diff options
Diffstat (limited to '31050-8.txt')
| -rw-r--r-- | 31050-8.txt | 1263 |
1 files changed, 1263 insertions, 0 deletions
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. Eaton and Peggy Lou Stewart + +*** END OF THIS PROJECT GUTENBERG EBOOK NEW ORDER OF FISHLIKE AMPHIBIA *** + +***** This file should be named 31050-8.txt or 31050-8.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. |