Article: The auditory region of Early Paleocene Pucadelphydae (Mammalia, Metatheria) from Tiupampa, Bolivia, with phylogenetic implications
Publication: Palaeontology
Volume:
50
Part:
5
Publication Date:
September
2007
Page(s):
1123
–
1154
Author(s):
Sandrine Ladevèze and Christian de Muizon
References
- AMRINE-MADSEN, H., SCALLY, M., WESTERMAN, M., STANHOPE, M. J., KRAJEWSKI, C. and SPRINGER, M. S. 2003. Nuclear gene sequences provide evidence for the monophyly of australidelphian marsupials. Molecular Phylogenetics and Evolution, 28, 186–196.
- ARCHER, M. 1976a. The basicranial region of marsupicarnivores (Marsupialia), interrelationships of carnivorous mammals, and affinities of the insectivorous marsupial peramelids. Zoological Journal of the Linnean Society, 59, 217–322.
- ARCHER, M. 1976b. The dasyurid dentition and its relationship to that of didelphids, thylacinids, borhyaenids (Marsupicarnivora) and peramelids (Peramelina: Marsupialia). Australian Journal of Zoology, Supplementary Series, 39, 1–34.
- ARCHER, M. 1982. A review of Miocene thylacinids (Thylacinidae, Marsupialia), the phylogenetic position of the Thylacinidae and the problem of priorisms in character analysis. 445–476. In ARCHER, M. (ed.). Carnivorous marsupials. Royal Zoological Society of New South Wales, Sydney, 804 pp.
- ARCHER, M. 1984. The Australian marsupial radiation. 633–808. In ARCHER, M. and CLAYTON, G. (eds). Vertebrate zoogeography and evolution in Australasia. Hesperian Press, Perth, 1203 pp.
- ASHER, R. J. 1999. A morphological basis for assessing the phylogeny of the ’Tenrecoidea’ (Mammalia, Lipotyphla). Cladistics, 15, 231–252.
- ASHER, R. J., HOROVITZ, I. and SÁNCHEZ-VILLAGRA, M. R. 2004. First combined cladistic analysis of marsupial mammal interrelationships. Molecular Phylogenetics and Evolution, 33, 240–250.
- BAKER, R. H. and DESALLE, R. 1997. Multiple sources of character information and the phylogeny of Hawaiian drosophilids. Systematic Biology, 46, 674–698.
- BREMER, K. 1988. The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42, 795–803.
- CARDILLO, M., BININDA-EMONDS, O. R. P., BOAKES, E. and PURVIS, A. 2004. A species-level phylogenetic supertree of marsupials. Journal of Zoology (London), 264, 11–31.
- CIFELLI, R. L. 1993. Early Cretaceous mammal from North America and the evolution of marsupial dental characters. Proceedings of the National Academy of Sciences, 90, 9413–9416.
- CLEMENS, W. A. 1966. Fossil mammals of the Type Lance Formation, Wyoming. Part II. Marsupialia. University of California, Publications in Geological Sciences, 62, 1–122.
- FARRIS, J. S., KÄLLERSJÖ, M., KLUGE, A. G. and BULT, C. 1994. Testing significance of incongruence. Cladistics, 10, 315–319.
- FLANNERY, T. E. and ARCHER, M. 1987. Hypsiprymnodon bartholomaii (Marsupialia: Potoroidae), a new species from the Miocene Dwornamor Local Fauna and a reassessment of the phylogenetic position of H. moschatus. 749–758. In ARCHER, M. (ed.). Possums and opossums: studies in evolution. Surrey Beatty and Sons and the Royal Zoological Society of New South Wales, Sydney, 788 pp.
- FLORES, D. A. and ABDALA, F. 2001. Diferencias morfologícas de cráneo y dentición en Didelphis albiventris y D. marsupialis (Didelphimorphia, Didelphidae) de Argentina y Bolivia. Comunica Museu de Ciências e Tecnologia PUCRS, Série Zoologia, Porto Alegre, 14, 99–207.
- FLYNN, J. J. and SWISHER, C. C. III. 1995. Cenozoic South American land mammal ages: correlation to global geochronologies. 317–334. In BERGGREN, W. A., KENT, D. V., AUBRY, M.-P. and HARDENBOL, J. (eds). Geochronology, time scales and global stratigraphic correlation. SEPM Special Publication, 54, 386 pp.
