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diff --git a/33560.txt b/33560.txt new file mode 100644 index 0000000..371f3a4 --- /dev/null +++ b/33560.txt @@ -0,0 +1,1415 @@ +The Project Gutenberg EBook of A New Genus of Pennsylvania Fish +(Crossoperygii, Coelacanthiformes) from Kansas, by Joan Echols + +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 Genus of Pennsylvania Fish (Crossoperygii, Coelacanthiformes) from Kansas + +Author: Joan Echols + +Release Date: August 28, 2010 [EBook #33560] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK NEW GENUS OF PENNSYLVANIA FISH *** + + + + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +and the Online Distributed Proofreading Team at +https://www.pgdp.net. + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS +MUSEUM OF NATURAL HISTORY + +Volume 12, No. 10, pp. 475-501, 7 figs. +October 25, 1963 + +A New Genus of Pennsylvanian Fish (Crossopterygii, Coelacanthiformes) +from Kansas + +BY + +JOAN ECHOLS + +UNIVERSITY OF KANSAS +LAWRENCE +1963 + + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Theodore H. Eaton, Jr. + + +~Volume 12, No. 10, pp. 475-501, 7 figs.~ +~Published October 25, 1963~ + +UNIVERSITY OF KANSAS +Lawrence, Kansas + +PRINTED BY +JEAN M. NEIBARGER, STATE PRINTER +TOPEKA, KANSAS +1963 + +[Transcriber's Note: Words surrounded by tildes, like ~this~ signifies +words in bold. Words surrounded by underscores, like _this_, signifies +words in italics.] + + + + +A New Genus of Pennsylvanian Fish (Crossopterygii, Coelacanthiformes) +from Kansas + +BY + +JOAN ECHOLS + + +INTRODUCTION + +In 1931 and 1932, H. H. Lane, C. W. Hibbard and W. K. McNown collected +the specimens that Hibbard (1933) described and made the basis of two +new species. These were from the Rock Lake shale member of the Stanton +formation, six miles northwest of Garnett, Anderson County, Kansas. In +1954, from a locality (KAn-1/D, see page 480) approximately one fourth +mile southwest of the first locality, specimens were quarried by F. E. +Peabody, R. W. Wilson and R. Weeks. In 1955 R. R. Camp collected +additional blocks of Rock Lake shale from this second locality. Study of +all of the materials from the above mentioned localities reveals the +existence of an hitherto unrecognized genus of coelacanth. It is named +and described below. + +I wish to thank Prof. Theodore H. Eaton, Jr., for suggesting the project +and for much helpful advice. I am indebted to Dr. E. I. White of the +British Museum (Natural History) for furnishing a cast of the +endocranium of _Rhabdoderma elegans_ (Newberry) for comparison, and to +Drs. Donald Baird (Princeton University), Bobb Schaeffer (American +Museum of Natural History) and R. H. Denison (Chicago Natural History +Museum) for loans and exchanges of specimens for comparison. I am +grateful to Dr. Bobb Schaeffer for advice on the manuscript. Mr. Merton +C. Bowman assisted with the illustrations. The study here reported on +was made while I was a Research Assistant supported by National Science +Foundation Grant G-14013. + + +SYSTEMATIC DESCRIPTIONS + +Subclass CROSSOPTERYGII + +Superorder COELACANTHI + +Order Coelacanthiformes + +Suborder DIPLOCERCIDOIDEI + +Family DIPLOCERCIDAE + +Subfamily ~Rhabdodermatinae~, new subfamily + + _Type genus._--_Rhabdoderma Reis_, 1888, Paleontographica, + vol. 35, p. 71. + + _Referred genus._--_Synaptotylus_ new, described below. + + _Horizon._--Carboniferous. + + _Diagnosis._--Sphenethmoid region partly ossified, and + consisting of basisphenoid, parasphenoid, and ethmoid + ossifications; paired basipterygoid process and paired + antotic process on basisphenoid; parasphenoid of normal + size, and closely associated with, or fused to, + basisphenoid; ethmoids paired in _Rhabdoderma_ (unknown in + _Synaptotylus_). + +_Discussion._--Because of the great differences in endocranial structure +between the Devonian and Pennsylvanian coelacanths, they are here placed +in new subfamilies. The two proposed subfamilies of the family +Diplocercidae are the Diplocercinae and the Rhabdodermatinae. The +Diplocercinae include those coelacanths having two large unpaired bones +in the endocranium (at present this includes _Diplocercides_ Stensioe, +_Nesides_ Stensioe and _Euporosteus_ Jaekel). The subfamily +Rhabdodermatinae is composed of coelacanths having reduced endocranial +ossification, as described in detail above, and now including +_Rhabdoderma_ Reis and _Synaptotylus_ n. g. + +Members of this subfamily differ from those of the subfamily +Diplocercinae in having several paired and unpaired elements in the +sphenethmoid region of the endocranium, instead of only one larger +ossification. They differ from those of the suborder Coelacanthoidei in +the retention of basipterygoid processes. + +_Synaptotylus_ is more closely related to _Rhabdoderma_ than to the +Diplocercines because the anterior portion of the endocranium contains +only a basisphenoid, parasphenoid, and probably ethmoids. The +sphenethmoid region was certainly not a large, unpaired unit as in the +Diplocercines. Probably the posterior part, the otico-occipital region +(not known in _Synaptotylus_), was much more nearly like that of +_Rhabdoderma_, which consisted of unpaired basioccipital and +supraoccipital, and paired prootics, exoccipitals, and anterior and +posterior occipital ossifications (Moy-Thomas, 1937: figs. 3, 4). +Moy-Thomas (1937:389) points out that in _Rhabdoderma_ the occipital +region is "considerably more ossified" than in any coelacanths other +than the Devonian forms. Berg (1940:390) thought that the Carboniferous +coelacanths should be placed in a separate family because they did not +have two large, unpaired bones in the endocranium. _Rhabdoderma_ and +_Synaptotylus_ represent another stage in evolution of the endocranium +in coelacanths, and, if classification is to be based on endocranial +structure, then this stage (represented by the two genera) may later be +given family rank as Berg suggested. Because _Rhabdoderma_ and +_Synaptotylus_ have only part of the sphenethmoid region ossified and +because they retain basipterygoid processes, they are considered to be +related and are included in the subfamily Rhabdodermatinae. + + +~Synaptotylus~, new genus + + _Type species._--_Synaptotylus newelli_ (Hibbard). + + _Horizon._--Rock Lake shale member, Stanton formation, + Lansing group, Missouri series, Upper Pennsylvanian. + +_Diagnosis._--Late Pennsylvanian fishes of small size, having the +following combination of characters: on basisphenoid, knoblike antotic +processes connected by a low ridge to