unless a copyright notice is included.  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.