Mammalian superorders

Transcription

1 2001. J. Linn. SOC.. 58: March 1976 Mammalian superorders ELI C. MINKOFF Department of Biology, Bates College, Lewiston, Maine 04240, U.S.A. Accepted for publicafion April 1975 The superordinal level can be used to great advantage in the taxonomy of mammals and other groups. Of the two cohorts of placental mammals, the Unguiculata is divided into six superorders: Archonta (Gregory s Archonta plus Lipotyphla), Glires, Ferae (including Deltatheridia and Carnivora), Paratheria (Edentata and Pholidota), Ganodonta. and an unnamed group for the Lagomorpha plus Anagalida. The cohort Ungulata is divided much as in Simpson (1945), with an additional superorder (Mutica) for the whales and with a superorder Amblypoda, separate from the Paenungulata, and corresponding to Romer s (1966) order of that name. CONTENTS Introduction Advantages of the category superorder Use of the superorder among the Mammalia Unguiculate superorders Ungulate superorders Formal classification References Appendix INTRODUCTION The arrangement of the placental mammals into superorders as well as orders is a convenient way of summarizing our present knowledge of their relationships. In this paper, I shall discuss first the advantages of using the superordinal rank generally. The remainder of the paper is devoted to the arrangement of the orders of placental mammals at this taxonomic level, a subject last thoroughly treated by Simpson (1945). Many important advances in our knowledge have been made since Simpson s classification was first published (among them, Van Valen, 1966; Emry, 1970; Szalay & McKenna, 1971; and many others), but no general scheme comparable with Simpson s has been proposed that might incorporate these advances. ADVANTAGES OF THE CATEGORY SUPERORDER Since the Superorder is not universally employed as a taxonomic category, it would seem quite proper to justify its use here. Some of the arguments on behalf of the Superorder could also be used to urge the use of, say, the 147

2 148 E. C. MINKOFF Superphylum (Valentine, 1973) or the Cohort (Minkoff, in press); other arguments, peculiar to the Superorder, stem principally from the fact that this category ranks immediately above the Order. Theories as to the evolutionary origins of various orders (currently a very active field of research) can often best be expressed taxonomically at the superorder level. Very early fossil genera may well be identifiable to superorder but not to order, especially if a critical portion of the anatomy is not preserved; the same may be true of incomplete remains of any geologic age. Classifications at the level of superorder are thus likely to be heuristic and stimulating to further research, in an effort either to substantiate or to discredit them. New orders in particular can most readily be comprehended if placed in a superorder; at least two such new orders have been proposed within the past decade. Students and others studying classification may learn quite readily the names of the larger orders, but the names of the smaller and less important orders are often easier for them to learn if they can be told to what superordinal group a particular order belongs. This is especially true if the members of the superorder share a common adaptive tendency, zoogeographic region, or the like: the Subungulata or Paenungulata of certain classifications, for example, contains four orders whose shared ancestry is presumably from a froup of large African swamp d well ers. At present the category of Superorder is used with some degree of regularity among the Mammalia, Amphibia, Teleostei, Crustacea, Arachnida, and other groups. Among the insects, the so-called orthopteroid complex is really a taxon of superordinal rank; the same is true of hemipteroid complex and panorpoid complex. Other groups which may to advantage adopt the practice of using superorders include the Aves (specifically the Neognathae), Chondrichthyes, Placodermi, and Cephalopoda. The advantages of the category are not by any means confined to the Mammalia. USE OF THE SUPERORDER AMONG THE MAMMALIA Although a number of 19th-century taxonomists (among them de Blainville, and Gill, 1870) used taxa which might now be considered superorders, the earliest definite and explicit use of this rank known to me is that of Weber (1904), and the earliest consistent use is that of Gregory (1910). Weber recognized three superorders within the placental Mammalia (his Unterklasse Monodelphia ), but not all orders were placed in superorders. All three of Weber s superorders (Edentata, Ungulata, Primates) were well-known taxa usually considered orders by his predecessors, but Weber wished to emphasize the distinctness of the various orders involved, while in each case still reaffirming their relationship at a higher taxonomic level. Gregory (1910) was the first taxonomist to divide all placental mammals into superorders. His seven superorders are listed below. Gregory used no taxonomic rank between the infraclass and the superorder, and in fact the writers of most 20th-century textbooks have despaired of using any rank between infraclass and order! A notable exception, and the only major revision of mammalian superordinal classification since Gregory, is Simpson (1949, who used both cohort and superordinal ranks. I have discussed in a separate

3 MAMMALIAN SUPE RORDERS 149 paper my reasons for re-arranging Simpson s cohorts, returning the rodents and carnivores to the Linnaean cohort Unguiculata, and removing the carnivores from but adding the Cetacea to the Linnaean cohort Ungulata. Four of the above-mentioned classifications are given in an appendix to the present work. Simpson s (1945) represents the best of his generation, Gregory s (1910) of the previous generation, and Gill s (1870) one of the most original and most far-sighted of the generation before Gregory. Each of these four classifications bears an important resemblance to the one here adopted. UNG UIC ULATE SUP E RO RD ERS Six superorders are here recognized within the cohort Unguiculata. The first is based on Gregory s (1910) Archonta, but with the important addition of the lipotyphlous insectivores. Gregory s name Archonta is quite available for this insectivore-bat-primate assemblage. The rodents are probably derived from within the Archonta (Wood, 1962; McKenna, 1961, 1966), but their remarkable adaptations and consequent evolutionary success would seem to merit for them superordinal distinction. The order Carnivora would seem to be derived from somewhere within or near the order Deltatheridia (including tenrecoids). The Linnaean name Ferae is available for this superorder, which may well include the ancestry of the next two superorders. The edentates were first clearly distinguished by Blumenbach ( ), who used Linnaeus name Bruta. A separate mammalian subclass for the edentates has been advocated by de Blainville (1 83 9), Gervais (1 858), Thomas (1887), and Scott (1903); Huxley (1872) also set aside the edentates sharply, and Gregory (1910) made his edentate assemblage a superorder. While I believe that a separate edentate subclass would exaggerate their distinctness, I still feel that the edentates are sufficiently different from other unguiculates to merit superordinal distinction at least. That they deserve to rank as a superorder coordinate with the rodents and the carnivores is indicated principally by the fact that the earliest fossil edentates are at least as dissimilar from contemporaneous insectivores as are the earliest carnivores or rodents, but also by the lack in the Edentata of an 0s penis (present in the other Unguiculata) and by the plasticity in the edentates of certain characters (number of cervical vertebrae, accessory zygapophyses, dermal armor, secondary isodonty, loss of enamel on crowns of teeth) usually constant among other mammals. The pangolins (Pholidota) were traditionally included in the Edentata until Weber (1891, 1904) emphasized their distinctness. The extinct palaeanodonts were studied most intensively by Matthew (19 1 8>, who emphasized their close relationship with both the pangolins and the true edentates (Xenarthra), and by Simpson (193 l), who affirmed their close relationship to the Xenarthra but minimized that with the pangolins. Emry (1970) argues that the newly discovered Patrioman& supports both Matthew s position, and the hypothesis that the Old World pangolins have descended from the North American palaeanodonts, geography notwithstanding. A pangolinedentate relationship above the ordinal level has been advocated by Weber (1904), Gregory (1910), and Matthew (1918), among others, but Emry (1970) denies this relationship. The relevant evidence is discussed by Simpson (1931), whose conclusions do

