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MEYER POSTILLA Published by the Peabody Museum of Natural History, Yale University Postilla includes results of original research on systematic, evolution- ary, morphological, and ecological biology, including paleontology. Syntheses and other theoretical papers based on research are also welcomed. Postilla is intended primarily for papers by the staff of the Peabody Museum or on research using material in this Museum. Editor: Zelda Edelson Postilla is published at frequent but irregular intervals. Manuscripts, orders for publications, and all correspondence concerning publications should be directed to: Publications Office Peabody Museum of Natural History New Haven, Conn., 06520, U.S.A. Lists of the publications of the Museum are available from the above office. These include Postilla, Bulletin, Discovery, and special publica- tions. Postilla and the Bulletin are available in exchange for relevant publications of other scientific institutions anywhere in the world. Inquiries regarding back numbers of the discontinued journal, Bulletin of the Bingham Oceanographic Collection, should be directed to: Kraus Reprint Co. Route 100 Millwood, New York 10546 A NEW OLIGOCENE HYRAX FROM THE JEBEL EL QATRANI FORMATION, FAYUM, EGYPT GRANT E. MEYER Peabody Museum of Natural History Yale University, New Haven, Connecticut 06520 (Received 5 June 1973) ABSTRACT A fossil hyrax from the Upper Fossil Wood Zone, Jebel el Qatrani formation is described. It represents a new genus and species Thyrohyrax domorictus. The type specimen as well as two referred specimens exhibit an internal man- dibular fenestra whereas four referred mandibular fragments do not. One nearly complete mandible with an internal mandibular fenestra has extremely inflated horizontal rami which enclose swollen ovoid chambers opening at the fenestra. This inflated ramus, which is believed to be unique among all mam- mals, and the internal mandibular fenestra are thought to be sexual char- acters. Evidence from other genera of the Saghatheriinae suggests these characters are found in females; not in males. POSTILLA 163: IIp. 31 DECEMBER 1973 2 POSTILLA 163 INTRODUCTION Yale University expeditions to the Fayum Province, Egypt, during 1961 to 1967 located fossil vertebrates in the Upper Fossil Wood Zone of the Jebel el Qatrani formation. Rich faunas were found at several localities in the strata of the upper part of the formation and seven Yale quarries were opened. The small hyracoid reported in this paper was found at Yale Quarry G near the base of the upper member of the Jebel el Qatrani formation and at Quarries M and I, near the middle of the upper member. The fauna found at Quarry G consists entirely of small-sized mammals or of small fragments of larger species and included Apidium moustafai, Propliopithecus cf. haeckeli, Phiomys andrewsi, Phiomys paraphiomyoides, Metaphiomys schaubi, Apterodon cf. altidens, proviverrid (new gen. and sp.). In Quarry I the hyracoid was found with Parapithecus sp., Apidium phiomense, Proplio- pithecus sp., Aegyptopithecus zeuxis, Aeolopithecus chirobates, Paraphiomys simonsi, Metaphiomys beadnelli, Apterodon cf. altidens, Apterodon macrog- nathus, Pachyhyrax crassidentatus, Bunohyrax fajumensis, Megalohyrax eocaenus, Titanohyrax ultimus and Bothriogenys gorringei. ABBREVIATIONS Abbreviations used in this paper are as follows: I incisor (I! = first upper incisor, I, = second lower incisor) C canine (C! = upper canine, C, = lower canine) M molar (M! = first upper molar, M. = second lower molar) P premolar (P? = third upper premolar, P, = fourth lower premolar) CGM_ Cairo Geological Museum, Cairo, Egypt YPM Peabody Museum of Natural History, Yale University, New Haven, Connecticut A NEW OLIGOCENE HYRAX 3 SYSTEMATICS ORDER HYRACOIDEA Huxley, 1869 FAMILY PLIOHYRACIDAE Osborn, 1899 SUBFAMILY SAGHATHERIINAE Andrews, 1906 Thyrohyrax, new genus TYPE. Thyrohyrax domorictus, new species. KNOWN DISTRIBUTION. Upper Fossil Wood Zone, Jebel el Qatrani formation, Fayum, Egypt. DIAGNOSIS. Small Saghatheriinae with a long narrow symphysis and brachy- 2.9 odont, selenolophodont dentition. The dental formula is rece I, is slightly larger than I, and its root is more rounded. Diastemata are between I, and I, and the lower canine. The lower canine is two-rooted. P,-P, molariform. Msg with a small, low hypoconulid. All cingula absent or very weak on lower teeth. P?