CHAPTER IX.

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THE AMERICAN COAL MEASURES BRANCHIOSAURIDÆ.

Definition Op the Family BranchiosauridÆ Fritsch, 1879. Coal Measures and Permian Op North America and Europe.

Fritsch, Fauna der Gaskohle, Bd. I, p. 69, fig. 30, 1879.

Lydekker, Cat. Fossil Reptilia and Amphibia, pt. IV, p. 210, 1890 (ProtritonidÆ;).

Stegocephalic, salamander-like animals, with broad, anteriorly truncate skull. Teeth smooth with large pulp-cavity. The parasphenoid narrowed anteriorly, posteriorly expanded to a shield-shaped plate. Vertebra; with the notochord persistent and intravertebrally expanded. Pelvis well developed, the ilium and ischium osseous with large cartilaginous margins, the pubis unknown, possibly hyaline cartilage. Ribs short, straight, present on almost all vertebrÆ. Skin with delicately ornamented scales. Eyes with sclerotic plates. Palatal elements toothless or with small tooth-like tubercles on pterygoids and palatines. Ventral armature on throat, chest, and abdomen, extending on to the limbs, consisting of small delicate scutellÆ arranged in a chevron pattern.

The above definition is modified from Fritsch (Fauna der Gaskohle, Bd. I, p. 69, 1879).

The North American species are: (?) Sparodus sp. indet. Dawson, Micrerpeton caudatum Moodie, Mazonerpeton longicaudatum Moodie, Mazonerpeton costatum Moodie, Eumicrerpeton parvum Moodie.

Genus MICRERPETON Moodie.

Moodie, Jour. Geol., 17, p. 39, figs. 1 to 6, 1909.

Type Micrerpeton caudatum Moodie.

The genus Micrerpeton, of which the single species is described below, was the first evidence of the occurrence of the Branchiosauria in America. There have been three other genera referred to the Branchiosauria from North American deposits, but there is good evidence that none of them belong there. The genus Amphibamus was originally referred to the Xenorhachia by Cope (105, pp. 134-137) on account of the supposed cartilaginous condition of the vertebrÆ and the absence of ribs. Later he abandoned this order and placed the form in the Branchiosauria, where it was retained by Zittel (642). Recently Hay has shown (316), and I am able to corroborate his statement, that ribs are present in the species, and that they are long and curved, not at all like the short ribs of the true Branchiosauria. These long, curved ribs undoubtedly exclude Amphibamus from the Branchiosauria and indicate its close affinities with the Microsauria. The genus Pelion has also been referred to this order on purely negative evidence (642, p. 375). This genus is excluded from the Branchiosauria by the well-ossified condition of the limb bones, in which the endochondral ossification is seen to be well developed, a condition not found, so far, among the true Branchiosauria. The form of the head and the elongate hind limb would also tend to exclude the genus from the Branchiosauria. In the Branchiosauria the fore limb is usually larger than the hind limb, the reverse of which is the case in Pelion. The genus Sparodus, as it occurs in North America, is uncertain. It is indicated by remains which are almost impossible of determination.

The genus Micrerpeton may be distinguished from other known Branchiosauria by the large size and anterior position of the orbits, absence of a posterior table to the skull, the short, heavy limb bones, the slender ilium, and the expanded, elongate, and laterally compressed tail. The genus may be defined as follows: Small forms, the known representative attaining a length of less than 2 inches; head broad and short; sclerotic plates present; interorbital space less than the least diameter of the orbit; occiput concave; pineal foramen in the line which cuts the posterior edge of the orbits; teeth small, pleurodont denticles; presacral vertebrÆ 20 or 21, of which probably 5 are cervical; 1 sacral; ribs short, straight, and heavy, central; scapula ovoid; limbs stumpy and heavy, fore limb exceeding the hind in size; endochondral ossifications distinctly absent; tail long, expanded, and flattened, probably provided with a thin expanded membrane; body covered with minute, ovoid or rounded scales which are ornamented with concentric lines; color markings vertical to the long axis of the body and abundantly present on the tail; lateral-line organs represented by the dorsal and median lateral lines on the tail, the sensory pits probably occurring in specialized darkened scales. Coal Measures of Mazon Creek shales near Morris, Grundy County, Illinois.

Micrerpeton caudatum Moodie.

Moodie, Jour. Geol., 17, p. 39, figs. 1-6, 1909.

Type: Specimen No. 12,313, Walker Museum, University of Chicago.

Horizon and locality: Mazon Creek shales, near Morris, Illinois.

The species is represented by very complete remains (plate 2), which are preserved on opposite halves of a nodule. The specimen was collected many years ago by Mr. W. F. E. Gurley at Mazon Creek, but it has never before been studied, although Dr. Newberry examined it and said in a note that Professor Cope should see it. Unfortunately Cope did not see it and it lay unknown for more than a quarter of a century. I am indebted to Dr. Stuart Weller for calling my attention to the specimen, as well as for the privilege of describing it.

The specimen is exceptionally perfect (plate 25, fig. 4). Nearly all the skeletal elements are present, and the general contour of the body, the character of the dermal covering, the color-markings, the lateral-line system, and many other features of interest have been detected. Such completeness of preservation is very uncommon even among the remains obtained from this locality. In this case the entire form was preserved, but the collector, in cracking the nodule, lost the chips containing the feet, so that only portions of the limbs remain. It is thus impossible to determine the phalangeal formula, but the feet were probably like those of Branchiosaurus amblystomus Credner, as given by Credner, to which species the present form is closely allied and indeed must be placed in the same family with Branchiosaurus, Pelosaurus, and Melanerpeton.