4 150 E. C. MINKOFF not directly support but do clearly allow such a relationship. Emry s (1970) arguments, that the Palaeanodonta cannot be directly ancestral to the Xenarthra, still do not refute Simpson s (1931) claim of a collateral relationship between the two. The palaeanodonts are more closely related, and probably directly ancestral, to the pangolins (Emry, 1970), but their relation to the Xenarthra, though weakened by Emry s revelations, is nevertheless real: if the Pholidota are regarded as the grand-children of the palaeanodonts, then the Xenarthra may well be their grand-nephews. Shrivastava (1963) has also shown that the shoulder musculature of pangolins is no more distinctive or peculiar than that of any of the xenarthrous families. All this might best be expressed taxonomically by accepting Emry s (1970) referral of the palaeanodonts to the order Pholidota, but also bracketing this order with the Edentata as a distinct superorder. For the pangolinedentate superorder, I use Thomas (1887) name Paratheria, first because other possible names (such as Effodientia Illiger, 181 1, or Bruta Linnaeus, 1758) originally excluded certain true edentates, second because Thomas used his term to emphasize the distinctness of edentates from other placentals (though on grounds which must now be rejected), and third because Gregory (1910: 323, 465) does confer some small dignity upon Thomas term. Another possibility would have been to use Edentata for the superorder and Xenarthra or some other name for the nuclear order (cf. Gregory, 1910: 465). The Taeniodonta are an extinct, enigmatic order, whose resemblances to the Edentata are surely superficial and convergent. (Such resemblances are always to the later, most specialized edentates, rather than to the earlier and more primitive ones.) Patterson (1949) has reviewed our knowledge of the order, concluding that taeniodonts are probably derived from unknown Cretaceous insectivores. The description of an Asian taeniodont by Dehm & Oettingen- Spielberg (1958) sheds no new light on the problem of taeniodont relationships. The convergent nature of their resemblance to the edentages argues against their inclusion in the edentate-pangolin assemblage (cf. Gregory, 1910). Their supposed derivation from unknown Cretaceous insectivores (quite possibly within the family Paleoryctidae) hardly qualifies the taeniodonts for inclusion in the Archonta, unless by the same reasoning we are also to include virtually every placental order in this same group. On the contrary, the early and sometimes extreme specializations of the Taeniodonta, and their lack of relationship to any other placental order, argue quite well for their inclusion as a separate superorder, for which Wortman s (1 896, 1897) name Ganodonta is available and appropriate. The Lagomorpha are manifestly not related to the Rodentia (see Wood, 1957; Moody, Cochran & Drugg, 1949), but their affinities are otherwise uncertain. Wood (1957) and Sych (1966) have favored ungulate affinities for this order, but if Van Valen (1964) and Szalay & McKenna (1971) are correct, their ancestry must be sought in or near the extinct order Anagalida, of Asian provenance. Their distinctness (or that of the Anagalida) from the Insectivora would seem to be of long standing (Cretaceous), thus meriting superordinal distinction in my opinion. I shall refrain from coining a name for this superorder (no existing name is available), largely because of the persistent uncertainty regarding rabbit ancestry.

5 MAMMALIAN SUPERORDERS 151 A comparison with Gill s (1870) classification reveals many similar groupings, including his Insectivorous series plus Primate series (my Archonta), his Rodent series (my Glires), his Feral series (my Ferae), his Edentate series (my Paratheria), and his Ungulate series (my Ungulata). To be sure, Gill classified the whales and lagomorphs differently, but in most respects his anticipation of modern gorupings is noteworthy. UNGULATE SUPERORDERS The arrangement of the Ungulata into superorders follows Simpson (1945) in recognizing the Protungulata (condylarths and their near descendents), Paenungulata (Proboscidea, Sirenia, and related orders, but see below), Mesaxonia (perissodactyls), and Paraxonia (artiodactyls). The Ferae (Carnivora) are here excluded from the Ungulata, while the whales are included as the superorder Mutica. A distinct superorder is recognized for the orders Pantodonta, Dinocerata, Pyrotheria, and Xenungulata. The tillodonts (Gazin, 1953) are an early, primitive order, bearing few affinities to any but the most primitive placental orders (hence, as Simpson and others have indicated, to the Insectivora). Van Valen (1963) and Romer (1966) have both suggested possible derivation from condylarths, and I have followed their lead in tentatively referring the tillodonts to the Protungulata. The Amblypoda were first recognized by Cope (1875, etc), who bracketed the pantodonts and uintatheres