CHAPTER XXI.

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THE MICROSAURIAN FAMILY SAUROPLEURIDÆ, FROM THE COAL MEASURES OF OHIO.

Family SAUROPLEURIDÆ Hay, 1902.

Hay, Bull. U. S. Geol. Surv., No. 179, p. 419, 1902.

The present family is an association of related forms due to similar structure of vertebrÆ, ribs, ventral scutellation, and limbs. There is no character in the skull which would indicate a separation of the genera here included, at least in the light of present knowledge.

The family may be characterized as: Subaquatic or terrestrial vertebrates with a typically amphibian development of the ventral armature; ribs intercentral, as in all members of the order; skull elongate and slender or broad and obtuse; cranial and dermal elements of the pectoral girdle sculptured; lateral-line canals indicated in one genus, Saurerpeton; limbs well developed, with well-developed digits and ungual phalanges claw-like; body usually slender, broad in Saurerpeton; the ribs broad and heavy; the vertebrÆ relatively stout; ventral armature highly developed, reaching the height of specialization among the Microsauria; scutellÆ consisting of rods, plates, or stout bristles. The family is represented by 5 genera: Sauropleura, Saurerpeton, Ctenerpeton, Leptophractus, and Eurythorax, the association of the last two genera being provisional. The genera may be distinguished as follows:

I. Pectoral elements sculptured, clavicles triangular, interclavicle diamond-shaped, ventral scutellÆ rods, arranged en chevron with anterior angle Sauropleura
II. Pectoral elements slightly sculptured, cranium broad, obtuse and sculptured, ventral armature broad imbricated plates extending on to the throat Saurerpeton
III. Limbs, skull, arches and dorsal vertebrÆ unknown, caudal vertebrÆ with fan-shaped neural and hÆmal spines which may indicate relationship with Ptyonius and Œstocephalus, but in those genera the ventral armature is weakly developed; ventral scutellÆ curved rod-like plates arranged en chevron with anterior angle, marked in abdominal region by distinct rounded pits Ctenerpeton
IV. Known only from fragments of the skull, teeth large and fluted; association in family provisional Leptophractus
V. Known only from a single interclavicle of peculiar form which resembles that of Saurerpeton; association in the family provisional Eurythorax

The members of this family are confined to the deposits of the Coal Measures at Linton, Ohio. Ctenerpeton, and possibly Sauropleura, were highly developed swimmers, but the strength of the limbs as exhibited, especially by Sauropleura and Saurerpeton, indicates that they had not entirely forsaken the land.

Genus SAUROPLEURA Cope, 1868.

Newberry, Proc. Phila. Acad. Nat. Sci., VIII, p. 98, 1856 (Pygopterus scutellatus).

Cope, Trans. Am. Phil. Soc., p. 22, 1869.

Cope, Geol. Surv. Ohio, II, pt. II, p. 402, 1875.

Hay, Bull. U. S. Geol. Surv., No. 179, p. 419, 1902.

Type: Sauropleura scutellata Newberry.

Vertebra and ribs well developed; limbs 4, rather large; 5 digits in the forefoot; carpus cartilaginous. Ventral armature of closely arranged rhomboidal scuta, arranged in lines, which are closely placed in chevrons, with the angle anterior.

In none of the species of this genus have the usual 3 thoracic shields (clavicles and interclavicles) been observed. The abdominal scutÆ, on the other hand, are much like those of Saurerpeton, being, however, smaller.

The species formerly described by Cope in the genus Colosteus are included in Sauropleura, where they find their closest allies.

There are 7 species belonging in this genus: Sauropleura digitata Cope, S. newberryi Cope, S. foveata Cope (Colosteus), S. scutellata Newberry (Colosteus) (type of genus), S. pauciradiata Cope (Colosteus), S. longidentata Moodie, S. enchodus Cope (Anisodexis). The species described by Cope as Sauropleura latithorax is regarded as belonging to a distinct genus, Saurerpeton.

Sauropleura scutellata Newberry.

Newberry, Proc. Phila. Acad. Nat. Sci., p. 98, 1856 (Pygopterus scutellatus).

Cope, Proc. Phila. Acad. Nat. Sci., p. 215, 1868.

Cope, Trans. Am. Phil. Soc., p. 22, 1869.

Cope, Geol. Surv. Ohio, II, pt. II, p. 402, 1875.

Type: Specimen No. 8669 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures. (Plate 14, fig. 3.)

This species was first described by Newberry as a fish belonging to the genus Pygopterus. Cope later placed it under the genus Colosteus and clearly showed its amphibian characters. The genus Colosteus was, however, based on a misconception. Cope in 1897 and Hay (317) referred the species to Sauropleura, where it is retained. The species is represented by a single individual preserved on a block of coal from Linton, and is also indicated by an interclavicle and its obverse; this element is of larger size than that of the type and was referred by Cope to Colosteus pauciradiatus. The characters of the plate are, however, so identical with those of the interclavicle in the type specimen that it is unhesitatingly referred to the present form.

The type specimen consists of the supero-lateral view of a crushed cranium with the anterior part of the body, exhibiting the interclavicle and the ventral scutellation. No limbs have been observed in this species. The mandibles are crushed across the cranium in such a way as to obscure its structure. The boundary of the left orbit is doubtfully determined as being a little back of the median line of the skull. There are small teeth present on the mandibles, but their number can not be determined. The cranial elements are sculptured with radiating grooves and ridges, but these are weakly developed. The snout is broad and but little narrower than the base of the skull. (Plate 21, fig. 5.)

The interclavicle, somewhat displaced, is the only element of the pectoral girdle preserved. It is peculiar in the possession of a backward extension which shows a beveled edge. The plate is ornamented by radiating grooves and ridges which are strongly developed. The larger specimen of an interclavicle shows the same characters as the one described, and it differs only in being about twice as large. There are no traces of limbs.