basipterygoid processes; entire +ventral surface of parasphenoid toothed; anterior margin of parasphenoid +notched and no evidence of hypophyseal opening. Dermal bones of skull +smooth or with low, rounded tubercles and striae; fronto-ethmoid shield +incompletely known but having one pair of large rectangular frontals +with posterolaterally slanting anterior margins; intertemporals large, +the lateral margins curving laterally; postorbital triangular, apex +downward; subopercular somewhat triangular; squamosal carrying sensory +canal that curves down posteriorly and extends onto a ventral +projection; opercular generally triangular; supratemporals elongate, +curving to fit lateral margin of intertemporals; circumorbital plates +lightly ossified. Palatoquadrate complex consisting of endopterygoid and +ectopterygoid (both toothed on medial surface), quadrate, and +metapterygoid, the latter smooth and having widened border for +articulation on anterodorsal margin. Pectoral girdle consisting of +cleithrum and clavicle (supracleithrum not seen); small projection on +medial surface of posterior portion of cleithrum; horizontal medial +process on clavicle. Pelvic plate bearing three anteriorly diverging +apophyses, and one denticulate ventromedian process for articulation to +opposite plate. Lepidotrichia jointed distally, but not tuberculated. +Scales oval, having posteriorly converging ridges on posterior exposed +parts. + +The name refers to the most distinctive character of the genus, the +connected antotic and basipterygoid processes on the basisphenoid, and +is derived from Greek, _synaptos_--joined, _tylos_ (masc.)--knob, +projection. + +_Synaptotylus_ is excluded from the advanced suborder Coelacanthoidei by +the retention of basipterygoid processes on the basisphenoid. +_Synaptotylus_ differs from _Rhabdoderma_ in several characters of the +basisphenoid, the most important being: knoblike antotic processes +(those of _Rhabdoderma_ are wider, more flattened and more dorsal in +position); small, lateral basipterygoid processes (in _Rhabdoderma_ +these are larger and farther ventral in position). + + +~Synaptotylus newelli~ (Hibbard) + + _Coelacanthus newelli_ Hibbard, 1933, Univ. Kansas Sci. + Bull., 21:280, pl. 27, figs. 2, 3. + + _Coelacanthus arcuatus_ Hibbard, 1933, Univ. Kansas Sci. + Bull., 21:282, pl. 26, fig. 8; pl. 27, fig. 1. + + _Rhabdoderma elegans_ Moy-Thomas, 1937 (in part), Proc. + Zool. Soc. London, 107(ser. B, pt. 3):399. + + _Type._--K. U. no. 786F. + + _Diagnosis._--Same as for the genus. + + _Horizon._--Rock Lake shale member, Stanton formation, + Lansing group, Missouri series, Upper Pennsylvanian. + + _Localities._--The specimens studied by Hibbard (K. U. nos. + 786F, 787F, 788) and no. 11457 were taken from the Bradford + Chandler farm, from the original quarry in SW-1/4, SE-1/4, + sec. 32, T.19S, R.19E. The remainder were collected from + University of Kansas Museum of Natural History locality + KAn-1/D, a quarry in sec. 5, T.19S, R.19E. Both of these are + approximately six miles northwest of Garnett, Anderson + County, Kansas. + + _Referred specimens._--K. U. nos. 786F, 787F, 788, 9939, + 11424, 11425, 11426, 11427, 11428, 11429, 11430, 11431, + 11432, 11433, 11434, 11449, 11450, 11451, 11452, 11453, + 11454, 11455, 11457. + + _Preservation._--Preservation of many of the specimens is + good, few are weathered, but most of the remains are + fragmentary and dissociated. One specimen (the type, no. + 786F) and half of another were nearly complete. Specimens + are found scattered throughout the Rock Lake shale (see p. + 498). + + _Morphology._--Terminology used for bones of the skull is + that of Moy-Thomas (1937) and Schaeffer (1952). + + +_Endocranium and parasphenoid_ + +[Illustration: FIG. 1. _Synaptotylus newelli_ (Hibbard). Restoration of +the basisphenoid, based on K. U. no. 9939, x 5. A, lateral view, B, +posterior view, C, ventral view.] + +The basisphenoid (see fig. 1) has been observed in only one specimen (K. +U. no. 9939) in posterodorsal and ventral views. The basisphenoid, +although somewhat crushed, appears to be fused to the parasphenoid. Both +antotic and basipterygoid processes are present, and are connected by a +low, rounded ridge. The antotic processes are large, bulbar projections. +These processes in _Rhabdoderma_ are wider and more flattened +(Moy-Thomas, 1937:figs. 3, 4). The antotic processes are at mid-point on +the lateral surface, not dorsal as in _Rhabdoderma_, and both the +processes and the ridge are directed anteroventrally. The basipterygoid +processes are smaller, somewhat vertically elongated projections, +situated at the end of the low connecting ridge extending +anteroventrally from the antotic processes, and are not basal as are +those of _Rhabdoderma_. The sphenoid condyles, seen in posterior view, +issue from the dorsal margin of the notochordal socket. The margins of +the socket are rounded, and slope down evenly to the center. A slight +depression situated between and dorsal to the sphenoid condyles is +supposedly for the attachment of the intercranial ligament (Schaeffer +and Gregory, 1961:fig. 1). The alisphenoids extend upward, +anterodorsally from the region above the sphenoid condyles, and may +connect to ridges on the ventral surface of the frontals. The lateral +laminae are not preserved, and their extent is unknown. + +In viewing the changes in the endocranium of Carboniferous and Permian +coelacanths, it would be well to consider the mechanical relationship of +the loss of the basipterygoid processes to the effect on swallowing +prey. Evidently many of the coelacanths, _Latimeria_ for example, are +predators (Smith, 1939:104); to such fishes a more efficient catching +and swallowing mechanism would be an adaptive improvement. Stensioe +(1932:fig. 14) presents a cross section of the ethmosphenoid moiety of +the endocranium of _Diplocercides kayseri_ (von Koenen) showing the +metapterygoid of the palatoquadrate loosely articulated to both the +antotic and basipterygoid processes. According to Tchernavin (1948:137) +and Schaeffer and Rosen (1961:190) the swallowing of large prey depends +on the ability of the fish to expand its oral cavity by allowing the +posteroventral portion of the palatoquadrate and the posterior end of +the mandible to swing outward. Where the palatoquadrate articulates with +the basisphenoid at the antotic and basipterygoid processes, as in the +Devonian coelacanths, it can not swing so far laterally as where it +articulates with only the dorsal, antotic process. Perhaps the loss of +the basipterygoid articulation