-P* molariform, lophodont with well-developed preprotocristae and prehypocristae which often come in contact with the ectoloph. Internal man- dibular fenestra present on some specimens, absent on others. Those speci- mens with fenestra have greatly expanded ramus with large inflated chamber within walls of ramus. ETYMOLOGY: from Thyra, Greek, a door, a window; hyrax, a term referring to the internal mandibular fenestra in the ramus. Thyrohyrax domorictus, new species TYPE. CGM 40001. Collected by G. E. Meyer in February, 1967 from Quarry M, Jebel el Qatrani formation, Egypt. Fragment of right horizontal Tamus with P,-P,, roots of M, and %2M.-M;. On the lingual side of the ramus approximately 5.5 mm below Ms, the top half of an internal mandib- ular fenestra is present. The roof of the internal chamber can be seen below, separate from the dentition. 4 HYPODIGM. Quarry M: 1967 Quarry G: 1962 1963 1964 1965 Quarry I: 1963 1964 1965 1966 POSTILLA CGM 40001 (Type specimen). YPM 23886, left ramus with P,-Mz, and part of the symphysis. YPM 18086, left M?. YPM 18037, left M?. YPM: 23827; left P+. CGM 40004, right Ps, I! and M?. YPM 23904, ?DP; ?7DP, CGM 4000S, right I°, C and ?P! in three associated fragments of maxillae. YPMe33153507 DR. YPM 23815, right Ms. YPM 23817, right Mo. YPM 33149, right ?P4, M! and P,. YPM 33150, left M3. YPM 23823, left P? and 4 P? in maxillary fragment. YPM 23825, fragment of left ramus with P,-'% P3. YPM 23824, fragment of right ramus with Msg. YPM 33149, left ramus with Ps-Ms. YPM 33152, 13 tooth fragments. YPM 23832, fragment of right ramus with M,-Mb. YPM 21488, fragment of left maxilla with P4-M?%. YPM 33151, right M? and P4,. YPM 33148, mandible with left I,-I,, Po-M, and right P.-M3, ascending ramus missing. YPM 23828, fragment of left ramus with Mo-Ms3. YPM 23829, right Mg. CGM 40002, left M, and Ms. YPM 23858, fragment of right ramus with M,-Mg, right tibia, five metatarsals, five phalanges and one distal 163 phalanx of right pes, proximal end of one metacarpal and five phalanges of right manus, one lumbar vertebra and several indeterminate fragments. YPM 23833, symphysis with left I,-I, and right I, and C. YPM 23939, nearly complete mandible, edentulous except for left ’2M. and Ms, jaw greatly expanded. LOCALITY AND HORIZON. YPM Quarries G, M and I, Upper Fossil Wood Zone, Jebel el Qatrani formation, Fayum, Egypt. Late Oligocene. A NEW OLIGOCENE HYRAX 5 DIAGNOSIS. Only known species of the genus. ETYMOLOGY: from domos, Greek, a house, a structure; rictus, Latin, jaws, a term referring to the chamber within the jaws. DESCRIPTION. Small Saghatheriinae with brachyodont, selenolophodont denti- tion. The symphysis is long and fairly narrow with a narrow shelf or ridge at its ventral border. I, and I, are procumbent, tricuspid when unworn; I, is slightly larger than I,. I, is separated from I, by a diastema and must be a small tooth, as determined by the alveolus. The lower canine is two-rooted, a feature that is unique to this species among the Saghatheriinae, but typical of the Pliohyracinae. The tooth is small, with a rudimentary paraconid and a large protoconid. The lower canine is in contact with P,. P, is submolari- form, with a centrally-positioned paraconid, separated from the protoconid by a short paracristid. The hypoconid is not V-shaped and there is an un- usually deep, narrow hypoflexid. P.