- GAUDIN, T. J., WIBLE, J. R., HOPSON, J. A. and TURNBULL, W. D. 1996. Reexamination of the morphological evidence for the cohort Epitheria (Mammalia, Eutheria). Journal of Mammalian Evolution, 3, 31–79.
- GODTHELP, H., WROE, S. and ARCHER, M. 1999. A new marsupial from the Early Eocene Tingamarra Local Fauna of Murgon, southeastern Queensland: a prototypical Australian marsupial. Journal of Mammalian Evolution, 6, 289–313.
- GOLOBOFF, P. A. 1993. NONA (a bastard son of Pee-Wee). Version. 2.0. Program and documentation. Available from http://www.cladistics.com, published by the author, Tucumán, Argentina.
- HERSHKOVITZ, P. 1982. The staggered marsupial lower third incisor (i3). Geobios, Mémoire Spécial, 6, 191–200.
- HOFFSTETTER, R. and PETTER, G. 1983. Paraborhyaena boliviana et Andinogale sallensis, deux marsupiaux (Borhyaenidae) nouveaux du Déséadien (Oligocène inférieur) de Salla (Bolivie). Comptes Rendus de l'Académie des Sciences, Paris, Série II, 296, 205–208.
- HOPSON, J. A. and ROUGIER, G. W. 1993. Braincase structure in the oldest known skull of therian mammal: implications for mammalian systematics and cranial evolution. American Journal of Science, 293, 268–299.
- HOROVITZ, I. and SÁNCHEZ-VILLAGRA, M. R. 2003. A morphological analysis of marsupial mammal higher-level phylogenetic relationships. Cladistics, 19, 181–212.
- JI, Q., LUO, Z.-X., YUAN, C.-X., WIBLE, J. R., ZHANG, J.-P. and GEORGI, J. A. 2002. The earliest known eutherian mammal. Nature, 416, 816–822.
- KIELAN-JAWOROWSKA, Z. and NESSOV, L. A. 1990. On the metatherian nature of the Deltatheroidea, a sister group of the Marsupalia. Lethaia, 23, 1–10.
- KIELAN-JAWOROWSKA, Z., CIFELLI, R. L. and LUO, Z.-X. 2004. Mammals from the age of dinosaurs: origins, evolution, and structure. Columbia University Press, New York, NY, i–xv + 630 pp.
- KIELAN-JAWOROWSKA, Z., PRESLEY, R. and POPLIN, C. 1986. The cranial vascular system in taeniolabidoid multituberculate mammals. Philosophical Transactions of the Royal Society of London, B, 313, 525–602.
- KIRSCH, J. A. W., LAPOINTE, F.-J. and SPRINGER, M. S. 1997. DNA-hybridisation studies of marsupials and their implications for metatherian classification. Australian Journal of Zoology, 45, 211–280.
- KIRSCH, J. A. W., REIG, O. A. and SPRINGER, M. S. 1991. DNA hybridization evidence for the Australian affinity of the American marsupial Dromiciops australis. Proceedings of the National Academy of Sciences, USA, 88, 10,465–10,469.
- KUHN, H.-J. 1971. Die Entwicklung und Morphologie des Schdels von Tachyglossus aculeatus. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 528, 1–192.
- KUHN, H.-J. and ZELLER, U. 1987. The cavum epiptericum in monotremes and therian mammals. 51–70. In KUHN, H.-J. and ZELLER, U. (eds). Mammalia depicta. Morphogenesis of the mammalian skull. Verlag Paul Varey, Hamburg, 140 pp.
- LADEVÈZE, S. 2004. Metatherian petrosals from the Late Paleocene of Itaboraí (Brazil), and their phylogenetic implications. Journal of Vertebrate Paleontology, 24, 202–213.
- LADEVÈZE, S. 2007. Petrosal bones of metatherian mammals from the Late Paleocene of Itaboraí (Brazil), and a cladistic analysis of petrosal features in metatherians. Zoological Journal of the Linnean Society, 150, 85–115.
- LUCKETT, W. P. 1994. Suprafamilial relationships within Marsupialia: resolution and discordance from multidisciplinary data. Journal of Mammalian Evolution, 2, 255–283.