reflects the development of a more +efficient mechanism for swallowing prey in these fishes. Schaeffer and +Rosen (1961:191, 193) show that in the evolution of the actinopterygians +several changes improved the feeding mechanism: some of these changes +are: (1) freeing of the maxilla from the cheek, giving a larger chamber +for the action of the adductor mandibulae; (2) development of a coronoid +process on the mandible; and (3) increase in torque around the jaw +articulation. In coelacanths, at least some comparable changes occurred, +such as: (1) loss of the maxillary, thus increasing the size of the +adductor chamber; (2) development of the coronoid bone, affording a +greater area for muscle attachment; (3) development of an arched dorsal +margin on the angular; (4) modification of the palatoquadrate complex, +with resultant loss of the basipterygoid processes. In _Synaptotylus_ +the basipterygoid processes are small, not basally located, and perhaps +not functional. A more efficient feeding mechanism developed rapidly +during the Carboniferous and has remained almost unaltered. + +[Illustration: FIG. 2. _Synaptotylus newelli_ (Hibbard). Restoration of +the parasphenoid, based on K. U. nos. 9939, 11451, x 5. A, ventral view, +B, dorsal view and cross sections.] + +The parasphenoid (see fig. 2) is a shovel-shaped bone having a wide +anterior portion and a narrower posterior portion of nearly uniform +width. Most of the ventral surface is covered with minute granular +teeth. The anterior margin is flared and curved posteromedially from the +lateral margin to a median triangular projection. The lateral margins +curve smoothly from the greatest anterior width to the narrow central +portion, where the margins become somewhat thickened and turned +dorsally. Posterior to this the lateral margins are probably nearly +straight. The external surface of the anterior section is nearly flat +and has a central depressed area the sides of which slope evenly to the +center. The internal surface is smooth and centrally convex. Because of +the fragmentary nature of all four observed specimens, total length was +not measured but is estimated to be 15 to 20 mm. The opening of the +hypophyseal canal was not present, possibly because of crushing. +Ethmoidal ossifications were not preserved in any of the specimens +studied. The parasphenoid differs from that of _Rhabdoderma elegans_ +(Newberry) in being more flared and widened anteriorly and more concave +centrally. + + +_Dermal bones of the skull_ + +Various portions of the cranial roof are preserved in several specimens +(see fig. 3). For comparisons with _Rhabdoderma elegans_, see Moy-Thomas +(1937:fig. 1). + +The premaxillaries and rostral elements are not preserved in any of the +specimens. Only one pair of relatively large frontals have been +observed; they are 5.5 to 9.0 mm. long and 2.0 to 3.5 mm. wide. These +are nearly flat bones, with the greatest width posteriorly 0.1 to 1.0 +mm. wider than the anterior portion. The midline suture is straight, the +lateral margins are nearly straight, the anterior margin slopes evenly +posterolaterally, and the posterior margin is slightly convex to +straight. The anterior margin in _R. elegans_ is essentially straight. +Ornamentation consists of sparse, unevenly spaced, coarse tubercles or +short striae. In one specimen both bones have small clusters of +tubercles near the lateral margins and about 2.0 mm. from the posterior +margin. None of these bones has alisphenoids or ridges on the ventral +surface, as Stensioe (1921:65, 97) described for _Wimania_ and _Axelia_. + +[Illustration: FIG. 3. _Synaptotylus newelli_ (Hibbard). Diagram of the +dermal bones of the skull, in lateral view, based on K. U. nos. 788 and +11432. x 2-1/2 approximately.] + +Only six supraorbitals have been preserved (see fig. 3). These are +nearly square, flat, thin bones lying nearly in place adjacent to a +frontal on K. U. no. 788. The smallest is anterior; the margins of all +are nearly straight. The bones are unornamented. Each bears a pore of +the supraorbital line just below the midline. The supraorbitals of _R. +elegans_ have a triangular outline and do not bear pores. + +Intertemporals (fig. 3) on several specimens vary from approximately 9.0 +to 15.0 mm. in length, 2.0 to 2.7 mm. in anterior width, and increase to +4.5 to 8.0 mm. in maximum posterior width. The midline suture is +straight, the anterior margin is concave and the lateral margin proceeds +laterally in a concave curve to the widest portion. In _R. elegans_ only +the anterior half of the corresponding margin is concave. The posterior +margin is slightly rounded and slopes anteriorly toward the lateral +margin. Ornamentation is usually of randomly oriented tubercles and +striae, although striae are more common in the posterior third and may +be longitudinal, whereas tubercles occur mainly on the anterior section. +No evidence of sensory pores, as seen on the intertemporal of _R. +elegans_, has been found. + +The supratemporals were observed on only one specimen (K. U. no. 788), +(fig. 3). Sutures were difficult to distinguish but the medial margin is +presumed to curve to fit and to articulate with the lateral margins of +the intertemporals. Lateral margins are smoothly curved but the anterior +and posterior margins were broken off. There appears to be no +ornamentation on this bone. The supratemporals are much more elongated +and curving than those in _R. elegans_. + +The cheek region is nearly complete in one specimen (K. U. no. 788), and +scattered parts occur in a few others (see fig. 3). The lacrimojugal of +no. 788 is elongate, with both ends curving dorsally. It differs from +the lacrimojugal in _R. elegans_, in which the anterior end extends +anteriorly and is not curved dorsally. The posterior and anterior +margins are not preserved; the greatest height appears to be posterior. +Pores of the suborbital portion of the infraorbital sensory canal are +seen on the dorsal surface of the bone. In _R. elegans_ the pores are on +the lateral surface. A section of the lacrimojugal on specimen no. +11425, broken at both ends, shows a thin layer of bone perforated by the +pores and covering a groove for the canal within the dorsal margin of +the bone. Both specimens are unornamented. + +A nearly complete postorbital (fig. 3) on specimen no. 788 is nearly +triangular, with the apex ventral. The concave anterior margin bears +pores of the postorbital part of the infraorbital line. Ornamentation +consists of widely spaced, coarse tubercles. + +Part of one squamosal is preserved. It