MOODIE
PLATE 2

Drawing × 3.5. of type specimen of Micrerpeton caudatum Moodie, from the Coal Measures of Mazon Creek, Illinois, showing skeletal elements, form of head and tail, the lateral-line sense-organs, banded color-markings, and ventral scutellÆ. On the edges of the tail impression are indications that in life the tail had a thin fold of skin above and below the fleshy portion, much as in the larvÆ of Amblystoma at the present day.

The remains here described represent a small, salamander-like form, and they are among the earliest geological evidence of the group, which, without doubt, gave rise to the modern salamanders. The parts preserved in the specimen are: the complete outline of the head with most of the cranial elements clearly distinguishable, as well as the black pigment of the choroid; the entire vertebral column, including pits in the tail region, where the vertebrÆ were without doubt entirely cartilaginous; parts of the pectoral girdle; the ilium; the left humerus; the ventral scutellation; the ribs of one side of the body and indications of ribs on the other; portions of both hind limbs; and a complete impression of the fleshy tail. On this impression of the tail are preserved small, horny scales, transverse color-markings, and the distinct impressions of the lateral-line system.

The bones of Micrerpeton caudatum, as in so many of the fossils from this locality, have been replaced by a white, friable mineral which is probably kaolin. The animal is preserved on its back and it is thus illustrated from the ventral side. The entire length of the animal is only 49 mm., of which the tail occupies nearly half.

Fig. 13. Restoration of Micrerpeton caudatum, a branchiosaur from the Coal Measures of Illinois. × 2.

The head has much the same shape as in the species of Branchiosaurus described and figured by Fritsch (251), Credner (181), and Thevenin (568). The eyes occupy relatively the same position as in that genus. The orbits are very large and broadly oval. Within the borders of the rim the stone is blackened as though by the black pigment of the iris, such as Cope has described in Amphibamus. Under a rather high power of magnification the cranial bones are seen to be represented by mere flakes of white mineral matter. The sutures separating the cranial elements are distinctly preserved on the main half of the nodule.

The openings of the skull are five the two orbits, the two minute nostrils, and the pineal foramen. A median suture separates the skull into halves and the pineal foramen lies slightly anterior to the posterior third of its length. The boundaries of the premaxillÆ are not distinct, but they are very small elements and form the inner border of the nostrils, which are clearly indicated by bosses of stone. The nasal element is nearly square and lies anterior to the frontal, which it borders broadly. The parietal is about the same size as the frontal and it apparently forms a portion of the inner border of the orbit, although this is not assured. The parietal is elongate and unites posteriorly with the postparietal. The postparietal, with the tabulare and the squamosal, form the posterior boundary of the skull, and they are hence not unlike the same elements in other Stegocephala. The pref rental forms the anterior border of the orbit. The lacrimal has not been detected. The maxilla is elongate and forms the antero-lateral border of the skull. The jugal forms an important element in the lateral border of the cranium and joins the quadratojugal posteriorly. The postfrontal is triangular and with the postorbital forms the posterior border of the orbit. Both of the elements are acuminate posteriorly, although the suture between them is indistinct, and they inclose between their posterior acuminations an anterior projection of the supratemporal. The squamosal has the usual relations and borders the supratemporal laterally. The latter element forms the quadrate angle of the cranium.

The entire length of the vertebral column is preserved, although the nature and structure of its elements can not be determined. The impressions of a few of the vertebrÆ show that some of the centra were amphicoelous, but other than this nothing is definite. The cavities which the centra occupied were filled by the white mineral matter and the force of the blow which cracked the nodule destroyed the form of the mold. It is possible that where the mineral matter has filled the cavities the centra were osseous or partly cartilaginous, and where the cavities were unfilled the centra were entirely cartilaginous. The length of the vertebral column from the base of the skull to the last impression of a cartilaginous centrum is 33 mm.

The number of centra between the sacral vertebra and the skull is 20 as they are preserved, but there may have been one more, the atlas. Fritsch has represented 21 in his restoration of Branchiosaurus salamandroides, and this is a further indication of an affinity between the two genera, although Credner has represented 26 presacral vertebra; in Branchiosaurus amblystomus. The presacral vertebrÆ are thus seen to vary within narrow limits, but the number of presacrals is near 20, and this may be taken as typical. It is interesting to notice that in modern forms of the salamanders the presacral vertebrÆ number about 20. There is but a single sacral centrum in Micrerpeton. The sacral rib has not been detected, but it is restored after the condition found in Branchiosaurus. The right femur partially covers the sacral vertebra, and its structure can not be determined. I count impressions of 17 caudal centra, of which at least 12 may have been partially ossified. In the cervical region there are distinct impressions of transverse processes on at least 5 vertebrÆ, and this number is assigned to the neck, although it is by no means certain that this is the correct number. The neck was at least short, if we may judge from the position of the remains of the pectoral girdle. No cervical ribs are definitely determined. There is a short rib lying between the fifth and sixth vertebrÆ, but to which it belongs is uncertain.

There are impressions of 10 ribs preserved on one side of the vertebral column and one on the other side. They are short, straight, and heavy, as are the same elements in Branchiosaurus. This character alone is sufficient to place Micrerpeton among the Branchiosauria, since no such ribs are known in other groups of the extinct amphibians. The ribs preserved lie next the seventh to the seventeenth vertebrÆ on the left side, and there is one on the right side which may belong to either the fifth or sixth vertebra. They are central in their attachment, and in this they agree well with the mode of rib attachment in the modern salamanders. All of the ribs are single-headed and are composed, for the most part, of perichondral tissue. The position of the ribs in the matrix inclined backwards, and, making a small angle with the vertebral column, is very suggestive of the condition in Branchiosaurus.