- LUO, Z.-X. 1994. Sister taxon relationships of mammals and the transformations of the diagnostic mammalian characters. 98–128. In FRASER, N. C. and SUES, H.-D. (eds). In the shadow of dinosaurs – early Mesozoic tetrapods. Cambridge University Press, Cambridge, 447 pp.
- LUO, Z.-X., JI, Q., WIBLE, J. R. and YUAN, C.-X. 2003. An early Cretaceous tribosphenic mammal and metatherian evolution. Science, 302, 1934–1940.
- LUO, Z.-X., KIELAN-JAWOROWSKA, Z. and CIFELLI, R. L. 2002. In quest for a phylogeny of Mesozoic mammals. Acta Palaeontologica Polonica, 47, 1–78.
- MacPHEE, R. D. E. 1981. Auditory regions of primates and eutherian insectivores: morphology, ontogeny, and character analysis. Contributions to Primatology, 18, 1–282.
- MAIER, W. 1987. The ontogenetic development of the orbitotemporal region of the skull of Monodelphis domestica (Didelphidae, Marsupialia), and the problem of the mammalian alisphenoid. 71–90. In KUHN, H.-J. and ZELLER, U. (eds). Mammalia depicta. Morphogenesis of the mammalian skull. Paul Parey Verlag, Hamburg, 140 pp.
- MARSHALL, L. G. 1981. Review of the Hathlyacyninae, an extinct subfamily of South American dog-like marsupials. Fieldiana: Geology, 7, 1–120.
- MARSHALL, L. G. 1982. Systematics of the South American marsupial family Microbiotheriidae. Fieldiana: Geology, New Series, 10, 1–75.
- MARSHALL, L. G. 1987. Systematics of Itaboraían (Middle Paleocene) age opossum-like marsupials from the limestone quarry at São José de Itaboraí, Brazil. 91–160. In ARCHER, M. (ed.). Possums and opossums: studies in evolution. Surrey Beatty and Sons and the Royal Zoological Society of New South Wales, Sydney, 799 pp.
- MARSHALL, L. G. and KIELAN-JAWOROWSKA, Z. 1992. Relationships of the dog-like marsupials, deltatheroidans, and early tribosphenic mammals. Lethaia, 25, 361–374.
- MARSHALL, L. G. and MUIZON, C. de 1988. The dawn of the Age of Mammals in South America. National Geographic Research, 4, 23–55.
- MARSHALL, L. G. and MUIZON, C. de 1995. Part II. The skull. 21–90. In MUIZON, C. de (ed.). Pucadelphys andinus (Marsupialia, Mammalia) from the Early Paleocene of Bolivia. Mémoires du Muséum National d'Histoire Naturelle, Paris, 164 pp.
- MARSHALL, L. G., CASE, J. A. and WOODBURNE, M. O. 1990. Phylogenetic relationships of the families of marsupials. 433–505. In GENOWAYS, H. H. (ed.). Current mammalogy. Plenum Press, New York, NY, 567 pp.
- MARSHALL, L. G., SEMPERE, T. and BUTLER, R. F. 1997. Chronostratigraphy of the mammal-bearing Paleocene of South America. Journal of South American Earth Sciences, 10, 49–70.
- MENG, J. and FOX, R. C. 1995a. Osseous inner ear structures and hearing in early marsupials and placentals. Zoological Journal of the Linnean Society, 115, 47–71.
- MENG, J. and FOX, R. C. 1995b. Therian petrosals from the Oldman and Milk River formations (Late Cretaceous), Alberta. Journal of Vertebrate Paleontology, 15, 122–130.
- MUIZON, C. de 1992. La fauna de mamíferos de Tiupampa (Paleoceno inferior, Formación Santa Lucía), Bolivia. 575–624. In SUÁREZ-SORUCO, R. (ed.). Fósiles y fácies de Bolivia, Vol. I. Vertebrados. Revista Técnica, Yacimientos Petrolíferos Fiscales Bolivianos, Santa Cruz, 718 pp.
- MUIZON, C. de 1994. A new carnivorous marsupial from the Palaeocene of Bolivia and the problem of marsupial monophyly. Nature, 370, 208–211.