The ventral armature of the body is rather weak as compared to that of Sauropleura pauciradiatus, but it is still composed of closely packed scutes arranged en chevron. The character of the ventral armature and the sculpturing of the interclavicle are taken as the principal specific characters. From the other species of the genus the present form differs in its more slender elements of the ventral armor and in the form of the skull. The ribs are not clearly defined.

Measurements.

mm.
Median length of skull 70
Posterior width of skull 40
Anterior width of skull 20
Length of jaw 50
Anterior width of jaw 7
Posterior width of jaw 12
Length of mandibular tooth 1.5
Length of entire specimen 150
Width across belly, maximum 47
Length of interclavicle 27
Width of interclavicle 13
Width across posterior extension 4

Sauropleura digitata Cope.

Cope, Proc. Phila. Acad. Nat. Sci., p. 216, 1868.

Cope, Trans. Am. Phil. Soc., XIV, p. 15, 1869.

Cope, Geol. Surv. Ohio, II, pt. II, p. 403, pl. xxxvii, fig. 1, 1875.

Type: Specimen No. 8004 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures. (Plate 20, fig. 4.)

This species is represented by a single individual which is quite distinct from other members of the genus. The specimen has been spread over a surface of the coal, and exhibits ventral armature, dorsal region with ribs, and anterior and posterior limbs. Skull and caudal region not present.

The ventral armature is arranged in parallel lines directed obliquely forwards and continuous on the median line, forming there a chevron. The individual elements are oat-shaped, and acuminate at both ends. They are moderately imbricate in an antero-posterior section. On the pectoral region between the fore limbs the series of scutellÆ assume different directions, forming chevrons directed backwards, and forming with those of the belly a complete X.

The humerus, ulna, and radius are rather stout, and of a size relative to the body, as in common types of existing lizards; the ulna and radius separate. The carpus is cartilaginous; the digits are 5 well-developed fingers having phalanges in the following numbers, commencing on the inside: 3, 4, 5, 6, 5. The last phalanx of the second is obscured, and it is not positive that the number is as given; it is more probable that it was 4 than 3. The outer toe was more slender than the others; the distal phalanges of all the toes are stout, as in modern Caudata.

The ribs are long and curved as in reptiles, and judging by their distances the vertebrÆ are short; the latter are not well-defined, but there is no indication of prominent spines of any kind. The pelvic bones and portions of the hind limbs are present, but so obscured and confused as not to be easily made out. Enough remains to show that the hind limbs were longer than the fore. Thirteen ribs on one side and twelve on the other are preserved, with short ribs in the sacral region. The specimen is very indistinct and it is difficult for one to be sure of all the characters described by Cope.

Measurements of the Type of Sauropleura digitata Cope.

mm.
Length of specimen 115
Greatest width 80
Diameter of ventral scute .75
Length of rib 15
Width of rib 1
Length of humerus 20
Length of radius 10
Length of ulna 9
Length of metacarpal 4
Length of fourth digit of hand 22
Length of interclavicle 20
Width of interclavicle 11

The interclavicle is diamond-Shaped, with surface punctate and edges radiately grooved at a distance of 1.5 mm. from the edge. The hand on the right side of the specimen contains 3, 3, 3, 6, 3 phalanges. The vertebrÆ are all imperfectly preserved.

Other specimens of this species are 2567 (48) , 8376 G, 8704 G, American Museum of Natural History. Coal Measures of Linton, Ohio. Collected by Dr. J. S. Newberry.

Sauropleura newberryi Cope.

Cope, Geol. Surv. Ohio, II, pt. II, p. 404, pl. xxxvii, figs. 2 and 3; pl. xli, fig. 5, 1875.

Type: Specimen No. 8612 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

The type specimen of the species exhibits a portion of the posterior part of the skull, a considerable part of the body, with the fore limbs and abdominal scutellÆ. No vertebrÆ can be definitely discovered and the ribs are not distinctly visible. The cranial fragment is the upper surface of the tabulare and adjacent elements, and a broad band of the posterior parts of these is seen to be smooth, and is preceded by a slightly roughened surface. The abdominal scutÆ are diamond-shaped, and are thin and light, not massive, as in S. scutellata, and are sometimes marked with a median longitudinal keel. The fore limb is large, especially the humerus, which is much dilated distally, and has a strong crest on the outer side from near the proximal end. The ulna and radius are much shorter, and more dilated proximally than distally; they are well separated. No phalanges are preserved.

The species is represented by other specimens, all of which are unsatisfactory in determining the structure of the form. A skull, apparently complete, is crushed out flat, so that little can be said of structure. The teeth are rather long, straight, acute, and striate at the base. The orbits are long and narrowed in front.

Measurements of Sauropleura newberryi Cope.
(No. 8612 G, and two unnumbered specimens,
American Museum of Natural History.)
mm.
Length of humerus 35
Proximal width of humerus 8
Distal width of humerus 14
Length of ulna 19
Proximal width of ulna 8
Posterior width of skull 67
Width at anterior angle of orbits 40
Interorbital space 13
Width of orbit 13
Length of orbit 27
Length of a maxillary tooth 4
Diameter of same at base 2

Sauropleura pauciradiata Cope.

Cope, Trans. Am. Phil. Soc., XV, p. 275, 1874.

Cope, Geol. Surv. Ohio, II, pt. II, p. 408, pl. xl, figs. 1-2, 1875.

Type: Specimen No. 8671 G, and obverse, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

The species was founded on a median and a lateral plate of 2 individuals. The clavicle is here associated with S. scutellata and the interclavicle thus remains as the type specimen. There are other remains associated by Cope with S. scutellata which are here shown to have closer affinities with the S. pauciradiata.

