MEMOIRS CAKNEG-IE MUSEUM. VOL. IV. NO. 5. DESCRIPTION OF NEW CARNIVORES FROM THE MIOCENE OF WESTERN NEBRASKA. By O. A. Peterson. While prospecting in the layer of fossil bones, which is exposed around "Car- negie Hill" and "University Hill" in the Miocene formation on the upper Niobrara River in Sioux County, Nebraska, with a view to opening additional quar- ries,1 the writer, assisted by Messrs. T. F. Olcott and A. A. Dodd, was extremely fortunate in finding in what is now called Quarry No. 3 (see Fig. 1, 3) various FIG. 1. Agate Spring Fossil Quarries. 1-2, quarries Nos. 1 and 2, on Carnegie Hill ; 3, quarry No. 3 ; AM, American Museum quarry on Carnegie Hill ; NU, Nebraska State University quarry on University Hill ; A, quarry A. (From a photograph by the author.) remains of carnivora, among which is a nearly complete skeleton of a canid. The material having been entirely freed from the matrix and prepared for mounting by the skillful work of Mr. S. Agostini, the writer was entrusted by Director \V. .1. Holland with the task of describing and illustrating the new material. 1 No quarries except quarry A hail at that time (1904) been opened in these hills. 205 206 MEMOIRS OF THE CARNEGIE MUSEUM In the quarry designated as No. 3 in the accompanying illustration, there were found from five to nine individuals of Daphcenodon superbus, of which No. 1589 is practically a complete skeleton. The latter was found imbedded together with another individual of the same species not fully adult. The soft sandstone in which the two were found was taken out in a large block and transported to the Museum. On examination it was found that the adult individual was partly articulated, especially the vertebral column and portions of the feet. The tail was found attached to the sacrum and thrown backward in a graceful curve so that the tip was lying close to the pelvis and abdominal region. The anterior nine vertebrae and the anterior portion of the tenth were preserved in position. At this point the caudal region had been exposed and the vertebrae had dropped out. By carefully measuring the natural curve, in which the tail was undoubtedly placed, its length was ascertained and the missing vertebrae were either restored in plaster, or by other vertebrae which were found in the talus below the place where the skeletons were dug out. The ribs of No. 1589 are not well preserved while those of the younger individual, No. 1589a, are quite complete. The posterior portion of the sternum is not present. One hind limb and foot of No. 1589 was found articulated. The posterior portion of the pelvis is weathered away, as is also the extreme anterior portion, leaving only the middle region. When material other than the type is used in the following description it is only for the purpose of elucidation and the catalog number will be mentioned in connection with the specimens referred to in order to prevent confusion. Scattered remains of a number of individuals, apparently of the same species, were found in all of the Agate Spring Fossil Quarries, but in Quarry No. 3 were found the most abundant remains of Daphcenodon superbus. Besides the type of Nolhocyon anneclens Peterson and another small carnivore described later in this paper there was little else found in Quarry No. 3 except remains of Daphcenodon superbus. This is interesting from the fact that hardly more than a stone's throw to the north and practically on the same level, the very rich and classic quarries, No. 1, No. 2, and the "University Quarry" are located (see Fig. 1). The lithological characters of the bone layer of Quarry No. 3 are similar to those in the large quar- ries and it was observed that the same barren pinkish bedrock found in the main quarries is also present in this quarry. The irregularity of the fossil-bearing stratum was observed to be quite like that in the quarries immediately to the north. As has been stated elsewhere, the stratum containing fossil bones in the Agate Spring Fossil quarries was most probably laid down by a stream full of shallow channels which shifted from time to time during its existence. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 207 Mucli field-work has been done and a considerable amount of material repre- senting the carnivora has been found in the lower and middle Miocene of South Dakota, western Nebraska, eastern Colorado and Wyoming during the past few years by the different museums of North America.2 Since further explorations of this region are likely to occur in the near future it has seemed best to postpone a more detailed and systematic work upon the carnivora of these localities and for the present only to publish the new forms recently found by the museum parties and especially to accurately describe the skeleton of Daphomodon superbus. It is believed that the study of the lower Miocene Canidae will be facilitated by the publication of an account of the nearly complete skeleton of one individual. In the following description the exceptionally well preserved and almost com- plete skeleton of Daph.osnus felinus Scott (No. 492, Carnegie Museum Catalog of Vertebrate Fossils) which Mr. J. B. Hatcher described (Memoirs Carnegie Museum, Volume I, pp. 65-108) will be used for comparison. In the first place it seems quite certain that Daphwnus is in the direct ancestral line of Daphosnodon, and secondly, the skeleton from the Oligocene now in the Carnegie Museum, represents the most complete individual of that genus as yet discovered. Valuable aid in this work was rendered by the authorities of the American Museum of Natural History and of Princeton University Museum, who courteously sent to the Carnegie Museum a number of types for comparison. The illustrations are from drawings made by Mr. Sydney Prentice and photographs by Mr. A. S. Coggeshall and the author. Family CANIDiE. Subfamily Amphicyonin.e. Daphoenodon superbus (Peterson). Amphicyon superbus Peterson, Annals Carnegie Museum, Vol. IV, pp. 51-53, PL XVIII, 1908. Daphanodon superbus (Peterson), Science, N. S., Vol. XXIX. pp. 620-621. Type: Skeleton nearly complete, No. 1589, Carnegie Museum Catalog of Vertebrate Fossils. Horizon : Miocene (Lower Harrison Beds). Locality: Agate Spring Fossil Quarries (Quarry No. 3), Sioux County, Nebraska. JJislhirtii'i ■characters. — In my previous notes in SbieriCfllgave the following distinctive characters : Cranium, comparatively short, broad, and low; muzzle large; 1 Professor F. B. Loomis, of AmlierBt, recently discovered two miles east of the Agate Spring Fossil Quarries the greater portion of a line skeleton whioh he regards as belonging to the same genus described below. 208 MEMOIRS OF THE CARNEGIE MUSEUM sagittal crest prominent ; brain-case small ; incisors heavy and short; canines compara- tively small and oval in cross-section ; P4 with antero-internal cusp of moderately large size and located relatively close to the main body of the teeth ; 3P and SP large and broad; M3 present, though small, practically one-rooted and aligned with the internal border of M1 and 3P. To the above cited characters should be added : 7f C\ Pf il/f ; the symphysis of the inferior ramus separated or slightly coossified ; the tubercles of all the molar-premolar teeth blunt; the heel of the lower carnassial tooth large: metatarsals not appressed at their proximal ends ; feet sub-dig itigrade ; tail long and heavy. The Skull. (Plates LXXIV-LXXV1I.) In the articles cited above attention was paid to the cranial and dental charac- ters of Daphcenodon superbus. In the present paper, therefore, I will only give a short review of my previous description and add characters which I regard as of interest and importance to the student. The cranium of this new genus represents an animal very nearly as large as a fully adult gray wolf, but the skull is proportionally shorter and broader, the brain- case smaller, the occipital condyles smaller, the basioccipital and basisphenoid of greater transverse diameter, the paroccipital process further separated from the tympanic bulla and directed more backward, the mastoid larger, the tympanic bulla smaller, and the postglenoid process heavier. When compared with Daphoenus felinus the skull is distinctly shorter and broader but the brain-case is evidently not much enlarged and there is a large frontal sinus ; the muzzle is heavier ; the pterygoids are shorter, and the region back of the pterygoids is apparently also somewhat shortened. The postglenoid and paroccipital processes are of the same relative size and position while the mastoid process is a truncated rounded knob, and is of proportionally greater size. In fact the entire mastoid region of Daphomodon seems to have increased, which is a distinct step away from the line of Canis. The space between the paroccipital process and the condyles is shorter, so that the occipital condyles are more sessile in the specimen under description than in the Oligocene genus.3 The tympanic bullae are present on both sides in the type specimen ; they are of moderately large size and triangular in general outline. The tubular process, which encloses the external auditory meatus is quite prominent and is protected posteriorly by the heavy mastoid, though not so completely coossified with the latter bone as in the recent dogs. On the postero-internal angle of the bulla there is a sharp vertical ridge which unites with a similar though much 3 In Daphtrnus felinus (No. 492, C. M. Cat. Vert. Foss. ) the base of the skull in the region of the condyles is some- what injured and the separation of the condyle from the paroccipital process may, in part, be due to this fact. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 209 heavier ridge on the antero-internal angle of the paroccipital process, thus separating the large pit for the tympanohyal externally, the jugular foramen, and the foramen lacerum posterius internally. The bulla seems to be fairly well fused to the tym- panic region while in the Oligocene genus it apparently always slipped off. From a study of the tympanic bulla in Daphwnodon superbus it is quite evident that though quite solidly attached along the antero-external angle and along the border of the basioccipital, the bone is not nearly so well ankylosed by the paroccipital process and the mastoid as for instance in Canis. In Baphasnus felinus the bulla appears to be more loosely attached to these processes than in the type under description. This fact apparently supports Mr. Hatcher's contention that the ankylosed auditory bulla in recent dogs should be regarded as a specialized character.4 It is interesting to note that the tympanic bulla in Cynodesmus brachypus (Cope) is, in the first place, larger ; and, secondly, much better ankylosed with the sur- rounding bones than in the form under description, and more nearly approaches that of Canis. Professor Cope 5 and Dr. Matthew 6 have observed that the paroccipi- tal process in Cynodesmus brachypus projects " backwards nearly as far as the pos- terior face of the occipital condyles." After a careful study of this region in the type of Cynodesmus brachypus, now before me, it would seem that the backward pro- jection of the paroccipital process, regarded as a primitive character by Matthew, is in reality due very largely to crushing, there being no part of the base of the skull which is left undisturbed. Besides, if the present position of the paroccipital process of the skull of C. brachypus is a natural one it would interfere with the free move- ment of the occipital condyle upon the atlas. Unfortunately the paroccipital proc- ess of the left side of C. brachypus has been broken off, which otherwise might have furnished good verification. From Dr. Matthew's statement and illustrations (/. c.) it is very evident that the paroccipital process in Cynodesmus ihomsoni is closely connected to the tympanic bulla and points almost directly downward at the tip, while in Cynodesmus thooid.es Scott 7 the process is heavy and projects more back- ward. It is quite likely that in a more perfect specimen of C. brachypus the condi- tion of the paroccipital process would be found to be more like that in C. ihomsoni and certainly not more backwardly directed than in C. ihooides. The postglenoid process of C. brachypus has proportionally a less transverse diameter than in Daphamdon superbus and is more nearly similar to that of Canis, while the foramen opticum occupies a similar position, i. e., more in advance of the exit of the foramen rotundum than in recent dogs. "'Oligocene Canidic/'JMBMoiKH CARNEOIK Muhkum, Vol. 1, pp. 72-7:1, 1902. "Bull. U. 8. Oeological awl (ieographical Survey of the Territories, Vol. VI, p. 389, 1881. "Ball. Am. Mus. Nat. History, Vol. XXIII, p. 186, 1907. 'Trans. Are. Pliilos. Society, Vol. XVII, p. 66, PI, I, rig. 1893. 210 MEMOIRS OF THE CARNEOIE MUSEUM In the type under description the greater number of the foramina are well pre- served. With the exception of the more internal position of the stylomastoid fora- men8 and the slightly more anteriorly placed foramen opticum, the size and position of the foramina are quite similar to those of Canis, which is well illustrated in Plate LXXV. The more internal position of the stylomastoid foramen is due to the large development of the mastoid process and limited backward extension of the tympanic bulla, while the position of the optic foramen may be due to the more advanced position of the orbit in Daplicenodon than in Canis. The alisphenoid canal is appar- ently larger in proportion than in the Oligocene genus. Fig. 2. Skull of young individual of Daphcetwdon auperbus. g nat. size. No. 1589a, C. M. Cat. Vert. Foss. Per, mastoid portion of periotic ; Gp, postglenoid process ; .7, jugal ; PI, palatine ; Stj, squamosal ; Pa, parietal ; Fr, frontal ; As, alisphenoid ; Os, orbitosphenoid ; I, lachrymal with foramen ; Mr, maxillary; io , infraorbital foramen ; op, optic foramen ; fr, foramen rotundum ; sf, sphenoidal fissure. (Note: The occipital plate, the jugal, the nasals and the premaxillaries are missing and a portion of the zygomatic arch is removed in order to give a better view of the dif- ferent elements of the skull in this region. ) In Fig. 2 is given a side view of the skull of the younger individual (1589a) which was found together with the type. The premaxillary, nasal, malar, supra- occipital, and the base of the skull are slipped off and lost, but what remains furnishes an admirable idea of each element and their relative proportions in com- parison with those in Canis. It is seen that the parietal and frontal are longer, lower, and less convex ; the alisphenoid of apparently the same proportion ; the orbitosphenoid as long, but of less vertical diameter ; the lachrymal larger ; and the maxillary shorter than in the recent form. From the type specimen the process of the premaxillary, which extends upward between the maxillary and the nasal, is much shorter and weaker than in Canis and in this respect more nearly similar to what is observed in Daphcenus. 8 The stylomastoid foramen is not nearly so well separated from the large pit for the tympanohyal as is generally the case in recent dogs. PETERSON: NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 211 Lower jaw. — The mandible is quite similar to that of Can is lupus in its general characters, but may better be compared with such a form as Lycaon on account of its broader symphysis and condyles. The alveolar border is, however, more completely taken up by the premolars in the latter form than in the fossil. There are two mental foramina : one directly below P2 and the other below PTi. The inferior border of the horizontal ramus is less curved fore-and-aft than in DaphcewLS, while the line of the alveolar border is about the same. The temporal fossa is broad and deep, the coronoid process large and the process of the angle very robust and more inflected than in Daphccnus or the recent forms. Aside from the large pit, which lodged a considerable sized tympanohyal in the skull, the hyoid arch is not known in Daphomodon superbus. Measurements of Skull and Jaws- Mm. Greatest length of skull 253 Distance from occipital condyle to incisors 228 Distance from occipital condyle to M3 112 Distance from M3 to and including the incisors 117 Greatest transverse diameter of the skull 148 Transverse diameter of occipital condyle , 46 Greatest transverse diameter of the brain case 58 Transverse diameter of the frontals at postorbital processes 68 Transverse diameter of muzzle at canine _ 51 Greatest length of mandible jg2 Height of mandible at coronoid process 82 Dentition. — The dentition (If Cj Pf Mf) of Daphcenodov superbus is completely preserved. The first and second upper incisors are laterally compressed and resemble those of Daphccnus more than those of recent forms. I3-, however, displays a rapid approach to the characters found in Canis, i. e., it has attained a much larger size and a more nearly caniniform structure. The tooth (P) is isolated by diastemata; the one in front being quite short, while the one behind is equal to the greatest transverse diameter of the tooth. As a whole the incisors are proportionally some- what larger than those of Daphccnus felinus and 1 - is distinctly further advanced toward conditions found in Canis. As has been stated elsewhere, the canine, though quite robust, is not so large, nor of the shape seen in Amphwyon giga ulcus of Europe, to which genus it was wrongly referred, tn the present genus the tooth is of moderately large size, oval in cross-section and recurved quite as in Chnis occir dentalis, though somewhat less compressed than in that species. The l>ase of the tooth and the fang are rounder, which causes a greater swelling on the side of the muzzle than is seen in the latter form. The molar-premolar teeth have rather blunt cuspa 1" is single-rooted and 212 MEMOIRS OF THE CARNEGIE MUSEUM has a simple crown ; it is separated from the canine and P— by short diastemata. The latter tooth is also isolated, but the diastema back of it is very short. This tooth has two roots ; its crown is comparatively simple, there being no anterior or posterior basal cusps. P- is simple-crowned, like the anterior premolar, and other- wise has about the same relative proportions as in Daphcenus felinus. Its position in the alveolar border is oblique, which makes it appear of small antero-posterior diameter in the side view of the skull. In P- (carnassial) a slight change from what is observed in the Oligocene genus has taken place. The deuterocone is slightly reduced in size and is closer to the main body of the tooth than in either Daphcenus or any of the allied John Day forms, and in this respect is more like the corre- sponding tooth in Amp>hicyon major of Europe. The deuterocone is, however, sup- ported by a much heavier root than either in Amphicyon or Canii and its oblique position in the alveolar border is totally unlike the latter genera. The smaller antero-posterior diameter and otherwise primitive features of the tooth places the genus rather closer to the Oligocene form and removes it further from Canis. M- and M- are larger than in any of the allied John Day forms [Temnocyon, Mesocyon, Para- daphwrvus) and have also undergone some changes from the corresponding teeth in the Oligocene genus, while Ma is apparently the same. This change of the molars is principally due to the greater development of the postero-internal angle, which gives these teeth a slightly greater antero-posterior diameter internally than is usually seen in the Oligocene species, and the tooth is otherwise apparently inter- mediate between that of these early forms and Canis occidentalis, which has a very prominent, sharp, and crescentic postero-internal tubercle, especially on M-. The development of the postero-internal angle is unusually like what we find in Arn- phicyon major and it is quite curious to find the conical structure of the different tubercles so nearly alike in the two genera. M- and M- of the European genus are, however, proportionally larger, the posterior intermediate cusp (metaconule), especially on Mj, is more strongly developed and the antero-external cusp (para- cone) is the larger, while in Daphcenodon para- and metacones are subequal and of the primitive type met with in Daphcenus; in fact the tubercles are more depressed than in the latter genus, as is seen by referring to the illustrations, PI. LXXV. The internal border of the grinding surface of M-3- extends slightly below that of the preceding tooth. This character, which is well shown on PI. LXXIV, may, in part at least, be due to slight displacement. Ma is, as stated, practically a duplication of that tooth in Daphcenus, i. e., a small tooth with transversely oval and very low crown with a very short root, the alveolus which is posteriorly formed by a thin and very delicate border in the maxillary. PETERSON: NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 213 The lower incisors are laterally compressed. IT is quite small in comparison with that of Canis, while I* and I,, though small, are in better proportion. The canine is robust and resembles that of Canis ocadentalis very closely, the cross-section of the crown being somewhat more regularly oval and less excavated on the antero- internal angle. PT is a small single-rooted tooth with the crown extended well forward from the base, the apex being nearly directly over the anterior face of the root. The tooth is isolated by diastemata, of which the one in front of it is the longer. The next three premolars are also well separated in the jaw. P* and P¥ have simple conical crowns with larger posterior than anterior bases and their shape and relative diameters are quite similar to those in Daphamus, while PT is somewhat enlarged and has a strong posterior cusp and prominent cingulum similar to some species from the Oligocene, Daphanus dodgei Scott and Temnocyon ahigenis Cope from the John Day formation of Oregon. In MT (carnassial) is observed the greatest change from what appears in the Oligocene form in the direction of the more recent type. It is at once seen that this tooth has increased in its antero-posterior diameter, the metaconid is situated lower down,9 and the hypoconid is bluntly trenchant and is greatly developed, while the entoconid is rather small. The tooth is not, however, modified into the perfect shearing blade met with in Canis, as it still retains to a greater or less extent the conical structure of the tubercles so characteristic of Daphcenus and its ancestors, the ihacidx. M2 is quite large ; the trigonid is raised only very slightly above the heel ; the proto-, para-, and metaconids are fused into a more or less solid and obtuse mass' the protoconid being the largest of the three, while the paraconid is represented only as a basal heel on the antero-internal angle. The heel is quite large and is occupied principally by an antero-posteriorly placed ridge (hypoconid) which is low and located nearer the external than the internal border. The entoconid is represented by a low and oblique ledge on the postero-internal angle of the crown. Mj is single- rooted and is reduced to a small oval-shaped crown with a slight elevation on the anterior half, while postero-internally there is a shallow valley produced, to some extent by trituration. The size of the tooth with reference to the one preceding it is proportionally the same as in Dapluvnm felinus here used for comparison. - It is interesting to note that Cynoimnm traoKypn f,„m «,, „iocene of the Laramie Peak M. advanced towards the recent dog, in having the premolars .arger, MJ reduced and the iover i-K lei transverse diameter. 