The pectoral girdle is represented by three distinct elements of the left side, which are identified as scapula, clavicle, and coracoid, following the nomenclature given by Woodward (631), although Credner (186) would name them otherwise. The scapula is represented by an ovoid fragment lying next to the vertebral column. The clavicle was probably spatulate, as in Melanerpeton, but the inner end of the element is not visible. The coracoid is represented by its outer end only, and its inner pointed extremity is not visible. The interclavicle has not been detected.

The humerus lies somewhat to one side of the pectoral girdle, as if there had been a large amount of epiphyseal cartilage. Its position may be due to post-mortem shifting, but there is little other evidence of any movement after deposition. The humerus is a large, heavy bone in comparison with the rest of the skeleton. It is expanded at each end, and its ends show concavities, proving that the bone is formed principally of perichondral tissue, as would be expected from such an early Branchiosaurian. The endochondrium has not yet developed in this form, which is evidently adult. There is no other element of the arm present.

There is but a single element of the pelvis preserved, a slender elongate rod which is undoubtedly the ilium, since it has the usual position for that element and is much too large for a sacral rib. It has much the same shape as the ilium in the modern Salamandra, and is not expanded as is the ilium of Branchiosaurus. This element, like the humerus, seems to have been but a hollow cylinder of bone and undoubtedly had cartilaginous ends, as in the ilium of the recent Salamandra. The two femora are preserved nearly entire, the right one lying upon and partly obscuring the sacral vertebra. The femur is much more slender than is the humerus, with slightly expanded ends, and, like the humerus, shows the concavities at the ends, indicative of the perichondral character of the tissue composing it. There are two elements of the leg preserved more or less entire. The larger bone may represent the tibia and the smaller the fibula. They both present characters similar to those of the femur and humerus. They are simple rods of bone tapering at the distal end. The feet have been lost, though doubtless they were present at one time.

The ventral surface of the body, as in other members of the Branchiosauria, was covered and protected by a series of small scutes arranged in the regular chevron pattern. The form of the scutes and their number can not be determined. The lines which represent them are, however, distinct. Some of the scutes are missing and some of them are obscured by lying over the vertebral column. They are all somewhat shifted to the left. The lines are very small and close together. I count 16 of them in a distance of 2 mm. In length the longest line preserved is a little more than 4 mm., measuring from the point of the chevron. The lines representing the scutes come to a point in a median ridge which is now represented by a line. The dermal scutes on the abdomen were probably the forerunners of the abdominal ribs of the reptiles (fig. 9).

The impression of the tail contains some of the most interesting features in the entire specimen. Scattered over it and in places laid in mosaic are impressions of small dermal scales, which may have covered the entire body. In form the scales are ovoid, being half as wide as long, and the markings on the scales partake of the nature of radiating lines, much after the pattern of the sculpturing of the cranial bones in the Microsauria. The scales are less than 0.5 mm. in diameter and their character can only be ascertained under high magnification. Near the middle of the tail there are preserved distinct transverse bands of dark color, which are more or less evident throughout the entire tail impression, but they are elsewhere not so distinct as in the central region. The lines are evidently due to rows of pigmented scales, and in all probability the animal's body was transversely striped.

The most interesting and important single structure discovered on the specimen is the impression of the lateral-line system, which is clearly evident as two dark lines on the impression of the fleshy part of the tail. The sense-organs are represented by two longitudinal rows of pigmented scales, one beginning at the tip of the tail, the other taking its origin from the median line somewhat further forward. I am indebted to Dr. Katashi Takahashi for calling my attention to the similarity of this arrangement to that found in the recent Necturus. The arrangement and disposition of the lines containing the sense-organs is practically the same in the two forms. The median lateral-line takes its origin from the extreme tip of the tail and is continued to the base, where the impression is broken. The dorsal lateral-line has its origin rather abruptly from the median lateral-line at a distance of 6 mm. from the tip of the tail. The sense-organs were undoubtedly located beneath specialized pigmented scales on the surface, and to this pigment is due the preservation of the lines.

The fact that the arrangement of the sense-organs of Micrerpeton corresponds so exactly to the condition found in Necturus is of considerable interest. Necturus alone among the modern tailed Amphibia has the arrangement described for the lateral-line system of Micrerpeton. All other forms of the Caudata, as also the larval forms of the Salientia, have an arrangement of the lateral-line system which is perfectly distinct from that found in Necturus, although the basis of the same arrangement is found in all. In Amblystoma, for instance, the median lateral-line is not present on the tail, and the dorsal line is incompletely developed. The close similarity of the arrangement of the systems of sense-organs in the two forms, Micrerpeton and Necturus, may be of genetic significance with regard to the latter form. The lateral-line sense-organs are of a very fundamental significance, and it is not at all improbable that the same arrangement of the lines has existed from the Carboniferous period or earlier. We know that such has been the case in a great many of the fishes. The ancestors of the modern Caudata must have originated somewhere in the basal Carboniferous or earlier periods, and, in the writer's opinion, the Branchiosauria represent the ancestral group of the Caudata. This suggestion is by no means new, since Baur and others have held the same view. This topic has been discussed at length elsewhere (459) by the writer.

The relations of the form Micrerpeton caudatum are readily determined. The number of the presacral vertebrÆ, the form and position of the ribs, the shape of the skull, the arrangement of the cranial elements, the structure of the pectoral girdle and the character of the ventral armature all clearly bespeak a close relationship with Branchiosaurus, Melanerpeton, Pelosaurus, and other European branchiosaurian forms from the Upper Carboniferous and Lower Permian.