The clavicle is a right-angled triangle, the inner and thin edge concave posteriorly, the posterior convex. The ridges and grooves are well developed on the specimen. The visceral side of the element is smooth.

Cope referred to S. scutellata the larger part of an individual with strongly developed ventral scutellation. This he figured on plate XXXVI, fig. 2 (123). Even a cursory glance will, however, suffice to show that the sculpturing of the clavicle just described and that exhibited by the specimen there figured are identical. Closer examination shows important differences between the species, the principal distinctions being the strongly developed ventral scutellÆ in Sauropleura scutellata, the difference in form and sculpture of the interclavicles, and the posterior extension of the interclavicle in S. scutellata which is wanting in S. pauciradiata. The sculpturing of the interclavicle in the form figured by Cope as S. scutellata, just referred to, is identical with the sculpture of the clavicles, as would be expected. The ridges on all of the pectoral elements of S. pauciradiata are strong and are rather few in number, while in S. scutellata the sculpturing is more in the form of interrupted grooves.

The specimen (plate 14, fig. 3) exhibits the great part of the body with one fore limb. The skull is wanting. The belly was very broad and strongly protected by broad, long scutes arranged en chevron. The scutes are close together and form a compact ventral armor for the animal. The fore limb is very weak. The humerus is represented by its distal end only. The radius and ulna are very short and weakly developed in comparison to the size of the animal. The limbs could not have supported the animal on land and served, probably, merely as organs of equilibration, for the animal was undoubtedly aquatic. The fingers are not all preserved and there is no carpus.

Fig. 34.
A. Interclavicle of Sauropleura pauciradiata Cope. × 1. (After Cope.)
B. Left clavicle of Sauropleura pauciradiata Cope. × 1. (After Cope.)

Numerous other remains formerly associated with S. scutellata are here referred to S. pauciradiata, on account of the strongly developed ventral armor, which is different from that of the type of S. scutellata. The remains do not, however, add to our knowledge of the anatomy of the forms, as they are very fragmentary.

Two skulls are provisionally associated with this species. One of these skulls is figured by Cope (123) on plate XXXIII, fig. 1. It is there referred to S. scutellata. That it can not, however, be referred to that species is manifest when the teeth are observed. In the type of S. scutellata the teeth are very small, sharp denticles, while in the skull under discussion the teeth are well developed and their bases are longitudinally grooved. The teeth are elongate in the anterior part of the skull and are shorter posteriorly. They are, however, all strong. The skull is acuminate and the orbit is located about midway of its length. The jaw is slightly longer than the cranium. The structure of the cranium can not be determined in either skull, and in one only the position of the orbits and the teeth.

Measurements of the Specimens of Sauropleura pauciradiata Cope.

mm.
Length of clavicle, left 43
Width of clavicle 25
Space between ridges 2
Large specimen No. 8657 G and obverse (13)
described us Colosteus scutellatus Newb.

mm.
Length of specimen, as preserved 117
Width of specimen, maximum 61
Length of forearm 9
Length of hand 10
Length of metacarpus 5
Length of clavicle 4
Width of clavicle 18
Length of interclavicle 40
Width of interclavicle 23
Width of a ventral scute 1.5
Length of scute from angle to end 35
Skull No. 8666 C, plate xxiii, fig. 1 (Cope, 123), Colosteus scutellatus.
mm.
Length of skull, as preserved 65
Width of skull 40
Diameter of orbit 6
Length of mandible 7
Width of mandible, maximum 10
Length of longest tooth 5
Width of tooth at base 1.5
Skull Nos. 8602 G and 8608 G.
mm.
Length of skull, as preserved 50
Width of skull 60
Diameter of orbit 15
Length of mandibular tooth 4.5

Other specimens associated with this species: Nos. 8668 G, 86746, 85540, 8661 G. All specimens in the American Museum and all from Linton, Ohio.

Scutes of Sauropleura.

There are found associated with the remains of the genus Sauropleura a number of heavy scutes or scales. There are three of them on the same block as the specimen of S. longidentata and are provisionally referred to the genus. There are a number of scutes preserved separately, but they agree in their characters with those discovered on the specimens. The scutes are elongate and usually acuminate at one end and having a broad base at the other. The acuminate end is slightly bent to one side so as to present the appearance of a hook. Others are shield-shaped and are quite large, while the majority of the hook-shaped ones are small. The shield-shaped elements have a rounded boss near the center of the plate and the edges are imbricated. Their nature and their proper location on the animal are a puzzle. They may not belong to the genus, but have been noticed with the remains of at least 3 species.

Measurements (in Millimeters) of Scutes Associated
with the Specimens of Sauropleura.


No. 4513 U. S. National Museum:
Length 27
Width, maximum 14
Width, minimum 3
Scutes associated with the specimen of Sauropleura longidentata:
Width across base 3
Width across tip 1

Additional specimens are: Nos. 3, 8673 G and 8470 G, American Museum of Natural History.

Sauropleura longidentata Moodie.

Moodie, Jour. Geol., 17, No. I, pp. 74-76, figs. 18, 19, 1909.

Type: Specimen No. 8619 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

MOODIE
PLATE 22

1. Type of Leptophractus lineolatus Cope, from the Coal Measures of Linton, Ohio. Portions of maxilla and mandible of left side with teeth. × 1. Original in American Museum of Natural History.

2. Type of Proterpeton gurleyi Moodie, from the Coal Measures of Illinois, near Danville. Cervical of an otherwise unknown vertebrate. Neural spine to the right. Original in Walker Museum, University of Chicago × 2.

3. Amphibian phalanx from the Coal Measures near Breeze, Illinois, of an unknown species. The probable form of the element is represented. × 3.