- MUIZON, C. de 1998. Mayulestes ferox, a boryaenoid (Metatheria, Mammalia) from the early Palaeocene of Bolivia. Phylogenetic and palaeobiologic implications. Geodiversitas, 20, 19–142.
- MUIZON, C. de 1999. Marsupial skulls from the Deseadan (Late Oligocene) of Bolivia and phylogenetic analysis of the Borhyaenoidea (Marsupialia, Mammalia). Geobios, 32, 483–509.
- MUIZON, C. de and BRITO, I. M. 1993. Le bassin calcaire de São José de Itaboraí (Rio de Janeiro, Brésil): ses relations fauniques avec le site de Tiupampa (Cochabamba, Bolivie). Annales de Paléontologie, 79, 233–268.
- MUIZON, C. de and CIFELLI, R. L. 2001. A new basal ‘didelphoid’ (Marsupialia, Mammalia) from the Early Paleocene of Tiupampa (Bolivia). Journal of Vertebrate Paleontology, 21, 87–97.
- MUIZON, C. de, CIFELLI, R. L. and CÉSPEDES PAZ, R. 1997. The origin of the dog-like borhyaenoid marsupials of South America. Nature, 389, 486–489.
- MUIZON, C. de and LANGE-BADRÉ, B. 1997. Carnivorous dental adaptations in the tribosphenic mammals and phylogenetic reconstruction. Lethaia, 30, 353–366.
- NILSSON, M. A., GULLBERG, A., SPOTORNO, A. E., ARNASON, U. and JANKE, A. 2003. Radiation of extant marsupials after the K/T boundary: evidence from complete mitochondrial genome. Journal of Molecular Evolution, 57, 3–12.
- NIXON, K. C. 2002. WinClada, Version 1.00.08. Program and documentation. Available from http://www.cladistics.com, published by the author, Ithaca, NY.
- NOVACEK, M. J. 1986. The skull of leptictid insectivorans and the higher-level classification of eutherian mammals. Bulletin of the American Museum of Natural History, 183, 1–112.
- NOVACEK, M. J. 1993. Patterns of diversity in the mammalian skull. 438–545. In HANKEN, J. and HULL, B. K. (eds). The skull: patterns of structural and systematic diversity. University of Chicago Press, Chicago, IL, 566 pp.
- OSGOOD, W. H. 1921. A monographic study of the American marsupial, Caenolestes. Publications of the Field Museum of Natural History, 14, 1–232.
- PATTERSON, B. and MARSHALL, L. G. 1978. The Deseadan, Early Oligocene, Marsupialia of South America. Fieldiana: Geology, 41, 37–100.
- PHILLIPS, M. J., LIN, Y.-H., HARRISON, G. L. and PENNY, D. 2001. Mitochondrial genomes of a bandicoot and a brushtail possum confirm the monophyly of australidelphian marsupials. Proceedings of the Royal Society of London, Series B, 268, 1533–1538.
- PRESLEY, R. 1980. The braincase in Recent and Mesozoic therapsids. Mémoires de la Société Géologique de France, Nouvelle Série, 139, 159–162.
- REIG, O. A., KIRSCH, J. A. W. and MARSHALL, L. G. 1987. Systematic relationships of the living and Neocenozoic American ‘opossum-like’ marsupials (suborder Didelphimorphia), with comments on the classification of these and of the Cretaceous and Paleogene New World and European metatherians. 1–89. In ARCHER, M. (ed.). Possums and opossums: studies in evolution. Surrey Beatty and Sons and the Royal Zoological Society of New South Wales, Sydney, 788 pp.
- ROUGIER, G. W. and BONAPARTE, J. F. 1988. La pared lateral del cráneo de Vincelestes neuquenianus (Mammalia, Eupantotheria) y su importancia en el estudo de los mamíferos mesozoicos. Resumenes V Jornadas Argentina de Paleontología de Vertebrados, 14–15.
- ROUGIER, G. W., WIBLE, J. R. and HOPSON, J. A. 1992. Reconstruction of the cranial vessels in the Early Cretaceous mammal Vincelestes neuquenianus: implications for the evolution of the mammalian cranial vascular system. Journal of Vertebrate Paleontology, 12, 188–216.