The above-described species, with others given below, is the earliest geological evidence of the Branchiosauria, since the oldest European forms are from the Stephanian (Upper Carboniferous) , which probably lies somewhat above the horizon of the Allegheny series of North America. The presence of the Branchiosauria in America is of considerable interest in the bearing it has on the distribution and migration of the Paleozoic animals. Knowledge of how the group came to occur in such widely separated localities in approximately contemporary geological strata is an unsolved problem of paleontology. It is possible that the piscian ancestors of the Amphibia migrated across or along the borders of the seas and began the amphibian phase of development independently in the two continents. That evolution should, in this case, have followed almost exactly parallel lines seems incredible.

Measurements of the Type.

mm.
Length of entire animal 49
Length of head in median line 6.5
Width of head at posterior border 8
Length of orbit 2.5
Width of orbit 2
Interorbital space 2
Length of the vertebral column 33
Length of the vertebral centrum in dorsal series 0.5
Length of trunk from base of skull to sacrum 22
Length of rib 1.5
Length of scapula 3
Maximum width of clavicle 2
Length of humerus 2.5
Length of ilium 1.5
Length of femur 2
Length of tibia 1.5
Length of tail impression 21.5
Width of tail impression at base 4

Genus EUMICRERPETON Moodie.

Moodie, Kans. Univ. Sci. Bull., VI, No. 2, p. 330, 1912.

Type: Eumicrerpeton parvum Moodie.

The genus is established on three well-preserved specimens representing nearly the entire anatomy. The generic characters are found in the very broad posterior table of the skull, with its short length, reduction of tympanic notch, and shortness of body. The body-length of Eumicrerpeton (plate 5, fig. 1) is less proportionately than that of other closely allied genera. Other generic characters are found in the sharp postero-lateral angle of the skull, and it is to be distinguished from Micrerpeton, especially, by the short, stumpy limb bones. The narrow, elongate eye, placed close to the edge of the skull, is a character not observed hitherto in the Branchiosauria. The genus is closely allied to Branchiosaurus of Europe.

Eumicrerpeton parvum Moodie.

Moodie, Proc. U. S. Nat. Mus., 40, p. 430, fig. 1, 1911.

Moodie, Kans. Univ. Sci. Bull., VI, No. 2, pp. 331-336, pl. 3, figs. 3 and 4; pl. 4; pl. 5, fig. 1; pl. 6, figs. 1 and 2, 1912.

Moodie, Amer. Nat., 44, pp. 367-375, figs. 1-4, 1910.

Moodie, Science, n. s., XXXI, No. 789, p. 233.

Type: Specimen No. 803, Yale University Museum. Other specimens, No. 802, Yale University Museum, and No. 4400, U. S. National Museum.

Horizon and locality: Mazon Creek shales, near Morris, Illinois.

The impression of the outline of the entire body is preserved (plate 3, figs. 1 and 2) in three specimens, and in all are found molds and impressions of the alimentary canal, which, in one specimen (471), are remarkably complete and instructive. The three specimens will be discussed separately, since they show different features.

The impression of the larger animal (No. 803, Yale University Museum), which is probably an adult, presents the following elements: the entire skull, both humeri, impressions of posterior and anterior ventral armature, portions of the alimentary canal, one femur, portions of a fibula and tibia, and the entire impression of the tail, on which, as in Micrerpeton caudatum, there occur two definite dark lines, one beginning at the tip of the tail and running obliquely along the tail to where the impression is broken at the anal region; the other beginning at a distance of 4.5 mm. from the tip and running almost parallel with the median line. These two lines undoubtedly represent the lateral-line system.

The skull is especially noted for its shortness and the great posterior width, as well as for the almost entire absence of the tympanic notch. The pineal foramen is located on a line with the posterior border of the orbits. The eyes themselves are narrow and acuminate at each end, with a pronounced convexity inwards and a flattening on the outer margin. They are located on the very border of the skull, but relatively more posterior than in Micrerpeton. No sclerotic plates are evident. The median suture can be indistinctly observed running the entire length of the skull. The sutures bounding the outside of the f rentals and the squamosals are partially evident, but not satisfactorily preserved. The mandible is represented by a mold which in wax impression shows short, stumpy teeth.

Posterior to the skull at a distance of a millimeter there are two sharp impressions which may represent the anterior edges of the interclavicle or they may be branchial elements. They are distinctly curved, however, and probably represent portions of the interclavicle. A wax impression does not show a discrete structure, but the boundaries of some larger element. No other remains of the pectoral girdle can be discerned. The humeri are short and relatively thick. Wax impressions show them to have had truncate or slightly concave ends, thus indicating the absence or slight development of the endochondrium. No other elements of the arm are preserved.

The impression of an elongate femur and the heads of the tibia and fibula of the left side are preserved.

The ventral armature is preserved in two small patches, and these show the chevron-shaped rods to have been very fine much more delicate than in Micrerpeton.

The body impression is very instructive and interesting, both in showing the form of the body and because in it are preserved the larger portions of the alimentary canal. The form of the body can best be discerned by reference to the figures (plate 3, figs. 1 and 2; plate 5, fig. 1).

The portions of the alimentary canal preserved consist of the greater portion of the stomach, three coils or loops of the small intestine, the rectum, and a pit which undoubtedly represents the anal opening. The anus is found at a distance of 16 mm. from the tip of the tail and is somewhat removed from the body portion, as in modern salamanders. On each side of the posterior end of the rectum there occur a pair of enlargements which probably represent the oviducts at their posterior ends (fig. 15, C).