214 MEMOIRS OF THE CARNEGIE MUSEUM Measurements of Dentition. Mm. Length of superior premolar 6eries 55 Length of superior molar series 34 Autero-posterior diameter of superior caDineat hase 13 Transverse diameter of superior canine at base 10 Anteroposterior diameter of PL 6 Transverse diameter of P1 5 Antero-posterior diameter of P^ 11 Transverse diameter of P1 5 Antero- posterior diameter of P3 13 Transverse diameter of P^ 7 Autero-posterior diameter of P4 22 Transverse diameter of P4, anteriorly 14 Antero-posterior diameter of Ml- 19 Traosverse diameter of M1 24 Antero-posterior diameter of M- 13 Transverse diameter of M1 18 Antero-posterior diameter of BX5 5 Transveree diameter of M3- 8 Length of inferior premolar series 47 Length of inferior molar series 46 Antero- posterior diameter of inferior canine at base 12 Transverse diameter of inferior canine at base 10 Antero-posterior diameter of PT 6 Transverse diameter of PT 4 Antero-posterior diameter of P2 19 Transverse diameter of P2 5 Antero-posterior diameter of PT 13 Transveree diameter of P5 6 Anteroposterior diameter of P.r 16 Traosverse diameter of P? 8 Antero-posterior diameter of Mr 25 Transverse diameter of MT 11 Antero-posterior diameter of Ms 14 Transverse diameter of M2 10 Antero-posterior diameter of Mj 9 Transverse diameter of M3 7 Although there are a number of apparently strong specific characters in the dentition of the second specimen (No. 1589a) found with the type, I refrain from establishing a second species on account of the immaturity of the specimen. In this second specimen the upper and lower premolars are much more crowded in the alveolar border in comparison with those in the type specimen. The upper carnas- sial has a better developed postero-external cutting ridge (tritocone) and the cusps are generally more trenchant. PT is relatively somewhat larger with a higher crown, the metaconid of the lower carnassial slightly smaller and M5 two-rooted. PETEKS0H : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 215 The remains indicate an animal of slightly larger size when fully adult (see Figs. 2 and 3). Fig. 3. Lower Jaw of Yoang Iudividu.il of Dnphrenodon supnbux. 'i nat. size. No. 1589a. In section 17 of Quarry No. 1 was found a fragment of a mandibular ramus with the deciduous sectorial in position and the permanent pT buried in the jaw. The latter tooth is only partially developed. The differ- ent cusps of the deciduous tooth are quite sharp and the heel is of considerable size and completely basin- shaped. The fangs are excessively spread in the an- tero-posterior direction, due to the constant pressure from below by the permanent tooth, and furnish a sure indication of the juvenility of this tooth. Except for its slightly smaller size, the tooth most nearly resembles that of "Cam's" anceps Scott and is here figured for the purpose of warning the student. The specimen may or may not represent Dapkcsnodon swperbus. Flo. 4. Daphamodon superbus? Fragment of lower jaw of young individual with deciduous sectorial in place and p, bnried in the jaw (No. 2201). Nat. size. The Vertebral Column. The formula of the vertebral column is as follows: cervicals seven, dorsals thir- teen, lumbars seven, sacrals three, and caudals twenty-eight? Witli the exception of the missing region of the caudals, already referred to in the introduction, the vertebral column of the type specimen is in excellent preservation. Atlas. — The atlas is quite long antero-posteriorly. Its large attenuated and wing-like transverse process gives the bone a transverse diameter proportionally equal to that in Cams. Ill general outline the bone is quite similar to that of the latter genus, hut there area number of di defences shown on closer examination which 216 MEMOIRS OF THE CARNEGIE MUSEUM reveal conditions more nearly identical with those found in Daphamus. Thus the posterior exit of the vertebrarterial canal is not superior as in the dogs, but more nearly posterior as in the cats;10 the anterior cotyles are small; the transverse Fig. 5. Atlas of D. superbus. J nat. size. 1, dorsal view ; 2, lateral view ; 3, ventral view. diameter of the articulation for the axis is proportionally less in size ; and the bone, with the exception of the transverse process, is, as a whole, rather heavier than in Cards. The atlanteo-diapophysical notch is deep and as in the dogs is not covered with a bony bridge to form a foramen. This character was observed in Daphanus by Professor Scott," while in some of the genera of the Mia- .f^HI , cidx (Oodectes) this notch was already converted into a fora- men.12 The foramen in the arterial groove, which pierces the wall of the neural arch on the ventral face of the trans- Fia. 6. Atlas of Ctinoties- . ,. , ,, , . i . i .-, . , „ KT Q1 .„ verse process, is exceedingly small, winch is also the case in mus brachypusCope. No. 8140, r ' © J cope Collection, Am. Museum the atlas of Daphamus here used for comparison. In Canis Natural History, i nat. size. , . e • », this foramen is ol larger size. In the atlas of Cipiodesmus brachypus the posterior exit of the vertebrarterial canal is more dorsally located (see Fig. 6) ; its atlantal notch is not nearly so deep and the foramen piercing the neural arch on the ventral face of the transverse proc- ess is as large as in Canis. Measuekments of Atlas. Mm. Greatest antero-posterior diameter 52 Greatest transverse diameter 99 Antero-posterior diameter at cotyles 39 Transverse diameter of cotyles anteriorly 51 Transverse diameter of cotyles posteriorly 36 Axis. — In comparison with Canis the axis is short and high and according to Professor Scott's description of the axis in Daphamus it appears also to differ from that genus in some important and rather surprising details. Scott says : " The cen- 1,1 Wortraan, J. L., " Osteology of Patriofelis," Bull. Am. Mus. Nat. Hist., Vol. VI, p. 137, 1894. 11 "Notes on the Canidie of the White River Oligocene," Trans. Amer. Philos. Society, Vol. XIX, p. 337, 1894. "Matthew, W. D., " Carnivore and Insectivora of the Bridger Basin," Memoirs Am. Mns. Nat. Hist., Vol. IX, p. 367, 1909. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 217 trum is elongate, narrow, and depressed, with a thin and inconspicuous hypopophy- sial keel." In Daphecnodon superbus the centrum, though somewhat depressed, is short in comparison with that of Canis and the hypopophysial keel is more strongly developed than in the latter. The ventral face of the centrum in the present genus is not nearly as concave antero-posteriorly as in the dog, and the ventral keel ter- minates posteriorly in a broader area, which is extended downward very slightly so as to form of its posterior three-fourths a slight antero-posterior concavity, but there is no prominent tubercle as in Canis. The articu- lation for the atlas is more convex and rises x • i .1 • i .1 fi j • Fig. 7. Lateral and Posterior Views of Axis of higher upon the sides than in Cants and is ■»**.*■ ° r D. superbus. \ nat. size. apparently not materially changed from that in the Oligocene form. The odontoid process also agrees with Professor Scott's description of that in the older form, which he says is " a long, slender, bluntly pointed peg, with a heavy, rounded ridge upon its dorsal surface, which is continued back along the floor of the neural canal." The transverse process is heavier, shorter, and is more nearly parallel to the centrum than in Canis, which is also characteristic of Daphcenus. On the lower border of the transverse process is developed an inferior branch or costal element (see Fig. 7) which is rather unusual on the axis among the dogs and is not mentioned by Scott in his descrip- tion of this process in Daphcenus. The vertebrarterial canal is apparently slightly further forward than in Daphamus, but it is still further back and slightly longer than in Canis. The neural canal is large, as in Daphcenus, and does not contract so much towards the hinder end as in Canis. The spinous process is a long high keel which is produced into a laterally compressed process anteriorly, rapidly ex- pands posteriorly, and is proportionally lighter than in the cats, c. g., Felis leo. The posterior face of the spine is excavated somewhat as in Felis, though in a less degree, and forms a different attachment for the cervical muscles than is seen on the con- vex and rugose surface in Canis. According to Scott, the neural spine in Dapha mis corresponds practically to the description here given. In Cynodeam.ua brachypua the centrum of the axis is long and narrow and in its general features, including the inferior keel and the odontoid process, answers more nearly to that of Canis than to that of Daphwims and Daphwnodon , while the neural spine is extremely long and overhanging both anteriorly and posteriorly. The latter process is proportionally longer than that in Leo, and tapers more gradually from the base of the postzygapophyses to the free end as in the Eocene creodonts. 218 MEMOIRS OF THE CARNEGIE MUSEUM Measurements of Axis. Mm. Antero-posterior diameter of the centrum, odontoid process included 54 Greatest transverse diameter of centrum, anteriorly 36 Greatest transverse diameter of centrum, posteriorly 22 Antero-posterior diameter of spine 53 Transverse diameter across postzygapophyses 34 Third Cervical Vertebra. — The general characters of the third cervical vertebra are similar to those of the same bone in Cants, though certain features are rather feline. Thus the inferior branch of the transverse process is strongly developed, the tubercle superior to the postzygapophysis is large and points directly backwards, and the neural spine is considerably developed, while in dim's the transverse process is smooth, the tubercle above the postzygapophysis points more upwardly and the neural spine is less developed. The centrum is short and has a strong keel, which terminates posteriorly in a broad area, but does not form a hypopophysial tubercle as in the dogs ; the transverse process is shorter and broader, and the vertebrar- terial canal is longer. Measurements of Tbird Cervical Vertebra. Mm. Antero posterior diameter of centrum 31 Antero-posterior diameter of neural arch including zygapophyses 46 Transverse diameter across transverse processes, posteriorly 52 Transverse diameter of centrum posteriorly 22 Vertical diameter of centrum posteriorly 14 Fourth and Fifth Cervical Vertebra:. — The fourth and fifth cervical vertebrae are heavier than the preceding vertebra. The postero-inferior branch, or costal t element of the transverse process, B is also more developed ; the neural ^fc^JI^ arches are more deeply emarginated IhL #r *n fr'011* and behind ; the neural <^JxL^ ■-^f'W-^a sPmes are higher and the tubercles j|r* WkSWiiW^ above the postzygapophyses are 1 3 smaller. There are otherwise no Fig. 8. Fifth Cervical Vertebra of D. mperbus. I nat. size. 1, marked differences between the posterior view ; 2, lateral view ; 3, anterior view. , . third, fourth, and fifth cervical ver- tebrae. In the two latter vertebras the ventral keel of the centrum does not termi- nate posteriorly in a tubercle as in Canis, but laterally has a broad area with small tubercles, similar to what is observed in Felis leo. PETERSON: NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 219 Measurements of Fourth and Fifth Cervical Vertebra. C. 4 C. 5 Mm. Mm. Antero-posterior diameter of centra 31 30 Transverse diameter of centra, posteriorly 22 22 Vertical diameter of oentra, posteriorly 15 16 Transverse diameter across transverse processes, posteriorly 55 59 Transverse diameter across prezygapophyses, posteriorly 43 43 Transverse diameter across postzygapopbyses 41 39 Sixth Cervical Vertebra. — The sixth cervical vertebra has the faces of the centrum more oblique than in any others of the cervical series, but not more opisthoccelous. The centrum is otherwise depressed and rather broad, as are the centra of all the cervicals. The ventral keel is quite prominent in the anterior region, but fades away to an evenly convex border posteriorly and is quite similar to that in Cards. The transverse process is very prominent, as in the latter genus, but the inferior lamella is divided into two parts, a small anterior and a larger posterior por- tion (see fig. 9). In the thooids this lamella forms a solid fig. 9 Sjxth cervical v'er- and thin blade, while in the alopecoids it is generally tebra of D- vperbus. J nat. slightly divided. In the cats it is divided in a greater or less degree. The vertebrarterial canal is proportionally longer than in Canis. The neural arch is broad, but does not form the connecting ledge between the pre- and postzygapophyses as prominently as in Canis ; there is no excessory process devel- oped on this connecting ledge as in the latter genus, though it would appear from Professor Scott's description (/. c, p. 338) that these ledges are perhaps more promi- nent on the arches of the cervical vertebrae in the present genus than in Dapkasnus. The anterior and posterior emar- ginations on the neural arches which separate the zygapophyses in front and behind the neural spines of the cervical vertebrae are deeper than in Canis : they are not unlike those in Viverra africana, and in (his respect are similar to Flo. 10. Cervical Vertebric of ('yiiwlntmu* hmehiipua Cope ( No. , „ , . _ a,.n„ „ .. *■ . „, ,nt. m. , i . • what was found by Scott to be the SHOCope Collection, Am. Museum of NaturalHistory). \ nat. size. J case in the OligOCene genus. The neural spine of the sixth cervical in the type is slightly damaged, but its entire length is represented, and it is shown to be robust and high. 220 MEMOIRS OF THE CARNEGIE MUSEUM In the sixth cervical vertebra of Cynodesmus brachypus the inferior branch of the transverse process is divided, but the anterior portion is much larger proportionally than that in Daphamodon. In all other respects the bone is more nearly like that in Canis than Daphwnodon. In fact the entire cervical series in C. brachypus is strikingly similar to that of the recent form except that the posterior exit of the vertebraterial canal of the atlas is further back, the neural spine of the axis more produced posteriorly, the inferior lamella of the transverse process of the sixth cervical is divided, and the neural spine of the seventh cervical is more robust and higher. Measurements of Sixth Cervical Vertebra. Mm. Antero-posterior diameter of the centrum 29 Transverse diameter of the centrum posteriorly 23 Vertical diameter of the centrum posteriorly 27 Transverse diameter at transverse processes 59 Transverse diameter of prezygapophyses 43 Transverse diameter of postzygapophyses 57 Seventh Cervical Vertebra. — The seventh cervical is characterized by its long and heavy transverse processes and neural spine, which are proportionally quite large when compared with those of Canis. From Professor Scott's observation it is quite clear that Daphcenvs also had high neural spines on the cervical vertebrae, which is gen- erally characteristic of the felines. It is further noticed that the anterior and posterior faces of the neural spine of the seventh cervical in Daphcenodon have conspicuous ex- cavations at the base, which are not present in Canis and Felis, while in Viverra zivetta they are represented to a slight fig. ii. Seventh Cervical extent. These excavations have rough surfaces for muscular vertebra of d. superb™, j attachment, and the one posterior is deeper than the one uat. size. anterior. The four vertebras preceding the one under de- scription have similar rough excavations (see Fig. 8) and these excavations also ap- pear in the case of the anterior dorsals. (See Fig. 13.) Measurements of Seventh Cervical Vertebra. Mm. Antero-posterior diameter of centrum 27 Transverse diameter of centrum posteriorly 25 Vertical diameter of centrum posteriorly 18 Transverse diameter of transverse processes 67 Transverse diameter of prezygapophyses 43 Transverse diameter of postzygapophyses 43 Height of neural spine 47 PETERSON: NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 221 First Dorsal Vertebra. — The centrum of the first dorsal is, as in DapKasnus, broad and depressed, which seems to be true of the dogs generally. The trans- verse process is heavier than in Cams lupus. The postzy- gapophyses point forward and outward in a greater degree ; their zygapophysial articulations are larger and present a more obliquely concave appearance than in the later genus, while the postzygapophyses are quite similar in every re- spect. The neural spine is robust and high, but on the whole proportionally lighter than in Cams. The excava- tions on the spine referred to in the description of the pre- ceding vertebra are also continued to the dorsal series, and in his description of Daphoenus (I. c, p. 339) Scott speaks of this condition as follows : " the notch between them [prezy- gapophyses] is very deeply incised, invading the base of the spine, a very different arrangement from that seen in Canis." Fio. 12. First Dorsal of V. muperbus. h Dat. size. Measurements of First Dorsal Vertebra. Mm. Antero- posterior diameter of the centrum 25 Transverse diameter of the cenlrnm 29 Vertical diameter of the centrum :... 16 Transverse diameter across the transverse processes 63 Transverse diameter across prezygapophyses 45 Transverse diameter across postzygapophyses 30 Height of neural spine 60 Second Dorsal Vertebra. — The centrum of the second dorsal is less depressed than that of the first, its anterior face being of more nearly circular outline and quite convex. Ventrally there are two prominent ridges on either side of the median keel, which extend nearly the entire length of the centrum. The transverse process and the zygapophyses are lighter than on the preceding verte- bra. The prezygapophyses face inward and slightly upward ; at their lower margins they are abruptly concave, due to sharp ridges at their bases which ex- tend diagonally upward and inward, uniting with the anterior border of the neural spine a short dis- tance above the arch. The triangular area on the anterior face of the spine, which is bounded laterally by these ridges, is quite deep and has a rugose surface identi- Fig. 13. Lateral and Posterior ViewB of Second Dorsal of D. superbus. % nat. size. 222 MEMOIRS OF THE CARNEGIE MUSEUM cal with that which has been described in the preceding vertebra, and is well shown in the illustration, Fig. 13. Measurements of Second Dorsal Vertebra. Mm. Anteroposterior diameter of tbe centrum 23 Transverse diameter of the centrum, posteriorly 27 Veitical diameter of the centrum, posteriorly 16 Transverse diameter at the transverse processes 50 Transverse diameter at the prezygapophyses 55 Transverse diameter at the postzygapophyses 22 Height of neural spine 57 Third and Fourth Vertebrz. — The third and fourth dorsals have their centra similar to that of the second. The transverse processes are lighter, the prezygapophyses are less complicated, facing directly upward, and the neural spines are less expanded at the top than in the preceding vertebra ; there are otherwise no characters which require a separate description. Fig. 14. Lateral Views of Third and Fourth Dorsals of D. sitperbw. J nat. size. Measurements of Third and Foukth Dorsal Vertebrae. Antero-posterior diameter of centra 23 Transverse diameter of centra 25 Vertical diameter of centra 16 Transverse diameter at transverse processes 47 Transverse diameter at prezygapophyses 24 Transverse diameter at postzygapophyses 20 Height of neural spine approximately 57 Fifth, Sixth, Seventh, Eighth, and Ninth Dorsal Vertebrae. — This series of dorsals are so similar to one another that they do not each require a separate description. The cen- tra are quite uniformly concave on the sides and have a ventral keel more or less indicated, while the individual in- crease in the antero-posterior diameter from the fifth to the ninth is hardly perceptible. The articular surfaces for the ribs are large and the intervertebral notches are low but FlG lg Ni of considerable antero-posterior dimensions. The transverse d. mperbus. i ith Dorsal of nat. size. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NERRASKA 223 processes decrease but very slightly from the fifth to the ninth, the articular sur- faces for the ribs are extensive on the free and expanded ends, but the excessory processes on the superior faces, which are so prominent in the recent dogs, are very inconspicuously developed. The neural spines in the whole series are heavy, but gradually reduced in height in proceeding backward, the ninth being quite thin transversely though broad antero-posteriorly. Measurements of Fifth and Ninth Dorsal Vertebrae. D. j D. 9 Mm. Mm. Anteroposterior diameter of centra 24 25 Transverse diameter of centra 24 26 Vertical diameter of centra 16 17 Transverse diameter at transverse processes 50 47 Height of neural spines 60 43 Tenth Dorsal Vertebra. — The centrum of the tenth dorsal vertebra is similar to that of the ninth, with the exception of the absence of facets for the capitulum of the rib. The transverse process is still quite heavy and has a large facet for the tuberculum of the rib, as is well shown in Fig. 16. The prezy- gapophysis is like that in the preceding vertebra, while the postzygapophysis is placed high and faces downward and out- ward, similarly to that in the recent dogs. The neural spine, however, is very characteristic in the present genus and differ- ent from that in the recent form. In Canis lupus the spine is thin transversely, quite broad antero-posteriorly, very low and fig. 16. Tenth Dorsal upright in position — a character which was apparently of D' mPerbus- I nat- size- already established in some of the early tertiary forms (Miacidse)-n In Daphwnodon superbus the neural spine of this vertebra is more backwardly inclined, and. with the exception of the postzygapophysis and the sudden reduction in size, it is like that in the true dorsals. In Canis latrans the neural spine of the tenth dorsal is ap- parently quite similar to the one here described, though much shorter in proportion. Measurements of Tenth Dorsal Vertebra. Mm. ADtero-posterior diameter of centrum 25 Transverse diameter of centrum, posteriorly 25 Vertical diameter of centrum 16 Transverse diameter at transverse processes 47 Height of neural spine, approximately 35 "Matthew, W. D., " Carnivora and Insectivora of the Bridger Basin," Memoirs Amer. Mug. Nat. Higt., Vol. IX., p. 367, 1909. 224 MEMOIRS OF THE CARXEGIE MUSEUM Elcrenth Dorsal Vertebra. — The centrum .of the eleventh dorsal vertebra is longer and also of slightly greater vertical diameter than is the case in the preced- ing vertebra and, as in tbe latter, there are no facets for ribs on the posterior face. The eleventh and twelfth dorsals in the skeleton of Daphasnus felinus (No. 492) have no facets on the posterior faces, although there are instead little rounded processes which extend slightly behind the faces of the cen- Fio. 17. Eleventh Dorsal tra, but which apparently never touched the heads of the of D. .uperbus. j nat. size. ^ ^ ^^ ^ nQ articular facetg ,loticed Oil them. The characters of the neural arch, zygapophyses, and spinous process are sud- denly changed in this vertebra, and are on the whole more like those of the true lumbars. The bone is injured in the region of the transverse process, but enough is preserved to indicate its rather small size, and it is quite doubtful whether it car- ried an articular facet for the tuberculum of the rib as in Canis and other recent Carnivora. The meta- and anapophysis are more prominent than in Cards; in fact the vertebra as a whole is heavier. Judging from the base of the neural spine it was thin transversely and rather broad antero-posteriorly ; it did not attain a great height, and was perhaps more or less anticlinal. Measurements of Eleventh Dorsal Vertebra. Antero-posterior diameter of centrum 26 Transverse diameter of centrnm 24 Vertical diameter of centrnm, approximately 17 Transverse diameter at transverse processes 37 Height of neural spine, approximately 15 Twelfth and Thirteenth Dorsal Vertebrae. — The twelfth and thirteenth dorsal vertebrae increase rapidly in size from the one preceding them. The twelfth is characterized by the practical absence of a transverse process, there being only a trace of a tubercle present near the posterior border of the pedicel immediately above the intervertebral notch. On the thirteenth dorsal is a decided knob-like transverse process which is shifted lower down than on the eleventh. The ventral keel is a heavy rounded ridge and is more prominently developed on the thirteenth than on the last vertebra mentioned. The meta- and anapo- physes are seen to be decidedly more robust in the two vertebrte here described Fig. 18. Twelfth and Thirteenth Dorsal Vertebrce of D. mpcrbus. \ nat. size. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 225 and the zygapophyses and neural spines, especially of the thirteenth dorsal, are typically lumbar in character. Measurements of Twelfth and Thirteenth Dorsal Vertebrae. d. 12 D. 13 Mm. Mm. Antero-posterior diameter of centra 28 29 Transverse diameter of centra posteriorly 26 27 Vertical diameter of centra posteriorly 17 19 Transverse diameter at transverse processes 31 33 Height of neural spine 25 First Lumbar Vertebra. — The first lumbar vertebra is slightly larger than the last dorsal. Besides the absence of facets for ribs it also differs from the latter in having a sharper ventral keel, bet- ter developed lateral tubercles on the ventral posterior face of the centrum, larger transverse processes, which are placed lower down, and a slightly heavier neural spine. In Dajihcc- nus felinus the lumbar vertebras have relatively longer and narrower centra, with the ventral keel and lateral posterior Fl°- 19- First Lumbar Ver- , • i -i n , . t . tebraof D.superbus. ' nat.size. tubercles of greater prominence, longer transverse processes, and the neural spines possibly somewhat higher than in the present genus. Measurements of First Lumbar Vertebra. Mm. Antero-posterior diameter of centrum 33 Transverse diameter of centrum 27 Vertical diameter of centrum 19 Transverse diameter at transverse processes, approximately 45 Height of neural spine 26 Fig. 20. Lumbar Vertebras of D. mperbus. J nat. size. 1, second lumbar ; 2, third lumbar ; 3, fifth lumbar. Second, Third, Fourth, and Fifth Lumbar Vertebrx. — This series of the lumbars may conveniently be described together. Their centra are broad and heavy, not unlike those in Canis, while the transverse processes, though quite heavy and directed forward, are much shorter than in the latter genus. The latter character 226 MEMOIRS OF THE CARNEGIE MUSEUM is a curious feature of this genus which seems to have departed from Daphcenus, Temnocyon,u and also from the recent canids generally, while, according to observa- tions by Dr. Matthew (/. c, p. 367) upon Miacis, its affinity with regard to the short transverse processes would seem to recall the earlier tertiary types. As in Daphcemis the metapophyses, anapophyses, and neural spines are proportionally more robust than those in the recent dogs. The meta- and anapophyses are heaviest on the first lumbar and gradually decrease in size backwards, the anapophysis of the fifth lumbar being quite small. Except on the first, second, and seventh lumbars the neural spines are broken off superiorly, but enough remains to ascertain their great anteroposterior diameters, which are not unlike those in canids generally. Measurements of Second and Fifth Lumbab Vertebra. l. 2 L. 5 Mm. Mm. Antero-posterior diameter of centra 36 40 Transverse diameter of centra posteriorly 27 29 Vertical diameter of centra posteriorly 19 ]9 Transverse diameter of transverse processes 50 72 Length of transverse processes from base (posterior) to tip end 14 24 Height of nearal spine 18 ? Sixth Lumbar Vertebra. — The sixth lumbar ver- tebra is characterized by the absence or very rudi- mentary development of anapophyses, and also by a considerable transverse broadening of the centrum posteriorly. There is otherwise very little difference between this vertebra and the one preceding it. Flo. 21. Sixth Lumbar Vertebra of D. superbus. \ nat. size. Measurements of Sixth Lumbar Vertebra. Mm. Antero- posterior diameter of centrum 40 Transverse diameter of centrum 31 Vertical diameter of centrum 19 Transverse diameter at transverse processes, approximately 84 Height of neural spine, approximately 45 Seventh Lumbar Vertebra. — The centrum of the seventh lumbar vertebra is as usual quite depressed. The transverse diameter of the posterior face is nearly equal to its antero-posterior diameter and the ventral keel and lateral ridges are promi- nent, especially on the anterior half of the centrum. The transverse process is MEyerman, J. "The Genus Temnocyon and a New Genus thereof, etc." Amer. Geol., Vol. XVII, pp. 273~ 274, 1896. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 227 unfortunately not completely preserved ; enough remains, however, to determine that it is fully as heavy as in Canis, but probably did not attain the length seen in the latter. The pre- and postzygapophyses, especially «*lfce 5- \ §Sm»tvi the latter, are much expanded in comparison with those of the preceding lumbars, and the neural spine is also quite light, although comparatively heavier than in the recent dogs. In the type of Gynodesmus brachypus the lumbar series (seven) is apparently completely represented. The centra, especially in the anterior part of the series, are somewhat FlG- 22- Seventh Lumbar less depressed than in Daphcenodon. The transverse pro- Vertebra of a s"perbus- * nat. size. cesses, though relatively longer than in Daphcenodon, are not attenuated to the same degree as in recent dogs. The metapophyses, anapophyses, and neural spines are proportionally lighter than in Daphcenodon and on the whole more nearly resemble the lumbar vertebrae of Canis. Measurements of Seventh Lumbar Vertebra. Mm. Antero-posterior diameter of centrum 34 Transverse diameter of centrum posteriorly .... 31 Vertical diameter of centrum posteriorly 18 Transverse diameter at transverse processes, approximately 72 Height of neural spine 33 Fig. 23. Dorsal, Lateral, and Ventral Views of Sacrum of D. super, i nat. size. Sacrum. — The sacrum is greatly developed, and, as usual, it consists of three well coossified vertebrae. In its relative size it is fully equal to that of Daphcenus described and figured by Professor Scott (I. c, p. 341, PI. XX, fig. 14), and is very much larger than in the recent dogs and proportionally somewhat more robust than in the lion or the tiger. The centra are unusually massive and not nearly so depressed as in the recent dogs, but resembling more those of the tiger. The transverse proc- esses however, are, very differently arranged, only the first sacral supporting the ilium by its very heavy pleurapophyses ; the second has the transverse process rep- 228 MEMOIRS OF THE CARNEGIE MUSEUM resented by a comparatively delicate roughened ridge, while the third has a promi- nent process which extends laterally and slightly back of tbe posterior face of the centrum. In tbe recent forms the transverse processes of the sacrum are more strongly developed, the first and second sacrals, especially, taking a more prominent part in the support of the ilium. In comparison with the heavy centra the neural canal is rather small, thus agreeing with what is observed in the Oligocene genus and differing from recent forms. The neural spine of the first sacral is relatively smaller than in the recent dogs or cats, while that of the last sacral is much heavier, the spine on the median vertebra consisting only of a sharp low ridge, which uninter- ruptedly connects with the spines in front and behind. In Cijnodesmus brachypus the sacrum is shorter and broader, the second sacral helps to support the ilium more prominently and the anterior neural spines were apparently proportionally more robust than in Daphcrnodon. The spinous process on the last sacral in C. brachypus however, is, quite prominent, and is more com- pletely separated from the spine in front of it than in either Daphcenodon or the recent dogs. Measurements of Sacrdm. Mm. Antero-posterior diameter of centra 77 Greatest transverse diameter across the pleurapophyses 59 Greatest transverse diameter at transverse processes of last sacral 49 Caudal Vertebrx. — The tail is very long and heavy ; it has undergone practi- cally no change from that of the Oligocene form and is practically as well developed as in the lion or the tiger. The anterior face of the centrum of the first caudal is more con- V'N^P/^ vex and less depressed than the posterior. The transverse process is very heavy and pro- jects well backwards. The pre- and postzyga- pophyses have perfectly formed articular facets and are well expanded transversely. The neural canal is of moderately large size and ^g3 ^T^?C2?28 tne spinous process is quite robust. fig. 24. candai Vertebra of d. mperbm. j The length of the centra of the succeeding nat. size. (1-28 indicate their respective posi- two Caildals agrees with that of the first. The tions in t e i . ) transverse process on the second however, is, broader antero-posteriorly than in the first, but the neural spine is very much reduced, while on the third caudal the neural spine is represented only by a low ridge. The centrum of the fourth caudal is four or five millimeters longer than PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 229 that of the first, while back of this vertebra the increase in length is more noticeable. The transverse processes are single up to the sixth, and the seventh caudal still bears the pre- and postzygapophyses with perfect articular facets in front and behind, while on the eighth and ninth the postzygapophyses are more or less imperfect, but the neural canal is complete and extends throughout nearly the entire length of the vertebrae. The last mentioned vertebra* is the longest in the series which was at- &*HA?a tached to the sacrum when the specimen was found, but there is reason for believ- ing that the vertebra continued to increase in length up to the eleventh or twelfth caudal. There are seven chevron bones present which were found more or less in position in the series of caudals here described. At the anterior portion of the tenth caudal the skeleton was exposed and a section of the tail dropped out. As mentioned in the introduction, the natural curve in which the tail undoubtedly was lying would allow room for six or possibly seven vertebrae. Working on this hypothesis the next vertebra which was found in position in the skeleton would be the seventeenth or eighteenth. This vertebra has the pre- and postzygapophysial prominences, anterior transverse processes, and ventral tubercles for chevrons well represented, and the centrum is of considerable length and robustness. The ten succeeding vertebras gradually decrease in length and robustness, the last being only seven millimeters long. In the type of Cipwdcsm.us brachypits the anterior four caudal vertebrae are pres- ent together with various others which represent different parts of the tail. The transverse process on the first is very long and robust, while on the second it is much shorter and indicates a greater decrease backward than is seen in Daphienodon. The centra are less depressed than in the latter genus and approach more closely those in the dogs. It is quite likely that the tail in C. brachypus when completely known will be found to be proportionally shorter than in Daphcenodon superbus. Measurements of Caudal Vertebra. C. 1 C. 2 C. 9 C. 17 C. 28 Mm. Mm. Mm. Mm. Mm. Antero-posterior diameter of centra 23 24 46 35 7 Transverse diameter of centra posteriorly 22 20 16 8 2 Vertical diameter of centra posteriorly 15 14 14 9 2 Transvere diameter at transverse processes 57 67 26 12 Ribs and Sternum. There are very few ribs represented with the type specimen, while in No. 1589a (the young individual found with the type) they are quite complete. The series as a whole is rather short and stout with much expanded ventral ends. In the ante- 230 MEMOIRS OF THE CARNEGIE MUSEUM rior region of the thorax the ribs have large capitular and tubercular articulations and rather flat shafts as in Canis, while pos- teriorly they are more cylindrical ; the elev- enth, twelfth, and thirteenth having no tuber- cular articulations. Sternum. — The sternum is represented only by the manubrium, which is more nearly like that in the tiger than in the dogs. Thus its size is proportionately large and it is greatly produced in front and much expanded trans- versely at the attachment for the first pair of ribs. In front of the rib facets the superior face has a subtriangular area which is concave from side to side and also slightly concave an- tero-posteriorly. In front the bone terminates in an obtuse, laterally compressed process, which projects forward and slightly down- Fig. 25. Ribs of Left Side of D. superbus. J nat. size. 1, first rib, type No. 1589 ; 2, second rib No. 1589a; 3, third rib No. 1589a ; 5, fifth rib No. 1589a ; 6, sixth rib No . 1589a. ward. The ventral face is produced into a heavy rounded antero-posterior ridge, which gives the bone a strong keel-like appearance. The posterior half of the presternum is more angular on cross-section. Fig. 26. Presternum of D. superbus. * nat. size, dorsal view ; 2, ventral view ; 3, lateral view. Measurements of Presternum. Greatest length ' Transverse diameter at first rib facets Vertical diameter at first rib facets The Os Penis. The os penis which is so well developed and characteristic in Daphcenus is not less so in Daphamodm. With the type and the younger skeleton there was found the anterior portion of an os penis (Fig. 27, 2) which I judge might belong with the young individual. The portion of the shaft present is sub-cylindrical m cross- section, with a sinuous curve and deeply grooved near the bisected end. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 231 In the material belonging to a skeleton of Daphcenodon in Amherst College, Massachusetts, now in the Carnegie Museum,15 there is an os penis of very large size, and also different in its general shape from that described above (see Fig. 27, 1). Fig. 27. Os Penis of D. superbus. h nat. size. 1, superior and lateral views of os penis, Amherst specimen ; 2, superior and lateral views of os penis with skeleton No. 1589a, Car. Mus. Cat. Vert. Foss. Thus its shaft is more elliptical in cross-section, especially toward the posterior end, and it has a less complex curvature at the anterior end. It is, however, deeply grooved near the anterior end as in the smaller specimen, though not bisected, but suddenly expanded to a round spout with many small tubercles on the anterior border. How much longer this bone was is rather difficult to say, but it would seem that it is broken off immediately in front of the rugose surface for the attach- ments of muscles to the pubes. Whether the larger bone represents the fully adult stage of development or pertains to another species cannot now be fully determined. If, on the other hand, the two different bones represent fully adult forms there is apparently a great range of variation of this bone in this genus. The Fore Limb. (Plates LXXVIII-LXXX.) Scapula (Plate LXXVIII). The scapula is well preserved and completely represented in the type specimen as well as the younger specimen found with it. This important bone of the fore limb displays a number of rather surprising features and probably sheds considerable light on the characters of the scapula in Daphwnus, which is not yet known. The bone as a whole reveals a curious combination of the features of the scapula of the tiger and the bear, and shows less similarity to that of the Canidae. As in the tiger, the bone is elongated, with sub-equal pre- and post- scapular fossse, the coracoid border is suddenly expanded and again contracted "On further exoavation by the writer in the quarry where Prof. Looinis of Amherst found the skeleton, there were found various parts which belong to the Amherst speoimen and will be forwarded to that institution. 232 MEMOIRS OF THE CARNEGIE MUSEUM before the suprascapular border is reached, and the acromion process on the spine is well developed, while the postscapular fossa is unequally divided by the prominent and sharp glenoid border, identically as in the bears. In the fox the scapula has a small area developed for the teres major muscle at the superior portion of the glenoid border, but it is quite small in comparison with that in the present genus, in which the scapula as a whole is proportionally broader transversely. The spine is high and terminates in an overhanging acromion, which, however, does not project below the glenoid cavity. Judging from the heavy acromion process there was undoubtedly a clavicle of perhaps considerable development. The glenoid cavity is not so large proportionally as in the large cats, and in this respect is rather more like that in Canis. The coracoid process also has about the same proportionate development as in Canis and is relatively much shorter than in the tiger. Professor Scott states (/. c, p. 381) that in Cynodktis "the coracoid process is unusually large ..." that the " prominent acromion descends below the level of the glenoid cavity ..." and that in general character the scapula " is rather viver- ine or raccoon-like than canine," which is not true of the scapula in Daphcenodon. In Tcmnocyon ferox, according to Eyerman's description w " the greatest vertical length [of the scapula] is from the coracoid to the posterior upper edge and not along the spine, as in C. familiari.s. This is not the case in the present genus, nor is the spine "falling short of the supra-scapular border," as in the John Day genus, but in this respect it is more nearly like the dogs. It is interesting to note that the area for the teres major muscle on the upper part of the glenoid border of the scapula in Hoplophonews is also quite large, and is quite similar to that in Daphcenodon. Measurements of Scapula. Mm. Greatest height from gleDoid cavity to suprascapular border 167 Greatest antero-posterior diameter 104 Antero-posterior diameter of glenoid cavity, including the coracoid process 39 Transverse diameter of glenoid cavity 23 Humerus (Plate LXXIX). — The humerus has apparently changed very little in its general characters from that of the Eocene forms. The slight shortening of the bone and the less developed supinator ridge, together with the somewhat greater development of the internal epicondyle in the present form are, the only noteworthy differences from those in the Oligocene genus Daphccnus. The tendency to a reduction of the supinator ridge in Daphcenodon is a distinct step from the early "Eyerman, J. The American Geologist, Vol. XVII, p. 274, 1896. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 233 types " toward the recent canids, while the bone on the whole tenaciously retains features met with in the early progenitors ; viz., the presence of the entepicondylar foramen, the heavy deltoid ridge which extends well down on the shaft, and many other minor details. In comparing the humerus with that of the tiger the similarity is surprisingly close, the deltoid crest of the recent form being proportionally even less developed, while the distal trochlea is transversely somewhat broader in proportion. The Fig. 28. Radius and Ulna of D. superbus. $ nat. size. humerus of Amphicyon major of Europe has also a greater transverse diameter distally and the supinator ridge is more rounded and heavier than in the genus under description. Measurements of Humerus. Mm. Greatest length 211 Anteroposterior diameter of head 52 Transverse diameter of head at tuberosities 44 Greatest transverse diameter of distal end 55 Greatest transverse diameter of distal trochlea 34 Greatest anteroposterior diameter of distal end 32 Radius and Ulna (Plate LXXIX). — The radius is proportionally longer and the humerus shorter in Dap>hocnodon than in Daphcenus. It is also observed that the shaft of the bone in the former genus has a less sinuous curvature than in the latter ; in other respects there are only minor details of difference in the two genera, and the feline characters are practically as prominently shown in the later as in the earlier form, i. e., the inner margin of the head overhangs the shaft as in the cats, the articular surface with the lesser sigmoid cavity of the ulna extends well around on the internal side of the head, and the tubercle for the biceps muscle is very prominent, while the distal end is quite broad transversely and displays well marked tendinal grooves, which, however, are not as well developed as those in the tiger. The radius in Amphicyon major is also quite cat-like, though the tendinal grooves on the inner angle of the distal end are apparently less developed than in Daphccnodon. " Wortman, J. L., "Study of Eocene Mammalia," etc., American Journal of Science, Vols. XI-XIV, p. 348, Fig 9, p. 438, 1902. 234 MEMOIRS OF THE CARNEGIE MUSEUM The olecranon process of the ulna is very short, truncated, and more inwardly produced than in the Oligocene genus. The tendinal sulcus, though well denned and wide, is not so deep as in Canis lupus, and consequently much less developed than in the cats. The principal feature of the greater sigmoid notch is its heavy coronoid process, which character is feline rather than canine. The shaft is very heavy and not much curved. The tuberosity for the flexor profundus digitorum muscle on the external border of the shaft is very heavy, as is well shown in Figs. 5 and 6, PL LXXIX. Distally the shaft is more trihedral in cross-section due to the sharp ridge which separates the internal and posterior surfaces as in the tiger. The articulation for the radius on the distal end is a heavy rounded knob much as in the large cats and entirely unlike that in the dogs. The styloid process ex- tends well down and has a decided constriction above the articulation for the cuneiform. In Cynodesmus brachypus both ulnae are represented. The olecranon process is short and truncated, the tendinal groove is deep as in Canis, but the coronoid process is larger in proportion than in the recent dogs, as is also the shaft. The distal end, however, lacks the heavy and broad development of .the rounded tubercle which articulates with the radius and is so conspicuous in the genus under descrip- tion. The feature of this reduction of the ulna in C. brachypus is apparently a decided step towards conditions found in Canis. Measurements of Radius and Ulna. Mm. Radius, greatest length 184 Radius, antero-posterior diameter of proximal end 16 Radius, transverse diameter of proximal end 23 Radius, transverse diameter of shaft medially 15 Radius, antero-posterior diameter of shaft medially 12 Radius, greatest antero-posterior diameter of distal end 23 Radius, greatest transverse diameter of distal end 34 Ulna, greatest length 223 Ulna, antero-posterior diameter at coronoid process 32 Ulna, transverse diameter at coronoid process 25 Ulna, transverse diameter of distal end at tubercle 20 Ulna, antero-posterior diameter of distal end at tubercle 11 Ihe Manus (PI. LXXX). — With the exception of the trapezoid the manus of the type specimen is quite completely represented. As the forefoot was found more or less disarticulated the respective positions of the phalanges as determined by the writer may be somewhat conjectural ; their positions are, however, suffi- ciently correct for a general description. In comparing the scapho-lunar bone with that of Da/phcen/us felinus it is found PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 235 that the similarity is remarkably close. The dorsal face of the bone in the present genus has a less vertical diameter than in the Oligocene form, which is due to the fact that tbe articulation for the radius is carried further downward on the anterior face than in the older type, in fact lower than in the lion and tiger, consequently far removed from conditions in the recent dogs, in which the dorsal face of the scapho-lunar is very large, the bone being much higher throughout. The articular facet for the radius also extends far downwards on the palmar face £g?$ij near the ulnar angle, as in the lion and tiger, ^\fe%- which indicates that there was even more power of flexure between these two bones (radius and scapho-lunar) in the present genus than in Dapluxnus, and is a condition strikingly similar to what is seen in the large cats. The radio- palmar process is well developed, but has not the vertical diameter seen in Canis, and is more nearly like that in the cats. It is, how- t.rlST ever, more distinctly separated, by a constric- „ ,, , , r ,, , . . Fig. 29. Scapho-lunar (left sides) of D .mper- tion, from the main body of the bone than m . „ . . . , , . „ , J bus. Nat. size. 1, dorsal view ; 2, palmar view; the felines, which is due to the greater devel- 3, radial view; 4, ulnar view; 5, inferior view; opment of the palmar face of the scapho- 6' 8UPeHor view- lunar in the latter. The ulnar face has no facet for the cuneiform, which is identical with that in Daphcenus, in which the cuneiform and scapho-lunar did not come in touch with one another sufficiently to establish facets. Distally there are well defined facets for the unciform, magnum, trapezoid, and trapezium respectively. The facet for the unciform is deeply excavated as in Daphcenus, while that for the magnum is extended downward somewhat less near the dorsal border, the beak of the lunar being less produced downward in the present genus. The facet for the trapezoid is a large, irregularly shaped, and obliquely placed surface, indicating a trapezoid of considerable size. The facet for the trapezium is located on the extreme radial angle, but is poorly demarcated from the facet for the trapezoid. Measurements of Scapho-lunab. Mm. Greatest antero-posterior diameter, including the radio-palmar process 29 Greatest transverse diameter 29 Greatest vertical diameter near dorsal face 19 Cuneiform. — As in Daphcenus the cuneiform is broad laterally, much depressed in the vertical direction, and has a large process which occupies the dorso-ulnar 236 MEMOIRS OF THE CARNEGIE MUSEUM face of the bone, and furnishes muscular attachment for Mc.V, but does not articu- late with the latter as in the dogs. The facets for ti@$>0m KS mj the ulna and pisiform on the proximal face are sub- equal in size and are separated by a prominent ridge, J,\ the former being more concave antero-posteriorly sp Mr than the latter. In the dogs the facet for the ulna is 4 different, being more sinuously curved, with an antero- posterior cavity on the ulnar angle and an antero- gs^ss«a posterior convexity on the proximo-dorsal angle of fig. 30. cuneiform (left side) of i). the bone. The distal face is taken up by a large superb™. Nat. size. i, dorso-nioar facet which is concave in all directions and articu- view ; 2, dorsal view ; 3, inferior view ; lates ^ the unciform. The bone as a whole is not 4, superior view ; 5, palmar view ; 6, paimo-radiai view. Ullllke the cuneiform of the tiger. Measurements of Cuneiform. Mm. Greatest antero-posterior diameter 14 Greatest transverse diameter 20 Greatest vertical diameter of dorsal face 8 Pisiform. — In its general proportions the pisiform l^v^^^^ is quite robust ; it is proportionally slightly heavier than in the tiger, but has a relatively shorter shaft. "^P"*"- ° 2 The articulations for the ulna and cuneiform are sub- equal in size and thus quite unlike the corresponding surfaces in Canis, in which the articulation for the cuneiform is much the larger of the two. As stated IG' l- ls'orm D. superbus. Nat. size. 1, ulnar above the shaft is short and heavy and terminates in and anpeiior views; 2, view of ar- an expanded and truncated free end with the ver- ticuiar facets of cuneiform and ulna ; , ■,. .-, .1 . c.u.cuneiform and ulua facets, re- tical diameter greater than the transverse. . , c spectrvely. Measurements of Pisiform. Mm. Greatest diameter 23 Antero-posterior diameter of articular facets 14 Vertical diameter at articular facets 9 Vertical diameter at free end 11 Trapezium. — The trapezium is of moderately large size. Its dorsal or dorso- radial face is considerably exposed (when viewed from in front), and its general out- lines are pentagonal, while on the palmar or postero-ulnar angle there is a heavy PETERSON : NEW CARNIVORES PROM MIOCENE OF WESTERN NEBRASKA 237 process for attachment of muscles. On the ulnar face is a large facet for the trapezoid, which extends well back, cov- ering the base of the palmar process, referred to above. On the proximal radial angle there are well formed facets for the scapholunar ; the one on the ra- dial angle is scarcely separable from the one on the palmar process. Distally there is a large obliquely placed facet for Mc. I which is concave from before backwards Fig. 33. Trapezium (lm-side)of D. superbm. N size. 1, dorsal view ; 2, nrmnr view ; 3, rrfenwf view; KwV'CA- pdnan view ; lufenui ' Una/' Measurements of Trapezium. Mm. Greatest autero-posterior diameter 12 Greatest transverse diameter 7 ; vertical diameter 10 E*vv; 1-4 Trapezoid. — The trapezoid is not known, but judging from the space in the car- pus when the trapezium is placed in position it would seem that the trapezoid is larger than the trapezium which is the reverse of what is seen in Daphasnits, studied by Mr. Hatcher (I. c, p. 86) and compared by the present writer. Magnum. — The magnum has a small dorsal face, but is in general a much larger bone than is indicated by this sur- face. The dorso-radial angle has a small facet for the trapezoid and this region of the bone is much depressed, while the radial face has a great vertical diameter, due to the sharp ridge which extends up- ward, articulating radially with the scapho-lunar, and on the ulnar face with the unciform. In Cam's the ulnar face has a large dorsal and a smaller palmar facet for the unciform ; the two facets being separated by a deep groove, while in Daphccno- don the facet is uninterrupted and surrounds an excavated area in the middle and posterior portions of the ulnar face. The radial face of the magnum is also deeply excavated. The palmar tuberosity is of considerable vertical diameter, while trans- versely it is compressed. Distally the bone is occupied by two facets, the larger for Ltrt FlG. 32. Magnum (right side) of D. superbus. Nat. size. 1, dorsal view; 2, p»d4rf-view ; 3, superior view; 4, pootorior view ; 5, »4imr view ; 6, inferior view. Palmar \r&ejvr- Scpi^or- 238 MEMOIRS OF THE CARNEGIE MUSEUM Mc. Ill and the smaller for Mc. IV. In Daphasnus the latter facets are divided in about the same proportions as in the present genus. Measurements of Magnum. Mm. Greatest antero-posterior diameter 20 Transverse diameter of dorsal face 11 Greatest vertical diameter 16 Unciform. — The facet for the cuneiform does not extend so far down on the ulnar angle as is the case in Daphcenus, otherwise I cannot see any difference in the unciform in the two genera. On the dor- sal face the bone has a transverse exca- n. ,JL lit H "It vation and lacks the heavy convex ru- gosity for muscular attachments seen in the tiger, nor is the palmar tuberosity so 'ifjSUli well developed ; otherwise the bones in t'"ih ( 1111 the two forms are not greatly dissimilar. From Canis the unciform in Daphoenodon is at once distinguished by the lateral Fig. 34. Unciform (left side) of D. superbm. Nat. ... „ ., „ . » ., .. , , , , . „ , , position of the facet lor the cuneiform, the size. 1, dorsal view ; 2, palmar view; 3, superior view; r 4, ulnar view ; 5, radial view ; 6, inferior view. latter facet being more nearly superior on the bones in the true dogs. The radial face is deeply excavated by a sulcus which corresponds to the one described on the ulnar face of the magnum. Around this sulcus is a large continuous, although ir- regularly shaped, facet for the magnum, which is altogether different from the four separated facets for the magnum in Canis. Immediately above the facet for the magnum there is a second facet, which articulates with the scapho-lunar. The ante- rior portion of this continues into the facet for the magnum in an imperceptible manner, while further back there is a decided ridge separating the two facets. Dis- tally the bone is entirely taken up by the large facets for Mc. IV and V. Measurements of Unciform. Mm. Greatest antero-posterior diameter 16 Greatest transverse diameter dorsally 13 Greatest vertical diameter 17 Metacarpal I. — In their general characters the metacarpals are feline rather than canine. The first metacarpal is of approximately the actual length of that in an adult specimen of Canis lupus, but it is heavier, its proximal end is more expanded, PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 239 and the articulation for the trapezium is broader. In comparative size the bone is very little smaller than in Daphcenus and consequently much heavier and longer than in Cards. On the ulnar face the rugosity for the ligamentary attachment to Mc. II is quite prominent and the tuberosity on the radial side of the head is as large proportionally as in recent cats, though the bone as a whole however, is, lighter and longer than Mc. I of the pollex in the latter. The distal end has a well formed facet which is separated palmarly by a prominent keel. The shaft is slightly curved, so that the distal end points a little in the ulnar direction. Measurements of Metacarpal I. Fig. 35. Dorsal and Ra- dial Views of Left Metacar- pal I of D. supeibus. Nat. size. Greatest length 32 Greatest antero-posterior diameter of head 10 Greatest transverse diameter of head 11 Metacarpal II — While the actual length of the first metacarpal is equal to that in Canis the second, third, fourth, and fifth are much shorter. This gives a char- acteristically low and broad appearance to the manus, as in Meles or perhaps even more similar to that in Arcticfis. Thus the second and fifth are of subequal length and are considerably shorter than the third and fourth. The head of Mc. II has the antero-posterior diameter much greater than the transverse, which is due to the comparatively small extent in the ulnar direction of the flange which carries the facet for Mc. III. In the cats this flange is very prominent on Mc. II and III, and forms a more completely interlock- ing condition of the metacarpals than is seen in the present Fio. 36. Dorsal and Radial genus and in the recent clogs generally. The shaft is sub- Views of Left Metacarpal ii cylindrical in cross.section and strongly arched in the dor- of D. superbus. Nat. size. sal direction. The distal end is suddenly expanded and the trochlea is more cat- than doe-like. Measurements of Metacarpal II. Mm. Greatest length 51 Greatest antero-posterior diameter of head 16 Greatest transverse diameter of head 16 240 MEMOIRS OF THE CARNEGIE MUSEUM Metacarpal III. — The third metacarpal is slightly longer than the fourth and is consequently the longest in the series. The proximal articulations of this meta- carpal as well as all the others are so placed that when articulated the distal ends spread very much more than in the recent dogs. This is also true of the metacarpals in Daphmius. As in the latter genus the dorsal face of the proximal end has a rugose excavation for tendinal attachments, while that area in Canis is raised higher up and in place of the excavation there is a considerable eminence. On Mc. Ill in Cynodesmus brachypus this eminence is also well developed. The shape of the dif- ferent facets of the proximal end are identical with those in Daphcenus ; i. e., the facet for the magnum is narrow and deep, that for Mc. II larger and more oblique Fig. 37. Dorsal and Radial Views of Right Metacarpal Fig. 38. Dorsal and Radial Views of Right Metacarpal III of D. superbus. Nat. size. IV of D. mperbui. Nat. size. than in Canis, and the facet for Mc. IV larger and more angular near the proximal border. As in Mc. II the shaft is round and arched forward. The distal trochlea is narrower than in the dogs and the dorsal portion is more spherical, while the carina is very prominent. Measurements of Metacarpal III. Mm. Greatest length 62 Greatest antero-posterior diameter of head 14 Greatest transverse diameter of head 11 Metacarpal IV. — The general features of the head of Mc. IV are not unlike' those of the third. The shaft, in fully adult forms however, is, characterized by the prominent rugosity for tendinal attachment on the ulnar angle, while in young individuals it is less noticeable. The shaft is arched in the dorsal direction and the distal end is similar to that of Mc. III. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 241 Fig. 39. Dorsal and Radial Views of Left Metacarpal V of D. super- bus. Nat. size. Measurements of Metacarpal IV. Mm. Greatest length 60 Greatest antero-posterior diameter of head 15 Greatest transverse diameter of head 12 Metacarpal V. — The fifth metacarpal is as heavy as the fourth, but it is shorter, as has been stated before. The proximal end is much expanded, especially in the ulnar direction, which is due to the heavy and rough tuberosity for the attachment of ligaments and muscles. The articulation for the unciform is regularly convex from before backward, the transverse diameter of the palmar area being greater than that near the dorsal face. In Daphoemis this articulation is pro- portionally broader, but the ulnar face of the head is not as prominently, developed as in the genus under consideration (see Fig. 39). The dorsal face of the shaft faces to the ulnar side of the manus as well as forward, and the shaft is arched forward in approximately the same proportion as that of Mc. I. The distal end is expanded and points slightly in the ra- dial direction when the bone is in position in the manus. The articulation for the proximal phalanx is similar to those on the metacarpals described above, except that the facet on the ulnar side of the pal- mar carina is placed higher up and is of larger size than the one on the radial side. Measurements of Metacarpal V. Mm. Greatest length 47 Greatest antero-posterior diameter of head 13 Greatest transverse diameter of head 13 Phalanges. — The proximal phalanges are proportionally somewhat shorter than in the Oligocene genus, otherwise they bear a close similarity in every respect. They are broad, depressed, strongly arched in the dorsal direction, com- paratively gently convex dorsally, and rather flat on the plantar faces. The proximal ends are deeply pitted for the distal ends of the metacarpals and the tubercles for ligamentary attachments are very prominent. The distal troch- lea is well formed, but is confined almost entirely to the plantar face. Fio. 40. Phalanges of Digit V (left forefoot) of D. superbw. Nat. size. 1, dorsal aud radial views of first phalanx ; 2, dorsal and radial views of second phalanx ; 3, plantar and radial views of terminal phalanx. 