-P, are molariform with differentiated metaconids and entoconids. The metaconid exhibits a slight metastylid on its distal edge. There is a preparacristid on P.. The protocristids and hypo- cristids form complete lophs. The lower molars have a small paraconid, slightly buccal to the centerline, near the protoconid. The metastylid is better FIG. 1. Occlusial and medial views of the type specimen of Thyrohyrax domorictus, CGM-40001. The internal mandibular fenestra can be seen beneath Mg in the medial view. 6 POSTILLA 163 FIG. 2. Occlusial views of Thyrohyrax domorictus, above, YPM-23886, a maxilla with P4-M® and below, YPM-21488, a ramus with the symphysis, roots of both I, and P,, and P, to M;. This ramus does not have an internal mandibular fenestra nor is it inflated. developed than in the premolars and the entoconid projects mesially, partly closing the valley between it and the metastylid. The protocristids and hypo- cristids are complete, forming lophs. Ms has a small, low hypoconulid. All cingula are absent or very weak in the lower cheek teeth. P? has a very small mesostyle and is much longer than broad, whereas P? possesses a well-developed mesostyle and is slightly longer than broad. The remainder of the upper cheek teeth are broader than long. P?-M? are molari- form, selenolophodont with well-developed preprotocristae and prehypo- cristae which often come in contact with the ectoloph. The parastyles and metastyles are well developed and there is a slight fold on the buccal side of the ectoloph opposite the paracone. The mesial and lingual cingula are well developed. There is a round to subround internal mandibular fenestra on the lingual side of the ramus below Mz in some of the specimens; this occurs also in Megalohyrax and Pachyhyrax. Each of these specimens has a greatly inflated ramus with a large swollen chamber in the horizontal ramus, beneath the den- tal series, which continues at least into the base of the inflated ascending ramus. There are three specimens known that possess the internal mandibular fenestra and four specimens that possess no fenestra and do not have inflated rami. Other than this difference, there is no morphological difference that can be used to separate the fenestrated and nonfenestrated specimens into differ- ent species. New collections made by Yale University in the Upper Fossil Wood Zone have confirmed that there are also specimens of two species of A NEW OLIGOCENE HYRAX 7 TABLE 1. Statistical data on teeth of Thyrohyrax domorictus. Abbreviations: L = length, M = molar (M! = first upper molar, M. = second lower molar), n = sample size, OR = observed range, P = premolar (P; = third lower premolar, P+ = fourth upper premolar), s = standard deviation, V = coefficient of variation, W = width, xX = mean. Measurements for length and width are in mm. n OR x 5 V PL 1 4.7 4.7 = = W 1 wl 3.1 = as PoL 5 5.6-6.0 5.82 1658 2.85 W 6 3.5-4.2 3.82 2720 712 Pa 5 elegal 6.58 4153 6.31 W 5 4.2-4.8 4.46 2179 4.89 P,L 5 6.5-7.3 6.82 3428 5.02 W 5 4.8-5.5 5.20 3240 6.23 M,L 7 7.4-8.4 WE 3536 4.57 W 7 4.6-5.8 5.14 5099 9.92 M.L 12 7.9-9.3 8.65 4253 4.92 Ww 12 5.0-6.5 5.66 5018 8.87 M,L 9 10.7-12.8 11.85 18515 7.19 W 9 5356.7 6.15 4637 7.54 P2L 1 5.8 5.8 a ass Ww 1 5.3 5.3 = = P3L 1 6.4 6.4 a a W 1 6.1 6.1 _ = PAL 2 6.4-7.0 6.70 4243 6.33 W 2 7.0 7.0 = a M!L 3 FAT6 7.33 2550 3.48 WwW 3 WIERD 7.93 4183 52a M2L 3 7.9-8.8 8.46 4950 5.85 W 3 8.6-9.2 8.93 3082 3.45 MEL 2 9.9-10.0 9.95 a = W 2. 9.8-10.0 9.90 1414 1.42 Bunohyrax and one species of Megalohyrax that possess an internal mandib- ular fenestra and specimens that do not. It therefore seems more logical to consider the possession of an internal mandibular fenestra as a sexual char- acter than to construct a phylogeny of twinned species, with and without fenestrae. From the known specimens of Thyrohyrax domorictus it cannot be deter- mined which sex possesses the fenestra since none of those specimens with the fenestra also possesses the sexually diagnostic incisors. In Bunohyrax faju- mensis and Megalohyrax eocaenus however, those specimens without the 8 POSTILLA 163 FIG. 3. Occlusial and basal view of Thyrohyrax domorictus, YPM-23939, showing the greatly inflated rami and in the basal view, the expanded chamber within the ramus. fenestra have large procumbent tusk-like incisors typical of extant males whereas specimens that do possess the fenestra have smaller more rounded incisors typical of extant females. These latter specimens are also 10 to 15 percent smaller than the specimens without the fenestra. It is therefore suggested that the females of Thyrohyrax domorictus possess an internal mandibular fenestra and inflated ramus; the males do not. The functional significance of this fenestra and chamber is unknown. A fenestra and