is somewhat triangular and its +apex is ventral. This bone is associated with the postorbital, +subopercular and lacrimojugal on no. 788. The preopercular sensory line +passes down the curving ventral margin of this bone, and extends +ventrally onto a narrow projection. A low ridge, nearly vertical, passes +dorsally from about mid-point of the canal to the dorsal portion. The +anterior margin is nearly straight, the ventral margin is concave, and +the dorsal margin is convex dorsally but may be incomplete. Perhaps the +squamosal and preopercular are fused. The surface appears smooth; the +view may be of the medial side. The squamosal of _R. elegans_ is nearly +triangular and notably different from that of _Synaptotylus newelli_. + +The subopercular (fig. 3) shows closely spaced tubercles on the lateral +surface. The bone is an elongated, irregular triangle with the apex +pointing anterodorsally. The margins are incomplete, except for the +concave, curving anterior margin. + +Numerous operculars (fig. 3) occur in the suite of specimens, both +isolated and nearly in place. Each is subtriangular; the apex of the +triangle is ventral. A slight convexity projects from the anterodorsal +border. The posterior margin is broadly but shallowly indented. +Otherwise the margins are smooth. Maximum height ranges from 8.0 to 11.0 +mm., and maximum width from 8.0 to 13.0 mm. Ornamentation varies from a +few widely spaced, randomly oriented tubercles to closely spaced +tubercles merging posteriorly into striae. On some specimens these are +parallel to the dorsal border, and oblique in the central portion. On +the posterior margins of several operculars the striae break up into +tubercles. A few operculars have closely spaced tubercles over much of +the surface. The internal surface is smooth. + + +_Visceral skeleton_ + +The palatoquadrate complex, best seen on K. U. no. 9939 (fig. 4), +consists of endopterygoid, ectopterygoid, metapterygoid and quadrate. No +trace of epipterygoids, dermopalatines or autopalatines, such as +Moy-Thomas (1937:392, fig. 5) described for _Rhabdoderma_, has been +observed. + +The endopterygoid has a long, ventral, anteriorly-directed process, and +an anterodorsal process that meets the metapterygoid in forming the +processus ascendens. The suture between the endopterygoid and +metapterygoid, seen in lateral view, is distinct in some specimens and +has an associated ridge; these bones appear to be fused in others, +without regard to size. This suture curves dorsally from a point +anterior to the quadrate and passes anterodorsally to the extremity of +the processus ascendens. The suture is visible on the medial side only +near the processus ascendens, for it is covered by a dorsal, toothed +extension of the endopterygoid. The endopterygoid has a smooth lateral +surface; the medial surface is covered with tiny granular teeth, in +characteristic "line and dot" arrangement. The teeth extend onto the +ventral surface of the ventral process. + +[Illustration: FIG. 4. _Synaptotylus newelli_ (Hibbard). Restoration of +the palatoquadrate complex, based on K. U. no. 9939, x 5. A, medial +view, B, lateral view.] + +Two long, narrow, splintlike bones covered on one surface with granular +teeth are interpreted as ectopterygoids. These are 13.0 and 16.0 mm. +long and each is 1.5 mm. wide. Orientation of these is unknown, but they +probably fitted against the ventral surface of the ventral process of +the endopterygoid (Moy-Thomas, 1937:fig. 5). + +[Illustration: FIG. 5. _Synaptotylus newelli_ (Hibbard). A, ceratohyal, +lateral (?) view, based on K. U. nos. 11429 and 11457, x 5. B, urohyal, +based on K. U. no. 11457, x 5.] + +The metapterygoid has a smooth surface in both views. The dorsal edge +has a thickened, flared margin that presumably articulated with the +antotic process of the basisphenoid. No articular surface for the +basipterygoid process has been observed. + +The quadrate is distinct and closely applied to the posteroventral +margin of the complex. In medial view the margin is nearly straight and +continues to the ventral edge. The ventral surface is thickened and +forms a rounded, knoblike articular surface. In lateral view the surface +is smooth; the anterior margin is irregular (or perhaps broken on all +specimens), and proceeds in an irregular convex curve from the posterior +to the ventral margin. + +The general shape of the palatoquadrate complex is most nearly like that +of _Rhabdoderma elegans_ (Moy-Thomas, 1937:fig. 5). The orientation of +the complex in the living fish was probably oblique, with the processus +ascendens nearly vertical, the quadrate oblique, and the ventral process +of the endopterygoid extending dorsoanteriorly and articulating with the +parasphenoid. + +Of the hyoid arch only the ceratohyals (see fig. 5A) are preserved in +several specimens. These are long, curved bones with a posteroventral +process and widened, flaring posterior margin. The medial (?) surface is +concave in one specimen. The lateral (?) surface displays a distinct +ridge on several specimens, arising on the dorsal surface opposite the +posteroventral process and extending diagonally to the anteroventral end +of the anterior limb. The impression of one other specimen appears to +have a central ridge because of greater dorsal thickness and narrowness. +Both surfaces are unornamented. + +The urohyal (see fig. 5B) is an unornamented, Y-shaped bone, with the +stem of the Y pointing anteriorly. Orientation with respect to dorsal +and ventral surfaces is uncertain. In one view a faint ridge, also +Y-shaped, occurs on the expanded posterior portion, and the surface is +convex. The anterior process has a convex surface, sloping evenly off to +the lateral margin; the opposite side of the process has a concave +surface. The posterior portion has a slightly depressed area (see fig. +5B) at the junction of the "arms" of the Y. + +The five branchial arches are represented by the ceratobranchials, +several of which are preserved on K. U. no. 11431. These are long bones +with anteriorly curving ventral ends. The medial surfaces are partly +covered with minute granular teeth; only the dorsal part is without +teeth. The dorsal articular surface is convex dorsally and rounded. + +The mandible (fig. 3), the best specimens of which are K. U. nos. 788 +and 11425, is seen only in lateral and ventral views, with only angular, +splenial and dentary visible. + +The angular forms the main body of the mandible, and is similar to that +of _Spermatodus_. The dorsal margin of the angular is expanded in the +central