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+The Project Gutenberg EBook of A New Species of Heteromyid Rodent from the
+Middle Oligocene of Northeast Colorado with Remarks on the Skull, by Edwin C. Galbreath
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: A New Species of Heteromyid Rodent from the Middle Oligocene of Northeast Colorado with Remarks on the Skull
+
+Author: Edwin C. Galbreath
+
+Release Date: November 23, 2010 [EBook #34412]
+
+Language: English
+
+Character set encoding: ASCII
+
+*** START OF THIS PROJECT GUTENBERG EBOOK NEW SPECIES OF HETEROMYID RODENT ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci
+and the Online Distributed Proofreading Team at
+https://www.pgdp.net.
+
+
+
+
+
+
+
+A New Species of Heteromyid Rodent from the Middle Oligocene of
+Northeast Colorado with Remarks on the Skull
+
+BY
+
+EDWIN C. GALBREATH
+
+University of Kansas Publications
+Museum of Natural History
+
+Volume 1, No. 18, pp. 285-300, 2 plates
+August 16, 1948
+
+University of Kansas
+LAWRENCE
+1948
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman; H. H. Lane, Edward H. Taylor
+
+Volume 1, No. 18, pp. 285-300, 2 plates
+August 16, 1948
+
+UNIVERSITY OF KANSAS
+Lawrence, Kansas
+
+PRINTED BY
+FERD VOILAND, JR., STATE PRINTER
+TOPEKA, KANSAS
+1948
+
+[Illustration]
+
+22-3342
+
+
+[Transcriber's Note: Words surrounded by tildes, like ~this~ signifies
+words in bold. Words surrounded by underscores, like _this_, signifies
+words in italics.]
+
+[Illustration: PLATE 2. _Heliscomys tenuiceps._ Univ. Kans. Mus. Nat.
+Hist., Vert. Paleo. Coll. No. 7702. A, dorsal view; B, lateral view; C,
+ventral view. All views approximately x 5.]
+
+[Illustration: PLATE 3. _Heliscomys tenuiceps._ Univ. Kans. Mus. Nat.
+Hist., Vert. Paleo. Coll. No. 7702. A, lateral view of right side of
+skull showing structures in orbital area. ALS, alisphenoid. FR, frontal.
+MAX, maxillary. OS, orbitosphenoid. PAL, palatine. PC, presphenoid
+canal. SF, sphenoidal fissure. SFr, sphenofrontal foramen. SPal,
+sphenopalatine foramen. Approximately x 9.3; B, occlusal view of P4-M3.
+Approximately x 23.4.]
+
+
+
+
+A New Species of Heteromyid Rodent from the Middle Oligocene of
+Northeast Colorado with Remarks on the Skull
+
+By
+
+EDWIN C. GALBREATH
+
+
+Heretofore our knowledge of the osteology of _Heliscomys_ Cope has been
+extremely limited; this genus previously was known by its teeth,
+fragmental maxillaries, incomplete palatine bone and mandible, and part
+of one forelimb. In the summer of 1946 the writer, as a member of the
+University of Kansas Museum of Natural History field party, discovered
+the anterior part of a skull of _Heliscomys_ in the middle Oligocene
+deposits of Logan County, Colorado. This specimen, representing a new
+species, yields a welcome, and greatly desired addition to our fund of
+information about the genus.
+
+The writer is indebted to Dr. Robert W. Wilson for advice and helpful
+criticism in the course of this study, and to Mr. Bryan Patterson of the
+Chicago Natural History Museum for the loan of comparative material.
+Mrs. Bernita Mansfield of the Geology Department, University of Kansas,
+prepared the plates.
+
+
+Family HETEROMYIDAE
+
+~Heliscomys tenuiceps~, new species
+
+_Holotype._--Anterior part of a skull with left P4-M3, No. 7702,
+Vertebrate Paleontological Collection, Museum of Natural History,
+University of Kansas.