chamber enclosed within the ramus are known in Bunohyrax, Megalohyrax and Pachyhyrax as well as in Thyrohyrax. A fossa, roughly similar in shape to the internal chamber, is found on the lingual side of the ramus of Geniohyus, Meroehyrax and Pliohyrax. No such structure is known in Saghatherium, Titanohyrax, Gigantohyrax or the living hyraxes. No anal- ogous feature is to be observed among other mammals nor has any other clue been found in the living hyraxes that has shed any light on this unique and bizarre feature. Andrews (1907) suggested that the cavity formed by this hollow was used as a resonating chamber for sound production. It seems odd, however, if this were the case, that the feature would have been located under the tongue. There is a guttural pouch in the Eustachian channel in living hyraxes that is A NEW OLIGOCENE HYRAX 9 used in sound production but it is not restricted to females. One fossil spec- imen, a juvenile Pachyhyrax crassidentatus shows a very deep round fossa which, however, does not connect with the internal cavity as it does in the adults. This character may suggest that the fenestra is formed by the invagi- nation of the structure it housed causing resorption of the bone of the ramus with its enlargement during adolescent growth. If this is the case, then the fenestra cannot act as a place of entrance for blood vessels or nerves. There are no holes or perforations of the chamber other than the one internal mandibular fenestra, and this chamber is not connected in any way to the dental battery. There is, however, a beveled edge at the dorsolateral side of the fenestra in all species that show fenestrae and there is a similar angulation at the same position of the sulcus of Pliohyrax. This beveled edge can only be interpreted as a passageway for something that entered the fenestra. If the fenestra was indeed formed by resorption of the wall of the ramus, then only something of a glandular nature that enlarged as the female reached sexual maturity could have passed through the fenestra into the chamber. No such organ is otherwise known. The tibia measures 97.4 mm in length and except for its larger size is virtually identical to the tibia of Procavia and the other extant species. The fibula was not fused to the tibia as in extant forms. The bones of the pes and manus are, except for their greater size, nearly identical to those of Procavia. Thyrohyrax domorictus, therefore, possessed feet and limbs remarkably sim- ilar to the extant hyraxes, even though the two forms are separated by at least 30 million years. It seems that the general character of the hyrax postcranium was established in early Tertiary times. PHYLETIC POSITION OF Thyrohyrax domorictus Thyrohyrax domorictus is assigned to the subfamily Saghatheriinae on the basis of the morphology of the teeth and possession of the internal mandib- ular fenestra. Internal mandibular fenestrae are presently known only in the hyracoids of the Oligocene and Miocene of North and East Africa that are included in the Saghatheriinae. These forms all possess the full eutherian dentition, have premolars that are submolariform or distinctly molariform, have hypoconulids on Mz and bunodont to selenolophodont molars. Thyrohyrax differs from Saghatherium by the possession of an internal mandibular fenestra and inflated ramus in some forms, its less inflated cusps, its more molariform premolars with more centrally placed paraconids, the metastylids on the lower cheek teeth, mesostyles on the upper premolars, the lack of spurs on the lingual side of the ectoloph of the molars and the relatively broader and shorter upper cheek teeth. The sole species of the genus Thyrohyrax differs from those of Megalohyrax and Pachyhyrax by its 10 POSTILLA 163 much smaller size, its shorter snout, its inflated ramus, its more selenodont teeth, its more molariform premolars, its metastylids and by its differently shaped upper molars. Dentally Thyrohyrax is very similar to Meroehyrax and may be close to, although not on, the direct line of ancestry of Meroehyrax. The two-rooted lower canine of Thyrohyrax, a feature unique to the Saghatheriinae, and the overall shape and selenolophodont nature of