region, with some variation. One specimen has an expanded +portion slightly anterior to that of the opposite angular. The articular +surface near the posterior end has not been observed; the posterior end +of the angular slopes off abruptly. The anterior sutures are seen in +only two specimens, K. U. nos. 788, 11425. The dentary meets the angular +in a long oblique suture; the dentary gradually tapers posterodorsally +and ends on the dorsal surface of the angular. The splenial fits into a +posteriorly directed, deep V-shaped notch on the ventral surface. The +lateroventral surface of the angular contains sensory pores of the +mandibular line. The ventral surface extends medially into a narrow +shelf, approximately 1.0 mm. wide, which extends the full length of the +bone; the external surface of this shelf is smooth and slightly concave +dorsally. Ornamentation of the angular consists of tubercles and +longitudinal or oblique striae, occurring mostly on the expanded +portion. The medial surface is not seen. Several broken specimens show +a central canal filled with a rod of calcite; in one of these the +sensory pores are also calcite-filled and appear to be connected to the +rod. Thus the pores originally opened into a central canal. + +The dentary is an unornamented bone with the anterior half curving +medially; the greatest height is anterior. This bone in specimen K. U. +no. 11425 bears irregularly spaced, simple, recurved, conical teeth; +nine were counted, but there is space for many others. One other +specimen, no. 11429, seems to have tiny tubercles on the surface. The +dentary meets the splenial dorsally in a straight suture. + +The splenial also curves medially, and as stated, meets the dentary in a +straight suture. Ornamentation on this bone was not observed. The +posterior margin is V-shaped and fits the notch in the angular. The +ventral surface bears three or more sensory pores of the mandibular +line. + +The gular plates are oval. The medial margin is straight to slightly +curved, the lateral margin curved crescentically, the posterior end is +blunt, and the anterior end somewhat rounded. Ornamentation varies +greatly; some bones show only a few tubercles, whereas others exhibit an +almost concentric pattern of closely spaced striae. Typically there are +some tubercles in the anterior quarter or third of the total length; +these pass into longitudinally oriented striae in the posterior section. +A few have only randomly oriented, widely-spaced striae. The internal +surface is smooth. + +The coronoid (K. U. no. 11428) is a triangular bone, with the apex +pointing dorsally. The lateral surface is smooth; no teeth were +observed. Moy-Thomas (1937:292, 293) mentions several tooth-bearing +coronoids in _Rhabdoderma_, but as yet these have not been seen in +_Synaptotylus_. + + +_Axial skeleton_ + +Only three specimens (K. U. nos. 786F, 787F, 11450) show parts of the +vertebral column, but isolated neural and haemal arches are numerous. +All are of the coelacanth type, having Y-shaped neural and haemal +arches, without centra. A total count of 38 was obtained, but this was +incomplete; the actual number was probably near 50. Counts of 10 and 16 +haemal arches were obtained in two of the specimens. Total height of +neural arches ranges from 7.5 to 12.0 mm., and of haemal arches, from +9.0 to 12.0 mm. The shorter arches are anterior and the height increases +gradually to a maximum in the caudal region. Height of the spines varies +from 4.0 to 9.0 mm., or from twice the height of the arch in the +anterior to three times the height in the caudal region. Total width of +the base, measured in isolated specimens because lateral views in other +specimens prevented measuring width, ranges from 0.7 to 4.2 mm. The +short, broad arches having short spines occur at the anterior end of the +spinal column; the narrower arches having tall spines occur toward the +caudal end. Broken neural and haemal arches show a thin covering of bone +with a central, calcite-filled cavity, which in life may have been +filled with cartilage (Stensioe, 1932:58, fig. 20). + +No ossified ribs have been observed, either isolated or in place. + +For further description of the axial skeleton, see Hibbard (1933). + +[Illustration: FIG. 6. _Synaptotylus newelli_ (Hibbard). Paired fin +girdles. A, pectoral girdle, lateral view, based on K. U. no. 11433, x +3.5. B, pelvic girdle basal plate, medial (?) view, based on K. U. no. +788, x 8. Anterior is toward the left.] + + +_Girdles and paired fins_ + +A nearly complete pectoral girdle on specimen K. U. no. 11433 (see fig. +6A) has only a cleithrum and clavicle. No evidence of an extracleithrum +or supracleithrum has been observed, but the extracleithrum may be fused +to the cleithrum. The two bones form a boot-shaped unit, with the +anteroventral part turned medially to form a horizontal process which +meets the opposite half of the girdle. In lateral view the surface is +unornamented, and convex in the ventral half. The suture between the +cleithrum and clavicle begins on the expanded posterior portion, the +"boot-heel," at a point immediately below the greatest width on the +posterior margin, passes anteriorly, then turns sharply and parallels +the anterior margin. The shape of the cleithrum resembles that in +_Rhabdoderma_ and the internal surface is not ridged (see Moy-Thomas, +1937:fig. 9). The exact orientation in the fish is uncertain, but if the +median extension is really horizontal, then the posterior expansion is +directed caudally. The medial surface is concave, steepest near the +anterior margin, and then slopes outward evenly. In medial view one +specimen (K. U. no. 11426) shows a small, caudally directed projection +of bone, evidently for articulation of the fin-skeleton, at the widest +portion of the cleithrum. Sutures on several specimens were indistinct. +Broken specimens show sutural faces, but many nearly complete specimens +show little or no indication of sutures, without regard to size of the +girdles. The internal structure of the fin was not observed. + +Numerous isolated basal plates of the pelvic girdle have revealed +details of structure but no information on the orientation. Presumably +the basal plates of _Synaptotylus_ had essentially the same orientation +as those of other coelacanths (Moy-Thomas, 1937:395). The most complete +basal plate is K. U. no. 788 (see fig. 6B). The three apophyses diverge +anteriorly; the horizontal one is best developed and the dorsal one is +least well developed. A median process (Schaeffer, 1952:49), denticulate +on several