MOODIE
PLATE 3

1. The larger specimen of Eumicrerpeton parvum Moodie. × 1.

2. The smaller specimen of Eumicrerpeton parvum Moodie. × 1.

3. Type specimen of Erpetobrachium mazonensis Moodie. × 1.

4. Type specimen of Erierpeton branchialis Moodie. × 1.

5 and 6. Type specimen of Mazonerpeton longicaudatum Moodie. × 1.

7. A copy of Cope's drawing of the type specimen of Amphibamus grandiceps Cope. The original was destroyed by fire. × 3.

The tail impression is more acuminate than in Micrerpeton, but shows the same structures as in that form, i.e., the lateral lines which have already been mentioned. Micrerpeton was a more rapid swimmer than the present form on account of the greater development of the tail.

The second specimen of the species (No. 802, Yale Museum) shows much the same character as the specimen already described, except that there are impressions of small, blunt teeth on the mandible. The two humeri and the femur of the left side are preserved and the interclavicle is represented by an identical impression, as in the first-described specimen. The tail impression, though similar in form, does not exhibit so much of the structure of the lateral lines (fig. 15, B).

The matter of especial interest in connection with this second specimen is the remarkably perfect preservation of the alimentary canal, which is entire, except for the very anterior end of the oesophagus. The posterior portion of the oesophagus, which measures 3.5 mm., is clearly preserved. Its anterior end is thrown around posteriorly and indicates that this end was loosened after death and became displaced before fossilization. The length preserved may represent the entire oesophagus. The oesophagus is constricted before it enters the stomach, which shows the usual curvature found in modern salamanders. The stomach measures 6 mm. in length by 2 mm. in breadth, and consists of a single enlargement as in the modern Amblystoma punctatum. It increases in size somewhat toward the pyloric end and then very gradually constricts to the pylorus. Three divisions of the small intestine can be seen. The most anterior one, corresponding to the duodenum, is segmented, as though the intestine had been filled with food before interment. The remainder of the intestine, corresponding to the ilium, is looped in the form of two figures 8 which are superimposed, with the upper portions of the 8 at right angles to each other. The rectum is clearly discernible, though its lower portion is somewhat obscured by having the lower part of the upper loop of the intestine lying over it. The anus lies at a distance of 1.5 mm. posterior to the transverse line from, the upper end of the femur, and is quite well back on the tail, as in modern salamanders. In this specimen also occur two oval bodies which may be identified as the lower ends of the oviducts, thus indicating, in all probability, that the animal was a female.

Fig. 14. Mazon Creek Amphibia.

A. Third specimen of Eumicrerpeton parvum Moodie, which exhibits the alimentary canal well preserved. × 2. Original in United States National Museum. a, anus; f, femur; h, humerus; ic, interclavicle; in, intestine; m, mandible; or, orbit; st, stomach; t, tibia and fibula.

B. Type specimen of Amphibamus thoracatus Moodie. Original in United States National Museum. × 2 (For description see P. 132.)

A dissection of several species of modern caudates has resulted in the discovery that the adult condition of the alimentary canal of all the species dissected (Amblystoma punctatum, Necturus maculosus, Diemyctylits torosus, D. viridescens, etc.) is much more complex than that exhibited by the specimen under discussion. A very near approach to the condition found in Eumicrerpeton parvum is found in an immature branchiate individual of Diemyctylits torosus, 56 mm. in length, from a fresh-water pond on Mount Constitution, on Orcas Island, Puget Sound, Washington.

The similarity of the intestinal structure is of considerable importance to our understanding of the relationship existing between the Carboniferous Branchiosauria and the modern Caudata, and only confirms other arguments, offered in another place (459), concerning their immediate relationship.

The branchiosaurian affinities of the present species are almost too evident to need discussion. The entire structure is essentially similar to that of other genera of the order.

The third specimen of this species (No. 4400 of the U. S. National Museum) is almost as perfectly preserved as were the other two specimens. The skull structure, the intermediate position of the pineal foramen, the epiotic notch, and the shape of the skull are essentially similar to the described specimens of the species. The present specimen is more developed than the other two and probably represents an adult. The alimentary canal is perfectly preserved.

It is nearly half again as long as the smallest of the above-described specimens, and the skull is proportionately longer and wider. There is preserved also an impression of the anterior ends of both clavicles. The right humerus is imperfectly preserved, as is also the right femur and tibia; other than these the fossil is merely an impression.

The skull is so similar to those described above that additional description is unnecessary. The pineal foramen is quite large and lies on a line which cuts the orbits into equal longitudinal parts. The interorbital space is about equal to the long diameter of the orbit. Traces of sclerotic plates are observed in the left orbit, but they are quite imperfect.

The alimentary canal (fig. 14a) is unlike the previously described structures, in that the intestine is longer and more convoluted. It lies in five longitudinal folds and ends in an enlarged cloaca, near which there are impressions of two glands, or the posterior ends of the oviducts, as was suggested for the Yale specimens. The creatures undoubtedly fed on small plants and animals much as do the recent salamanders.

Measurements of Eumicrerpeton parvum Moodie.