4. Large rib of a stereospondylous stegocephalan otherwise unknown. The rib may represent a species of Macrerpeton or may even belong with Macrerpeton deani Moodie, but exact identification will have to wait for future discoveries. From the Coal Measures of Linton, Ohio. Original in U. S. National Museum. × 1.

5. Type of Cope's species Tuditanus mordax referred by him to the cranium, on account of the sculpturing of the elements. We now know the specimen to be portions of the interclavicle and clavicles of Diceratosaurus punctolineatus, as Cope suggested they might be. Original in American Museum of Natural History. × 1.

6. Skull of Baphetes planiceps Owen from the Coal Measures of Nova Scotia. × 0.45. Original in the British Museum. South Kensington. After Owen.

This species may be distinguished from other members of the genus by the large size and shape of the cranium (462) and the broad mandible (plate 16, figs. 2, 3) with its very long teeth. The skull of Sauropleura digitata Cope is not known, but the body of that animal as preserved represents far too small a form for the skull to be referred to that species. The skull is fully half as long as the dorsal region of S. digitata Cope, so that an association of the remains would be incongruous. It differs from the skull of S. scutellata Newberry in size and proportions. The skull of S. scutellata is narrow, while in S. longidentata it is quite broad. The teeth of the latter are also characteristic of the species, since in all other known species of this genus in which the skull is preserved the large anterior tooth is wanting.

The bones of the skull show the coarse sculpturing of the larger species of Microsauria. It consists more of radiating grooves than of pits. The skull, as restored (462), is broadly ovate, with the posterior border truncate. The muzzle is broad and the nostrils are, apparently, located near the anterior margin.

The posterior border of the orbits lies near the median transverse line of the skull. They are circular and are removed some distance from the margin of the cranium. Only the frontal and parietal can be determined with certainty.

The mandible is heavy and is provided with pleurodont, heterodont teeth. Near the anterior end of the mandible there is a very long fang-like tooth, longitudinally striated and slightly recurved, which arises from a broad base and attains to considerable prominence. The other teeth are smaller, though the next succeeding one is still of considerable size. All of the teeth preserved are longitudinally striated, but only the two anterior ones are recurved to any extent.

Measurements of the Type.

mm.
Length of the skull in median line 75
Width of skull at posterior border, estimated 80
Width of skull across orbits 60
Width of orbit 10.6
Length of orbit 12
Interorbital space 16
Length of jaw, as preserved 48
Width of jaw, maximum 16
Width of jaw, minimum 5
Length of largest tooth 11
Width of longest tooth at base 4.5
Length of shortest tooth 3
Width of shortest tooth at base 1

Sauropleura foveata Cope.

Cope, Trans. Am. Phil. Soc., XIV, p. 24, 1869.

Cope, Geol. Surv. Ohio, II, pt. II, p. 406, pl. xxxvi, fig. I, 1875.

Type: Specimen No. 8676 G and obverse No. 8675 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

This species is represented by an interclavicle and its impression, which show a beautiful sculpturing entirely distinct from that of the other species of this genus. In size it is intermediate between the largest of the interclavicles of S. scutellata and S. pauciradiata. The pattern of the sculpturing is, however, its main distinction. The plate is finely pitted and there are few evidences of grooves. Near the posterior border of the plate the pits become somewhat defined by ridges which take on a radiating pattern with the center of the plate as the center. The beveled margins are rugose, except at the edges.

Measurements of the Type Specimen of Sauropleura foveata Cope.

mm.
Median length of interclavicle 43
Width of interclavicle, maximum 23

Sauropleura enchodus Cope.

Cope, Proc. Am. Phil. Soc., p. 406, 1885 (Anisodexis).

Type: Specimen No. (51) 2558, American Museum of Natural History, Newberry Collection.

Horizon and locality: Discovered by Sam Huston at the Linton, Ohio, Coal Measures.

An examination of the type specimen (plate 16, fig. 4) of this form resulted in no new facts. The reference of the species to the Permian genus Anisodexis by Cope is probably incorrect. Our knowledge of the two known species of this genus is not sufficient to separate them, but for the sake of convenience the Linton species is placed in the genus Sauropleura. It is located here principally on account of the form and structure of the teeth. Cope's original description is given below:

"The generic characters are apparent in the very unequally sized teeth with round section. The portion upon which the species is based is a part of the right ramus of the mandible, which is in the specimen viewed from the inner side. The jaw is obliquely and smoothly truncated from below, for the symphysis and surface of the bone is smooth. There is a very large tooth near the extremity of the dentary bone. Behind it is an interval equal to three times the diameter of its base, which is followed by a tooth of about one-third the length of the first tooth. Posterior to this one are two teeth of the same size as the second, all being separated from each other by about a tooth's diameter. These are followed by three subequal teeth of about two-thirds the length of the first tooth, and separated by about their own diameter from each other. They are all perfectly straight, very acute, and without the trace of a cutting-edge. The inflection-grooves extend to or a little beyond the middle of the length."

The present species is smaller than the type of Anisodexis imbrecarius from the Texas Permian, to which genus Cope originally referred the present species, and the apices of the teeth do not display the opposite cutting-edges seen in the Texas form.

Measurements of the Type Specimen.

mm.
Length of jaw, including 7 teeth 31
Depth of ramus at second tooth 10
Length of first tooth 10.5
Length of third tooth 3.5
Length of sixth tooth 7.5

Skin of Sauropleura Sp.

Moodie, Bull. Am. Mus. Nat. Hist., XXVI, p. 355, pl. lx, fig. 1, 1909.

The present specimen is interesting on account of the presence of what I take to be a portion of the skin, which is preserved as a smooth mold over the ribs and ventral scutellÆ. The skin was undoubtedly that of the back, since the creature is preserved on its belly, and is interesting in not showing the slightest trace of scales or other hard plates. The ventral scutellÆ are characteristic of the species of the genus Sauropleura. With one species of this genus, Sauropleura scutellata Newberry, the writer has found associated scutes of some size, and the same fact has been noted by Cope.

