initial commit of ebook 33560HEADmain

1 files changed, 1415 insertions, 0 deletions

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.  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 1.F.3.  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.