+
+_Geological Age and locality._--Silts of Orellan age in the Cedar Creek
+facies of the Brule formation in "Chimney Canyon," Sec. 3, T. 11 N, R.
+54 W, Logan County, Colorado.
+
+_Diagnosis._--Size larger than any known species; P4 with
+posteroexternal cusp (metacone) anterior to central (hypocone) and
+lingual (entostyle) cusps, which are connected by a cingulum; internal
+cingula of molars undivided, and as high as paracone and metacone; style
+of each cingulum opposite the straight median valley; rostrum deep and
+laterally compressed.
+
+_Description._--The type consists of the preorbital and interorbital
+parts of a skull. Its size is comparable to that of the Recent
+heteromyid, _Liomys pictus_ Merriam. _L. pictus_ is the species referred
+to in the comparisons below when only the generic name _Liomys_ is
+mentioned. Both incisors have been broken off. The right tooth-row is
+missing, but the left row is complete, and its orientation indicates
+that the tooth rows were parallel. The zygomata are broken off close to
+the rostrum, which is relatively narrow in comparison with its length
+and depth. In this narrowness, the specimen resembles _Florentiamys_
+Wood more than it does such Recent heteromyids as _Liomys_ or
+_Heteromys_, where the rostrum is much wider at the dorsal surface than
+at the ventral surface (correlating with the wide interorbital
+dimension). In No. 7702 the rostrum is not appreciably expanded on the
+dorsal surface. The wide interorbital dimension also gives a tapering
+appearance to the rostrum of the Recent heteromyids, when viewed
+dorsally, which is not seen in the fossil specimen. Like those of most
+heteromyids, the nasals and premaxillaries project forward beyond the
+incisors.
+
+_H. tenuiceps_ has a distinctly heteromyidlike appearance, and it is
+obvious that the features of the anterior part of the skull, which
+characterize the heteromyids, had been established by middle Oligocene
+time.
+
+The nasal bone extends caudad as far as does the premaxillary; they
+terminate at the anterior border of the orbit. The nasal is widest
+anteriorly where it curves downward on the side to meet the anterior
+projection of the premaxillary bone beyond the incisor. Posteriorly, the
+two nasals have practically parallel lateral borders much as in
+_Liomys_.
+
+The frontal bone dorsally is relatively narrower than in any Recent
+heteromyid, and closely resembles that of the geomyids. There is a
+slight depression in the midline of the skull where the two frontals
+unite, but no evidence of a ridge for the attachment of the temporal
+muscle. In lateral view, the ledge seen in _Liomys_ at the dorsal
+surface is absent, nor is this surface rounded as in _Geomys_.
+Preservation around the nasolacrimal canal is poor, but traces of
+sutures indicate that the frontal bone is not involved in the
+posteromedial wall of that canal. The orbital plate is broad,
+comparatively flat, and extends farther ventrad than in _Liomys_, and
+enters into the composition of the sphenopalatine foramina. Ventrally
+the frontal bone meets the orbital processes of the palatine and
+maxillary bones, and posterolaterally meets the orbitosphenoid.
+
+In the anterodorsal angle of the rim of the orbit the lacrimal bone
+rests against the frontal and maxillary bones, where the body of the
+lacrimal contributes to the formation of the posteromedial wall of the
+nasolacrimal canal. Only a slight part of the maxillary process of the
+lacrimal remains on each side.
+
+The premaxillary bone, which constitutes most of the anterior part of
+the rostrum, is typically heteromyid in shape. The frontal process is
+long and slender. On the side of the rostrum the premaxillary forms the
+anterointernal border of the infraorbital foramen. The ventrolateral
+border of the bone is expanded slightly and aids in the formation of the
+tuberosity made by the maxillary bone at the ventroposterior border of
+the foramen. Ventrally the premaxillary makes up the anterior two-thirds
+of the lateral wall of the incisive (anterior palatine) foramen. It is
+not possible to establish what part of the median septum between the
+foramina is made up of premaxillary bones. The incisor arches through
+the premaxillary in a manner similar to that in _Liomys_, with the upper
+wall of the root canal being formed by the upper surface of the bone.