Genus SAURERPETON Moodie, 1909.

Moodie, Jour. Geol., XVII, No. 1, p. 80, fig. 23, 1909.

Type: Saurerpeton latithorax Cope.

This generic name is erected for the reception of a single species described by Cope in 1897. The name is made necessary by the wide divergence of the characters exhibited by the present species from those of the species of the genus (Sauropleura) to which Cope (176) referred this species. The form is not a member of the genus Sauropleura, for reasons given below.

The species of the genus Sauropleura have a lanceolate head with homodont dentition or nearly so. The orbits are located well back in the skull. The form of the body is elongate and slender and the limbs where known are long and attenuated. The ventral scutellation consists of oat-shaped scutes arranged in a chevron series. The form here described as Saurerpeton latithorax Cope has nearly the opposite of all of these characters, and it is incongruous to locate the form under the former genus. The skull of Saurerpeton latithorax Cope is broad and heavy. The teeth are heterodont. The body is broad and stout and the limbs are of unusually strong proportions. The character of the ventral armature is also of a very different type. In Saurerpeton it consists of very broad imbricating scutes which form a single piece across the abdomen and are angulated to form the chevron pattern which is so common among the Stegocephalia.

Saurerpeton latithorax Cope.

Cope, Proc. Am. Phil. Soc., XXXVI, p. 86, pl. iii, fig. 4, 1897.

Moodie, Jour. Geol., XVII, No. 1, p. 80, fig. 23, 1909.

Type: Specimen No. 4471, U. S. National Museum.

Horizon and locality: Linton, Ohio, Coal Measures. Collected by R. D. Lacoe.

This species is indicated by remains of the anterior half of a large amphibian (plate 17) preserved on a block of bituminous coal from the Linton mine. The form is unusual in the proportions of the head and the width of the thoracic region. In these characters it stands alone among the Amphibia from this locality, where the forms are for the most part of rather slender build and tapering, pointed head.

The skull is represented in a fairly complete condition and shows the usual stegocephalian arrangement of the skull elements, as well as the sculpturing of the bones, which is similar to that found in other members of the Microsauria. The skull is broadly rounded, with the posterior border incised, the broad tympanic notches thus rendering the shape of the skull somewhat like that of the branchiosaurs. The orbits are broad ovals and lie well forward in the skull. They are separated by a space which is greater than the greatest diameter of the orbit. The pineal foramen lies well back and is clearly indicated as a circular opening which lies in the median suture in the posterior half of the parietals. The nostrils seem to be elongate and have an oblique position, as is represented in the diagram (fig. 35), but this character is not ascertained definitely.

The borders of the premaxillÆ and the anterior suture of the nasal can not be determined, though they may have had some such arrangement as suggested. The nasal is represented, so far as is determinable, by an oblong element lying between the frontal and the anterior border of the skull. The f rentals are very large and form a portion of the inner border of the orbit. The parietal is probably the largest bone in the cranium and together the two elements form a large quadrangular space in the posterior half of the skull. They inclose the circular pineal foramen. The postparietal is a small element lying on the posterior border of the skull and with the tabulare and a part of the supratemporal forms the projection. The prefrontal is probably a small element, especially if the lacrimal is present. There seems t o be an indication of a suture separating the lacrimal from the prefrontal, but this character is not assured. The boundaries of the maxillÆ are clear anteriorly. They indicate that this element was elongate, as is usual, and impressions of teeth borne on the mandible would indicate the probability that the teeth were heterodont. In the premaxillary region there is a long, strong tooth preserved, and on the maxillary near the posterior extremity of this bone there are impressions of teeth which are no more than one-fourth as large as the premaxillary one. The borders of the jugal can not be ascertained, since the skull is injured on both sides in this region. Likewise the quadratojugal is conjectural. The boundaries of the postfrontal and the postorbital are clearly ascertained. They together form the entire posterior border of the orbit and send prolongations along the lateral borders of these openings. They are both acuminate posteriorly and these points are inclosed by the tabulare and supratemporal for the postfrontal and by the squamosal and prosquamosal space for the postorbital. The boundaries of the tabulare show this element to be quite large and extending forward into an acumination which is inclosed by the parietal and the postfrontal. The sutures bounding the squamosal have been obscured by injury in removing the specimen and are indeterminable.

Fig. 35.—Outline drawing of the skull and skeleton, as preserved, of Saurerpeton latithorax (Cope), from the Linton, Ohio, Coal Measures. Original in U. S. National Museum. × 0.75.
Skull: fr, frontal; j, jugal; mx, maxilla; n, nasal; no, nostril; or, orbit; par, parietal; pof, postfrontal; po, postorbital; pp, postparietal; pf, prefrontal; pmx, premaxilla; qj, quadratojugal; m, mandible; spa, supraorbital lateral-line canal; spt, supratemporal; sq, squamosal; tab, tabulare; y, pineal foramen.
Skeleton: cl, clavicle: c, carpus; rb, ribs; h, humerus; ic, interclavicle; met, metacarpals; psc, pectoral scutellÆ; r, radius; vs, ventral scutellÆ; u, ulna.

The vertebral column is represented by a ridge showing along the median plane of the specimen. Its characters can not be ascertained. There are a few evidences of ribs. They represent long, slender, non-alate elements, their mode of articulation with the vertebrÆ being obscured by the plate-like ventral armature which covered the abdomen of the animal.