under this name. Flower and Lydekker (1891) associated this group with the Hyracoidea, Proboscidea, Condylarthra, and Toxodontia (= Notoungulata) under the name Subungulata. Schlosser (1923) and Weber (1928) refined the Subungulata by adding the Sirenia and the newly discovered Embrithopoda, and by excluding the condylarths, notoungulates, and the Amblypoda. As thus restricted, the Subungulata form a closely-knit group of generally heavy, primitively swamp-dwelling ungulates of African origin, distinct on zoogeographical as well as other grounds from the Amblypoda and Pyrotheria. The latter order, in which the xenungulates were usually included, were often associated (e.g., by Ameghino) with the Proboscidea on the basis of resemblances now known to be convergent. Simpson ( 1945) included seven orders in his superorder Paenungulata, using the four primitively African orders Proboscidea, Sirenia, Hyracoidea, and Embrithopoda (i.e., Schlosser s Subungulata ) as the nucleus of this superorder. He expressed misgivings as to the frankly hypothetical inclusion of the Pantodonta, Dinocerata, and Pyrotheria, noting that there probably was not a single primitive specifically paenungulate phylum that split up into the seven orders I place here... Doubting the strict monophyly of his assemblage, Simpson added, It is probable that there was some protungulate division that tended toward graviportal body build..., and that all these seven orders came from such a protungulate group... Paula Couto s (1952) study of Xenungulata, and Wheeler s (1961) study of Dinocerata, both substantiate the existence of resemblances between these two orders, some of them also shared by the Pantodonta and Pyrotheria, but not by the Subungulata of African ancestry. Wheeler specifically hinted at the possibility of an amblypod assemblage equivalent to the present one, but it remained for Romer (1966) to associate together the Pantodonta, Dinocerata,

6 152 E. C. MINKOFF Pyrotheria, and Xenungulata as suborders of an enlarged order Amblypoda. He further emphasized the distinctness of this group by excluding them from his chapter on Subungulates. On page 244, he declares: In the early Tertiary..., a number of archaic ungulate groups... rapidly tended... to attain large size and ponderous build.... One series of forms of this sort, the elephants and other subungulates, appears to have had its center in Africa.... A number of other groups of archaic character... were predominantly American in their history. These... included the Pantodonta, Dinocerata, Xenungulata, and Pyrotheria. Romer s emended Amblypoda seems to be a convenient, defensible, and probably also a natural group. That the four included groups do each merit ordinal rank has been emphasized by Simpson (1945: 241): The possibly polyphyletic origin of the [Paenungulata], the ancient and deeply defined separation of its constituents, and their long histories and great differentiation are such that each must logically be given ordinal rank. Superorders as now used are theoretical or even hypothetical groupings. Orders are the major divisions of placentals that can reasonably be said to have an established common origin, and by these criteria these are orders, and their collocation is superordinal. Thus, Romer s Amblypoda are given superordinal rank, and his four suborders restored as orders. The name Amblypoda is long familiar for this taxon. The order Desmostylia may well be associated with either the Amblypoda or Paenungulata; on zoogeographical considerations alone, the former association would seem the more likely. The parallelism of the Pyrotheria with the Proboscidea, of the Desmostylia with the Sirenia, and in some ways of the Dinocerata with the Embrithopoda all tend to suggest that the Amblypoda and Paenungulata occupied broadly the same adaptive zone in different parts of the world. Their resemblances need not, although they may nevertheless, indicate a common (but remote) protungulate ancestry, nor can a perissodactyl ancestry (cf. Van Valen, 1971) be ruled out either. FORMAL CLASSIFICATION In the following classification, the asterisk (*) denotes extinct taxa. Class MAMMALIA Linnaeus 1758: 14. Subclass PROTOTHERIA Gill 1872: vi sensu Hopson 1970: 7. *Infraclass EOTHERIA Kermack & Mussett 1958: 212. *Order TRICONODONTA *Order DOCODONTA Infraclass ORNITHODELPHIA Blainville 1834 Cfide Palmer 1904: 780). Order MONOTREMATA *Infraclass ALLOTHERIA Marsh 1880: 239. *Order MULTITUBERCULATA Subclass THERIA Parker & Haswell 1897: 448 (=Eutheria Gill 1872: 1, now used in a restricted sense).