+Due to the narrowness of the rostrum, the root of the incisor is
+prominently outlined on the side of the rostrum, both in the
+premaxillary and maxillary bones. With this modeling of the side of the
+rostrum because of the incisor root canal, and the flaring of the
+posterior and ventral edges of the infraorbital foramen, the side wall
+of the premaxillary appears as a depressed area. Anterior to the incisor
+root the tip of the premaxillary projects forward, and parallels its
+opposite, laterally, instead of turning inward as in _Liomys_. This
+condition, together with the prominence of the root canal, makes the
+anterior tip project as a flange. The premaxillary extends downward as a
+plate of bone, and embraces the posterior and lateral sides of the
+incisor as in Recent heteromyids. The interpremaxillary foramen, if
+present, is obscure. However, there appears to be a foramen posterior to
+the incisor, which possibly has taken over the function of the
+interpremaxillary foramen.
+
+Both maxillary bones are incomplete, and lack the zygomatic processes.
+The rostral part of the maxillary is compressed laterally, as is the
+premaxillary. The anterior border of the maxillary contributes to the
+formation of the border of the anterior opening of the infraorbital
+canal where, at the posteroventral border of the opening, the bone is
+produced into a prominent tuberosity which projects laterally
+approximately one millimeter on each side. The infraorbital foramen
+(anterior opening of the infraorbital canal) lies about midway between
+the anterior end of the skull and the root of the zygoma. High on each
+side of the rostrum, and beneath the dorsal edge of the masseteric
+plate, is an area containing small foramina. The zygomasseteric plate is
+inclined forward at the dorsal end, and extends anteriorly almost to the
+highest part of the arch of the canal for the root of the incisor. The
+posterior end of the infraorbital canal lies on the median side of the
+zygomatic root as it does in _H. hatcheri_ Wood. Ventrally the zygomatic
+root rises above the fourth premolar as in _H. gregoryi_ Wood, _H.
+hatcheri_, and in Recent heteromyids. The ventral part of the orbit,
+containing the sphenopalatine foramen, presphenoid foramen, and the
+sphenoidal fissure, is not constricted as in _Liomys_, but is open like
+that of the squirrels. This condition is emphasized by the narrowness of
+the interorbital part of the skull and the more vertical position of the
+orbital plate.
+
+The alisphenoid bone is large and forms part of the posteromedial wall
+of the orbit. The sphenofrontal foramen lies in the suture between the
+extreme anterior margin of this bone and the frontal bone.
+
+The orbitosphenoid bone makes up little of the orbital wall. It occupies
+the posterior area of the orbit between the alisphenoid and palatine,
+and is in contact with these bones and the frontal. The presphenoid
+canal between the orbits is large, and the entrance at each end is well
+separated from the sphenoidal fissure. Damage to the sphenoidal fissure,
+which occurred prior to preservation, obscures its relationship to the
+optic foramen. No bar was found that would indicate that the two
+openings were widely separate. Anteroventrally the sphenoidal fissure is
+bounded by the orbitosphenoid bone, and dorsolaterally by the
+alisphenoid bone. Between the presphenoid foramen and the
+orbitosphenoid-frontal suture there is a distinct ridge, and the suture
+between the two bones lies in an elongate pit or trough formed by the
+anterior sloping side of the ridge and the impressed lateral wall of the
+frontal bone.
