{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,17]],"date-time":"2026-03-17T08:29:33Z","timestamp":1773736173414,"version":"3.50.1"},"reference-count":56,"publisher":"American Association for the Advancement of Science (AAAS)","issue":"5311","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Science"],"published-print":{"date-parts":[[1997,4,18]]},"abstract":"<jats:p>Fossils from a large-bodied hominoid from early Miocene sediments of Uganda, along with material recovered in the 1960s, show features of the shoulder and vertebral column that are significantly similar to those of living apes and humans. The large-bodied hominoid from Uganda dates to at least 20.6 million years ago and thus represents the oldest known hominoid sharing these derived characters with living apes and humans.<\/jats:p>","DOI":"10.1126\/science.276.5311.401","type":"journal-article","created":{"date-parts":[[2002,7,27]],"date-time":"2002-07-27T09:37:56Z","timestamp":1027762676000},"page":"401-404","source":"Crossref","is-referenced-by-count":156,"title":["A Hominoid Genus from the Early Miocene of Uganda"],"prefix":"10.1126","volume":"276","author":[{"given":"Daniel L.","family":"Gebo","sequence":"first","affiliation":[{"name":"D. L. Gebo, Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, USA."},{"name":"L. MacLatchy, Department of Anatomical Sciences, State University of New York, Stony Brook, NY 11794, USA."},{"name":"R. Kityo, Department of Zoology, Makerere University, Kampala, Uganda."},{"name":"A. Deino, Berkeley Geochronology Center, Berkeley, CA 94709, USA."},{"name":"J. Kingston, Department of Geology and Geophysics, Yale University, New Haven, CT 06520, USA."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Laura","family":"MacLatchy","sequence":"additional","affiliation":[{"name":"D. L. Gebo, Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, USA."},{"name":"L. MacLatchy, Department of Anatomical Sciences, State University of New York, Stony Brook, NY 11794, USA."},{"name":"R. Kityo, Department of Zoology, Makerere University, Kampala, Uganda."},{"name":"A. Deino, Berkeley Geochronology Center, Berkeley, CA 94709, USA."},{"name":"J. Kingston, Department of Geology and Geophysics, Yale University, New Haven, CT 06520, USA."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Robert","family":"Kityo","sequence":"additional","affiliation":[{"name":"D. L. Gebo, Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, USA."},{"name":"L. MacLatchy, Department of Anatomical Sciences, State University of New York, Stony Brook, NY 11794, USA."},{"name":"R. Kityo, Department of Zoology, Makerere University, Kampala, Uganda."},{"name":"A. Deino, Berkeley Geochronology Center, Berkeley, CA 94709, USA."},{"name":"J. Kingston, Department of Geology and Geophysics, Yale University, New Haven, CT 06520, USA."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alan","family":"Deino","sequence":"additional","affiliation":[{"name":"D. L. Gebo, Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, USA."},{"name":"L. MacLatchy, Department of Anatomical Sciences, State University of New York, Stony Brook, NY 11794, USA."},{"name":"R. Kityo, Department of Zoology, Makerere University, Kampala, Uganda."},{"name":"A. Deino, Berkeley Geochronology Center, Berkeley, CA 94709, USA."},{"name":"J. Kingston, Department of Geology and Geophysics, Yale University, New Haven, CT 06520, USA."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"John","family":"Kingston","sequence":"additional","affiliation":[{"name":"D. L. Gebo, Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, USA."},{"name":"L. MacLatchy, Department of Anatomical Sciences, State University of New York, Stony Brook, NY 11794, USA."},{"name":"R. Kityo, Department of Zoology, Makerere University, Kampala, Uganda."},{"name":"A. Deino, Berkeley Geochronology Center, Berkeley, CA 94709, USA."},{"name":"J. Kingston, Department of Geology and Geophysics, Yale University, New Haven, CT 06520, USA."