The pectoral girdle is represented by the remains of three elements which are interpreted as being the interclavicle and the two clavicles. The interclavicle is broad and is rounded posteriorly. There is no evidence of the usual acumination posteriorly. The element is nearly as wide as long. There is a prominent longit udinal keel on the ventral surface of the interclavicle and radiating lines which may indicate the courses of blood-vessels or nerves or may be the ornamentations of the element, probably the latter. The clavicle has the usual microsaurian form. It has three points and is truncate exteriorly. It is ornamented with radiating grooves of a shallow and not strongly pronounced character. There is no evidence of the coarse sculpture of the later forms. If the scapula is represented it is merely by an indeterminate fragment insufficient for description.

The pectoral limbs are preserved nearly entire. The left fore limb lacks only a few phalangeal bones, and these were preserved with the remainder of the skeleton but were lost in the mining process. The humerus is an extraordinary element on account of its robust dimensions. It is very stoutly built and represents a powerful limb. It is expanded at each extremity and the width of its shaft is about equal to one-fourth of its length. The ulna and radius present the same characters as the humerus, i.e., in being robust, with stout shaft and expanded ends. The ulna is slightly longer than the radius and has an expanded upper end. The radius is short and does not have the proximal expansion. The carpus was cartilaginous. Its position is represented by a blank space on the coal. There are 4 digits preserved and in all probability this was the entire number. The metacarpals are elongate and expanded at the extremities. The first and second digits are represented nearly complete. The first digit is extremely interesting in the possession of a claw-like terminal phalanx which much resembles that of some lizards. There are 3 phalanges in the first digit and 4 in the second. The phalangeal formula may have been 3-4-4?.

The ventral scutellation of this species is of an unusual character. It consists of broad, imbricated scutes which are in a single piece and which are arranged in the usual chevron pattern. The scutes were, apparently, broadest in the middle and tapered somewhat at the extremities. This character alone is sufficient for separating the genus from that of any other known form.

The genus finds its nearest allies in the forms of the species of the genus Sauropleura, in which Cope formerly located the present species. The skull of the form described as Tuditanus radiatus Cope is quite similar to the present form, both in the sculpturing and arrangement of the elements. The characters wherein the present form resembles the species of Sauropleura are the possession of broad pectoral plates and strong, digitate limbs. The general form of the body and skull is different in the two groups. It is slender in Sauropleura and decidedly stout, short, and heavy in Saurerpeton.

Measurements of the Type.

mm.
Length of specimen 130
Median length of skull 51
Width of skull at posterior border 62
Width of skull across orbits 48
Length of orbit 14
Width of orbit 10
Interorbital space 16
Length of longest tooth preserve 14
Length of shortest tooth preserved 1
Length of the interclavicle 26
Width of interclavicle, maximum 23
Length of clavicle 22
Width of clavicle 18
Length of abdominal scutes 28
Length of humerus 19
Width of humerus at upper end 6.5
Length of ulna 11
Length of radius 9
Length of metacarpal 3
Length of first digit 12
Length of terminal phalanx 3.5
Length of lower jaw on the curve 70
Width of lower jaw, maximum 8
Length of rib 30

Genus CTENERPETON Cope, 1897.

Cope, Proc. Am. Phil. Soc., XXXVI, p. 83, pl. iii, fig. 1, 1897.

Type: Ctenerpeton alveolatum Cope.

The genus Ctenerpeton was founded by Cope for the reception of a single species which presents some very unusual characters. The form shows close relationships to the genera Urocordylus (334), Œstocephalus, and Ptyonius. These genera agree with the present one in the possession of very characteristic vertebrÆ which are signalized by the elongate and ornamented characters of the neural and hÆmal spines. These project prominently from the body of the vertebra and have the ends of the projections truncate and divided into fine points, thus causing the spine to have much the appearance of a comb. The surface of the neural spine is sometimes marked with a shallow groove. The spines are longer and more slender in the genera Œstocephalus, Urocordylus, and Ctenerpeton than they are in the species of the genus Ptyonius, where they are short, although the usual pectinations are present.

The character on which this genus rests is the shelf-like extension (plate 23, fig. 2) of the abdominal plates. This is of a very unusual character and entirely unknown in any other species of Carboniferous Amphibia. The term Ctenerpeton has reference to the fact that the ends of these shelf-like plates project in a pectination along the side of the abdomen.

Ctenerpeton alveolatum Cope.

Cope, Proc. Am. Phil. Soc., XXXVI, p. 83, pl. iii, fig. 1, 1897.

Moodie, Proc. U. S. Nat. Mus., 37, p. 24, pl. 10, 1909.

Type: Specimen No. 4475, U. S. National Museum, Lacoe Collection.

Horizon and locality: Linton, Ohio, Coal Measures.

The species rests on a single specimen (plate 19) from Linton, Ohio, and is preserved on a block of bituminous coal. It is in a very good state of preservation. There are present on the block of coal a part of the right fore limb, the greater part of the dorsal portion of the animal, and the anterior part of the tail. There are no evidences of hind limbs, although this may not be taken as evidence that they were not present on the animal. No thoracic plates have been observed. The chevron armature is present beyond the cloacal region, but there are no evidences of the specialized clasping organs which are apparently developed from the abdominal armature in some forms (251).

Each dermosseous rod of the abdominal scutellation consists of three pieces a median angulated portion and the two lateral parts which form the shelf-like projection along the side of the abdomen (plate 23, fig. 2). The marginal chevron differs in form from the other plate, aside from the fact that it is not angulated. The lateral shelf is composed of flattened plates which articulate with the median piece, and at the place of articulation there is a ridge present in the specimen. The exterior plates are curved backwards and are somewhat attenuated distally. They are broader than the median piece and differ also in the absence of the characteristic alveoli. The median plate is angulated and is of more slender proportions than the lateral pieces. Its ventral surface is ornamented with a single row of closely placed alveoli which resemble in a great degree the alveoli of the jaw of some small animal. The ventral scutellation is broad anteriorly, but becomes more slender posteriorly and shortly posterior to the cloacal region disappears.