+
+The palatine bone is represented by fragments joined to other bones of
+the skull. The maxillary process of the left palatine bone is united to
+the maxillary by a highly sinuous suture. The union of the palatines to
+the maxillaries make a suture in the shape of a "V" with the base
+forward and somewhat blunt. The canal for the palatine artery and nerve
+has a multiple opening on the palate. One major foramen opens on each
+side of the palatomaxillary suture, and two or possibly three smaller
+foramina open posteriorly on the palatine bone. Prominent on the
+palatine bone, posteromedial to the third molar, is the foramen
+(palatine pit) for the palatine vein. Collectively, this complex of
+foramina is often known as the posterior palatine foramina. Wood (1933)
+states that _H. gregoryi_ has two posterior palatine foramina as in
+Recent genera, the anterior one opening opposite the posterior end of
+M1, and the posterior one opposite the median part of M3. The orbital
+process of the left palatine bone lies inside (medial to) the palatine
+process of the maxillary. Anteriorly this orbital process meets the
+orbital process of the maxillary bone, and the sphenopalatine foramen is
+found in the suture between these two bones and the frontal.
+
+As previously mentioned, the preserved dentition of this specimen
+consists of the complete left row of cheek teeth and roots of the
+incisors.
+
+The incisor is compressed laterally, more so than in any Recent
+heteromyid. The anterior face is rounded, asulcate, and covered with a
+heavy band of enamel, whereas the posterior side, due to lateral
+compression, is drawn out into a thin blade. The root of the incisor is
+at the lateral border of the premaxillary, so it is obvious that the two
+incisors converged on each other at the midline to form a cutting
+surface. The writer has not examined the asulcate, laterally compressed
+incisors of _H. hatcheri_, and cannot say how they compare with this
+specimen.
+
+The most significant features of the cheek teeth are their size, and the
+undivided internal cingulum. The molars are well worn, but the pattern,
+as a whole, is easily discernable.
+
+P4 has an anterior cusp and three posterior cusps as in other members of
+the genus. However, the buccal cusp (metacone) of the metaloph is
+considerably anterior to the central (hypocone) and lingual (entostyle)
+cusps, and the three cusps do not form a curve as in other species. In
+size the central cusp is largest, the buccal cusp is practically as
+large, and the lingual cusp is small. A cingulum connects the lingual
+and central cusps at the posterior margin of the tooth. In the Pipestone
+Springs specimen of _Heliscomys_ reported by McGrew (1941) the central
+and buccal cusps were connected by a cingulum, and some _H. hatcheri_
+specimens have all three cusps connected in a similar manner. A low arm
+or ridge extends from the lingual cusp forward to the lingual side of
+the base of the anterior cusp. The valleys between the posterior cusps
+are shallow. There is no sign of the small cuspule on the anteroexternal
+base of the anterior cusp seen in _H. gregoryi_, _H. hatcheri_, and the
+Pipestone Springs specimen. However, when one sees the variability of
+the cuspules on P4 of _H. hatcheri_, the presence of a minor cuspule
+does not seem to be of taxonomic importance.
+
+M1 deviates from the pattern typical of _Heliscomys_ more than do any of
+the other molar teeth. However, it must be kept in mind that some of the
+differences may be due to wear. For example, the protocone and paracone,
+and the hypocone and metacone are united to form protoloph and metaloph
+respectively. If the height of the external border of the paracone and
+metacone is taken into account and compared with the worn inner parts of
+these two cusps and the equally well-worn protocone and hypocone, it
+appears that these cusps formed no more of a true bilophodont tooth than
+do the cusps in other species of _Heliscomys_; in each of the species
+the cusps generally are separate entities. _H. gregoryi_ is reported to
+have an "incipient tendency to form lophs," and _H. hatcheri_ does the
+same when worn, but by union with the anterior cingulum. If cusps in _H.