the lower teeth are features held in common with Meroehyrax. The fact that Thyrohyrax possesses an internal mandibular fenestra, places it in the Saghatheriinae while Meroehyrax with a fossa is placed in the Pliohyracinae. Thyrohyrax may, however, be more closely related to the form that gave rise to the Pliohyracinae than any other presently known member of the Saghatheriinae. GEOLOGICAL SETTING The Jebel el Qatrani formation, which is 110 to 270 m thick, north of the Birkel el Qarun, consists of terrestrial deposits of quartz sandstone, siltstones, claystones and pebble conglomerates. “The basal 100 meters [Lower Fossil Wood Zone] is characterized by a complex of large-scale trough cross- stratified channel-lag and point—-bar deposits and the lack of flood plain de- posits. These characteristics are indicative of a loosely sinuous, low gradient, medium velocity stream. Point-bar, levee, crevasse-splay, flood basin, and occasional transitional channel fill deposits are characteristic of the upper portion of the unit [Upper Fossil Wood Zone].’”’ (Bowen and Vondra, in press. ) Thyrohyrax is found at Yale Quarry G which consists of 0.3 m of mottled reddish orange and light olive gray saline claystone, which represents the upper portion of a point-bar deposit. It is also found at Quarries M and I which are located in 16 m of fine— to coarse-grained grayish yellow sandstone in the lower portion of a point-bar deposit. Quarries G and I are deposited 126.2 m and 75.8 m respectively below the Widan el Faras Basalt which has been variously dated at 24.7 + 0.4 million years and 27.3 + 2 million years (Simons, 1968). The contact between the basalt and the Jebel el Qatrani formation is disconformable. ACKNOWLEDGMENTS I am deeply indebted to Dr. Elwyn L. Simons of Peabody Museum of Natural History, Yale University, and Mr. Darwish el Faar of the Cairo Geological A NEW OLIGOCENE HYRAX 11 Museum, Cairo, Egypt, for allowing me access to material in their collections. Helpful discussions were held with Dr. Simons and Dr. David Pilbeam of Yale University, and Dr. Carl Vondra and Mr. Bruce Bowen of Iowa State University. I am grateful for their many helpful suggestions. The photographs are by A.H. Coleman of Harvard University. The manuscript was typed by M. H. Meyer. Grants that enabled the collection of the fossils reported in this paper were Smithsonian Foreign Currency Program Grants nos. 5, 23 and 1841 and National Science Foundation Grants G-18102, GP-433 and GP-3547 made to Dr. E. L. Simons. LITERATURE CITED Andrew, C. W. 1906. A descriptive catalogue of the Tertiary Vertebrata of the Fayum, Egypt. Brit. Mus. [Nat. Hist.], London, xxxvili + 324 p. 1907. Note on some vertebrate remains collected in the Fayum, Egypt. Geol. Mag. 4: 97-100. Bowen, B. E. and C. F. Vondra. 1974. Paleoenvironmental interpretations of the Oligocene Jebel el Qatrani formation, Fayum Depression, Egypt. In press. Huxley, T. H. 1869. An introduction to the classification of animals. Churchill and Sons. London, viii + 147 p. Osborn, H. F. 1899. On Pliohyrax kruppi Osborn, a fossil hyracoid from Samos, Lower Pliocene, in the Stuttgart collection. Proc. 4th Internat. Congr. Zool. [1898], Cambridge, p. 172-173. Simons, E. L. i968 African Oligocene mammals: Introduction, history of study, and faunal succession. In Elwyn L. Simons and Albert E. Wood, Early Cenozoic mammalian faunas[,] Fayum Province, Egypt. Bull. Peabody Mus. Nat. Hist., Yale Univ. 28. 105 p. REVIEW STYLE FORM TITLE ABSTRACT OMENCLATURE ILLUSTRATIONS FOOTNOTES TABLES REFERENCES JTHOR’S COPIES PROOF COPYRIGHT INFORMATION FOR AUTHORS The Publications Committee of the Peabody Museum of Natural History reviews and approves manuscripts for publication. Papers will be published in approximately the order in which they are accepted; delays may result if manuscript or illustrations are not in proper form. To facilitate review, the original and one carbon or xerox copy of the typescript and figures should be submitted. The author should keep a copy. Authors of biological papers should follow the Style Manual for Biological Journals, Second Edition (Amer. Inst. Biol. Sci.). 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