242 MEMOIRS OF THE CARNEGIE MUSEUM The phalanges of the second row are also proportionally shorter than those in the older genus, but display the same curious depression and asymmetrical outlines. Perhaps the most curious feature of these phalanges are their articulations with the terminal phalanges. In the dogs these articulations are decidedly concave from side to side and convex from the dorsal to the plantar faces, while in the present form they are very much less concave in the plantar region and inclined to be convex from side to side at the dorsal faces, like those in the cats. The ungual phalanges are high, not very long, and compressed laterally. There is a plainly marked groove on the plantar faces near the distal ends of the claw- plates. The latter rise to a sharp ridge dorsally and are of slightly greater trans- verse diameter on the plantar faces. Proximally the claw-plate is surrounded by a heavy hood, which terminates palmarly in a heavy mass together with the subungual process. The latter extends well back under the distal articulation of the middle phalanx, when articulated with the latter, and supported a strong tendon of the flexor profundus digitorum. Dorsally the hood does not project backward so as to overhang the articulation for the median phalanx as in the cats, but the hood in this region is nevertheless of considerable size and has apparently a rugose area for the attachment of some retractal ligaments. The articulation for the median phalanx is biconcave as in the felines. After a cai'eful study, there does not seem to be any reason to doubt that this animal had some power of retracting its claws, and in this respect it is. identical with Daphccnus. In Amphicyon major Mc. III? is proportionally short, the shaft is more greatly arched dorsally, the palmar tuberosity is heavier and the distal trochlea is more hemispherical on the dorsal face than in Ddphcenodon, which altogether gives even a more feline look to the manus, so far as we know it, in the European species. In Cynodesmus brachypus the only bones present which represent the manus are the magnum and Mc. III. The former is decidedly more canine in character than the corresponding bone in Daphasnodon, being less depressed in front, having the ascend- ing ridge, which articulates with the scapho-lunar, of much less vertical and greater transverse diameter, and the distal articulation more sharply notched on the radial border. Mc. Ill in Cynodesmus is shorter and lighter than in Canis latrans, but the bone is otherwise quite like the latter, especially the proximal end, while its length is about 12 mm. shorter than in Daphanodon and also considerably lighter. A second phalanx, which undoubtedly belongs with the type of Cynodesmus brachypus, is quite depressed, but its distal trochlea is concave from side to side and convex from in front backward, as in Canis. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 243 Fig. 41. Internal View of Right Half of Pelvis of D. superbus. I nat. size. The Hind Limb. (Plate LXXVIII and Plates LXXXI-II.) The pelvis of the type is represented by the greater part of the ilium, while the ischium and pubes are represented only by the portions surrounding the acetabu- lum. In a larger specimen of Daphcenodon which belongs to the Amherst College the innominate bone is more nearly complete (see PI. LXXVIII), but the posterior portion of the ischium is also lacking in this specimen. In comparing the pelvis with Professor Scott's description of that in Daphasnus (I. c, pp. 349-350) it is evident that the similarity in the two genera is quite close. As in the Oligocene genus the gluteal surface of the ilium is divided by a prominent longitudinal ridge, unlike that in recent dogs, or cats where the surface is more evenly concave. The ilium is less expanded at the supra-iliac border than in Canis, but probably somewhat more so in comparison with that in Daphcenus. The ilium, as a whole, is relatively broader than in the tiger or lion, and is not unlike that in Viverra zivetta, as was also observed by Professor Scott in his study of Daphasnus. The attachment for the sacrum is quite heavy and is situated well back. The ischial border and the great sacro-sciatic notch is short. The iliac sur- face is not well defined on account of the short pubic border ; the ilio-pectineal eminence however, is, well developed. The acetabular border is very prominent and terminates in a large tubercle at the anterior border of the acetabulum. The latter is of moderate size and depth, the center of the floor being occupied by a large depressed and roughened area, which indicates the attachments of a large cushion of fat as well as the ligamentum teres of the femur. The cotyloid notch is quite broad and excavates the ischium deeply. As in Daphcenus the ischium is long in proportion to that in the recent dogs and even longer proportionally than in the tiger or lion; the obturator foramen consequently has a more oblong outline than in the above mentioned genera and the pelvis as a whole has a greater length. 244 MEMOIRS OF THE CARNEGIE MUSEUM In Cynodesmus brachypus the pelvis is practically complete and again displays characters which are entirely unlike those in Daphamodon and are more like those in the recent canids. The ilium, while not as much expanded at the supra-iliac border as in Canis, has a tendency to become more generally concave on the external face, the attachment for the sacrum is more anterior, and the ischium and pubes are decidedly shorter than in the genus under description. Measurements of Pelvis. Mm. From supra iliac border to acetabulum, approximately 105 Width of ilium at posterior inferior spiDe 41 Widtb of ilium at great sacro-sciatic notch 30 Antero-posterior diameter of acetabulum 28 Vertical diameter of acetabulum 25 Femuj LXXXI). The femur of the type is not represented except by surface fragments which may or may not belong to this individual. The younger specimen (No. 1589a) which was found with the type has the shafts of both femora with the proximal ends lost and the distal ends slipped off at the epiphysial su- tures (see fig. 42). A third .and larger specimen, which belongs to Amherst College, has the femur well preserved. This complete femur will here be used in connection with the description of the type and the material of the younger individual in the Carnegie Museum (see PI. LXXXI, figs. 1 and 2). The length of the femur cannot be accurately ascertained, but I judge that it is approximately equal to that in an adult specimen of Cam's lupus. The head is rather small and hemispherical, and the neck is moderately long. The pit for the ligamentum teres is deep and located on the posterior half of the head nearer the inferior than the superior bor- der. When the bone is held in position the head is directed upward and inward in approximately the same proportion as is the case in the recent dogs, and as is also the case in Daphcenus. The transverse bridge from the head to the great tro- chanter is quite heavy on its posterior face, which causes the digital fossa to Fig. 42. Dorsal and Fibular Views of Left Femur of Young Individual No. 1589a. ' nat. size. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 245 appear quite narrow transversely and crowded over to the fibular side of the posterior face. The fossa is not very deep and is partly surrounded by the great trochanter. The latter is not as prominent as in Canis, which may be due to individual variation, as I find in the Amherst specimen that the digital fossa and greater trochanter are more nearly of the proportionate size seen in the recent form. The lesser trochanter is prominent and located on the posterior face near the tibial angle as in Canis, while in the Amherst specimen it is situated more directly poste- riorly. These differences may be due to individual or possibly to specific varia- tions. The third trochanter is not present ; while in Daphoenus it is apparently represented, though small ; in fact Mr. Hatcher in his description 1R did not regard this eminence as being the third trochanter, but in his illustration (I. c, PL XIX, fig. 1) he faithfully indicates the tubercle as it appears on the specimen. Dr. Eyer- man definitely states that Daphoenus has a third trochanter,19 while Professor Scott -° did not have material of this genus sufficiently complete to fully demonstrate its pres- ence or absence. The shaft of the young individual referred to above is rather slender, which is due to the immaturity of the animal and must not be regarded as an estab- lished character. The shaft of the Amherst specimen is long, slender, arched forward, and has a very prominent and rugose ridge on the fibular angle, which extends nearly throughout the entire length of the shaft, as in the lion. The linea aspera is also well defined. In No. 1589a the lower portion of the shaft broadens more gradually toward the distal end than is the case in the Amherst specimen, which again may be due to the immaturity of the specimen in the Carnegie Museum. The distal end of the latter has slipped off, as stated above, and as also indicated in the accompany- ing figure (fig. 42). The condyles in No. 1589a are moderately expanded laterally and posteriorly, which is a feline rather than canine character, while in the Amherst specimen the condyles are well extended back of the vertical line of the shaft, as in Canis. These variations may possibly be due to the differences in age of the two specimens. The rotular trochlea is no wider in proportion than in Canis, but is somewhat shorter. The intercondylar notch is quite deeply excavated and slightly oblique in position. Patella. — This bone is not very well represented in the material of the Carnegie Museum. One patella which was found among the surface fragments is proportion- ately narrower and thicker than that bone in Daphoenus and, though larger, is quite similar to that in Canis. I8"01igocene CanidaV' Memoirs Carnegie Museum, Vol. I, pp. 88-89, 1902. 19 American Geologist, Vol. XIII, p. 279, 1896. t0 Trans. Amer. Pliilos. Society, Vol. XIX, p. 350, 1898. 246 MEMOIRS OF THE CARNEGIE MUSEUM Tibia (PL LXXXI). — The tibia is represented by the distal ends and the greater portions of the shaft of the right and left, but the proximal ends were unfortu- nately broken off and lost. The shaft is rather heavy and was apparently quite short. The cnemial crest extends well down and the shaft immediately below this crest becomes triangular in cross-section, with the acute apex of the triangle directed forward. The areas for the different muscles are well defined by distinctly marked ridges, the one on the posterior face which separates the flexor longus digitorum from the flexor longus hallucis being especially prominent. On the distal end the groove for the tendon of the tibialus posticus is well developed, and the internal malleolus is very large as in Daphmius. The astragalar articulation is not so deep as in the recent dogs ; it is also more oblique, and does not have the transverse sulcus which interrupts these facets in Canis. On the fibular face is a well formed facet for the distal end of the fibula. Measurements of Tibia. Mm. Greatest length, approximately 200 Antero-posterior diameter of shaft at lower end of cnemial crest 25 Transverse diameter of shaft at lower end of cnemial crest 17 Greatest antero-posterior diameter of distal end 19 Greatest transverse diameter of distal end 34 Fibula (PI. LXXXI). — Asin the tibia the proximal end of thefibula isalso broken ofi and lost. In proportion the shaft is somewhat more slender than is the case in Daphanus, and has a double curvature ; its proximal portion sweeping toward the shaft of the tibia, while more distally it again curves outward and presents a rather short arch before meeting the articular facet for the tibia. As on the latter bone the surfaces for the different muscles are well defined by prominent ridges. The distal end is much expanded. The groove for the peroneus longus is well defined, as is also that for the peroneus brevis and tertius. The two grooves are separated by a prominent tubercle (outer malleolus) similar to that in the lion and quite unlike that in Canis where the groove for the peroneus brevis and tertius is very much less developed and the malleolus enlarged. On the tibial face there is a large round facet for the astragalus, which is gently convex in all directions. Immedi- ately above this large facet for the astragalus is a smaller facet for the tibia, which is plane and fits perfectly against a corresponding facet on the external face of the tibia. Measurement of Fibula. Mm. Greatest length, approximately 190 Antero-posterior diameter of distal end 19 Transverse diameter of distal end 10 PETERSON : NEW CARNIVORES FROM MIOCENE OP WESTERN NEBRASKA 247 Pes (PI. LXXXII). — The right hind foot of the type was found almost completely articulated. Consequently there is practically no doubt as to the true position of each bone in the pes. Astragalus. — While the main features of the astragalus are in general quite similar to those in Daphasnus felinus there are some detailed differences which are of considerable importance, and which Wk^^ mark the transition from the conditions found in the older type to the more modi- | H jfff'/ ^ec^ structures in the recent forms. It is ^^IffltliMi seen that the articular trochlea for the /ff wW|p||"^ tibia is more perfectly grooved in the pres- ent genus than in Daphamus, the internal and external condyles being more nearly Fig. 43. Anterior and Posterior Views of Astragalus of d. superbus. Nat. size. subequal, as in Ganis. The articulation of the tibial face of the internal condyle, however, is, widely interrupted by a broad vertical sulcus, which is not present in the recent form, nor does it entirely bisect this articulation on the astragalus of Daphostms, while in the tiger there is a similar broad area on the inner side of the internal condyle. The neck is relatively shorter, though constricted as in Daphosnus, which is due to the laterally expanded head, as in the cats. The head is inclined sharply inward and on the distal face is a large facet for the navicular, which is convex in all directions. The calcaneal facets are separated by a deep, narrow groove, and are subequal in size, the external being more oblique and less strongly concave than in Ganis. In the latter genus there is a minute articular facet for the calcaneum on the fibular face of the head. In the present genus this small facet is also present, but instead of articulating with the distal end of the calcaneum, as in the dogs, it comes in contact with the dorso-tibial angle of the cuboid, as in Daphanus felinus. Measurements of Astragalus. Mm. Greatest height 38 Transverse diameter of trochlea 21 Transverse diameter of head ... 21 Antero-posterior diameter of head 12 Calcaneum. — On comparing the calcaneum with that of Dapha-nus felinus there is very little difference detected except in size. The palmar face of the tuber calcis is more concave antero-posteriorly on its upper portion and slopes more gradually toward the distal end than in the Oligocene genus, which is due to a 248 MEMOIRS OF THE CARNEGIE MUSEUM Fig. 44. Ant rior and Fibular Views of Calcaneum of J), superbus. Nat. size. greater developed eminence on the lower half of the palmar face in Daphosnodon. The free end of the tuber is suddenly expanded, especially laterally, and there is a well defined groove for the tendon of Achilles, as in Daphanus and the recent dogs and cats generally. There is on the outer angle of the dorsal border a long and rugose groove, which, however, is not so deep as that in the Oligocene genus. This groove is also quite conspicuous in the recent lion, while in Canis it is entirely wanting. The articular facets for the astragalus are separated as usual by the groove for the interosseous ligament, the external facet being quite oblique and more gently con- vex antero-posteriorly than in Canis, while the sustentaculum is more promi- nent, the internal facet being slightly con- cave infero-superiorly and carried well over the superior edge, not unlike what is seen in Canis. The greater process of the calcaneum is much abbreviated, so that the head of the astragalus extends well be- low the distal end, as in Dapliamus, also observed by Mr. Hatcher (/. c, p. 91). The peroneal tubercle is well developed, causing the fibular face of the bone to be quite concave supero-inferiorly. The distal end has a large facet for the cuboid, which is gently concave in all directions and pear-shaped in its general outline. The astragalus and calcaneum of Amphicyon major are represented in our col- lection by accurate casts, which display well marked differences from the present genus. In the first place the astragalus in the European form is broader and shorter in proportion ; secondly, the head is more sessile, due to the shorter neck ; thirdly, the sulcus which separates the articular facet for the tibia on the tibial face of the inner condyle in Daphosnodon is absent in Amphicyon ; and fourthly, the external articular facet for the calcaneum is less concave supero-inferiorly than in the Amer- ican form. The tuber of the calcaneum in Amphicyon is proportionally shorter, and the palmar border is much heavier than the dorsal, while in Daphosnodon the two borders are subequal in thickness, as is also the casein Daphasnus. In the European form there is no well defined groove for the tendon of Achilles and the free end of the tuber calcis is much enlarged, as in the bears. The sustentaculum is prominent, as in Daphosnodon, but the greater process of the calcaneum is more abbreviated. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 249 Measurements of Calcaneum. Mm. Greatest length 67 Greatest transverse diameter 33 Length of tnber from sustentaculum to free end 40 Navicular. — The navicular differs from that of Daphamus by having a propor- tionally greater antero-posterior diameter, which is due almost entirely to the development of the palmar border in the Miocene genus. The tuberosity on the tibial face is also fully as prominent as in the Oligocene form, entirely un- (^ '' £12 like that in Canis, and is not dissimilar to what is found in the recent tiger or 4 r g lion. The palmar face is, as stated, greatly produced, and formed into a Ellllp^l/ ^df-% ' , I' ^Kfe large tuberosity, which, however, is not reduced transversely by the deep notches FlG- 45- Navicular (right side) of d. superb™. Nat. , . . ,, size. 1, dorsal view ; 2, superior view ; 3, fibular view ; on the tibial and fibular sides as in La- ... . , . , . . „ .... , . 4, plantar view ; 5, inferior view ; 6, tibial view. rns, but presents a more nearly perfect triangular outline of the entire bone when viewed from above or below. On the fibular face are two small facets, one palmar and the other dorsal. The one lo- cated dorsally is sometimes wanting, but in the type it is faintly indicated and touches the lower portion of the facet on the proximal dorsal angle of the cuboid when extreme rotation takes place, while the palmar facet articulates with a cor- responding facet on the proximal palmar angle of the cuboid. The facet for the head of the astragalus is concave in the usual manner, while the facet for the en- tocuneiform is quite large and is not crowded posteriorly in the same degree as in Canis or Felis. Measurements of Naviculae. Mm. Greatest antero-posterior diameter 21 Greatest transverse diameter 24 Greatest vertical diameter 11 Entocufieiform. — The entocuneiform, though of considerably smaller size pro- portionally than in Daphoenus, is still of large size, and occupies a more dorsal posi- tion in the tarsus than in recent forms. The bone is high, laterally compressed, having the transverse diameter greater on the plantar side than on the dorsal side, and possesses a large plantar hook, which causes the articulation for Mt. I to be 250 MEMOIRS OF THE CARNEGIE MUSEUM quite concave in the fore-and-aft direc tion. The proximal articulation for the navicular ascends rapidly in the plan- tar region, due to a heavy process on the posterior face which points upward and backward. On the fibular face are two facets, a large one proximally for Entocaneiform (right side) of d. superius- the mesocuneiform and a smaller one 1, dorsal view ; 2, inferior view ; 3, plantar distally, which articulates with Mt. II. view; 4, fibular view ; 5, tibial view. Measurements of Entocuneiform. Mm. Greatest antero-posterior diameter 13 Greatest transverse diameter 7 Greatest vertical diameter 16 Mesocuneiform. — The mesocuneiform has relatively a greater antero-posterior and a lesser transverse diameter, especially of the distal face, than is the case in Daphosnus. Consequently the bone, which is triangular in outline when viewed from below, has a longer apex directed backward than is seen in the Ohgocene genus. The facet for the navicular is (g, /J slightly less oblique than in Daphcenus, but £ has the same antero-posterior concavity. On the tibial face there is a large facet for 4 the entOCUneiform which is located On the FlG- 47- Mesocuneiform (right side) of D. superbvs. .. r . , -, -, . Nat. size. 1, dorsal view; 2, fibular view ; 3, tibial upper portion of the bone and is concave . ..... „ . . 