+tenuiceps_ do form lophs, the process is definitely not by union of the
+cusps with the anterior cingulum. The transverse median valley is deep
+and divides the tooth on the buccal side. The anteroposterior valleys
+are shallow and hanging, and can be said to exist only as indentations
+between the two sets of cusps. The paracone and metacone are much higher
+than the other two cusps, but much of this disparity in height may be
+the result of greater wear on the protocone and hypocone; _H. gregoryi_
+agrees with _H. tenuiceps_ in these respects. Possibly the protocone and
+hypocone were much larger than the paracone and metacone. The internal
+cingulum of M1 exhibits only one large cusp opposite the medial end of
+the transverse valley, and shows no evidence of having been divided into
+two cusps. It is barely possible that there may have been two cusps and
+that wear makes it appear that there was only one. I doubt that there
+were two cusps because the cingulum is still so high (as high as the
+outer edges of the paracone and metacone) as to suggest that it is only
+slightly worn. Posteriorly this single cusp in the cingulum is united
+with the hypocone. Anteriorly the cusp is confluent with an anterior
+cingulum that is small, but, nevertheless, plainly visible as it crosses
+the occlusal face of the tooth to the paracone. There is some reason to
+believe that there was a posterior cingulum, but wear, which has
+obliterated even the posterior wall of the hypocone, prevents my being
+certain about this. This cingulum is absent in _H. gregoryi_ and present
+in _H. hatcheri_.
+
+M2 compares favorably with M1 except for the following differences: The
+protocone and hypocone are equal to the paracone and metacone in area,
+but not in height; although the internal cingulum is undivided, there is
+no evidence of a cusp as in M1. Here, too, the cingulum is as high as
+the paracone and metacone. Possibly the cingulum was confluent with the
+hypocone. The internal cingulum continues around the margin of the tooth
+to the paracone as an anterior cingulum which is sharper and plainer
+than the anterior cingulum on M1. There is no evidence of a posterior
+cingulum.
+
+M3 shows a great amount of wear, and the occlusal pattern is not too
+clear. The median transverse valley is reduced almost to a pit, and the
+paracone and metacone are divided by a small notch. The protocone and
+paracone, the latter being much higher, are larger than the metacone
+which is reduced in size, and not all this difference in size can be the
+result of wear. The hypocone is absent. The internal cingulum is as high
+as the paracone and shows no evidence of division into two cusps, but in
+M3 this character is apparently variable for _H. gregoryi_ does not have
+the internal cingulum divided and _H. hatcheri_ has it markedly so. A
+slight anterior arm of the internal cingulum may have reached forward to
+the anterior face of the protocone. Wear prevents knowing whether a
+crest surrounds the tooth completely, or only on three sides.
+
+In size the teeth of _H. tenuiceps_ average twenty per cent larger than
+any of the upper teeth of _H. gregoryi_, _H. hatcheri_, or the Pipestone
+Springs specimen, and exceed any of the known lower teeth including
+those of _H. vetus_ and _H. senex_ by twenty-five per cent or more.
+Inasmuch as the upper teeth rarely exceed the lower in length in all the
+related genera of heteromyids, it is assumed that a similar relationship
+existed between the upper and lower molars of _H. tenuiceps_ and,
+therefore, that this species can be distinguished by its large size. The
+relative size of the premolars and molars is the same in _H. tenuiceps_
+as in other species of _Heliscomys_. However, within the framework of
+this similar relationship there are two differences. P4 of _H.
+tenuiceps_ is relatively larger than the P4 of _H. gregoryi_, and
+relatively smaller than the P4 of _H. hatcheri_. The width of the molars
+is relatively greater in _H. tenuiceps_ and _H. gregoryi_ than in _H.
+hatcheri_.
+
+
+MEASUREMENTS
+
+(In millimeters)
+
+                                                  U. K. M. N. H.
+                                                  (Vert. Paleo.)
+                                                     No. 7702
+Height of skull at M2                                  7.48
+Length from anterior end of nasals to rear of M3      15.41
+Length of nasal bones                                 10.50
+Width of rostrum at highest point of root canal        3.97
+Interorbital width                                     4.39
+Estimated length of skull                             25.00
+I, anteroposterior length                              1.56
+I, transverse width                                    0.63
+P4-M3 crown length                                     3.75
+P4-M3 alveolar length                                  3.80
+P4, anteroposterior length[A]                          1.05
+P4, transverse width                                   1.08
+M1, anteroposterior length                             0.93
+M1, transverse width                                   1.17
+M2, anteroposterior length                             0.93
+M2, transverse width                                   1.14
+M3, anteroposterior length                             0.78
+M3, transverse width                                   0.93
+
+[Note A: This and the following measurements at occlusal surface.]