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"David","family":"Pilbeam","sequence":"additional","affiliation":[{"name":"D. L. Gebo, Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, USA."},{"name":"L. 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J., Nature 360, 641 (1992).","journal-title":"Nature"},{"key":"e_1_3_1_11_2","first-page":"218","volume":"54","author":"Filler A. G.","year":"1981","unstructured":"Filler A. G., Am. J. Phys. Anthropol. 54, 218 (1981).","journal-title":"Am. J. Phys. Anthropol."},{"key":"e_1_3_1_12_2","doi-asserted-by":"crossref","unstructured":"C. V. Ward ibid. 92 291 (1993).","DOI":"10.1002\/ajpa.1330920306"},{"key":"e_1_3_1_13_2","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1006\/jhev.1994.1012","volume":"26","author":"Sanders W. J.","year":"1994","unstructured":"Sanders W. J., Bodenbender B. E., J. Hum. Evol. 26, 203 (1994).","journal-title":"J. Hum. Evol."},{"key":"e_1_3_1_14_2","doi-asserted-by":"crossref","first-page":"669","DOI":"10.2475\/ajs.267.6.669","volume":"267","author":"Bishop W. W.","year":"1969","unstructured":"Bishop W. W., Miller J. A., Fitch F. J., Am. J. Sci. 267, 669 (1969).","journal-title":"Am. J. Sci."},{"key":"e_1_3_1_15_2","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/S0047-2484(81)80026-7","volume":"10","author":"Pickford M.","year":"1981","unstructured":"Pickford M., J. Hum. Evol. 10, 73 (1981).","journal-title":"J. Hum. Evol."},{"key":"e_1_3_1_16_2","unstructured":"M. Pickford B. Senut D. Hadoto J. Musisi C. Kariira C. R. Acad. Sci. Paris Ser. II 9 681 (1986)."},{"key":"e_1_3_1_17_2","unstructured":"One sample from each of the capping basaltic lava flows in the vicinity of Moroto localities I and II was dated by the 40 Ar\/ 39 Ar incremental heating technique. The sample from Moroto I is relatively unaltered (MO-747) whereas the Moroto II flow (MO-744) shows partial replacement of plagioclase and pyroxene by fine-grained alteration products. Two separately prepared 40-60 mesh fractions of intergrown plagioclase-pyroxene microphenocrysts from Moroto I yielded 40 Ar\/ 39 Ar incremental release spectra characterized by a single age plateau (Fig. 1). These two samples yielded plateau ages of 20.68 \u00b1 0.08 and 20.65 \u00b1 0.07 Ma. An inverse isochron treatment indicated ages of 20.63 \u00b1 0.08 and 20.60 \u00b1 0.06. The more altered sample from Moroto II (plagioclase concentrate) yielded a discordant spectrum characterized by an initial increase in apparent age followed by a rapid decrease to about 20 Ma in the remaining 60% of the cumulative 39 Ar release (Fig. 1). This initial stair-step pattern is interpreted as reflecting recoil loss from fine-grained alteration phases whereas the plateau that follows may represent Ar release from relatively unaltered plagioclase cores. Although this sample is inconclusive by itself agreement of the plateau age of this plagioclase with that of the matrix material from MO-747 is supporting evidence for the 20.6 Ma age obtained at the Moroto I locality."},{"key":"e_1_3_1_18_2","unstructured":"G. T. Cebula et al. Terra Cognita 6 139 (1986)."},{"key":"e_1_3_1_19_2","unstructured":"S. D. Samson and E. C. Alexander Chem. Geol. Isot. Geosci. 6 27 (1987)."},{"key":"e_1_3_1_20_2","unstructured":"We used femoral head radius midshaft cortical area and bicondylar width to estimate femur length using separate ape and monkey regressions. The range was from 255 to 291 mm with a mean length of 270 mm [see also (20)]."},{"key":"e_1_3_1_21_2","doi-asserted-by":"crossref","unstructured":"C. B. Ruff Folia Primatol. 48 9 (1987).","DOI":"10.1159\/000156283"},{"key":"e_1_3_1_22_2","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1016\/S0047-2484(05)80006-5","volume":"22","author":"Rose M. D.","year":"1992","unstructured":"Rose M. D., Leakey M. G., Leakey R. E. F., Walker A. C., J. Hum. Evol. 22, 171 (1992).","journal-title":"J. Hum. Evol."},{"key":"e_1_3_1_23_2","doi-asserted-by":"crossref","unstructured":"L. M. MacLatchy and W. H. Bossert ibid. 31 425 (1996).","DOI":"10.1006\/jhev.1996.0070"},{"key":"e_1_3_1_24_2","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1002\/ajpa.1330910103","volume":"91","author":"Ruff C. B.","year":"1993","unstructured":"Ruff C. B., Trinkaus E., Larsen C. S., Am. J. Phys. Anthropol. 