- ROUGIER, G. W., WIBLE, J. R. and HOPSON, J. A. 1996a. Basicranial anatomy of Priacodon fruitaensis (Triconodontidae, Mammalia) from the Late Jurassic of Colorado, and a reappraisal of mammaliaform interrelationships. American Museum Novitates, 3183, 1–38.
- ROUGIER, G. W., WIBLE, J. R. and HOPSON, J. A. 1996b. Middle-ear ossicles of the multituberculate Kryptobaatar from the Mongolian Late Cretaceous: implications for mammaliamorph relationships and the evolution of the auditory apparatus. American Museum Novitates, 3187, 1–43.
- ROUGIER, G. W., WIBLE, J. R. and HOPSON, J. A. 1998. Implications of Deltatheridium specimens for early marsupial history. Nature, 396, 459–463.
- SÁNCHEZ-VILLAGRA, M. R. 2001a. Ontogenetic and phylogenetic transformations of the vomeronasal complex and nasal floor elements in marsupial mammals. Zoological Journal of the Linnean Society, 131, 459–479.
- SÁNCHEZ-VILLAGRA, M. R. 2001b. The phylogenetic relationships of argyrolagid marsupials. Zoological Journal of the Linnean Society, 131, 481–496.
- SÁNCHEZ-VILLAGRA, M. R. and WIBLE, J. R. 2002. Patterns of evolutionary transformation in the petrosal bone and some basicranial features in marsupial mammals, with special reference to didelphids. Journal of Zoological Systematics and Evolutionary Research, 40, 26–45.
- SEGALL, W. 1970. Morphological parallelisms of the bulla and auditory ossicles in some insectivores and marsupials. Fieldiana: Zoology, 51, 169–205.
- SORENSON, M. D. 1999. TreeTot, Version 2. Boston University, Boston, MA.
- SPRINGER, M. S. and WOODBURNE, M. O. 1989. The distribution of some basicranial characters within the Marsupialia and a phylogeny of the phalangeriformes. Journal of Vertebrate Paleontology, 9, 210–221.
- SPRINGER, M. S., KIRSCH, J. A. W. and CASE, J. A. 1997. The chronicle of marsupial evolution. 129–161. In GIVNISH, T. J. and SYTSMA, K. J. (eds). Molecular evolution and adaptative radiation. Cambridge University Press, New York, NY, 621 pp.
- SPRINGER, M. S., WESTERMAN, M., KAVANAGH, J. R., BURK, A., WOODBURNE, M. O., KAO, D. J. and KRAJEWSKI, C. 1998. The origin of the Australasian marsupial fauna and the phylogenetic affinities of the enigmatic monito del monte and marsupial mole. Proceedings of the Royal Society of London, Series B, 265, 2381–2386.
- SWOFFORD, D. L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods), Version 4. Sinauer Associates, Sunderland, MA, 142 pp.
- SZALAY, F. S. 1982a. A new appraisal of marsupial phylogeny and classification. 621–640. In ARCHER, M. (ed.). Carnivorous marsupials. Royal Zoological Society of New South Wales, Sydney, 804 pp.
- SZALAY, F. S. 1982b. Phylogenetic relationships of the marsupials. Geobios, Mémoire Spécial, 6, 177–190.
- SZALAY, F. S. 1994. Evolutionary history of the marsupials and an analysis of osteological characters. Cambridge University Press, Cambridge, 481 pp.
- SZALAY, F. S. and SARGIS, E. J. 2001. Model-based analysis of postcranial osteology of marsupials from the Paleocene of Itaboraí (Brazil) and the phylogenetics and biogeography of Metatheria. Geodiversitas, 23, 139–302.
- SZALAY, F. S. and TROFIMOV, B. 1996. The Mongolian Late Cretaceous Asiatherium, and the early phylogeny and paleobiogeography of Metatheria. Journal of Vertebrate Paleontology, 16, 474–509.
- VAN DER KLAAUW, C. J. 1931. The auditory bulla in some fossil mammals, with a general introduction to this region of the skull. Bulletin of the American Museum of Natural History, 62, 1–352.
- VILLARROEL, C. and MARSHALL, L. G. 1982. Geology of the Deseadan (Early Oligocene) age Estratos Sallas in the Salla-Luribay Basin, Bolivia, with description of new Marsupialia. Geobios, Mémoire Spécial, 6, 201–211.