No. 803 (222), Yale University Museum: mm.
Length of animal 37.5
Length of skull 4.5
Posterior width of skull at table 6
Long diameter of eye 1.75
Transverse diameter of eye 65
Length of left humerus 1.50
Number of ventral armature rods in 1 mm 10
Length of femur 1.75
Width across base of tail impression 3.5
Length of tail from base to tip 17
No. 802 (471) Yale University Museum:
Length of animal 30
Length of skull 4
Posterior width of skull 5
Length of oesophagus 3.5
Length of stomach 6
Width of stomach 1.33
Estimated length of intestine 18
Width across base of tail impression 2.5
Length of tail from base to tip 7
No. 4400, U. S. National Museum:
Length of entire animal 45
Length of skull 6
Width of skull 9
Transverse diameter of orbit 1.50
Long diameter of orbit 2.25
Interorbital space 2.50
>Diameter of pineal foramen 50
Length of body from back of skull to pelvis 22
Greatest width of body 9
Length of tail 16
Width of tail at base 5
Length of humerus 3
Length of femur 2.50
Length of tibia (fibula?) 1.75
Length of stomach 3
Length of intestine (estimated) 5.6
Width of intestine 1

1 and 2. VertebrÆ of Spondylerpeton spinatum Moodie. × 1.

3. Type specimen of Mazonerpeton costatum Moodie. × 1.

4. Type skeleton of Cephalerpeton ventriarmatum Moodie. × 0.9.

5 and 6. The halves of the nodule containing a practically complete skeleton of Amphibians grandiceps Cope. × 1.

Originals of above figures in the Yale University Museum.

Genus MAZONERPETON Moodie.

Moodie, Kans. Univ. Sci. Bull., vol. VI, No. 2, p. 336, 1912.

Type: M. longicaudatum Moodie.

The genus is distinguished from other known branchiosaurian genera by the great length of the dorsal region, the elongate tail (plate 5, fig. 2), with its well-developed caudal ribs, the reduction of the tympanic notch, the broad nature of the scapula, the elongate interclavicle, and the slender ilium. The number of dorsal vertebrÆ is identical with that of Branchiosaurus of Saxony.

Mazonerpeton longicaudatum Moodie.

Kans. Univ. Sci. Bull., VI, No. 2, p. 337, pl. 3, figs. 1-2; pl. 7, fig. 3; pl. 10. 1912.

Type: Specimen No. 795 (1234), Yale University Museum.

Horizon and locality: Mazon Creek shales, near Morris, Illinois. (Plate 3, figs. 5 and 6.)

The remains consist of the following elements: an incomplete skull; nearly the entire vertebral column, consisting of cervical, dorsal, sacral, and caudal vertebrÆ, 36 in number; several ribs preserved on each side of the vertebral column; a portion of the ventral armature; the scapulÆ; a clavicle; the interclavicle; both humeri; the radius and ulna of one side and the ulna of the other; portions of both hands; the ilium of the right side; both femora, and a partial impression of the left tibia.

The skull is, unfortunately, very poorly preserved. Enough remains, however, to determine the essential characters. The skull bones, unlike any other American branchiosaurian, have an ornamentation consisting of sharp pits and elevations which in places have a quincuncial arrangement and in others take the form of definite lines of pits or tubercles similar to the condition found in many of the Microsauria. The orbits are large and are situated back of the median transverse line of the skull. They are almost circular in form and contain 6 elongated sclerotic plates very closely arranged around the borders of the right orbit. The plates are twice as long as wide. The interorbital width is 1.25 times the transverse diameter of the orbit.

Not many of the sutures of the skull are discernible. Portions of the frontals, the nasals, the prefrontals, the parietals, and the supratemporals can be identified. Their arrangement is shown in figure 14a. There is a decided posterior table to the skull, with truncate posterior border. The tympanic notch is shallow, with its outer border not so well protected as in Branchiosaurus.

The cervical vertebrÆ are incomplete, but their number was 4 or 5, as in Micrerpeton. The structure of the dorsal vertebrÆ is also uncertain, although the shape can be discerned. The vertebrÆ are short and thick, very unlike the long, cylindrical vertebrÆ of Cephalerpeton. The heavy transverse process is quite evident on the best preserved vertebrÆ. This process recalls that described by Credner for the Saxony Branchiosauria. Several of the vertebrÆ show the articulation of the ribs with this process. The ribs of the caudal region recall very strongly those of Branchiosaurus. They are quite heavy in the anterior caudal region and then diminish rather rapidly to the point where the tail is broken and lost.

The ventral armature is represented by a patch of chevron rods 21 mm. in length. The rods take a very peculiar form, being short crescentic bundles of fine rods, hair-like in appearance. In one of these bundles I count 5 smaller rods. The bundles are arranged in rows similar to the pattern so characteristic of the Carboniferous Amphibia, as described elsewhere. The patch of ventral armature preserved belongs to the abdominal region. A single row of the crescentic bundles measures 11 mm.

Both scapulÆ are preserved in their entire form. They are quite different from those of any other genus, being broadly crescentic with a posterior concavity and an anterior protuberance. The anterior surface of both scapulÆ is obscured. Vascular foramina occur near the base of both scapulÆ; there being three of them in the right scapula, arranged in the form of an isosceles triangle. The morphology of these foramina is uncertain. They have never been observed among the Carboniferous Amphibia, and, so far as I am aware, they are entirely unknown among higher vertebrates.

The temnospondylous Amphibia of the Carboniferous and Permian possess, in the coÖssified scapula-coracoid, three foramina, very similar to the ones in the present form, but they are confined to the coracoidal region and, in the Branchiosauria, as identified by Credner, the coracoid is a free element, although I have never been sure of its identity among American forms. Williston has called these foramina the glenoid, the supraglenoid, and the supracoracoid foramina (Journal Geology, XVII, No. 7). They are not, however, to be correlated with the three foramina above mentioned, since in the Temnospondylia the foramina belong with the coracoid and not with the scapula. The condition of the Temnospondylia occurs in the bony fish Xiphactinus audax Leidy, and an analogous condition obtains in the reptiles, as in the Mosasaurs and Dinosaurs.