MOODIE
PLATE 23

1. Left leg and pelvis of Ichthycanthus platypus Cope, from the Coal Measures of Ohio. × 1.7. Original the property of the Department of Geology of Columbia University. c=centrale; F=fibula; fi=fibulare; Fe=femur; i=intermedium; Il=ilium; T=tibia; t, 1-5=distal tarsalia; ti=tibiale; V=caudal vertebrÆ; I-V=digits.

2. Ventral scutellÆ of Ctenerpeton alveolatum Cope, from the Coal Measures of Ohio. × 3.5. Original in U. S. National Museum.

The fore limb is represented by the upper portions of the ulna and radius and 2 digits of the right hand. The digits are long and slender and seem to represent digits I and II, since they show evidences of 3 and 4 phalanges respectively. The portions of the forearm preserved are too meager for description.

The vertebrÆ have already been characterized as of the type first described in Urocordylus. The neural fans are not much, if any, wider than the hÆmal fans. They are both situated on an elongate spine with a slender base. The edges of the two fans are pectinated and the dorsal spine is distinguished by the presence of a longitudinal groove in the center of the spine. The length of the tail may have been considerable, judging from the character of the vertebrÆ preserved

Measurements.

mm.
Length of specimen 150
Width of belly, maximum 28
Length of lateral chevron plate 7
Width of lateral chevron plate 1.75
Length from tip of lateral chevron to median angle 10
Length of a caudal vertebra 10
Width of the same vertebra with spines 20
Height of neural spine 8
Height of hÆmal spine 8
Width at distal end of hÆmal spine
Width at distal end of neural spine 3.5
Length of foot, as preserved 12

Genus LEPTOPHRACTUS Cope, 1873.

Cope, Proc. Phila. Acad. Nat. Sci., p. 340, 1873.

Cope, Geol. Surv. Ohio, II, pt. II, p. 399, 1875.

Cope, Proc. Am. Phil. Soc., XX, p. 461, 1882.

Type: Leptophractus obsoletus Cope.

The genus was established on various parts of the cranium of a large amphibian. The only parts which can with certainty be referred to the genus are the upper and lower jaws of 3 specimens. These bear large teeth, round in section at the base, but with acute compressed apex, with a cutting-edge on the anterior face; the enamel is delicately grooved, as an external indication of the labyrinthic structure. A characteristic feature is seen in the presence of a large elongate tooth in the upper jaw, in the position of a canine which much exceeds in length any of the others. The sculpture of the cranium is but little marked in the known specimens. In the type the lower jaw is marked with inosculating grooves. Three species are known, which are among the largest of the Linton Amphibia.

Leptophractus obsoletus Cope.

Cope, Proc. Phila. Acad. Nat. Sci., p. 340, 1873.

Cope, Geol. Surv. Ohio, II, pt. II, p. 399, 1875.

Cope, Proc. Am. Phil. Soc., XX, p. 461, 1882.

Type: Specimen Nos. 55 G and 57 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

The species was about as large as an adult Florida alligator, and probably exceeded or at least equaled in size any of the Carboniferous Amphibia. The following account is taken directly from Professor Cope's "Batrachia of the Ohio Coal Measures" (123). The description has been verified from an examination of the type material.

"The teeth are rather distantly grooved for some distance above the base. They are of different sizes; the smaller are compressed, and with fore and aft cutting edges. The external surface of the dentary bone is marked with short oblique grooves along its middle region; above these are grooves which inosculate, forming a figure like an open net dragged in the long direction.

"Excepting the grooves the teeth are smooth. The smaller ones are close together and their crowns are curved backwards; the larger ones are at more remote intervals; both have enlarged bases. Whether both forms are in the same series I can not determine. There are from four to five of the smaller to an inch.

Measurements of Type of Leptophractus obsoletus Cope.

mm.
Depth of fragment of jaw 75
Length of smaller teeth 19
Length of longer tooth 23
Width of vertex at middle scuta 176
Width of paired median scuta 56
Width of single scute 36
Length of single scute 48

"Some vertebrÆ were found at the same locality, but there is no evidence as to the species to which they may have pertained. They are short, concave on one end, and probably so on the other. The centrum of one is 12 mm. in diameter; neural spines injured." (Geol. Surv. Ohio, II, pt. II, pl. 39, fig. 3.)

"A third and larger specimen was found by Professor Newberry during the field season of 1874. It includes an oblique view of one side, and the top of the cranium from the posterior part of the orbits to the end of the muzzle, with the corresponding part of the alveolar region of the dentary bone, with teeth. The bones of the skull appear to have been rather light, and though the surface is irregular, the sculpture consists only of shallow impressions of varying size and intervals. The orbits are also badly defined, but appear to have been large, and separated by a narrow frontal bone. The premaxillary bone is preserved, and shows clearly the sutures that separate it from its fellow and from the maxillary. A large foramen—perhaps the nostril—separates it from the maxillary, so that it forms an irregular crescent. It supports two teeth, of which the anterior is the larger, but there were perhaps others in advance, as the alveolar border is imperfect towards the end of the muzzle. The anterior two teeth of the maxillary bone are followed by a strong groove which rises towards the sides of the muzzle. At first sight this gives the impression of the maxillo-premaxillary suture, and makes it appear that both the premaxillary bones are preserved, and that the foramen above described separates the premaxillary spines, instead of representing the nostril. The cutting edges of the teeth of these bones have, however, one direction, whence they represent one side of the cranium only; were both sides represented, the directions of the tooth axes would be reversed.