- VOSS, R. S. and JANSA, S. A. 2003. Phylogenetic studies on didelphid marsupials II. Nonmolecular data and new IRBP sequences: separate and combined analyses of didelphine relationships with denser taxon sampling. Bulletin of the American Museum of Natural History, 276, 1–82.
- WIBLE, J. R. 1984. The ontogeny and phylogeny of the mammalian cranial arterial pattern (internal carotid artery). Unpublished PhD thesis, Duke University, NC, 705 pp.
- WIBLE, J. R. 1986. Transformations in the extracranial course of the internal carotid artery in mammalian phylogeny. Journal of Vertebrate Paleontology, 6, 313–325.
- WIBLE, J. R. 1987. The eutherian stapedial artery: character analysis and implications for superordinal relationships. Zoological Journal of the Linnean Society, 91, 107–135.
- WIBLE, J. R. 1989. Vessels on the side wall of the braincase in cynodonts and primitive mammals. 406–408. In SPLECHTNA, H. and HILGERS, H. (eds). Trends in vertebrate morphology. Fortschritte der Zoologie, 35, 647 pp.
- WIBLE, J. R. 1990. Petrosals of Late Cretaceous marsupials from North America, and a cladistic analysis of the petrosal in therian mammals. Journal of Vertebrate Paleontology, 10, 183–205.
- WIBLE, J. R. 1991. Origin of Mammalia: the craniodental evidence. Journal of Vertebrate Paleontology, 11, 1–28.
- WIBLE, J. R. 2003. On the cranial osteology of the short-tailed opossum Monodelphis brevicaudata (Didelphidae, Marsupialia). Annals of Carnegie Museum, 72, 137–202.
- WIBLE, J. R. and HOPSON, J. A. 1993. Basicranial evidence for early mammal phylogeny. 45–62. In SZALAY, F. S., NOVACEK, M. J. and McKENNA, M. C. (eds). Mammal phylogeny: Mesozoic differentiation, multituberculates, monotremes, early therians, and marsupials. Springer-Verlag, New York, NY, 591 pp.
- WIBLE, J. R. and HOPSON, J. A. 1995. Homologies of the prootic canal in mammals and non-mammalian cynodonts. Journal of Vertebrate Paleontology, 15, 331–356.
- WIBLE, J. R. and ROUGIER, G. W. 2000. Cranial anatomy of Kryptobaatar dashzevegi (Mammalia, Multituberculata); and its bearing on the evolution of mammalian characters. Bulletin of the American Museum of Natural History, 247, 1–124.
- WIBLE, J. R., ROUGIER, G. W., NOVACEK, M. J. and DASHZEVEG, D. 1995. A mammalian petrosal from the Early Cretaceous of Mongolia: implications for the evolution of the ear region and mammaliamorph interrelationships. American Museum Novitates, 3149, 1–19.
- WIBLE, J. R., ROUGIER, G. W., NOVACEK, M. J. and McKENNA, M. C. 2001. Earliest eutherian ear region: a petrosal referred to Prokennalestes from the Early Cretaceous of Mongolia. American Museum Novitates, 3322, 1–44.
- WOODBURNE, M. O. and CASE, J. A. 1996. Dispersal, vicariance, and the Late Cretaceous to Early Tertiary land mammal biogeography from South America to Australia. Journal of Mammalian Evolution, 3, 121–161.
- WROE, S. 1997. A reexamination of proposed morphology-based synapomorphies for the families of Dasyuromorphia (Marsupialia): Part I, Dasyuridae. Journal of Mammalian Evolution, 4, 19–52.
- WROE, S. 1999. The geologically oldest dasyurid, from the Miocene of Riversleigh, north-west Queensland. Palaeontology, 42, 501–527.
- BRAMMALL, J. and COOKE, B. N. 1998. The skull of Ekaltadeta ima (Marsupialia, Hypsiprymnodontidae?): an analysis of some marsupial cranial features and a re-investigation of propleopine phylogeny, with notes on the inference of carnivory in mammals. Journal of Paleontology, 72, 738–751.
- WROE, S., EBACH, M., AHYONG, S., MUIZON, C. D. and MUIRHEAD, J. 2000. Cladistic analysis of dasyuromorphian (Marsupialia) phylogeny using cranial and dental characters. Journal of Mammology, 81, 1008–1024.