specimens, articulates with the corresponding process of the +opposite plate. The expanded part that articulates with the skeleton of +the fin extends caudally. The posterior expanded part is nearly square +in outline, resembling the dorsal, rectangular projection. One side +bears ridges leading to the extremities of the apophyses, and faint +crenulations on the median process. This may be the medial view. The +other view displays a smooth surface, usually without indication of the +ridges seen in the reverse view. These specimens differ somewhat from +the basal plates of _Rhabdoderma_ and appear to be intermediate between +_Rhabdoderma_ and _Coelacanthus_ (Moy-Thomas, 1937:fig. 10A, B). The +apophyses are not free as in _Rhabdoderma_ but webbed with bone almost +to their extremities, as in _Coelacanthus_. + +The pelvic fin is seen in only two specimens (K. U. nos. 786F, 788). +That on no. 788 is lobate and has 25 lepidotrichia, jointed for +approximately the distal half, and 2.5 to 13.0 mm. in length. Total +length of the fin is 25.0 mm. There is no trace of the internal skeletal +structure or of the articulation to the basal plate in either specimen. +For a description of the fin on no. 786F, see Hibbard (1933:281). + + +_Unpaired fins_ + +A few isolated bones on K. U. no. 788 (fig. 7) are interpreted as basal +plates of the unpaired fins. For additional description of the unpaired +fins on the type, K. U. no. 786F, see Hibbard (1933). + +Two of these bones are flat, smooth and oblong, bearing a diagonal ridge +that extends in the form of a projection. Orientation is completely +unknown. These may be basal plates of the anterior dorsal fin. The fin +on no. 786F that Hibbard (1933:281) interpreted as the posterior dorsal +fin is now thought to be the anterior dorsal fin. + +[Illustration: FIG. 7. _Synaptotylus newelli_ (Hibbard). Basal plates of +unpaired fins. A, anterior dorsal fin, based on K. U. no. 788, x 10. B, +posterior dorsal fin, based on K. U. no. 788, x 12. C, anal fin, based +on K. U. no. 11450, x 5. Anterior is toward the left.] + +One distinctive bone may represent the basal plate of the posterior +dorsal fin. This incomplete specimen shows two projecting curved +processes, bearing low but distinct ridges, which diverge, probably +anteriorly. The central portion is narrow. The two ridges continue onto +the posterior portion. This has been broken off, but shows that the +ridges diverge again. The surface is smooth, except for the ridges. As +before, orientation is uncertain. On no. 786F this fin was interpreted +by Hibbard (1933:281) as the anal fin. + +Only part of one basal plate of the anal fin was preserved on K. U. no. +11450. That plate is oblong and has an expanded anterior end. The +narrow, constricted part bears two oblique ridges and a few tubercles. +The posterior part has nearly straight margins (represented by +impressions) and the posterior margin is oblique, sloping +anteroventrally. The flared anterior part has a smooth surface. This +basal plate is more nearly like those of _Coelacanthus_, according to +the descriptions given by Moy-Thomas (1937:399). The basal plate is +associated with seven apparently unjointed, incomplete lepidotrichia. +The anal fin on no. 786F is interpreted as the anterior dorsal fin +(Hibbard, 1933:281). + +The caudal fins are preserved on K. U. nos. 786F, 787F, and have a total +of 24 lepidotrichia, 12 above and 12 below. These are jointed for the +distal half or two-thirds, and are up to 16.0 mm. in length. In specimen +no. 787F the supplementary caudal fin has at least seven lepidotrichia, +the longest of which is 11.0 mm. but incomplete. Anterior lepidotrichia +appear unjointed but the posterior ones are jointed for the distal +two-thirds (?) (these are broken off). The supplementary caudal fin is +approximately 1.5 mm. long and 8.0 mm. or more wide. The supplementary +caudal fin on K. U. no. 786F described by Hibbard (1933:281) could not +be observed; this part of the caudal fin is missing. + + +_Squamation_ + +In the suite of specimens isolated scales are numerous, but patches of +scales are rare. Only two specimens (K. U. nos. 786F, 787F) are complete +enough for scale counts, but preservation permits only partial counts. +In general the scales resemble those of _Rhabdoderma elegans_ +(Newberry). + +The scales are oval. The exposed posterior part of each bears +posteriorly converging ridges; the anterior part is widest and shows a +fine fibrillar structure. There are at least six scale-rows on either +side of the lateral line. Lateral line scales show no pores, and except +for slight irregularities in the orientation and length of the posterior +ridges, closely resemble the others. Central ridges on the lateral line +scales are shorter and tend to diverge from the center of the impression +of the canal. The lateral line canal shows only as the impression of a +continuous canal 0.7 mm. in diameter. Preservation is poorest in scales +along the line of the neural and haemal arches; therefore lateral line +scales are rarely preserved. Isolated scales are of two types: those on +which the posterior ridges converge sharply and form the gothic arch +configuration mentioned by Hibbard (1933:282), and those which do not. +Both types of scales can be present on one fish, as shown by specimen +no. 788. This is not apparent on nos. 786F and 787F; all of the scales +on these specimens appear to be much alike. Both Moy-Thomas (1937:385) +and Schaeffer (1952:51, 52) have remarked on the variation of the scales +on different parts of the same fish. Because the number of ridges and +amount of convergence of the ridges is not related to size of the scale, +it is concluded that these characters are not of taxonomic significance. + +The strong resemblance of the scales of the Garnett specimens to those +of _Rhabdoderma elegans_ (Newberry) caused Moy-Thomas (1937:399) to add +Hibbard's two species to the synonymy of _R. elegans_. But at that time +only the scales could be adequately described. If the shape of the scale +and the number and pattern of ridges can vary with age, size and shape +of the scale, it follows that assignment of isolated scales to a species +should not be attempted. Assignment to genus should be made only with +caution. + +_Discussion._--The relationship of _Synaptotylus_ to other coelacanths +is obscure at present. The knoblike antotic processes on the +basisphenoid are unlike those of any other known coelacanth. The +palatoquadrate complex is shaped like that of _Rhabdoderma elegans_ but +consists of fewer bones, probably because