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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/34412-8.txt b/34412-8.txt new file mode 100644 index 0000000..77e8cfb --- /dev/null +++ b/34412-8.txt @@ -0,0 +1,854 @@ +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: ISO-8859-1 + +*** 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 × 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 × 9.3; B, occlusal view of P4-M3. +Approximately × 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. 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Galbreath. + </title> + <style type="text/css"> + + p { margin-top: .75em; + text-align: justify; + margin-bottom: .75em; + } + h1,h2,h3,h4,h5,h6 { + text-align: center; /* all headings centered */ + clear: both; + } + hr { width: 33%; + margin-top: 2em; + margin-bottom: 2em; + margin-left: auto; + margin-right: auto; + clear: both; + } + + table {margin-left: auto; margin-right: auto;} + + body{margin-left: 10%; + margin-right: 10%; + } + + .pagenum { /* uncomment the next line for invisible page numbers */ + /* visibility: hidden; */ + position: absolute; + left: 92%; + font-size: smaller; + text-align: right; + } /* page numbers */ + + .linenum {position: absolute; top: auto; left: 4%;} /* poetry number */ + .blockquot{margin-left: 5%; margin-right: 10%;} + + + .center {text-align: center;} + .smcap {font-variant: small-caps;} + + .caption {font-weight: bold;} + + .figcenter {margin: auto; text-align: center;} + + .figleft {float: left; clear: left; margin-left: 0; margin-bottom: 1em; margin-top: + 1em; margin-right: 1em; padding: 0; text-align: center;} + + .figright {float: right; clear: right; margin-left: 1em; margin-bottom: 1em; + margin-top: 1em; margin-right: 0; padding: 0; text-align: center;} + + .footnotes {border: dashed 1px;} + .footnote {margin-left: 10%; margin-right: 10%; font-size: 0.9em;} + .footnote .label {position: absolute; right: 84%; text-align: right;} + .fnanchor {vertical-align: super; font-size: .8em; text-decoration: none;} + + .poem {margin-left:10%; margin-right:10%; text-align: left;} + .poem br {display: none;} + .poem .stanza {margin: 1em 0em 1em 0em;} + .poem span.i0 {display: block; margin-left: 0em; padding-left: 3em; text-indent: -3em;} + .poem span.i2 {display: block; margin-left: 1em; padding-left: 3em; text-indent: -3em;} + .poem span.i4 {display: block; margin-left: 2em; padding-left: 3em; text-indent: -3em;} + + </style> + </head> +<body> + + +<pre> + +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: ISO-8859-1 + +*** 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. + + + + + + +</pre> + + + + +<h1>A New Species of Heteromyid Rodent from the Middle Oligocene of +Northeast Colorado with Remarks on the Skull</h1> + +<h3>BY</h3> + +<h2>EDWIN C. GALBREATH</h2> + +<p class="center"> +University of Kansas Publications<br /> +Museum of Natural History<br /> +<br /> +Volume 1, No. 18, pp. 285-300, 2 plates<br /> +August 16, 1948<br /> +<br /> +University of Kansas<br /> +LAWRENCE<br /> +1948<br /> +<br /> +<br /> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman; H. H. Lane, Edward H. Taylor<br /> +<br /> +Volume 1, No. 18, pp. 285-300, 2 plates<br /> +August 16, 1948<br /> +<br /> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +PRINTED BY<br /> +FERD VOILAND, JR., STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1948<br /> +<br /> +<br /> +22-3342<br /> +</p> + +<hr style='width: 45%;' /> + +<div class="figcenter" style="width: 428px;"> +<img src="images/i_003.jpg" width="428" height="650" alt="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 × 5." title="" /> +<span class="caption">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 × 5.</span> +</div> + +<hr style='width: 45%;' /> + +<div class="figcenter" style="width: 499px;"> +<img src="images/i_004.jpg" width="499" height="650" alt="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 × 9.3; B, occlusal view of P4-M3. +Approximately × 23.4." title="" /> +<span class="caption">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 × 9.3; B, occlusal view of P4-M3. +Approximately × 23.4.</span> +</div> + + + +<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_289" id="Page_289">[Pg 289]</a></span></p> +<h2> New Species of Heteromyid Rodent from the Middle Oligocene of +Northeast Colorado with Remarks on the Skull</h2> + +<h4>By</h4> + +<h3>EDWIN C. GALBREATH</h3> + + +<p>Heretofore our knowledge of the osteology of <i>Heliscomys</i> 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 <i>Heliscomys</i> 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.</p> + +<p>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.</p> + + +<h4>Family HETEROMYIDAE</h4> + +<h4><b>Heliscomys tenuiceps</b>, new species</h4> + +<p><i>Holotype.</i>—Anterior part of a skull with left P4-M3, No. 7702, +Vertebrate Paleontological Collection, Museum of Natural History, +University of Kansas.