+
+_Discussion._--_Heliscomys tenuiceps_ shows beyond any doubt that the
+heteromyid pattern of skull was developed by mid-Oligocene times, and in
+this species was already undergoing lateral compression. The major
+change later made in heteromyid skulls is broadening of the dorsal
+surface of the skull in the interorbital area.
+
+The complete confirmation of Wood's (1939) statement that the
+"sciuromorph" zygomasseteric structure had been developed by this time
+in the heteromyid rodents as it had been in the early Eomyids is
+demonstrated in this specimen. Further, it is to be noted that the
+infraorbital canal is not sciuridlike, but has been forced forward on
+the rostrum, as in the Geomyoidea.
+
+In some ways this skull shows similarities to _Florentiamys loomisi_
+Wood, of the early Miocene, which aid in determining the relationship of
+that unusual genus to _Heliscomys_ and to the heteromyids in general.
+When Wood described _Florentiamys_ the peculiar combination of
+characters found in this animal prompted him to speculate that: (1) It
+was a typical heteromyid which had secondarily developed cingula; (2)
+its cheek teeth were nearer the primitive pattern than were those of any
+other known fossil heteromyid, and that _Heliscomys_ represented a
+simplification in the reduction of the cingula; or (3) it was not a
+heteromyid, but a parallel development from the "Paramys" stock. Wood
+favored the second possibility. Now that a part of the skull of one
+species of _Heliscomys_ is known, the undivided internal cingulum that
+is confluent with the hypocone, the lateral compression of the deep
+rostrum, and the general similarity to the heteromyids appear as points
+in common between the two skulls, and demonstrate the closeness of
+_Florentiamys_ to the heteromyids. However, the specimen does not
+contribute anything new to use in choosing between Wood's first two
+postulates. In the writer's opinion the undivided internal cingulum is a
+primitive condition that has survived in _Florentiamys_ and _Heliscomys
+tenuiceps_. This common character together with the laterally compressed
+rostrum leads me to think that structurally, _H. tenuiceps_ is a link
+between _Florentiamys_ and the ancestral form of _Heliscomys_.
+Admittedly P4 of _Florentiamys_ seems far from the _Heliscomys_ pattern,
+but I think that this highly specialized structure could have been
+derived from _Heliscomys_ or a common ancestor.
+
+
+
+
+LITERATURE CITED
+
+
+MCGREW, PAUL O.
+
+1941. Heteromyids from the Miocene and Lower Oligocene. Geol. Ser. of
+Field Mus. Nat. Hist., vol. 8, pp. 55-57, 1 fig.
+
+WOOD, ALBERT E.
+
+1933. A New Heteromyid Rodent from the Oligocene of Montana. Jour.
+Mamm., vol. 14, pp. 134-141, 5 figs.
+
+1935. Evolution and Relationship of the Heteromyid Rodents with New
+Forms from the Tertiary of Western North America. Annals of the Carnegie
+Mus., vol. 24, pp. 73-262, 157 figs.
+
+1937. Part II. Rodentia, in The Mammalian Fauna of the White River
+Oligocene; by William Berryman Scott and Glenn Lowell Jepsen. Trans.
+Amer. Phil. Soc., n.s., vol. 28, pp. 155-269, figs. 8-70, pls. 23-33.
+
+1939. Additional Specimens of the Heteromyid Rodent Heliscomys from the
+Oligocene of Nebraska. Amer. Jour. Sci., vol. 237, pp. 550-561, 11 figs.
+
+_Transmitted March 1, 1948._
+
+22-3342
+
+
+
+
+
+End of the Project Gutenberg EBook of A New Species of Heteromyid Rodent
+from the Middle Oligocene of Northeast Colorado with Remarks on the Skull, by Edwin C. Galbreath
+
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