"The premaxillary and maxillary teeth exhibit a cutting edge on the outer posterior margin of the distal half; the base of the crown is subround in section. The line-like grooves are distinct but not numerous, their intervals measuring 75 mm. Beyond them the enamel is smooth. The second maxillary tooth is larger than the first, which is equal to the last premaxillary. The third and fourth maxillaries are equal to the second, but the fifth is larger and longer, exceeding all the others. The teeth of the dentary bone differ from those of the upper jaw in having the cutting edge of the crown on the anterior aspect, while the posterior border is obtuse. There is an obtuse cutting edge on the posterior margin of the anterior mandibular teeth.

"This description is derived from an adult animal, as the maxillary teeth in some instances are partially worn away by friction on their anterior and outer faces."

Measurements.

mm.
Length of maxillary bone preserved 146
Length of same supporting five teeth 73
Length of first maxillary tooth 15
Diameter of same at base 6
Diameter of second at base 8
Length of basis of fifteen teeth of the dentary 145

Leptophractus dentatus new species.

Type: Specimen No. 10850, American Museum of Natural History. Collected by Dr. J. S. Newberry.

Horizon and locality: Linton, Ohio, Coal Measures.

Fig. 36. Mandible of Leptophractus dentatus new species, from the Linton, Ohio, Coal Measures. × 1. Original in American Museum of Natural History.

The type is a single right mandible, nearly entire, of a rather large animal. The specific characters for the separation of the new form from the previously described L. obsoletus and L. lineolatus are the smaller size and slenderness of the mandible, associated with uniform teeth, which are slender and delicately fluted.

There are 17 teeth preserved, the largest of which is 8 mm. in length. From the posterior tooth the series gradually descends to half this length 0.5 inch from the anterior end of the mandible.

The exact form of the mandible can not be determined, but so far as can be seen it is very slender, coming almost to a point at the anterior end. The posterior portion is wide, but apparently not very heavy. There is a fragment associated with the specimen which discloses a few teeth, but its position in the cranium can not be determined.

Measurements of the Type of Leptophractus dentatus Moodie.

mm.
Length of mandible, as preserved 80
Anterior width of mandible 3
Posterior width 28
Length of most anterior tooth 4
Length of tenth posterior tooth 7
Width of this tooth at base 3

Leptophractus lineolatus Cope.

Cope, Proc. Am. Phil. Soc., XVI, p. 576, 1877.

Type: Specimen No. 1088 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures. (Plate 22, fig. 1.)

Cope's description of this species, to which I have nothing to add, is as follows:

"This species is based on portions of the skull of two individuals of large size. Both upper and lower jaw with teeth are represented and the teeth are of very large size. The deepest element preserved has been provisionally referred to the mandible and will be so described. This element bears two types of teeth in very heterodont fashion. The teeth in the back portion of the jaw are rather short and slender. The teeth more anteriorly are, some of them, very long, rather stout, with their bases longitudinally fluted, as are they all v The longest tooth in the jaw measures slightly less than an inch. The bone is not well preserved, but seems to have been ornamented with grooves of no great depth. The most anterior teeth of the jaw are smaller than the posterior ones.

"The upper jaw is set with teeth which are more uniform in size and there is but little tendency to heterodonty. These teeth are also striated at their base and all end in a sharp point. They are all, apparently, straight. There is no tendency to curve as there is in the genus Macrerpeton. The upper teeth are more closely set than are those of the lower jaw, which are rather distantly placed.

"Another specimen of a smaller individual presents the same portions of the skeleton and the same characters. It is possible that this skull will be found to belong to Ichthycanthus ohiensis Cope, which is based on very large vertebrÆ and limb bones. The remains described as Leptophractus lineolatus are, however, unlike any other skull re mains which are thus far known.

"This species represents one of the largest types of the Carboniferous Amphibia of Ohio. It probably attained a length of several feet. It was also the most carnivorous of any of the forms."

Measurements of the Type Specimen of Leptophractus lineolatus Cope.

mm.
Length of specimen, as preserved 98
Depth of dentary bone at the middle 30
Anteroposterior diameter of mandibular tooth at base 3.5
Length of a posterior mandibular tooth 9
Length of longest tooth in jaw 22
Anteroposterior diameter of the same at base 6
Length of small maxillary tooth 7
Anteroposterior diameter of same at base 2

Collected by Dr. J. S. Newberry. Other specimens of the species are Nos. 1086 G and 1087 G, American Museum.

Eurythorax sublÆvis Cope.[C]

[C] Dr. Hussakof now regards (Bull. Amer. Mus. Nat. Hist., vol. 35, p. 128, 1916) the type specimen of this species as an opercular element of one of the dipnoan fishes [Sagenodus sublÆvis (Cope)]. The discussion is left here, however, as a matter of historical interest.

Cope, Proc. Am. Phil. Soc., p. 177, 1871.

Cope, Geol. Surv. Ohio, II, pt. II, p. 401, pl. xl, fig. 4, 1875.

Fig. 37.—So-called interclavicle of Eurythorax sublÆvis Cope. (Sagenodus.) No. 75. (After Cope.)

Type: Specimen No. 8605 G, American Museum of Natural History, Collection of Dr. J. S. Newberry.

Horizon and locality: Linton, Ohio, Coal Measures.

The genus and species were established on a single, large, almost perfect interclavicle of peculiar form. It may or may not be distinct, but it can not be associated definitely with any known genus, so that it will be designated as Cope described it. It exhibits on its outer or lateral borders broad, smooth surfaces for the contact of the overlapping margins of the lateral plates. The form is subround, with a large excavation from the posterior margin on each side. The narrowed portion left in the middle behind has a convex outline. Some delicate radiating grooves occur on the exposed surface, but they are very shallow.

Measurements of Type.

mm.
Length of interclavicle 71.5
Greatest width 78
Width of lateral concavity 39

                                                                                                                                                                                                                                                                                                           

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