7 MAMMALIAN SUPERORDERS 153 *Infraclass TRITUBERCULATA Zittel 1893: 100 (or Osborn 1893). *Order PANTOTHERIA (or EUPANTOTHERIA) *Order SYMMETRODONTA Infraclass METATHERIA Huxley 1880: 654 (=Marsupialia Illiger 1811 : 75, =Didelphia Blainidle 1816: 117 [not p. 1091, sensu Blainville 1834). Order MARS UPICARNIVOR A (or POLY PROTODONTA) Order PAUCITUBERCULATA (or CAENOLESTOIDIA) Order PERAMELINA (or PERAMELIDA) Order DIPROTODONTA Infraclass EUTHERIA Gill 1872: 1 sensu Huxley 1880: 657 (=Monodelphia Blainville 1816: 117 [not p ). Cohort UNGUICULATA Linnaeus 1766: 21, 24. Superorder ARCHONTA Gregory 1910: 322. Order INSECTIVORA (with suborders Proteutheria, Macroscelidea, Dermoptera & Lipotyphla). Order CHIROPTERA Order PRIMATES Superorder GLIRES Linnaeus 1758: 56. Order RODENTIA Superorder FERAE Linnaeus 1758: 37. Order DELTATHERIDIA (or CREODONTA) Order CARNIVORA *Superorder GANODONTA Wortman 1896: 259. *Order TAENIODONTA Superorder PARATHERIA Thomas 1887: 459. Order EDENTATA (or XENARTHRA) Order PHOLIDOTA (incl. *Palaeanodonta as suborder). Unnamed superorder *Order ANAGALIDA Order LAGOMORPHA Cohort UNGULATA Linnaeus 1766: ; Superorder MUTICA Linnaeus 1766: 22, 24. Order CETACEA Superorder PROTUNGULATA Weber 1904: 587 (or Marsh 1884: 182). *Order CONDYLARTHRA Order TUBULIDENTATA *Order LITOPTERNA *Order NOTOUNGULATA *Order ASTRAPOTHERIA (incl. *Trigonostylopoidea). (?)*Order TILLODONTIA Superorder PAENUNGULATA Simpson 1945: 131 (=Subungulata Flower & Lydekker 1891: 274, 414 sensu Schlosser 1923, nec Illiger 1811: 92). Order PROBOSCIDEA Order HYRACOIDEA *Order EMBRITHOPODA Order SIRENIA

8 154 E. C. MINKOFF *Superorder AMBLYPODA Cope 1875: 24, 28 sensu Romer 1966: 385; new as superorder. *Order PANTODONTA * Order D INOC ERATA *Order PYROTHERIA *Order XENUNGULATA (?)*Order DESMOSTY LIA Superorder MESAXONIA Marsh 1884: 9, 177. Order PERISSODACTYLA Superorder PARAXONIA Marsh 1884: Order ARTIODACTYLA REFERENCES BLAINVILLE, H. M. de, Prodrome d une nouvelle distribution systimatique du rkgne animal. Bull. Sci. SOC. Philom. Paris (sbr. 31, 3: [ Pp are incorrectly numbered BLAINVILLE, H. M. de, Cours de la facult6 des sciences, [Cited by Palmer (1904: 780); not seen.] BLAINVILLE, H. M. de, Nouvelle classification des mammifkres. Ann. fr. btrang. anat. physiol. 2: 268. BLAINVILLE, H. M. de, Ostebgraphie ou description iconographique comparbe du squelene et du systkme den taire des mammif6res. 4 vok. BLUMENBACH, J. F., Handbuch der Naturgeschichte, 2 vols. Gottingen: Johann Christian Dietrich. COPE, E. D., Systematic catalogue of Vertebrata of the Eocene of New Mexico collected in Rept. Engineer Dept., U.S. Army, Geogr. Explor. Surv. (Lt. George M. Wheeler), 37 pp. DEHM, R. & OETTINGEN-SPIELBERG, T., Palaontologische Untersuchungen im Tertiir von Pakistan. 2. Die mitteleocanen Saugetiere von Ganda Kas bei Basal in Nordwest-Pakistan. Abh. buyer. Akad. Wiss. (N.F.), 91: EMRY, R. J., A North American Oligocene pangolin and other additions to the Pholidota. Bull. Am. Mus. nut. Hist., 142: FLOWER, W. H. & LYDEKKER, R., An introduction to the study of mammals living and extinct, 763 pp. London: A. & C. Black. GAZIN, C. L., The Tillodontia: an early Tertiary order of mammals. Smithson. misc. Collns., 121(10): GERVAIS, F. L. P., Histoire Naturelle des MammifGres, 2: 344 pp. Paris: H. Curmer. GILL, T., On the relations of the orders of mammals. Proc. Am. Ass. Advmt. Sci. 19: GILL, T., Arrangement of the families of mammals with analytical tables. Smithson. misc. Collns., 11(1): GREGORY, W. K., The orders of mammals. Bull.Am. Mus. nut. Hist., 27: HOPSON, J. A., The classification of nontherian mammals. J. Mammal., 51: 1-9. HUXLEY, T. H., A manual of the anatomy of vertebrated animals, 431 pp. New York: D. Appleton. HUXLEY, T. H., On the application of the laws of evolution to the arrangement of the Vertebrata and more particularly of the Mammalia. Proc. zool. SOC. Lond.: ILLIGER, C., Prodromus systematir mammalium et avium additis terminis roographicus utriudque classis. 301 pp. Berlin: C. Salfeld. KERMACK, K. A. & MUSSETT, F., The jaw articulation of the Docodonta and the classification of Mesozoic mammals. Proc. R. SOC. Lond. (Ser. B), 149: LINNEAUS, C., Systema naturae per regna tria naturae..., 1: 824 pp. Editio decima, reformata. Stockholm: Laurentii Salvii. LINNAEUS, C., Systema naturae per regna tria naturae..., 1: 532 pp. Editio duodecima, reformata. Stockholm: Laurentii Salvii. McKENNA, M. C., A note on the origin of rodents. Am. Mus. Novit., 2037: 1-5. McKENNA, M. C., Paleontology and the origin of the primates. Folia Primat., 4: MARSH, 0. C., Notice of Jurassic mammals representing two new orders. Am. J. Sci. (Ser. 3), 20: MARSH, 0. C., 1884 [? Dinocerata. A monograph of an extinct order of gigantic mammals. Monogr. U.S. Geol. Surv., 10:

9 MAMMALIAN SUPERORDERS 155 MATTHEW, W. D., Edentata. In W. D. Matthew & W. Granger, A revision of the lower Eocene Wasatch and Wind River faunas. Part V. Insectivora (continued), Glires, Edentata. Bull. Am. Mus. nut. Hist.. 38: MOODY, P. A,. COCHRAN, V. A. & DRUGG, H., Serological evidence on lagomorph relationships. Evolution, 3: OSBORN, H. F., On the rise of the Marnmalia in North America. Stud. Biol. Lab. Columbia CON. (ZOOL), l(2): 145. PALMER, T. S., Index generum mammalium: a list of the genera and families of mammals. N. Am. Fauna, 23: PARKER, T. J. & HASWELL, W. A A textbook ofzoology, 2: 683 pp. London: Macmillan. PATTERSON, B., Rates of evolution in taeniodonts. In G. L. Jepsen, E. Mayr & G. G. Simpson (Eds.), Genetics, paleontology, and evolution: Princeton, N.J.: Princeton Univ. Press. PAULA COUTO, C. DE, Fossil mammals from the beginning of the Cenozoic in Brazil. Condylarthra, Litopterna, Xenungulata, and Astrapotheria. Bull. Am. Mus. nat. Hist. 99: ROMER, A. S., Vertebrate paleontology, 3rd ed., 468 pp. Chicago: Univ. of Chicago Press. SCHLOSSER, M., Mammalia. In K. A. v. Zittel, F. Broili & M. Schlosser, Grundzuge der Palaontologie (Palaozoologie). vierte Auflage, zweite Abteilung, Vertebrata: pp Munich & Berlin: R. Oldenbourg. SCOTT, W. B., 1903 [ Mammalia of the Santa Cruz Beds. Part 1. Edentata. Rep. Princeton Univ. Exped. Patagonia, 5(1): SHRIVASTAVA, R. K., Deltoid musculature of the Edentata, Pholidota, and Tubulidentata. Okajimas Folia anat. jap., 38: SIMPSON, G. G., Meracheiromys and the Edentata. Bull. Am. Mus. nut. Hist., 59: SIMPSON, G. G., The principles of classification and a classification of mammals. Bull. Am. Mus. nut. Hbt.. 85: SYCH, L., Czy przodkami zajaca byly ssaki kopytne? Were the ungulated mammals ancestors of the hare? Przegl. zool.. 10: SZALAY, F. S. & McKENNA, M. C., Beginning of the age of mammals in Asia: the late Paleocene Gashato fauna, Mongolia. Bull. Am. Mw. nut. Hist., 144: THOMAS, O., On the homologies and succession of the teeth in the