1 r r view; 4, inferior view; 5, superior view. antero-posteriorly. On the fibular face, near the proximal angle, is a facet for the entocuneiform which extends throughout the entire fibular face, antero-posteriorly, but is of small vertical diameter and corresponds to a facet on the tibial face of the ectocuneiform. Distally the bone is taken up by the facet for Mt. II. Measurements of Mesocuneiform. Mm. Greatest antero-posterior diameter 13 Greatest transverse diameter 9 Greatest vertical diameter 9 Ectocuneiform. — When the ectocuneiform is compared with that of Daphoenus it is at once seen that this bone in Daphcenodon has not increased in size in the PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 251 same ratio as the entocuneiform has decreased. Both meso- and ectocuneiforms are therefore of approximately the same proportionate size as in the Oligocene form. The dorsal face has a more perfect diamond- shaped appearance than in Canis or Felis, \P """^%j fM^X^' -'l*1 ^ Wi'' ml ^T ' W which is due chiefly to the greater obliquity / ftt MOpTSfff di jK$ U ' -3^\ U^^ajw *&■ — *^ of the proximal and distal articular facets in ^_j^=2 Dapliecnodon. The facet for the navicular on the proximal face is especially oblique, due to a prominent ascending process on the fib- ular angle. This process has a large facet for iJtu %'SsA \Jr 5 JjtM 6 the cuboid on its fibular face, and near the Fig. 48. Ectocuneiform (right side), of D. distal end there are two minute facets for Mt. supirbus. Nat. size, l, dorsal view ; 2, fibular IV, one dorsal and the Other palmar. The ™w ; 3, tibial riew; 4, plantar view ; 5, inferior . • -i, view; 6,superior view. tibial face has three facets, one proximally for the mesocuneiform referred to above, and two distally for Mt. II. The distal face is more oblique and also more concave antero-posteriorly than in the recent forms, while the plantar process is reduced in size and is canine rather than feline in its general characters. Measurements of Ectocuneiform. Mm. Greatest anteroposterior diameter 21 Greatest transverse diameter 12 Greatest vertical diameter 18 Cuboid. — In comparison the cuboid is of a slightly greater transverse and less vertical diameter than that of Daphccnus. Otherwise there are no striking differences in this bone in the two genera. The dor- " - If ft 2^-ii£i^ StiiMw^ sa^ ^ace *s ^S^y concave in the proximo- distal direction, while on the fibular plantar face is found the deeply excavated groove 1*4JI| f°r tne peroneus longus tendon. Above this groove is a large tuberosity which forms a j -^ solid, broad, oblique ridge extending from the proximo-fibular to the distal tibial angle, Fig. 49. Cuboid (right side) of D. superbm. ° Nat. size. 1, dorsal view ; 2, superior view ; 3, plan- and not Separated by a Vertical Sulcus 3S ill tar view ; 4, inferior view ; 5, tibial view ; 6, fibular CaillS. Superior to this ridge the plantar face is taken up by a large, rugose, excavated area for ligamentous insertions. The fibular, face is nearly plane and has three artic- 252 MEMOIRS OF THE CARNEGIE MUSEUM ulations ; the two proximal ones are small indistinct facets which articulate, one with the head of the astragalus and the other (near the plantar surface) with the navicular. This curious feature of articulations in the tarsal bones is quite unusual and is also found in Daphcenus and to a less degree in Hoplophoneus, as Mr. Hatcher observed in his study ("The Oligocene Canidae," p. 91). In the middle region of the tibial face is the third facet, which is of considerable size ; it is triangular in outline and articu- lates with the ectocuneiform. Proximally the cuboid has a large surface for the calcaneum, which is convex in all directions in a greater degree than in Canis or Felis and even more than in Hoplophoneus, though not nearly so oblique as in the latter. Distally there is one large facet for metacarpals IV and V, which is concave antero-posteriorly and also slightly concave laterally, the extreme fibular border being slightly recurved upward to accommodate the lateral portion of the proximal facet of Mt. V. Measurements of Cuboid. Mm. Greatest antero-posterior diameter 20 Greatest transverse diameter 17 Greatest vertical diameter 21 Metatarsal I. — The first metatarsal has the appearance of being shorter than that of Daphcenus when placed in position in the pes, but it is in reality of very nearly the same proportionate length, this shortened appearance being chiefly due to the proportionally less vertical diameter of the entocuneiform in the Miocene genus. These bones (Mt. II) in the two genera are very similar in every respect. As in Daphcenus the head is much expanded both antero-pos- I(V,'IiSm teriorly and transversely, with two prominently developed I Mr and rugose tubercles on the plantar face, one on the tibial and t e other on the fibular angle. The facet for the ectocu- neiform is quite large and is convex antero-posteriorly, but is slightly more oblique than that of the Oligocene genus, so that (1*431 tne digit points more inward when the pes is articulated. The shaft is of good proportionate size and is arched forward. Fig. so. Fibular and Dor- The distal trochlea is hemispherical on the dorsal face and sal views of Right Metatarsal on the piantar ftlce it is divided by a strong keel. With the I of D. snperbm. Nat. size. . exception of the reduction of the entocuneiform and the position of the metatarsal there is no great change of the first digit of the pes from that of the Oligocene form, the digit on the, whole being more greatly developed than in recent dogs or cats, and more resembling that of Viverra zivetta. l'ETERSOX : NEW CARNIVORES FROM MIOCENE OE WESTERN NEBRASKA 253 Measurements of Metatarsal I. Mm. Greatest length 42 Antero-posterior diameter of head 11 Transverse diameter of head 11 Transverse diameter of distal trochlea 9 Metatarsal II. — As in Daphcenus the articular facet for the mesocuneiform on the head of Mt. II is oblique, slightly concave antero-posteriorly, and rises to a high and sharp angle along the fibular border. The fibular side has two large facets for the ectocuneiform, thus the head of Mt. II extends above those on Mt. I and III when articulated in the foot. The plantar tuberosity is of considerable size and is located on the fibular angle. Between the plantar tuberosity and the dorsal face there is on the tibial face an excavated and rough surface, against which lies the head of Mt. I, but there is no true articular facet for the latter. The shaft is much shorter and lighter than in the corresponding bones in Canis or Fells and is in proportion to the rest of the limb perhaps more nearly like that of the bears. Distally the bone is expanded transversely, the dorsal face of the trochlea be- ing spheroidal and the plantar face divided by a prominent carina as in the metacarpals. Views of Eisht Metatarsal n of I), superbus. Nat. size. Measurements of Metatarsal II. Mm. Greatest length 59 Antero-posterior diameter of head 16 Transverse diameter of head 10 Transverse diameter of distal end 13 Metatarsal III. — As was observed by Professor Scott in his study of Datphxmus (I. a, p. 356) the relative proportion of Mt. Ill is greater than in Dmictis, Hoplopho- neus or the viverines. It is also considerably longer proportionally than Mt. II in Canis, while actually the bone is much shorter in the genus under description. The head has a large antero-posterior diameter, the plantar tuberosity being quite large both transversely and vertically, as in Canis. The articular facet for the ecto- cuneiform is of large size ; it is quite oblique in position and is convex both antero- posteriorly and transversely. On the tibial face the head is deeply excavated, but there is no articular facet forMt. II, except a slight area on the extreme tibial angle 254 MEMOIRS OF THE CARNEGIE MUSEUM Fibular and Dorsal tight Metatarsal III of Nat. size. of the plantar tuberosity. On the fibular face there are two facets for Mt. IV, which are separated by a deep sul- cus. The dorsal one is located on the posterior face of the flange produced from the dorsal portion of the head, and extends in the fibular direction so as tooverlap Mt. IV, causing a strongly interlocking condition of these bones, as in the carnivora generally. The facet back of the sulcus is on the plantar tuberosity and articulates with a corresponding facet on the plantar tibial angle of Mt. IV. The shaft is quite heavy, considerably arched in the dorsal direction, and oval in cross-section. The distal end is similar to that on Mt. II, but has a more symmetrical trochlea. ;.i Measurements of Metatarsal III. Mm. Greatest length 70 Antero-posterior diameter of head 18 Transverse diameter of head 14 Transverse diameter of distal end 14 Metatarsal IV. — The fourth metatarsal is the longest in the series. Its head rises slightly above that on Mt. Ill, while the two bones extend equally far in the distal direction. The proximal face of the head has a large convex facet for the cuboid, while on the tibial side there is one facet situated on a prominent tubercle which fits into a corresponding pit on Mt. Ill, and the other facet for the same metatarsal is located on the plantar tibial angle, as stated above, thus causing these bones to firmly interlock, as in Daphosnus. On the plan- tar side there is a heavy tuberosity of cubical appear- ance and on the fibular side there is a deep pit sim- ilar to, though more pronounced than that for Mt. IV on the fibular face of Mt. III. Above this pit is a large facet which also articulates with Mt. V. The shaft is arched forward in the same degree as that on Mt. Ill, but is further differentiated from the latter by a prominent and rugose elevation for ligamentous attachments similar to that of Daphosnus. As in Mt.III the distal trochlea is quite symmetrical. Fig. 53. Fibular and Dorsal Views of Right Metatarsal IV of D. superbtts. Nat. size. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA Measurements of Metatarsal IV. Mm. Greatest leDgth 73 Antero-posterior diameter of head 17 Transverse diameter of head 12 Transverse diameter of distal end 13 255 Metatarsal V. — The fifth metatarsal is of about the same length as Mt. II though slightly lighter. The head is greatly expanded, especially in the transverse diameter, due to the large development of the ascending fibular pro- cess, which is much more developed than in Canis and nearly equal in proportion to that of Daphanus. On the posterior tibial side this process is succeeded by a heavy plantar tuberosity which extends throughout the remain- der of the plantar face and rises above the articular facet on the proximal end. The tibial face has a projecting eminence carrying an articular facet for a corresponding pit on Mt. IV already described. The facet for the cuboid is quite large ; it is slightly convex antero-posteriorly,and along the fibular border it is recurved upwards on the base of the ascending process referred to above, so that the facet is concave transversely and faces inward as well as up- ward like that of the older form. The shaft is well arched and has the usual transversely oval cross-section found in older types. The distal end is enlarged along the fibular border in about the same proportion as Mt. I is expanded on the tibial angle, giving the trochlea an oblique and asymmetrical appearance. Cynodesmus brachypus is an animal of considerably smaller size than Dapha- nodon, but Mt. V, which is represented in the type, is of very nearly equal length, though much lighter than that of Daphanodon. The ascending process on the fibular angle of the head is much larger than in Canis, but it is directed upward and outward pretty much as in the latter, and the articulation for the cuboid is small and faces directly upward as in the recent form, entirely unlike that of Daphanodon and Daphanus. Another modified character of importance in Mt. V of C. brachypus is seen in the shaft. The latter is more arched in the dorsal direction than in Canis, but it is plainly seen that the proximal half is trihedral, which is due to the more complete parallel arrangement of the metatarsals, approaching, condition found fcfee QW* in the modern Canidx, while in Daphanodon and its progenitors the upper por- tion of the shaft is transversely oval, due to a less appressed condition and a more Fig. 54. Fibular and Dorsal Views of Right Metatarsal V of D. superbus. Nat. size. 256 MEMOIRS OF THE CARNEGIE MUSEUM direct divergence from the proximal ends, which is a general feature of the early types of the Miacidse. Measurements of Metatarsal V. Mm. Greatest length 62 Antero-posterior diameter of head 13 Transverse diameter of head 19 Transverse diameter of distal end 12 Sesamoids. — The sesamoids are proportionally somewhat smaller than in the recent canids, but have otherwise no noteworthy features differing from the latter. Fig. 55. Phalanges of Digits III, IV, and V, of Right Hind Foot of D. superhus. Nat. size. 1, dorsal and tibial views of first phalanx of third digit ; 2, fibular and plantar view9 of terminal phalanx of third digit ; 3, dorsal view of first phalanx of fourth digit ; 4, tibial and dorsal views of second phalanx of fourth digit ; 5, fibular and dorsal views of first phalanx of fifth digit. Note: The hood of the ungual phalanx is partly broken off. Phalanges. — The proximal row of phalanges are distinctly shorter in propor- tion to those in Daphcenus, which is also true of the phalanges of the manus. They are otherwise quite similar in structure, the proximal articulation being a concave pit with a broad plantar groove, and the distal trochlea confined principally to the plantar face. The median row of phalanges is asymmetrical in the same proportion as those in the manus, and indicate that the unguals were drawn toward the fibular borders when the little retractility which they undoubtedly possessed, was exer- cised. The unguals are similar to those in the manus. Measurements of Phalanges. Mm. Greatest length of proximal phalanx, fourth digit 29 Greatest length of median phalanx, fourth digit 19 Greatest length of ungual phalanx, fourth digit 24 PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 257 Restoration of Daphcenodon superbus. (Plates LXXXIII and LXXXIV.) The unusually well preserved and symmetrical bones of the type of Daphceno- don superbus have caused comparatively little trouble in correctly articulating the different parts of the skeleton. The restoration has been very well executed by Mr. Serafino Agostini. The mounted skeleton furnishes many additional features of the animal which only in this way can be fully appreciated. The position of the head is as high in relation to the horizontal line of the vertebral column as the animal ordinarily held it in life. The long neural spines on the cervical vertebrae further indicate that the animal probably carried the head rather low. The curvature of the neck at the junction of the thorax is gentle, and the position of each dorsal vertebra increases in height until the lumbar region is reached, where a sudden downward curvature again takes place, not unlike that in the recent tiger. The twelfth and thirteenth dorsals have no facets for a tuberculum of the ribs as in Felis tigris and it is quite doubtful whether the eleventh dorsal of the present form carried a tubercular facet for the rib as the transverse process is poorly developed. The thorax was on the whole quite small, which is also charac- teristic of the Oligocende genus. The heads of the ribs of the left side (except the eleventh or twelfth) are all represented, while the shafts are restored after the more completely preserved ribs of No. 1589a. On the left side the ribs are poorly pre- served, there being present only the heads and portions of the shafts of seven. The fore limbs display a close similarity to those in Daphcenus. The deltoid and supinator ridges of the humerus are prominent, the former low down on the shaft, indicating the downward extent of the heavy muscles, while the development of the latter indicates the power of supination of the manus. Though the forearm is proportionally longer than in the latter genus the feet are remarkably little advanced, being short and broad, the pollex little reduced, and the position of the phalanges sharply angled as in the felines, but proportionally somewhat shorter than in Daphcenus. The power of retractility of the unguals was nearly as great as in the older form and no doubt still served the purpose of assisting in catching and holding the prey. As has already been stated the posterior portion of the pelvis, portions of the femora, and the proximal ends of the tibia? were unfortunately weathered out and disintegrated when the specimen was found, but from other material the compara- tive measurements were made, and the proportions, as they are in the skeleton repre- sented on Pis. LXXXIII-LXXXIV, are thought to be approximately correct. The 258 MEMOIRS OF THE CARNEGIE MUSEUM hind feet are also quite broad, but somewhat longer than the fore feet, while the ungual phalanges of the former had apparently a power of retractility equal, or nearly equal, to that of the latter. Altogether the skeleton shows a long and com- paratively slender body, long tail, and short and heavy limbs with broad and short feet. Measurements of Skeleton. Cm. Greatest length of vertebral column, the skull and all curvatures included 210 Length of vertebral column, atlas to posterior end of sacrum 92 Length of tail, approximately 88 Length of lumbar series 26.5 Length of dorsal series 33 Length of cervical series 21 Height of skeleton at first dorsal 57 Height of skeleton at first lumbar 61 Height of skeleton at supra-iliac border, approximately 58 Notes on Individual Variation of Daphcenodon. The superior dentition of the material at hand shows very little variation in general character, except in the young individual (No. 1589a) found with the type of Daphcenodon superbus and described on pages 210 and 215. The mandibular symphysis in some individuals however, is, longer, pT has been dropped out and the alveole closed up, while m-j is sometimes inserted by two strong fangs. In a series of cervicals (No. 1589b), the fourth, fifth, sixth, and seventh, which were found on the surface and in the talus at Quarry No. 3 (Agate Spring Fossil Quarries), are seen some differences from those described of the type of Daphcenodon superbus. These differences are most likely due to individual variation and sexual distinction, and will be thus treated in this connection pending the discovery of more material. This cervical series is in the first place considerably heavier and pertains to a larger individual than the type specimen. Secondly, the ventral keels appear to terminate posteriorly in a more condensed rugose tubercle approaching more nearly that of Canis in its characters than to that of the type of Daphcenodon. Thirdly, the anterior projection of the transverse process of the sixth cervical vertebra in the series under consideration is much smaller than the corresponding portion in this vertebra of the type specimen. One of the more significant differences from the latter is seen in the sixth and seventh cervicals. The base of the neural spine of the seventh cervical is more nearly oval in cross-section, with no such excavations in front or back of the spine as is characteristic of the type specimen described above. The base of the spine of the sixth cervical is also without these excavations, so that PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 259 it is quite safe to say that this individual did not possess this character of the verte- brae. Another important feature is seen in the prominent antero-posterior ridge in the floor of the neural canal, which is pierced by a transverse venous foramen, while in the type specimen this ridge is rather poorly developed and the area in the middle region, where the foramen is located in the larger specimen, is much greater and was not always bounded above by a bony bridge so as to form a foramen. The transverse process is proportionally much shorter, which would indicate a narrower neck in the larger individual. In the Amherst specimen, which is about equal in size to the one represented by the cervical vertebra? described above, there are some lumbar vertebrae preserved. One of these has the transverse processes preserved. These processes are propor- tionally longer than those in the corresponding region of the lumbars in the type, and more in accordance with the conditions found in Daphamus, Temnocyon, and the recent forms. The os penis of the Amherst specimen of Daphomodon is greatly different from that in No. 1589a (Cam. Mus. Cat. Vert. Foss.) and is described on pages 230 and 231. Relationships of Daphccnodon. That Daphasnodon had its true ancestor in Daphanus of the Oligocene forma- tion can hardly be doubted, from the study which has resulted in the above com- parative description. Daphasnus in turn is generally regarded as a descendant of the Miacidw of the earlier Tertiary of North America. In the Oligocene and early Miocene there were a number of diverging lines, which as a whole were neverthe- less quite homogeneous in their general structure, though apparently sufficiently diversified to show with more or less clearness their destiny in later times. These points have already been ably treated by Scott, Boule, Schlosser, Matthew, Merriam, Hatcher, Wortman, and others, and will not be discussed in the present paper, except in so far as to assign the phyletic position of Daphamodon. It would appear that Daphcenodon is not yet represented by any very closely allied species20 in the intervening formations from the lower Harrison beds down to the lower portion of the Oligocene where Daphamus, its predecessor, is found. The genera from the John Day are already too far advanced in different ways to be seriously considered as intermediate forms. Thus Temnocyon and Mesocyon have m- absent and differ in many other important respects, some species21 of the latter genus having a well developed entoconid on the lower sectorial, and being possibly 20 Mesocyon robmtm Matthew from the lower Miooene of South Dakota bears general similarities, but nn is apparently too much reduced and more trenchant, and m:1 is probably sometimes absent. "Mesocyon Josephi (?). See Merriam, Uuiv. of Cal. Publications, Bull. Dept. Geol., Vol. V, pp. 19-:0, 1906. 