Fig. 14a. Skeleton of Mazonerpeton longicaudatum Moodie. c, carpus; cl, clavicle; cr, caudal ribs; cv, caudal vertebrÆ; h, humerus; f, femur; or, orbit; r, radius; sp, sclerotic plates; sc, scapula; u, ulna: vs, ventral scutellÆ. From Mazon Creek. Original in Yale University Museum.

Near the outer edge of the right scapula there is a large fragment preserved, which, I think, must be the misplaced clavicle. It is obscurely triangular or, more exactly, spatulate. The interclavicle is represented by fragments only, and seems to have had a narrow form.

The humeri recall those of Micrerpeton. They are somewhat elongate and apparently cylindrical in their normal condition, although somewhat flattened in the fossil. The shaft is considerably constricted at the middle and the ends are expanded, in which expansion the lower end exceeds. The ends are abruptly truncate, indicating a small amount of endochondral ossification or its entire absence.

The mesopodial elements, unlike what is described for Cephalerpeton, are quite dissimilar in form, recalling the condition in Mesosaurus brasiliensis McGregor. The larger element is, apparently, the ulna. It has the lower end greatly expanded and the shaft is curved outward, resembling very much a reptilian ulna. The radius is much smaller than the ulna, lacks the lower expansion, and is shorter by 1 mm.

The carpus is represented merely by a blank space, with evidences of impressions of cartilage in the sandstone. The hand of the right side contains 4 phalanges. There are 2 phalanges preserved in the first digit, including a sharp-pointed terminal phalanx, and the second digit has only the metacarpal. The third has the metacarpal and the first phalanx, which does not differ in form, but only in size, from the metacarpal. The fourth digit contains only the metacarpal. Of the left hand there are portions of 3 digits preserved, including 3 metacarpals and a phalanx, which in structure are not different from those of the right hand.

The ilium of the left side is preserved, apparently entire. It is elongate and cylindrical, its upper end adjoining the twenty-eighth vertebra. The head of the femur lies close to the lower end of the ilium, so that that element must have been suspended in the flesh, much as in modern salamanders. It could not have been of much use as a support for the body. The form of the femur is not unlike that described for the humerus, save that its lower end is smaller than the upper, while in the humerus the extremities are of equal diameter. A portion of the right femur is preserved, extending in an opposite direction to the left.

Measurements of the Type.

mm.
Length of entire specimen 64
Length of portion of skull preserved 6.5
Posterior width of same 7
Width across orbits 11
Long diameter of orbit 3
Transverse diameter of orbit 1.75
Interorbital width 4.75
Length of dorsal vertebrÆ 48
Length of caudal series 11
Length of anterior dorsal centrum 2
Length of anterior dorsal rib 4
Length of anterior caudal rib 1.75
Length of scapula 5
Greatest width of scapula 4.25
Probable length of interclavicle 6
Width of same 3
Length of clavicle 4.5
Width of same 1.5
Length of right humerus 6
Distal width of same 2
Length of ulna 3.25
Distal width of same 1
Length of radius 3
Width of carpal space 2
Length of metacarpal 1.74
Length of first phalanx 1.75
Length of distal phalanx of right hand .35
Number of chevron rods in 1 mm. 4
Length of ilium 2.25
Length of femur 4
Proximal width of femur 1.50

Mazonerpeton costatum Moodie.

Moodie, Kans. Univ. Sci. Bull., VI, No. 2, p. 341, pl. 2, fig. 3; pl. 8, fig. 4; pl. 9, fig. 2; pl. 10. 1912.

The remains on which the present species is based are inclosed in a much fractured nodule. The parts of the animal which have been identified are: a part of the skull and left mandible, two clavicles, a humerus, impressions of several vertebrÆ, a portion of the dorsal region of the body with several ribs, two portions of the caudal region with several ribs and unidentified fragments. (Plate 4, fig. 3.)

The animal, from the shape and form of the ribs, is undoubtedly a branchiosaurian, since short, heavy, straight ribs have not yet been found to be associated with other than branchiosaurian structures. It is placed in the genus Mazonerpeton on account of the structure of the pectoral elements, the form of the humerus, and the length of the tail, all of which agree in structure with Mazonerpeton longicaudatum. The animal attained, perhaps, a length of 4.5 inches, while that of M. longicaudatum was about 3 inches. The tail in the present species is very long and slender, more elongate than in any other known branchiosaurian.

Fig. 14b. Skeleton of Mazonerpeton costatum Moodie. × 1.5. Original in Yale University Museum, dv, dorsal vertebra; ch, neutral spines; cl, clavicle; cv, caudal vertebrÆ; f, femur: h, humerus; m, mandible; rb, rib; sk, skull; v, vertebrÆ.

The part of the skull preserved is very unsatisfactory and, aside from the fact that it seems to represent the under side of the left half of the skull, little can be said. Three sutures can be observed, but what sutures they are is undetermined. The left mandible lies crushed on the edge of the skull and partially obscures what little there is of that structure. The slightly curved impression, from which the bone has been either broken or weathered, measures 13 mm. in length by 3 mm. in posterior diameter by 1 mm. in anterior diameter. These measurements show the element to have been slender and pointed anteriorly.

Very little accurate information can be derived from the study of the vertebral column of the specimen, nor can the dorsal vertebral formula be made out, since only a portion of the length of that region is preserved and only a few rather indefinite impressions can be discerned. These impressions show the vertebrÆ to be short and higher than in most Branchiosauria.