of fusion. The scales resemble +those of _Rhabdoderma_. With regard to general shape of fin girdles, the +pectoral girdle resembles that of _Eusthenopteron_ more than that of +_Rhabdoderma_, but the cleithrum is more nearly like the cleithrum of +_Rhabdoderma_. The pelvic girdle appears to be midway between those of +_Rhabdoderma_ and _Coelacanthus_ in general appearance. Regarding the +basal plates of the remaining fins, those of _Synaptotylus_ appear to +resemble basal plates of both _Rhabdoderma_ and _Coelacanthus_. +Considering the structure of the sphenethmoid region of the braincase, +_Synaptotylus_ is probably more closely related to _Rhabdoderma_ than +to other known coelacanth genera. + + +COMMENTS ON CLASSIFICATIONS + +Classification of Carboniferous coelacanths has been difficult, partly +because the remains are commonly fragmentary, and significant changes in +anatomy did not become apparent in early studies. In general, +coelacanths have been remarkably stable in most characters, and it has +been difficult to divide the group into families. As Schaeffer (1952:56) +pointed out, definition of coelacanth genera and species has previously +been made on non-meristic characters, and the range of variation within +a species has received little attention. For example, Reis (1888:71) +established the genus _Rhabdoderma_, using the strong striation of the +scales, gular plates and posterior mandible as the main characters of +this Carboniferous genus. Moy-Thomas (1937:399-411) referred all +Carboniferous species to _Rhabdoderma_, redescribed the genus and +compared it to _Coelacanthus_, the Permian genus. He cited as specific +characters the ornamentation of the angulars, operculars and gular +plates (Moy-Thomas, 1935:39; 1937:385). Individual variation in some +species has rendered ornamentation a poor criterion. This variation is +apparent in _Synaptotylus newelli_ (Hibbard), some specimens having +little or no ornamentation; others having much more. The number of +ridges and pattern of ridges on the scales also varies. Schaeffer +(1952:56) has found this to be true of _Diplurus_ also. Moy-Thomas +(1935:40; 1937:385) realized that the type of scale is a poor criterion +for specific differentiation. In the search for features useful in +distinguishing genera of coelacanths, Schaeffer and Gregory (1961:3, 7) +found the structure of the basisphenoid to be distinctive in known +genera, and thought it had taxonomic significance at this level. Higher +categories should have as their basis characters that display +evolutionary sequences. A recent classification (Berg, 1940), followed +in this paper, reflects two evolutionary trends in endocranial structure +of coelacanths: reduction of endocranial ossification and loss of the +basipterygoid processes. Because there has been little change in other +structures in coelacanths, Berg's classification is the most useful. +Berg (1940:390) includes _Rhabdoderma_ in the suborder Diplocercidoidei +because of the presence of the basipterygoid processes, and in the +single family, Diplocercidae, but remarks that because of the reduced +amount of endocranial ossification the Carboniferous Diplocercidae +"probably constitute a distinct family." In considering this concept of +classification, the subfamilies Diplocercinae and Rhabdodermatinae of +the family Diplocercidae are proposed above. The subfamily +Rhabdodermatinae includes at present _Rhabdoderma_ and _Synaptotylus_. +The principal characters of the subfamily Rhabdodermatinae, named for +the first known genus, are the retention of the basipterygoid processes +and the reduction of endocranial ossification. Application of this +classification based upon endocranial structure would probably change +existing groupings of species of Carboniferous coelacanths; the entire +complex of Carboniferous genera should be redescribed and redefined. It +will be necessary to consider endocranial structure in any future +classification. + +The greater part of the evolution previously mentioned appears to have +been accomplished during the Carboniferous; thereafter coelacanth +structure became stabilized. The trend progressed from Devonian +coelacanths which had two large unpaired bones in the endocranium, and +both antotic and basipterygoid processes on the basisphenoid, to +Carboniferous fishes in which ossification was reduced to a number of +paired and unpaired bones embedded in cartilage, and retaining both +processes, and then post-Carboniferous kinds with reduced ossification +and no basipterygoid processes. The Pennsylvanian was evidently the time +of greatest change for the coelacanths, and they have not changed +significantly since, in spite of the fact that since the Jurassic they +have shifted their environment from shallow, fresh water to moderate +depth in the sea (Schaeffer, 1953:fig. 1). The changes in endocranial +structure appear to be significant, and are perhaps related to higher +efficiency of the mouth parts in catching and swallowing prey (see p. +482). + + +ENVIRONMENT + +The coelacanth fishes from the Rock Lake shale are part of the varied +fauna collected from Garnett. Peabody (1952:38) listed many elements of +the fauna and flora, and concluded that the deposits are of lagoonal +origin. In addition to numerous invertebrates (including microfossils) +and arthropods, a number of vertebrates other than coelacanths have been +found. These include at least one kind of shark, _Hesperoherpeton +garnettense_ Peabody, one or more kinds of undescribed labyrinthodonts +and the reptiles _Petrolacosaurus kansensis_ Lane, _Edaphosaurus ecordi_ +Peabody, and _Clepsydrops_ (undescribed species). This is indeed a rich +vertebrate fauna, and the earliest known reptilian fauna. Much of the +rock contains plant remains. The flora that has been identified is +adapted to growing in a well-drained soil; although it contains some +elements considered characteristic of the Permian, it is of +Pennsylvanian age (Moore _et al._, 1936). Peabody (1952:38-39) discusses +the features of these lagoonal sediments. Much of the fauna and flora +suggests continental origin, but the many marine invertebrates at some +horizons indicate that at least some of the sediments were of marine +origin. + +Little can be said about the actual environment of the living fishes of +the genus _Synaptotylus_. Remains of these fishes occur in layers +containing marine invertebrates, as well as in those containing plant +remains and vertebrate skeletal parts, and in those nearly completely +composed of dark carbonaceous material. Most of the remains