91, 21 (1993).","journal-title":"Am. J. Phys. Anthropol."},{"key":"e_1_3_1_25_2","doi-asserted-by":"crossref","unstructured":"C. B. Ruff and A. C. Runestad Annu. Rev. Anthropol. 21 407 (1992).","DOI":"10.1146\/annurev.an.21.100192.002203"},{"key":"e_1_3_1_26_2","unstructured":"J. A. Runestad thesis Johns Hopkins University Baltimore MD (1994)."},{"key":"e_1_3_1_27_2","unstructured":"We estimated body size using hominoid regression equations for femoral head surface area midshaft breadth and cortical area and femoral epicondylar breadth using regressions from combined ape and monkey samples as well as published regressions (20 27 28). Size estimates range from 36 to 54 kg."},{"key":"e_1_3_1_28_2","doi-asserted-by":"crossref","first-page":"687","DOI":"10.1016\/0047-2484(88)90025-5","volume":"17","author":"Ruff C. B.","year":"1988","unstructured":"Ruff C. B., J. Hum. Evol. 17, 687 (1988).","journal-title":"J. Hum. Evol."},{"key":"e_1_3_1_29_2","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1002\/ajpa.1330950405","volume":"95","author":"Aiello L. C.","year":"1994","unstructured":"Aiello L. C., Wood B. A., Am. J. Phys. Anthropol. 95, 409 (1994).","journal-title":"Am. J. Phys. Anthropol."},{"key":"e_1_3_1_30_2","doi-asserted-by":"crossref","unstructured":"D. Roberts in Primate Locomotion F. A. Jenkins Ed. (Academic Press New York 1974) pp. 171\u2013200.","DOI":"10.1016\/B978-0-12-384050-9.50011-8"},{"key":"e_1_3_1_31_2","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1002\/ajpa.1330470308","volume":"47","author":"Gingerich P. D.","year":"1977","unstructured":"Gingerich P. D., Am. J. Phys. 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(Plenum New York 1983) pp. 325\u2013351.","DOI":"10.1007\/978-1-4684-8854-8_12"},{"key":"e_1_3_1_36_2","unstructured":"M. D. Rose in ibid. pp. 405\u2013417."},{"key":"e_1_3_1_37_2","unstructured":"___ in Postcranial Adaptation in Nonhuman Primates D. L. Gebo Ed. (Northern Illinois Univ. Press DeKalb IL 1993) pp. 252\u2013272."},{"key":"e_1_3_1_38_2","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1002\/ajpa.1330900106","volume":"90","author":"Ward C. V.","year":"1993","unstructured":"Ward C. V., Walker A., Teaford M. F., Odhiambo I., Am. J. Phys. Anthropol. 90, 77 (1993).","journal-title":"Am. J. Phys. Anthropol."},{"key":"e_1_3_1_39_2","unstructured":"Order Primates Linnaeus 1758; suborder Anthropoidea Mivart 1864; infraorder Catarrhini Geoffroy 1812; superfamily Hominoidea Gray 1825; family not assigned. Genus Morotopithecus gen. nov. Generic diagnosis: Large hominoid male weight around 40 to 50 kg most comparable in facial and dental morphology to Proconsul and Afropithecus. Differs from later Miocene and extant apes in having a longer midface and has less alveolar prognathism than extant large apes. Differs from Afropithecus later Miocene hominoids and extant apes in the greater degree of cingular development on cheek teeth especially molars. Differs from Afropithecus and Proconsul in having a narrower interorbital region and larger premolars relative to M1; differs from Proconsul in a smaller M2 and M3 relative to M1. Differs from Afropithecus in possessing a shorter premaxilla a higher face a broader nasal aperture (5) a P 3 that is much broader buccally and in which the paracone is situated closer to the protocone buccal wrinkling on the side of the molars and a larger M 3 . Judging from worn occusal surfaces the enamel was intermediate thin as in P. major (51). Further computed tomography scans on the skull of Afropithecus reveal a thick palate with a small incisive canal (52) an anatomical condition that is distinctly different from the large canal of the Moroto palate (53). The glenoid is rounder in shape and more like that of extant apes and atelines than can be inferred for Miocene hominoids other than Oreopithecus and possibly Dryopithecus (42). Lumbar morphology differs from that of Proconsul and resembles that of Oreopithecus possibly Dryopithecus and extant large apes and siamangs in transverse process position. Proximal femoral morphology resembles that of cercopithecines and primitive hominoids such as Proconsul differing from that of extant apes. Distal femoral anatomy resembles that of Proconsul Kenyapithecus and extant apes in mediolateral breadth but differs from hylobatids Proconsul and Kenyapithecus in the buttressing of the intercondylar notch. In 1962 L. S. B. Leakey referred the Moroto palate to Pseudogorilla (54) but did so without any species diagnosis. Pseudogorilla was created by Elliot in 1912 for ape specimens from the \u201cUpper Congo\u201d (55) now referred to Gorilla. Regardless of the validity of Pseudogorilla the Moroto fossil is clearly different from any extant ape. Type species: Morotopithecus bishopi sp . nov. Etymology: Moroto after Moroto township in Karamoja District in Uganda and pithekos from the Greek for ape; and after the late W. W. Bishop. Type specimen: UMP 62-11 (UMP Ugandan Museum of Paleontology) a palatofacial specimen with all teeth. Type locality: Moroto II. Distribution: Early Miocene Karamoja District NE Uganda. Hypodigm: Type and UMP 62-10 and UMP 66-01 which are probably associated mandibular fragments; UMP 62-12 left upper canine; UMP 67-28 middle lumbar vertebra; UMP 68-05 middle lumbar vertebral body; UMP 68-06 a last thoracic vertebra; UMP 68-07 the lamina and base of a spine of a lumbar vertebra; MUZM 80 right and left femoral pieces (all from the Moroto II locality); and MUZM 60 scapular fragment with glenoid (Moroto I locality). Specific diagnosis: As for genus."},{"key":"e_1_3_1_40_2","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/0047-2484(91)90062-Z","volume":"21","author":"Ward C. V.","year":"1991","unstructured":"Ward C. V., Walker A., Teaford M. F., J. Hum. Evol. 21, 215 (1991).","journal-title":"J. Hum. Evol."},{"key":"e_1_3_1_41_2","doi-asserted-by":"crossref","unstructured":"K. C. Beard M. F. Teaford A. 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B"},{"key":"e_1_3_1_53_2","unstructured":"S. C. Ward personal communication."},{"key":"e_1_3_1_54_2","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1002\/ajpa.1330610204","volume":"61","author":"___","year":"1983","unstructured":"___, Kimbel W. H., Am. J. Phys. Anthropol. 61, 157 (1983).","journal-title":"Am. J. Phys. Anthropol."},{"key":"e_1_3_1_55_2","unstructured":"L. S. B. Leakey Ann. Mag. Nat. Hist. 4 689 (1962)."},{"key":"e_1_3_1_56_2","unstructured":"D. G. Elliot Rev. Primates 3 225 (1912) (from Monographs of the American Museum of Natural History New York 1912)."},{"key":"e_1_3_1_57_2","unstructured":"We thank the many helpful individuals in Uganda including the Office of the President the National Research Council and the staff at the Zoology Museum at Makerere University as well as C. Chapman for the many efforts made on our behalf. We thank D. Aleper for assistance in the field; B. Masek for assistance in the laboratory; M. Mehrer and J. Flynn; and the American School of Prehistoric Research and the L. S. B. Leakey Foundation for contributing funding to this project."}],"container-title":["Science"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.science.org\/doi\/pdf\/10.1126\/science.276.5311.401","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,1,13]],"date-time":"2024-01-13T04:45:32Z","timestamp":1705121132000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.science.org\/doi\/10.1126\/science.276.5311.401"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[1997,4,18]]},"references-count":56,"journal-issue":{"issue":"5311","published-print":{"date-parts":[[1997,4,18]]}},"alternative-id":["10.1126\/science.276.5311.401"],"URL":"https:\/\/doi.org\/10.1126\/science.276.5311.401","relation":{},"ISSN":["0036-8075","1095-9203"],"issn-type":[{"value":"0036-8075","type":"print"},{"value":"1095-9203","type":"electronic"}],"subject":[],"published":{"date-parts":[[1997,4,18]]}}}