The caudal series is represented by two sections, one of which is, apparently, from near the base of the tail, judging from the size of the caudal ribs preserved; the other is from near the tip of the tail, and shows the constituents to have been long and slender. Ribs are apparently absent on this section. The position of the two caudal sections shows that when the animal died it was coiled up much like a snake, so that in the fractured nodule three sections of the body are visible. The tail was probably half as long again as the body.

The ribs throughout the body are short, heavy, and straight, with, in the dorsal series, a lateral and a distal expansion which is taken as a distinctive specific character. Judging from imperfect impressions in the dorsal series, the ribs were attached to a transverse process of the centrum, thus agreeing with other branchiosaurians in this respect. The ribs show a progressive decrease in length from the cervical region to the point of their disappearance on the tail.

MOODIE
PLATE 5

1. A reconstruction of the possible appearance in life of the Coal Measures branchiosaurian, Eumicrerpeton parvum Moodie, a small, primitive salamander, less than 2 inches in length, based on three specimens from the Mazon Creek Shales. The lateral-line organs are represented as dark bands on the tail, the sense-organs being, apparently, situated beneath specialized pigmented scales, to which is due the preservation of the lines.

2. A restoration, natural size, of the branchiosaurian, Mazonerpeton, based on two specimens. The form of the animal is quite salamander-like. It is shown when about to feed on a specimen of Acanthotelson stimpsoni, which is said to be a brackish-water crustacean. The branchiosaur and crustacean may possibly have inhabited the same body of water.

The pectoral girdle is represented by two elements, one of which is certainly the right clavicle, and the other is possibly the left clavicle, though its form is somewhat distorted by pressure. Both elements are in the form of an elongate spatula with the dorsal surface greatly concave and the inner end acuminate.

The right humerus is imperfectly preserved, though the impression allows one to gain an exact idea of its form. It lies under the right clavicle. Its ends are truncate with a contracted shaft and expanded extremities; the bone was apparently hollow.

Fig. 15.

A. Impression of Erierpeton branchialis Moodie. bb, basibranchial; hyp, hypohyals; m, mandible; d, body impression. × 3.

B. Eumicrerpeton parvum Moodie. a, anus; f, femur; h, humerus; in, intestine; l, liver; st, stomach; r, radius; u, ulna. × 3.3.

C. Larger specimen of Eumicrerpeton parvum Moodie. a, anus; d, dorsal lateral-line; h, humerus; in, intestine; ml, median lateral-line; st, stomach. × 2.6.

D. Skeleton of Erpetobrachium mazonensis Moodie. cl, clavicle; h, humerus; r, radius; sc, scapula; u, ulna. × 2.

E. Rib of Mazonerpeton costatum Moodie. × 2.5. Originals in the Yale University Museum.

In another nodule (No. 804, Yale Museum) there is a single bone preserved which resembles, to a great extent, a rib of the present species (fig. 15, E), although somewhat larger, and it has been provisionally identified as such. The element is very slightly curved, but shows the expanded head of the rib of this species.

Measurements of the Type of Mazonerpeton costatum Moodie.

No. 800 (777), Yale University Museum: mm.
Length of portion of skull preserved 14
Length of right clavicle 16
Width of right clavicle 4
Length of dorsal region represented 30
Length of cervical rib 8
Length of dorsal rib 6-5
Length of caudal rib 3
Length of caudal portion of body preserved 55
Length of mandible 15
Greatest width 6
Length of right humerus 10
Greatest width of humerus 2
No. 804 (332), Yale University Museum:
Length of rib 11
Width of head of rib 2
Diameter of shall 1

Sparodus sp. (?).

Dawson, Phil. Trans. Roy. Sue. London, pt. II, p. 643, pl. 40, figs. 52 to 56, 1882.

Dawson, Proc. and Trans. Roy. Soc. Canada, XII, p. 75. 195.

Type: Specimen in the Peter Redpath Museum of McGill University.

Horizon and locality: Coal formation at the South Joggins, Nova Scotia.

The material on which the above determination is based was collected in 1878 by Sir J. W. Dawson in the coal formation at the South Joggins, Nova Scotia. Nothing has been collected since that date that would give additional information as to the nature of the form represented. I give here Dawson's description of the remains:

Fig. 15a. Type material of Sparodus, consisting of a, a tooth (× 25); b, four of the smaller teeth (in maxilla?) (× 25); c, three bony scales (× 5); d, fragment of a limb bone (× 2); e, a vertebra (× 2). (After Dawson.)

"In the coaly matter or mineral charcoal at the base of tree No. 10 appeared a few fragments of an animal which may possibly belong to the above-named genus of Fritsch, though I am by no means certain of this identification or of the real nature of the animal.

"The skull is represented by a fragment of a maxillary or intermaxillary bone, with blunt conical teeth. It is smooth or marked merely with microscopic dots. There is also a fragment which may be a palatal bone studded with minute teeth.

"A few vertebrÆ associated with the above bones are long and narrow, with large zygapophyses and long neural spines. Length of body (i.e., of the vertebra) about 3 millimeters.

"With these remains are a few bony scales different from those of any other species found in these trees, and more resembling scales of Ganoid Fishes. They are somewhat rectangular in form, enameled on the surface and beautifully sculptured with waving lines.

"In the same trunk were found some teeth and bones referable to Hylerpeton dawsoni, and it is not impossible that the remains above referred to may have belonged to some creature devoured by that animal, and which would not otherwise have obtained admission to the interior of an erect tree. The tree itself had been removed by the sea, all but a little of the base, and this was in a very unsatisfactory state, so that doubt might even exist as to the limit between the deposit in the interior of the tree and that under its base."


                                                                                                                                                                                                                                                                                                           

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