are +fragmentary and consist of isolated bones, isolated scales, and +dissociated skulls; only one specimen and half of another are nearly +complete. Many published statements on _Rhabdoderma_, a related genus, +indicate both marine and fresh-water environments. Wehrli (1931:115) +regarded _Rhabdoderma elegans_ (Newberry) as a euryhaline species, and +cited its occurrence with both marine and fresh-water fossils. Aldinger +(1931:199) also found this to be the case with other species, and Fiege +(1951:17) quotes others as giving the same information. Keller +(1934:913) thought that few Carboniferous fishes were exclusively +marine, and stated that the majority of them became adapted to fresh +water during the late Carboniferous. Later, Schaeffer (1953:175) stated +that all Carboniferous and Permian coelacanths were fresh-water fishes, +and that many were from swamp deposits. If Keller is correct, then +members of the genus _Synaptotylus_ may have inhabited the lagoon, the +adjacent sea, or the streams draining into the lagoon. Perhaps these +fishes swam upstream, as modern salmon and tarpon do, although there is +no direct evidence for this. Possibly they lived in the lagoon at times +of scant rainfall and little runoff, when the salinity of lagoon water +approached normal marine values or the fishes may have lived in the +streams, and after death were washed into the lagoon. As numerous +remains of land plants and animals were washed in, perhaps this best +accounts for the presence of the fish in nearly all layers of the +deposits, not only the marine strata. + + +SUMMARY + +A new genus of Pennsylvanian coelacanths, _Synaptotylus_, is described +and a previously named species, _Coelacanthus newelli_ Hibbard, 1933 +(_C. arcuatus_ Hibbard, 1933, is a junior synonym), is referred to this +genus. All specimens of _Synaptotylus newelli_ (Hibbard) were collected +from the Rock Lake shale member of the Stanton formation, Lansing group, +Missouri series, six miles northwest of Garnett, Anderson County, +Kansas. _Synaptotylus_ is distinguished from all other coelacanths by a +basisphenoid having large, knoblike antotic processes each connected by +a low ridge to a small basipterygoid process. _Synaptotylus_ is most +closely related to _Rhabdoderma_, but is intermediate between +_Rhabdoderma_ and _Coelacanthus_ in shape of the fin girdles and basal +plates. Two new subfamilies, Diplocercinae and Rhabdodermatinae, of the +family Diplocercidae, are proposed. _Synaptotylus_ and _Rhabdoderma_ are +included in the subfamily Rhabdodermatinae, because both exhibit reduced +ossification in the endocranium and retain basipterygoid processes. + +Loss of the basipterygoid processes in post-Carboniferous coelacanths +may reflect the development of a more efficient feeding mechanism, by +allowing the palatoquadrate complex and mandible to swing farther +laterally and expand the oral cavity. + +_Synaptotylus newelli_ (Hibbard) may have occupied either the sea or +fresh water; these fishes occur in lagoonal deposits with reptiles and +amphibians, arthropods, marine invertebrates and remains of land plants. + +Because scale patterns on _Synaptotylus_ and _Rhabdoderma_ are so nearly +similar and vary with size of the scale and its location on the fish, it +is recommended that isolated scales not be assigned to a species, and to +a genus only with great caution. + + + + +LITERATURE CITED + + +ALDINGER, H. + +1931. Ueber karbonische Fische aus Westfaelen. Paleont. Zeit., +13:186-201. + + +BERG, L. S. + +1940. Classification of fishes, both Recent and fossil. Moscow and +Leningrad, 1940 (J. W. Edwards, Ann Arbor, Michigan, 1947, offset +reproduction, pp. 1-345, 197 figs., plus English translation of text, +pp. 346-517, 1947.) + + +FIEGE, K. + +1951. Eine Fisch-Schwimmspur aus dem Culm bei Waldeck. Neues Jahrb. +Geol. and Palaeont. Jahrgang 1951:9-31. + + +HIBBARD, C. W. + +1933. Two new species of _Coelacanthus_ from the middle Pennsylvanian of +Anderson County, Kansas. Kansas Univ. Sci. Bull., 21:279-287. + + +KELLER, G. + +1934. Fischreste aus dem oberkarbon des Ruhrgebiets. Gluckauf, +70:913-917. + + +MOORE, R. C., ELIAS, M. K., and NEWELL, N. D. + +1936. A "Permian" flora from the Pennsylvanian rocks of Kansas. Jour. +Geol., 44:1-31. + + +MOY-THOMAS, J. A. + +1935. A synopsis of the coelacanth fishes of the Yorkshire Coal +Measures. Ann. Mag. Nat. Hist., 15 (ser. 10): 37-46. + +1937. The Carboniferous coelacanth fishes of Great Britain and Ireland. +Proc. Zool. Soc. London, 107 (B): 383-415. + + +PEABODY, F. E. + +1952. _Petrolacosaurus kansensis_ Lane, a Pennsylvanian reptile from +Kansas. Kansas Univ. Paleont. Contrib., 1:1-41. + + +REIS, O. M. + +1888. Die Coelacanthinen mit besonderen Beruecksichtigung der im Weissen +Jura Bayerns verkommenden Arten. Palaeontographica, 35:1-96. + + +SCHAEFFER, B. + +1952. The Triassic coelacanth fish _Diplurus_, with observations on the +evolution of the Coelacanthini. Bull. Amer. Mus. Nat. Hist., 99:art. 2, +29-78. + +1953. _Latimeria_ and the history of the coelacanth fishes. New York +Acad. Sci. Trans., (2) 15:170-178. + + +SCHAEFFER, B., and GREGORY, J. T. + +1961. Coelacanth fishes from the continental Triassic of the western +United States. Amer. Mus. Novitates, 2036:1-18. + + +SCHAEFFER, B., and ROSEN, D. E. + +1961. Major adaptive levels in the evolution of the actinopterygian +feeding mechanism. Am. Zool., 1:187-204. + + +SMITH, J. L. B. + +1939. A living coelacanthid fish from South Africa. Trans. Roy. Soc. +South Africa, 28:1-106. + +STENSIOe, E. A. + +1921. Triassic fishes from Spitzbergen. Part I. Vienna, Adolf +Holzhausen: 1-307. + +1932. Triassic fishes from East Greenland. Meddel. om Gronland, +38:1-305. + + +TCHERNAVIN, V. V. + +1948. On the mechanical working of the head of bony fishes. Proc. Zool. +Soc. London, 118:129-143. + + +WEHRLI, H. + +1931. Die Fauna der Westfaelischen Stufen A und B der Bochumer Mulde +zwischen Dortmund und Kamen (Westfaelen). Palaeontographica, 74:93-134. + + +_Transmitted March 29, 1962._ + + + + + +End of the Project Gutenberg EBook of A New Genus of Pennsylvania Fish +(Crossoperygii, Coelacanthiformes) from Kansas, by Joan Echols + +*** END OF THIS PROJECT GUTENBERG EBOOK NEW GENUS OF PENNSYLVANIA FISH *** + +***** This file should be named 33560.txt or 33560.zip ***** +This and all associated files of various formats will be found in: + https://www.gutenberg.org/3/3/5/6/33560/ + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +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. 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