</p> + +<p><i>Geological Age and locality.</i>—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.</p> + +<p><i>Diagnosis.</i>—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.</p> + +<p><i>Description.</i>—The type consists of the preorbital and interorbital +parts of a skull. Its size is comparable to that of the Recent +heteromyid, <i>Liomys pictus</i> Merriam. <i>L. pictus</i> is the species referred +to in the comparisons below when only the generic name <i>Liomys</i> 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 <i>Florentiamys</i> +Wood more than it does such Recent heteromyids as <i>Liomys</i> or +<i>Heteromys</i>, where the rostrum is much wider at the dorsal surface than +at the ventral<span class='pagenum'><a name="Page_290" id="Page_290">[Pg 290]</a></span> 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.</p> + +<p><i>H. tenuiceps</i> 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.</p> + +<p>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 +<i>Liomys</i>.</p> + +<p>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 <i>Liomys</i> at the dorsal +surface is absent, nor is this surface rounded as in <i>Geomys</i>. +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 <i>Liomys</i>, 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.</p> + +<p>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.</p> + +<p>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 <i>Liomys</i>, 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<span class='pagenum'><a name="Page_291" id="Page_291">[Pg 291]</a></span> as in <i>Liomys</i>. 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.</p> + +<p>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 <i>H. hatcheri</i> Wood. Ventrally the zygomatic +root rises above the fourth premolar as in <i>H. gregoryi</i> Wood, <i>H. +hatcheri</i>, 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 <i>Liomys</i>, 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.</p> + +<p>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.</p> + +<p>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.</p> + +<p>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<span class='pagenum'><a name="Page_292" id="Page_292">[Pg 292]</a></span> 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 <i>H. gregoryi</i> 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.</p> + +<p>As previously mentioned, the preserved dentition of this specimen +consists of the complete left row of cheek teeth and roots of the +incisors.</p> + +<p>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 <i>H. hatcheri</i>, and cannot say how they compare with this +specimen.</p> + +<p>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.</p> + +<p>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 <i>Heliscomys</i> reported by McGrew (1941) the central +and buccal cusps were connected by a cingulum, and some <i>H. hatcheri</i> +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 <i>H. gregoryi</i>, <i>H. hatcheri</i>, and the +Pipestone Springs specimen. However, when one sees the variability of +the cuspules on P4 of <i>H. hatcheri</i>, the presence of a minor cuspule +does not seem to be of taxonomic importance.</p> + +<p>M1 deviates from the pattern typical of <i>Heliscomys</i> 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 <i>Heliscomys</i>; in each of the species +the cusps generally are separate entities. <i>H. gregoryi</i> is reported to +have an "incipient tendency to form lophs,"<span class='pagenum'><a name="Page_293" id="Page_293">[Pg 293]</a></span> and <i>H. hatcheri</i> does the +same when worn, but by union with the anterior cingulum. If cusps in <i>H. +tenuiceps</i> 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; <i>H. gregoryi</i> +agrees with <i>H. tenuiceps</i> 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 <i>H. gregoryi</i> and present +in <i>H. hatcheri</i>.</p> + +<p>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.</p> + +<p>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 <i>H. gregoryi</i> does not have +the internal cingulum divided and <i>H. hatcheri</i> 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.</p> + +<p>In size the teeth of <i>H. tenuiceps</i> average twenty per cent larger than +any of the upper teeth of <i>H. gregoryi</i>, <i>H. hatcheri</i>, or the Pipestone +Springs specimen, and exceed any of the known lower teeth including +those of <i>H. vetus</i> and <i>H. senex</i> 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<span class='pagenum'><a name="Page_294" id="Page_294">[Pg 294]</a></span> of <i>H. tenuiceps</i> and, +therefore, that this species can be distinguished by its large size. The +relative size of the premolars and molars is the same in <i>H. tenuiceps</i> +as in other species of <i>Heliscomys</i>. However, within the framework of +this similar relationship there are two differences. P4 of <i>H. +tenuiceps</i> is relatively larger than the P4 of <i>H. gregoryi</i>, and +relatively smaller than the P4 of <i>H. hatcheri</i>. The width of the molars +is relatively greater in <i>H. tenuiceps</i> and <i>H. gregoryi</i> than in <i>H. +hatcheri</i>.