260 MEMOIRS OF THE CARNEGIE MUSEUM more nearly in the ancestral line of Cynodesmits (C. brachypus). Philotrox, Hyxno- cyon, and Enhydrocyon are removed from this line by the reduction of the premolars and other important features. Paradaphapius is a small form with long narrow skull, deuterocone of pA well internally, M- aligned with the outer cusps of the anterior molars, heel of lower molars basin-shaped, and tympanic bulla of propor- tionally larger development. Of the later American forms which are perhaps most closely related to Dapha no- don, Amphicyon americanus Wortman is quite well preserved and furnishes the most satisfactory comparison. This species, like the European, A. major, has the canines enormously developed and the premolars reduced when compared with Daphwno- don. PT is less oblique in the alveolar border, M- and M- are apparently more nearly subequal in size, while ftP is aligned with the outside of the molars and is somewhat larger than in the latter genus. Amphicyon major of Europe is further differentiated from the present form by its short, broad, bear-like astragalus and sub-plantigrade hind feet. Amphicyon lemanensis is another well known European form bearing certain resemblances to Daphwnodon superbus, but which on closer study displays numerous differences, among which mention may be made of a larger size three-rooted and tri-cuspid M-, a smaller P-, proportionally longer, narrower, and higher skull, as well as a differently constructed axis, and other characters of importance. Amphicyon crucians, figured by Dr. Filhol (Ann. des Sciences Geologique, Vol. X, pi. 12, figs. 1-3, 1879) show only general similarity to Daphwnodon, so far as comparison can be made, the crowns of the premolars being higher and more pointed, and P3 having a basal tubercle, while that tooth in Daphwnodon has none. Cephalogale geoffroyi is, with regards to the general features of the skull, more like Daphwnodon, but the proportions and construction of the tubercular molars are dif- ferent, M- being absent, and this European form is apparently no nearer related to Daphwnodon than for instance the American genus JElwrodon. In fact the apparent relations observed between the European and American forms here compared should perhaps be regarded as purely superficial and due entirely to independent develop- ment without necessarily close relationship. With regard to the ancestry of the genus described above there can certainly be no doubt that we find in Daphwnus from the American Oligocene all the required characteristics pointing more clearly to Daphwnodon than they are so far found in any known European forms, the ancestral types of the latter genera having perhaps already occupied contemporaneously these European regions for a long time. PETERSON : NEW CARNIVORES 'FROM MIOCENE OF WESTERN NEBRASKA 261 From the study of the osteology of Daphosnodon superbus it is quite evident that this genus belongs to a phylum existing through the Oligocene and Miocene periods, with rather conservative adaptations when compared with such forms as Temnocyon, Mesocyon, Cynodesmus, Tephrocyon and in fact with many coexisting forms of Canidse. Although the genus should be, for the present at least, regarded as belonging to the subfamily Amphkyoninx, it is not altogether unlikely that, upon gaining more complete knowledge of all the different supposed allied forms, it will be regarded as representing a separate subfamily (Daphceninw) paralleling Temnocyon, Mesocyon, Cynodesmus, and Tephrocyon on the one hand and the true Amphicyon on the other, but less differentiated from Daphcenus than either of the genera regarded as descendants from the Oligocene forms. The phylum terminated, most probably, in some such forms as Dinocyon or Borophagus, or perhaps in some of the so-called Amphkyones in later formations of North America, while the Awphi- cyoninse of Europe were paralleling the line. This would seem to lend color to Mr. Hatcher's contention * that Protemnocyon and Proamphkyon are both valid genera, the former pointing to Temnocyon and Meso- cyon, while the descendants of the latter, if not "Amphicyon" as Hatcher believed, may not yet have been discovered. The reduction or apparent absence of M^, together with other features in Protemnocyon from the Oligocene, is suggestive of the later John Day form, while M^ in true Daphcenus still persists in an almost unchanged form in Daphosnodon from the Miocene. The bony structure of Daphosnodon, though extremely cat-like in many respects, is on the whole more closely related to the Canidm, of which family it was an aber- rant line not continued to the present time. A fairly good attempt has been made by Mr. Theodore A. Mills to construct the soft parts of Daphcenus felinus, from the skeleton which is now in the Carnegie Museum. And the curious combination of the characters of the cat and the dog are, if anything, even more striking when an allowance of flesh is represented upon the bony structure (see PI. LXXXV). The downward extent of the heavy muscles of the limbs, the broad and short feet, with the semi-retractile claws, the long body and tail are especially cat-like, while the head is dog-like in every respect. The tail is represented more or less like that of Felis concolor, there being no perceptible means of knowing whether this appendage was bushy, as in the dogs, or more slender as in cats generally. Altogether the model represented by the figure on PL LXXXV is instructive, as it furnishes at least a conception of a primitive form ancestral to cats and dogs. Of this primitive stem Daphcenus and especially Daphosnodon on the whole, appear to be far less specialized " "Oligocene Canidie," /. c, p. 105. 262 MEMOIRS OF THE CARNEGIE MUSEUM than most of their contemporaneous forms and consequently more nearly like the old types of the Miacidse from the Eocene formations. Canid (Undetermined Species). A humerus (No. 2400) of a very large canid, the size of a lion, was found in the lower Harrison beds on Whistle Creek, Sioux County, Nebraska, about eight or ten Fig. 56. Anterior View of Humerus of Canid. Sp. indt. J nat. size. No. 2400. miles east of the Agate Spring Fossil Quarries. With the exception of its greater size and the greater distal extent of the deltoid ridge this humerus most nearly resembles that of Daphcenodon. The bone is very nearly the size of the humerus of Amphicyon major of Europe and not unlike the latter, so far as comparisons can be made. In the Amer- ican species, however, the deltoid ridge extends lower down and the anconeal fossa is not so high as in the European species. The humerus possibly rep- resents some genus closely related to such large forms as Dinocyon from the later Tertiary of Montana de- scribed by Mr. E. Douglass23 or D. (Borophagus) gidleyi from the Miocene of Texas described by Dr. W. D. Matthew.24 The quarry in which the hu- merus (fig. 56) was found contains a similar fauna to that of the Agate Spring Fossil Quarries. A fifth metacarpal (No. 1897) of a large carnivore, which may belong to this species, was found by the writer in 1904 among the surface fragments in the quarry, which was worked the following years by Professor Barbour (Agate Spring Fossil «"New Vertebrates from the Montana Tertiary," Ann. Car. Mns., Vol. II, p. 192, 1903. ""A Skull of Dinocyon from the Miocene of Texas," Bull. Amer. Mus., Vol. XVI, p. 129, 1902. Fig. 57. Fibular and AnteriorViews of Calcaneum of Cauid sp. indet. i nat. size. No. 2211. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 263 Quarries, "University Hill" Quarry). This bone is very heavy, short, and much arched in the dorsal direction. A calcaneum of nearly an equally large carnivore was found in the same horizon north of the Niobrara River in this general locality and is illustrated on page 262, Fig. 57. The remains indicate the presence of these large carnivores in the fauna found in these quarries. Measurements. Mm. Greatest length of humerus 305 Greatest anteroposterior diameter of head 87 Greatest transverse diameter of head 68 Greatest length of Mo. V 63 Greatest length of calcaneum 90 Greatest transverse diameter at snsteutacular facet 45 Borocyon* robustum gen. et sp. nov. Type: (No. 1918, Cam. Mus. Cat. Vert. Foss.) Front of right ramus, caudal vertebra?, fragments of manus, part of hind limb and foot, and various other fragments. Horizon : Miocene, Upper Harrison beds. Locality : Whistle Creek, Sioux County, Nebraska. Mr. T. F. Olcott, while engaged with the Carnegie Museum party in 1905, dis- covered and collected this specimen about four or five miles southeast of the Agate Spring Fossil Quarries. As indicated above it was found in the upper Harrison beds. Although the type is fragmentary it enables me to give some characters which may be regarded as of generic value. The type is perhaps more closely allied to such forms as Amphicyon sinapius Matthew from later horizons than to A. arnericanus, A. ursinus, or any of the European forms with which it has been compared. The jaw is undoubtedly quite short and heavy, and the premolars are of considerable size when compared with A. ursinus. The portion of the jaw preserved compares fairly well with JElurodon taxoides Hatcher, but the canine is heavier and P2 has no pos- terior cusp as in that species. A. meandrinus had the jaw proportionally longer in front of PT than the present genus. It is, of course, quite provisionally that this genus is placed in the Daphanus-Daphwnodon phylum, but pending the discovery of more complete material it would seem most satisfactory from the study of the limb and foot structure to regard the genus as a later member of this aberrant line. The material represents an animal nearly the size of a fully adult lion. Generic characters. Canine proportionally large and oval in cross-section, PT single- rooted, P2 low-crowned with jrrotoconid well anteriorly, no posterior cusp, and a large * jiopOf = greedy ; /tkw = dog. 264 MEMOIRS OF THE CARNEGIE MUSEUM posterior base, M?f present, heavy and elongated tail, limbs and feet well proportioned and sub-digitigrade as in Daphamus and Daphomodon, little or no retractility of the unguals. General Description and Specific Characters. The material represents a fully adult animal judging from the much worn canine. The latter is oval in cross-section and is proportionally larger than that tooth in Daphomodon. Judging from the abraded anterior face of the canine it is quite evident that P was of large size as in Daphcenodon. The postero-external angle of the canine is also worn by the contact with the upper canine. As in Fig. 58. Right Side of Lower Jaw-fragment and Crown View of Teeth of Borocyon robustum. Nat. size. No. 1918. Fig. 59. Caudal Vertebra; of Borocyon robustum. J nat.? size. No. 1918. 1, lateral and ventral views of caudal 7 ; 2, lateral and ventral views of caudal? 9. Daphcenodon there is a short diastema back of the canine. PT is single-rooted. P^ is low-crowned and has the protoconid placed more anteriorly than in Daphosnodon, which results in a less developed anterior and greater posterior base than on Pg in the latter genus. There is no posterior tubercle on P^ as in JElurodon, but the cingulum is well developed ; it surrounds the tooth and is smooth on the external side, while internally it is slightly mammilated. There is a large mental foramen directly under the posterior portion of P^- The symphysis is quite heavy and extends well back as in Daphcenodon. The only parts present representing the vertebral column in the type are a number of caudal vertebras from different regions of the tail. The centra of these vertebrae are long and heavy, indicating a tail of approximately the same propor- tions as that of Daphcenodon. The manus is represented by a complete trapezoid, Mc. I, and other fragments. The trapezoid is depressed and wide transversely, indicating a broad manus. The PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 265 facet for the scapho-lunar is slightly convex from before backward near the dorso- radial angle, while further back and along the ulnar portion it is concave in the same direction. Radially there is a large facet for the trapezium, which takes up nearly the entire face. On the ulnar side the facet, for the lunar, meets that for the magnum at an acute angle so as to form a prominent ridge, which extends a short distance back from the dorsal border. Back of the ridge the ulnar face is taken up by a large rugose area for the at- tachment of muscles and ligaments. On the distal face there is a large subtriangular articulation for the second metacarpal. The free end of the pisiform indicates this bone to be quite like that in Daphomodon. Mc. I has the head less expanded transversely, the shaft less arched forward, and less curved outward at the distal end than that of Da pluenodon superbus ; there is otherwise little or no difference between this bone in the two forms. Other fragments of the metacarpals suggest heavy bones in this region. The phalanges are proportionally short when compared with Daphosnodon. Those of the proximal row are other- wise arched in the dorsal direction similar to the conditions found in the latter genus, while the median phalanx is depressed and broad. There is no ungual pha- lanx present with the type. The hind limb is represented by the lower half of the tibia and fibula, the astragalus, navicular, ento-, meso-, and ectocuneiforms, and other fragments. With the exception of the proportionately somewhat greater antero-posterior diameter of the distal end, the tibia offers little or no characteristic differences by which it may be distinguished from that of the smaller genus. The distal end of the fibula is more compressed laterally than in Daphoenodon ; otherwise there is also little difference in this bone in the two forms. The astragalus has a slightly longer neck and the articular surface of the trochlea descends lower than in Daphrenodon, otherwise this bone does not differ in the two genera. The principal difference of the navicular from that of Daphwno- don is seen in the smaller development of the plantar tuberosity ; the rugose area for muscular attachment slanting more upward and backward, resulting in the com- paratively small vertical diameter of the tuberosity. The entocuneiform is quite large, and of good proportionate vertical diameter. The tibial plantar angle is more Fig. 60. Foot Bones of Borocyon robualum. £ oat. size. No. 1918. 1, dorsal view of metacarpal I ; 2, dorsal view of a second phalanx ; 3, dorsal view of a first phalanx ; 4, proximal view of trapezoid ; 5, dorsal view of trapezoid ; 6, radial view of trapezoid ; 7, ulnar view of trapezoid. 266 MEMOIRS OF THE CARNEGIE MUSEUM concave in the vertical direction than in Daphamodon, which results from the greater development of the ascending plantar tuberosity of the present form. The meso- and ectocuneiforms are similar to those in Daphaenodon. Metatarsal III has the articular surface for the ectocuneiform less oblique and the tibial and fibular faces of the head less exca- vated than in Daphaenodon. Mt. V is quite unlike that of the latter genus in respect to the articular facets of its head. The tibial face has a somewhat smaller facet for Mt. IV and the facet for the cuboid is entirely proximal, while in Daphanus and Daphaenodon the facet for the cuboid extends well in the fibular di- rection, encroaching on the base of the ascend- ing process so as to form a lateral as well as a proximal contact with the cuboid. Another important difference from the older forms is seen in the vertical position of the ascending process on the fibular face of the head. In the Oligocene and Miocene forms, here used for comparison, this process is very heavy and projects upward and outward forming, with the excavation for the peroneus longus tendon on the cuboid, a transversely broad groove. In the present genus the ascend- ing process, though large, is directed more vertically and the space between it and the fibular face of the cuboid is comparatively narrow. Near the proximal ends the shafts of the meatarsals are slightly triangular in cross-section. The distal end of Mt. V? is somewhat less expanded transversely than in Daphaenodon, but the trochlea is quite similar, being very convex dorsally and separated by a prominent carina on the plantar aspect. Measurements. Mm. Antero-posterior diameter of canine at base 21 Transverse diameter of canine at base 15 Antero-posterior diameter of pj 14 Transverse diameter of p* 18 Vertical diameter of ramus at p2 35 Length of ? ninth caudal 58 Anteroposterior diameter of trapezoid 19 Greatest transverse diameter of trapezoid 20 Fig. 61. Fragments of Hind Limb and Foot of Borocyon robiistum. \ nat. size. No. 1918. 1 dorsal view of tibia and fibula ; 2, dorsal view of hind foot. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 267 Greatest vertical diameter of trapezoid 12 Length of Mc. I 49 Greatest transverse diameter of head Mc. I 12 Greatest length of proximal phalanx of 3rd or 4th digit 31 Greatest length of median phalanx of 3rd or 4th digit 21 Height of tarsus 86 Transverse diameter of tarsus, approximately 58 Height of astragalus 50 Greatest transverse diameter of astragalus 40 Greatest transverse diameter of trochlea of astragalus 28 Antero-posterior diameter of navicular 30 Greatest transverse diameter of navicular 36 Greatest vertical diameter of navicular 18 Antero-posterior diameter of entocuneiforra 17 Transverse diameter of entocuneiform, distal end 7 Vertical diameter of entocuneiform 25 Cynodesmus thomsoni Matthew. Bull. Amer. Mus. Nat. Hist., Vol. XXIII, p. 186, 1907. This species is represented by a considerable portion of a skeleton (No. 1529, Cam. Mus. Cat. Vert. Foss.) found in the upper Harrison beds on the Niobrara River, Sioux Co., Nebraska, and adds to our knowledge of that species. Accord- ing to Dr. Matthew's statements there are apparently no hind feet with the type, while in the present specimen both fore and hind feet are quite well preserved and indicate about the same proportionate size as those of Cynodesmus brachypus. In fact most of the skeletal features are quite similar to those of the latter species so far as they can be compared. P* is absent on both sides and considerable crushing has taken place at the base of the skull of C. brachypus which prevents a very accu- rate comparison, but it is quite evident that M- is fully as much reduced compara- tively as in G. thooides Scott, and is considerably smaller than in the type of C. thomsoni and the specimen in our collection referred to the latter species. These and other differences of the dentition no doubt indicate more progressive develop- ment in C. brachypus and C. thooides so far as certain features of the dental struc- ture are concerned. The structure of the feet and limbs certainly approaches very much nearer the conditions found in Canis than is the case in Daphoonodon. Among these may be mentioned the higher position of the deltoid crest, the less prominent supinator ridge of the humerus, the reduced distal end of the ulna, the laterally expanded supra-iliac border the concave gluteal surface and shorter ischium of the pelvis, the more completely grooved astragalus, the proportionally greater length of the metatarsals over the metacarpals, the tendency of the metatarsals to become appressed at their proximal ends, and the more reduced pollex and hallux. 268 MEMOIRS OF THE CARNEGIE MUSEUM Fig. 62. Pelvis aDd Feet of Cynodesmus thorn* 2, external view of pelvis ; 3, anterior view of pes. Nat. size. No. 1591. 1, anterior view of manns ; When the skull of the present specimen (No. 1591) is compared with Matthew's description and figures (I. c, pp. 186-188) it is seen that in practically all the details of structure, except the apparently smaller brains in our specimen, which may be due to crushing, the two specimens agree quite closely. P-j in the Carnegie Mu- seum specimen has no posterior cusp, while Dr. Matthew's figure indicates one on the type. The present specimen represents an old individual, as the tubercular teeth are much worn. Tephrocyon (Canis) temerarius Leidy. Proc. Acad. Nat. Sei. Phila., p. 21, 1858; Jour. Acad. Nat. Sci., Vol. VII, p. 29, PL I, fig. 12, 1869. A fragment of a lower jaw (No. 2404, Carn. Mus. Cat. Vert. Foss.) is provision- ally referred to this species as the measure- ments and general appearances are quite FIG. 63. Crown View of Teeth and External View ^^ ^ ^^ of ^ ^ ent of Jaw-fragment of Tephrocyon (Canis) temerariua Leid Nat size specimen was found on Whistle Creek, Sioux PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 269 County, Nebraska ; it was lying on the surface and possibly pertains to a later Miocene or Pliocene deposit which apparently thinly covers the lower Miocene section in certain places in this valley. The accompanying figures are given in order to show the close similarity of this species to T. rurestris Merriam and T. hip- pophagus Matthew and Cook.21 The only differences between the present form and T. hippophagus so far as comparison can be made, is the smaller size and the slightly slenderer jaw of T. temerarius. MUSTELIDJE. Paroligobum's (Brachypsalis) simplicidens gen. nov. Annals Carnegie Museum Vol. IV, pp. 44-46, 1906. Type: Right mandibular ramus (No. 1553, Cam. Mus. Cat. Vert. Foss.). Horizon: Lower Harrison beds (Miocene). Locality : Near Agate Spring Fossil Quarries, Sioux County, Nebraska. Plesiotype : Left mandibular ramus, greater portion of femur, fibula, and bones of fore and hind foot (No. 2389, Carn. Mus. Cat. Vert. Foss.). Horizon : Lower Harrison beds (Miocene). Locality : Agate Spring Fossil Quarries (quarry No. 3), Sioux County, Nebraska. Generic Characters. — Dentition, ?3,i.4.2. • Canine heavy and oblong in cross-section. Premolars crowded, heavy, rather bluntly crowned, no accessory posterior cusps except on p^. Carnassial comparatively large, approaching condition in recent forms (Gulo), but possesses a small entoconid which is located high up on the crown. M? much reduced in size. Femur longer than the tibia in the same proportion to that in Meles or Gulo. Feet broad and short; pes sub-plantigrade ; ungtcals not retractile. General Description of the Plesiotype and Specific Characters. The lower jaw is only very slightly larger than that in the type of this species previously described and is much larger than Meles taxus. The mandible below p-5 is very deep and the posterior mental foramen is located directly below the posterior fang of p3. Judging from the bottom of the alveolus for the canine, that tooth was strongly developed and the premolars appear to have been crowded, as in the type. The inferior border of the horizontal ramus is gently convex in the fore-and-aft direction, as in the type. Opposite the vertical ramus the inferior border is only slightly raised as in Gulo luscus and unlike that in Meles, in which the angle is higher. The posterior portion of the lower jaw as a whole is otherwise very like that of the latter genera ; possessing a large and deep temporal fossa, a large "Matthew, W. D., and Cook, Harold J., Bull. Amer. Mus. Nat. Hist., Vol, XXVI, p. 373, 1909. 