</p> + + +<h4><span class="smcap">Measurements</span></h4> + +<h4>(In millimeters)</h4> + + +<div class='center'> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td align='left'></td><td align='center'>U. K. M. N. H.</td></tr> +<tr><td align='left'></td><td align='center'>(Vert. Paleo.)</td></tr> +<tr><td align='left'></td><td align='center'>No. 7702</td></tr> +<tr><td align='left'>Height of skull at M2</td><td align='right'>7.48</td></tr> +<tr><td align='left'>Length from anterior end of nasals to rear of M3</td><td align='right'>15.41</td></tr> +<tr><td align='left'>Length of nasal bones</td><td align='right'>10.50</td></tr> +<tr><td align='left'>Width of rostrum at highest point of root canal</td><td align='right'>3.97</td></tr> +<tr><td align='left'>Interorbital width</td><td align='right'>4.39</td></tr> +<tr><td align='left'>Estimated length of skull</td><td align='right'>25.00</td></tr> +<tr><td align='left'>I, anteroposterior length</td><td align='right'>1.56</td></tr> +<tr><td align='left'>I, transverse width</td><td align='right'>0.63</td></tr> +<tr><td align='left'>P4-M3 crown length</td><td align='right'>3.75</td></tr> +<tr><td align='left'>P4-M3 alveolar length</td><td align='right'>3.80</td></tr> +<tr><td align='left'>P4, anteroposterior length[A]</td><td align='right'>1.05</td></tr> +<tr><td align='left'>P4, transverse width</td><td align='right'>1.08</td></tr> +<tr><td align='left'>M1, anteroposterior length</td><td align='right'>0.93</td></tr> +<tr><td align='left'>M1, transverse width</td><td align='right'>1.17</td></tr> +<tr><td align='left'>M2, anteroposterior length</td><td align='right'>0.93</td></tr> +<tr><td align='left'>M2, transverse width</td><td align='right'>1.14</td></tr> +<tr><td align='left'>M3, anteroposterior length</td><td align='right'>0.78</td></tr> +<tr><td align='left'>M3, transverse width</td><td align='right'>0.93</td></tr> +</table></div> + +<p>[Note A: This and the following measurements at occlusal surface.]</p> + +<p><i>Discussion.</i>—<i>Heliscomys tenuiceps</i> 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.</p> + +<p>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.</p> + +<p>In some ways this skull shows similarities to <i>Florentiamys loomisi</i> +Wood, of the early Miocene, which aid in determining the relationship of +that unusual genus to <i>Heliscomys</i> and to the heteromyids in general. +When Wood described <i>Florentiamys</i> the peculiar combination of +characters found in this animal prompted him to speculate that: (1) It +was a typical heteromyid which had secondarily developed<span class='pagenum'><a name="Page_295" id="Page_295">[Pg 295]</a></span> cingula; (2) +its cheek teeth were nearer the primitive pattern than were those of any +other known fossil heteromyid, and that <i>Heliscomys</i> 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 <i>Heliscomys</i> 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 +<i>Florentiamys</i> 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 <i>Florentiamys</i> and <i>Heliscomys +tenuiceps</i>. This common character together with the laterally compressed +rostrum leads me to think that structurally, <i>H. tenuiceps</i> is a link +between <i>Florentiamys</i> and the ancestral form of <i>Heliscomys</i>. +Admittedly P4 of <i>Florentiamys</i> seems far from the <i>Heliscomys</i> pattern, +but I think that this highly specialized structure could have been +derived from <i>Heliscomys</i> or a common ancestor.</p> + + + +<hr style="width: 65%;" /> +<h2><a name="LITERATURE_CITED" id="LITERATURE_CITED"></a>LITERATURE CITED</h2> + + +<p><span class="smcap">McGrew, Paul O.</span></p> + +<p>1941. Heteromyids from the Miocene and Lower Oligocene. Geol. Ser. of +Field Mus. Nat. Hist., vol. 8, pp. 55-57, 1 fig.</p> + +<p><span class="smcap">Wood, Albert E.</span></p> + +<p>1933. A New Heteromyid Rodent from the Oligocene of Montana. Jour. +Mamm., vol. 14, pp. 134-141, 5 figs.</p> + +<p>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.</p> + +<p>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.</p> + +<p>1939. Additional Specimens of the Heteromyid Rodent Heliscomys from the +Oligocene of Nebraska. Amer. Jour. Sci., vol. 237, pp. 550-561, 11 figs.</p> + +<h4><i>Transmitted March 1, 1948.</i></h4> + +<p><span class='pagenum'><a name="Page_296" id="Page_296">[Pg 296]</a></span>22-3342</p> + + + + + + + + +<pre> + + + + + +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. 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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. 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