270 MEMOIRS OF THE CARNEGIE MUSEUM coronoid process, transversely broad and well rounded condyle, heavy masseter ridge, and prominent hook-like angle. The condyle is on a horizontal line with the denti- tion and is slightly lower than in Meles taxus and more like that of Gulo luscus. The ramus is broken in the region of pT but this tooth is present in the type. P? is Fig. 64. Lower Jaw of ParoUgobunis simplicidens. j$ uat. size. 1, internal view of jaw ; 2, external view of jaw. of considerably large size judging from the posterior root, which is still in position in the jaw. P3 is robust, and, as in the type, has a great transverse diameter with the protoconid placed well anteriorly. Internally there is a smoothly rounded cingulum, which continues from the antero-internal angle of the tooth, as in Gulo luscus, and is somewhat more developed than in the type. The crown of the tooth is much abraded showing that the animal was fully adult. Back of p7 is a space of 11 mm. for P4. MT is unfortunately broken anteriorly so that the blade cannot be described. The metaconid is preserved and is rather small and placed high up on the crown. The heel is of moderately large size with the hypoconid well developed, forming the external edge of the heel, while the entoconid is probably absent.26 Back of mT there is a small alveolus, indicating a single-rooted tooth, which was even smaller than that tooth in the type. The contour of the lower jaw and especially the denti- tion are on the whole rather more suggestive of Gulo luscus than Meles taxus. !6The abraded condition of the tooth does not admit of a positive statement as to the presence or absence of an entoconid. PETERSON: NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 271 Measurements of Mandible. Mm. Total length of the ramus fragment 110 Antero-posterior diameter of PT 11 Greatest transverse diameter of P^ 8 Antero-posterior diameter of Mr approximately 18 Greatest transverse diameter of heel 8 Manus. — The manus is represented by a number of metacarpals of both sides, the unciform and trapezoid of the right foot and a phalanx ; the latter may equally well belong to the hind foot. The unciform is of very nearly the same proportionate height, but of less trans- verse diameter than in Meles taxus; the facet for the cuneiform descends on the ulnar side to about the same extent as in the recent form, while the facet for the scapho- Fia. 65. Right Mandibular Ramus of Paroligobunis simplicidens. § nat. size. Type No. 1553. l,external view ; 2, crown view of teeth. Fig. 66. Foot Bones of Paroligobunis simplicidens. f nat. size. No. 2398. 1, dorsal view of portion of right manus ; 2, dorsal view of a first and second phalanx ; 3, tibial view of mesocuneiform ; 4, dorsal view of niesocu- neiform. lunar occupies a liberal portion of the radial angle. The facet for the magnum is continuous from the distal to the proximal face, as in Daphanus and Daphosnodon, but the lunar facet is more distinctly separated from the facet for the magnum than in the latter genera. This is due to a prominent sinuous ridge, which extends from the dorsal border, obliquely upward and backward so that the upper portion of the facet for the magnum is concave supero-inferiorly, while the lower portion is plane antero-posteriorly ; the facet for Mc. Ill near the distal face is convex antero-pos- teriorly. The palmar tuberosity is compressed transversely ; it is more prominent than in Meles taxus or the recent otter, but does not extend so high up upon the palmar face as in Daphoenus or Daphosnodon. Distally the bone is much expanded and the articulations for Mc. IV and V are transversely broad and concave in the antero-posterior direction. The trapezoid is quite depressed and triangular in outline. Its proximal face 272 MEMOIRS OF THE CARNEGIE MUSEUM slants excessively downward in the ulnar direction, meeting the facet for Mc. II at an acute angle, so as to form a prominent and sharp ridge, which extends a short distance from the dorsal face backward almost identical with that in Melcs and Lutra. Back of this ridge on the ulnar face is a rugose area for muscular attach- ments, the bone terminating palmarly in a round obtuse point. Distally the sur- face for Mc. II is convex transversely, the radial side less strongly than the ulnar, while antero-posteriorly the articulation is very slightly concave. The radial face is taken up almost entirely by the large facet for the trapezium. The latter is undoubtedly of large size judging from the large facets on the trapezoid and on Mc. I. The first metacarpal is rather short when compared with that of Gulo, or even with that of the recent badger, and is more like that in the otter ; it is also proportion- ally shorter than that in Megalictis ferox Matthew from the lower Miocene of South Dakota. The proportionate length of the metacarpals as a whole agree quite closely, however, with those in the latter genus. The facet for the trapezium is, as stated, of large size, and it is obliquely convex antero-posteriorly, extending well down on the shaft as in Melcs, and is not unlike that of Megalictis feroxMatthevf and sEluro- cyon brevifacies Peterson.27 The shaft has received some injury in the process of col- lecting, but enough is preserved to show that it was heavy, well arched forward, and quite cylindrical. The distal trochlea is fully developed and carried the usual two phalanges. The second metacarpal is quite heavy and in its general proportion it compares well with that in Megalictis. The head rises above that on Mc. Ill, so that the ulnar face has a facet for the magnum similar to that in Meles, while in Megalictis the head is apparently raised higher (see illustration, fig. 15, of Dr. Matthew's paper, I. a). The facet for the trapezoid is concave transversely, the ulnar angle being higher than the radial. The palmar tuberosity is a hemispherical knob, while radially the head has an uneven surface for muscular attachment. On Mc. II of the right side there is no articular facet for Mc. I, while the one on the left side has a small facet near the dorsal angle. The origin of this facet may, in part, be due to injury received in this region, which is plainly shown by the rough surface of the proximal portion of the bone. The shaft is very heavy, slightly arched forward, and transversely oval in cross-section. The dorsal portion of the trochlea is not nearly as hemispherical as in Daphoenus, while plantarly it is divided by a prominent carina, the radial portion occupying a higher position than the ulnar, causing asymmetry of the trochlea. "See fig. 19, 5, Ann. Cam. Mus., Vol. IV, p. 71, 1906. PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 273 The third metacarpal is no heavier than the second, though longer. The articu- lar facet for the magnum slants inwardly, but the radial border is not raised as in Daphcenodon so as to form a transverse concavity, but instead it is very nearly plane transversely and evenly convex antero-posteriorly. On the radial side is a facet for Mc. II which is succeeded, posteriorly, by a deep sulcus, the palmar tuber- osity forming its posterior border. On the ulnar side the proximal end is ex- cavated in order to accommodate the head of Mc. IV and the articular facet for the unciform is concave due chiefly to the development of the dorso-ulnar angle of the head. The shaft is very slighty arched and transversely oval, due to the divergence of the metacarpals directly from the proximal end as in the mustelids generally. The trochlea is more symmetrical than that of Mc. II. Measurements. Mm. Unciform, greatest antero-posterior diameter 13 Unciform, greatest transverse diameter 12 Unciform, greatest vertical diameter 15 Trapezoid, greatest antero-posterior diameter 10 Trapezoid, greatest transverse diameter 9 Trapezoid, greatest vertical diameter 6 Metacarpal I, greatest length 26 Metacarpal II, greatest length 37 Metacarpal III, greatest length 41 Median phalanx, greatest length 17 A proximal phalanx of the pollex or hallux has the proximal end much ex- panded, the shaft strongly arched and of considerable length, while the distal trochlea is confined principally to the plantar face. A median phalanx found with the specimen is much depressed, quite long, and does not indicate retractility of the ungual. The Hind Limb. The greater portion of the femur is preserved. The head is well rounded and sits on a long neck, as in Meles, Lutra, and Gulo. The pit for the ligamentum teres is located on the posterior half near the lower border. Unfortunatety the great tro- chanter, together with the upper portion of the shaft, is lost on its external side, while internally the shaft is preserved, showing a large lesser trochanter. On the lower half of the shaft the fibular border is produced into a very prominent ridge, which is even more pronounced than in Meles taxus or Gulo luscus and terminates below in a rugose tubercle similar to that of Ganis. The lateral development of the shaft gives it the characteristic transversely broad appearance seen in Gulo luscus or Meles taxus. The rotular trochlea is shallow and broad, the condyles have great trans- 274 MEMOIRS OF THE CARNEGIE MUSEUM verse diameter and are separated by a rather shallow groove, the external condyle being quite oblique to the long axis of the bone. The fibula is heavy and has the proximal and distal ends much expanded. The shaft sweeps gently towards the shaft of the tibia and is also gently arched in the backward direction. The facet for the astragalus is nearly plane and the pos- terior face has the tendinal groove and external malleolus fully as well developed as in the recent badger or otter. Fia. 67. Limb Bones of Paroligobunis aimplicidens. 5 nat. size. 1, anterior view of femur ; 2, posterior view of femnr ; 3, fibular view of fibula ; 4, anterior view of fibula. Both astragali are represented. In comparison with the length of the bone the trochlea is somewhat narrower than that of Megaliclis fcrox. The trochlea is very shallow and is more oblique and imperfect than in either Meles or Lutra and rather more like that of Gulo luscus, the internal condyle being less developed, while the exter- nal is higher than in the two recent forms first mentioned. The tibial face of the inter- nal condyle is not interrupted by the sulcus, which is so characteristic of Daphosnodon, but is continuous as in Gulo and Meles. The pos- terior limit of the trochlea, as well as its down- 63. Astragalus of Paroligobunis simplici- nat. size. 1, anterior view; 2, superior view ; 3, tibial view. ward extent on the neck of the anterior face, is approximately in the same propor- PETERSON : NEW CARNIVORES FROM MIOCENE OF WESTERN NEBRASKA 275 tion to that in the recent forms. One of the more curious features of the bone is the deep and obliquely placed sulcus, which separates the groove for the flexor muscles from the astragalar trochlea. In this groove is located the astragalar fora- men (see fig. 68, 2), which pierces the bone and again appears in the groove be- tween the internal and external facets for the calcaneum on the plantar face, as in Meles taxus. The neck is perhaps somewhat longer than in Megalictis, which again reveals a tendency towards Gulo luscus. The head is also expanded and convex in the same way. A mesocuneiform which was found with the specimen and very likely belongs with it, is high and narrow when compared with Meles, and is apparently also slightly higher in proportion to that of Megalictis, but is otherwise quite like that of the latter, the tibial face possessing a large facet for the entocuneiform. Measurements. Mm. Femur, leDgth from head to distal end 174 Femur, transverse diameter of distal end 42 Femur, anteroposterior diameter of distal end 36 Fibula, greatest length 130 Astragalus, greatest length 33 Astragalus, greatest transverse diameter 26 Astragalus, greatest transverse diameter of trochlea 16 Mesocuneiform, greatest antero-posterior diameter 11 Mesocuneiform, greatest transverse diameter 6 Mesocuneiform, greatest vertical diameter 8 Affinity and Systematic Position of Paroligobunis. While some features of the limbs in Paroligobunis simphcidens suggest that of Lutra ; i. e., the oblique and limited posterior extent of the internal condyle of the femur, the backward sweep of the shaft of the fibula, the proportionate length and arrangement of the metacarpals; there are more important characters which very strongly suggest that the animal was probably fossorial rather than aquatic in his habits. The femur is much longer than the fibula and the shaft of the former bone is straight and is transversly expanded near the distal end as in Meles. The fibula is on the whole more nearly like that of the latter genus than that of the otter, while the astragalus strongly suggests that of Gulo luscus and has the astragalar foramen, a characteristic feature in Meles. The John Day form, Oligobunis, was regarded as an aberrant member of the Ccmidse by Professor Cope, while Dr. Matthew (Bull. Amer. Mus., Vol. XXIII, p. 193, 1907) after the re-examination of the type, questions Cope's reference and transfers the genus to the Mustelidas. After a compar- ison and careful study of the newly acquired material now in the Carnegie Museum 276 MEMOIRS OF THE CARNEGIE MUSEUM I fully agree with Matthew in the taxonomic position of Oligobunis. Furthermore it is quite evident that the present genus is closely related to the John Day form and quite likely in a direct line of descent which the generic name used implies. Oligobunis lepidus Matthew (I. c, p. 19-i) is a connecting type ; it is a smaller species than the John Day form, but indicates a type specialized differently from the present genus, having Mg very much less reduced, while PTis more reduced than the same tooth in the present genus, indicating greater tendency towards the loss of one tooth and the retention of the other in 0. lepidus than in Paroligobunis simplicideus. M-g- in 0. lepidus is one half the antero-posterior diameter of the carnassial, while that tooth in the present genus is only one third the antero-posterior diameter of the carnassial, plainly showing that in the present genus there is a more strongly marked advance toward modern forms. Potamotherium (= Stephanodon) of Europe has a general resemblance to the new genus here proposed, but it is quite clear that this European form is at least generi- cally different. In comparing the present type specimens with Dr. Schlosser's 2S figures, it is seen that PT in the American form is smaller, the premolar series increases more in size, from before backward in the jaw, the metaconid of MTis less developed and placed higher up on the crown, and the heel is smaller. Upon further comparison it is seen that the lower jaw is deeper and shorter in Paroligo- bunis. Of especial importance is the proportion of the hind limbs of the two genera. In Potamotherium the femur is shorter than the tibia, an aquatic feature, while in Paroligobunis the femur is longer than the tibia. The material at hand displays further comparison which may, however, be regarded as of less importance between these two genera ; namely, the more strongly developed cingula and accessory tubercles of the premolars, and the external position of PT in the European form, while in Paroligobunis simplicidens Py is internal due to the reverse obliquity of P-j in the latter. Paroligobunis (Potamotherium) lacota which was provisionally referred to the European genus by Matthew and Gidley23 appears to be quite closely related to Paroligobinus simplicidens having apparently somewhat smaller canines and a slen- derer jaw than the latter species. Dr. Matthew states (I. c, p. 254) that Cope's type of Potamotherium (Lutrictis) Igcopotamicus is lost and cannot be verified with Cope's reference, there being no illustrations of the type specimen. Stenogale (Potamctherium) robusta Cope was based on a lower jaw containing part of the dentition and was found in the Upper Miocene near Fort Niobrara, Nebraska. !S Beitrage Paleontologie Osterich-Ungars und des Orients, PI. VIII, figs. 18, 63, 64, 65, 1888. "Boll. Amer. Mas. Nat. Hist., Vol. XX, p 254, 1904. PETERSON! NEW CARNIVORES FROM MIOCENE OF, WESTERN NEBRASKA 277 Matthew and Gidley doubtfully referred this type to Potamotlierium. As this Ameri- can form now stands I cannot see that it is much nearer Potamotlierium than Steno- gale as the jaw is deeper in proportion ; it is also somewhat shorter and more ro- bust, the symphysis is lighter, the premolars are smaller and Ms larger than in the former genus, while the latter genus has the under border of the lower jaw very much more curved fore-and-aft, the premolars are too large in proportion to the car- nassial, the heel of the latter is entirely different in shape, and very much smaller, and the tubercles of the teeth are evidently sharper and higher than in the American specimen. The latter form should, I think, occupy a new generic position (Brachy- gate) somewhat near the Oligobunis phylum although the premolars are proportion- ally smaller than in the latter, especially in the transverse diameter. In Paroligo- bunis the jaw is pi-oportionally shorter and heavier, the alveolar border is more curved from before backward, the premolars are more crowded, and the carnassial has apparently a sborter heel than in the new genus here proposed. Fig. G9. Alveolar Border and External View of Jaw of Bruchypsalis pacijcepltatus Cope. Nat. size. (Cope col- lection, No. 8544, Am. Mns. Natural History.) Brachypsalis pachycephalia Cope was proposed (7. c, p. 951) on a rather inade- quate type, a left mandibular ramus, from the same locality in which Stenogale robusta was found. Professor Cope afterwards referred this type to Potamotheriwm of Europe, while Matthew and Gidley (/. c, p. 254) accept the type as valid and distinct from Potamotlierium. In a former paper30 the present writer regarded Paro- ligoburvis (Brachypsalis) simplicideiis as possibly an ancestral form to Brachypsalis, a view which is no longer tenable inasmuch as the latter species has M» proportionally larger, and the carnassial apparently smaller in proportion to the premolars. (Compare figs. 64 and 65 with fig. 09.) "Ann. Cam. Mus., Vol. IV, p. 45, 1906. SThe-nlcHs tow* 5A- 278 MEMOIRS OF THE CARNEOIE MUSEUM JElurocyon Peterson and Megalictis Matthew are more modified forms which, however, are apparently related to this group of the Mastelidtr. One of the chief generic differences between Mlurocyon, Megalictis, Oligobunis, and Paroiigobunis is the presence of the tnetaeonid on MT of the two latter and the absence of it in the case of the two former genera. In Paroligobwn/is the metaconid is situated nearly as high up as the summit of the protoconid, and quite closely fused to the latter. ; Some of the Siwalik mustelids (MelivorQdon) are also closely allied to this general group. 1 fully agree with Dr. Matthew's11 statements that Mellivora and Gvlo are the nearest living allies of Mlwrodon and Megalictis; I would in fact also include Oligobunis, Paroligobunis, and possibly the proposed new genus Bntrhgguh1 as not so very distantly related especially to JElurocyon and Megalictis. o'nenic+is " Bull. Amer. Mua. Nat. Hist., Vol. XXIII, p. 204, 1907. Memoirs Carnegie Museum, Vol. IV. Plate LXXV. Drawn 6y V'«"/ Prentu Daphr.nodon euperbui. Type. (No. 1589, C. M. Cat. Vest. Foss.) Palatal View of Cra- xit m. Nat. Size. Memoirs Carnegie Museum, Vol. IV. Plate LXXVI. Draum by Sydney I'mitice Daphcmodon superbiu. Type. (No. 15S!», ('. M. Cat. Vert. Foss.) Superior View op Cra- nium, Nat. Size. Memoirs Carnegie Museum, Vol. IV. Plate LXXVII. Di-auu bif S,,,l,irii Praitic- Dcuphoenodon superbus. Type, (No. lf>X9, C. M. Cat. Vert. Foss. ) Anterior axd Posterior Views of Skull. Nat. Size. Plate LXXVIII. Daphmnodon superbus. Size. External View of Right Scapula (No. 1589, C. M. Cat. Vert. Foss.) axd Left Side of Pelvis (Note : the illustration of the pelvis is from a specimen in the collection of Amherst College.) n by Sydney Prentice Nat. [EM01RS CARMEGIE MUSEUM, VOL. IV. Plate LXXIX. O — *" Drawn l>u Sydney Prentice Duphoenodon superbus. Type. (No. 1589, C. M. Cat. Vert. Fobs.) 1, Radial View of Humerus of Eight Side. 2 An- terior View of Humerus ok Right Side. 3, Radial View of Radius op Right Side. 4, Anterior View of Radius of Right Side. 5, Ulnar View of Ulna of Right Side. 6, Anterior View of Ulna ok Right Side. All Fig- ures Natural Sizi;. Memoirs Carnegie Museum, Vol. IV. Plate LXXX. 'J Drawn by Sydney Prattice Duphoenodon svperbvs. Type. (No. 1589, C. M. Cat. Vert. Foss. ) Dorsal and Palmar Views of Right Manus. Nat. Size. (The hoods of the ungual phalanges are more or less broken.) Memoirs Carnegie Museum, Vol. IV. Plate LXXXI Daphcenodon superb™. Type. (No. 1589, C. M. Cat. Vert. Fobs.) 1, Anterior View of Left Femur. 2, Tibial View of Left Femur. 3, Anterior View of Left Tibia. 4, Fibular View of Left Fibula. 5, Posterior View of Left Fibula. All Figures Natural Size. (The illustrations of the femur are from a specimen in the collection of Amherst College.) Memoirs Carnegie Museum, Vol. IV, Plate LXXXII. Daphwnodon superbus. Type. (No. 1589, C. M. Cat. Vert. Foss. ) Dorsal and Plantar Views of the Pes. Natural Size. \\ i |zq a H w is H * S5 m O PS