Dasyuridae, with an attempt to trace the history of the evolution of mammalian teeth in general. Phil. Tram. R. Soc. Lond., 18878: VALENTINE, J. W., Coelomate superphyla. Systematic Zool.. 22: VAN VALEN, L., The origin and status of the mammalian order Tillodontia. J. Mammal., 44: VAN VALEN, L., A possible ongin for rabbits. Evolution, 18: VAN VALEN, L., Deltatheridia, a new order of mammals. Bull. Am. Mus. nat. Hbt. 132: VAN VALEN, L., Adaptive zones and the orders of mammals. Evolution, 25: WEBER, MAX, Beitrige zur Anatomie und Entwicklung des genus Man&. Zoologische Ergebnisre einer Reise in Niederlandisch Ost-Indien, 2: Leiden: E. J. Brill. WEBER. MAX, Die Saugetiere. Einfuhrung in die Anatomie und Systematik der recenten und fossilen Mammalia, 866 pp. Jena: G. Fischer. WEBER. MAX, Die Saugetiere. Einfuhrung in die Anatomie und Systematik der recenten und fossilen Mamalia I, Anaromiacher Teil (1927), 444 pp.; 2, Systematischer Teil (1928). 898 pp. Jena: G. Fischer. WHEELER, W. H., Revision of the uintatheres. Bull. Peabody Mus. nut. Hbr., 14: WOOD, A. E., What, if anything, is a rabbit? Evolution, 11: WOOD, A. E., The early Tertiary rodents of the family Paramyidae. 7hns Am. Phil. Soc., 52: WORTMAN, J. L., Psittacotherium. a member of a new and primitive suborder of the Edentata. Bull. Am. Mus. nat. Hist.. 8: WORTMAN, J. L., The Ganodonta and their relationship to the Edentata. Bull. Am. Mus. nut. Hisr., 9: ZITTEL, K. A. v., Handbuch der Palaontologie. Abteilung I. Palaozoologie. Band I V, Vertebrara (Mammalia): 799 pp. Munich: R. Oldenbourg. 11

10 156 E. C. MINKOFF APPENDIX Synopsis of four influential classifications (Extinct taxa are indicated by an asterisk) Blainville ( ) Monodelphes [Placental mammals] Maldentb [Edentatal Bien dentis Primatis [Primates] Secundatks [Carnivores and insectivores] Tertiatks [Rodents] Quaternatks [Ungulates] Gravigrades [Proboscidea and Sirenia] On logrades fdoigts impairs [Perissodactyla and Hyracoideal A doigts pairs [Artiodactyla] Gill (1870) Subclass Ornithodelphia Order Monotremata Sub class Didelp hia Order Marsupialia Subclass Monodelphia [Placental mammals] Insectivorous series Order Insectivora Order Chiroptera Primate series Order Primates Feral series Order Ferae [Carnivora] Order Cete [Cetacea] Rodent series Order Glires [Rodents and lagomorphs] Edentate series Order Bruta, or Edentata Ungulate series Order Hyracoidea Order Proboscidea Order Sirenia Order Ungulata Suborder Perissodactyla Suborder Artiodactyla Class Mammalia *?Subclass Promammalia Subclass Prototheria Order Monotremata Gregory ( 19 10)

11 MAMMALIAN SUPERORDERS 157 Subclass Theria Infraclass Metatheria *Order Triconodonta *?Order Trituberculata (=Pantotheria) Order Marsupialia Infraclass Eutheria Superorder Therictoidea Order Insectivora [ Lipotyphlous insectivores] Order Ferae [Carnivora] Superorder Archonta Order Menotyphla Order Dermoptera Order Chiroptera Order Primates Superorder Rodentia Order Glires [Rodents and lagomorphs]?superorder Edentata (=Paratheria) *Order Taeniodonta Order Tubulidentata Order Pholidota Order Xenarthra Superorder Cetacea *Order Zeuglodontia Order Odontoceti Order Mystacoceti Superorder Ungulata *Order Protungulata [ Condylarthra plus Taligrada ] *Order Amblypoda [ Pantodonta and Dinocerata] *Order Barytheria Order Sirenia Order Proboscidea Order Hyraces *Order Embrith opoda *Order Notoungulata [sensu lato] Order Mesaxonia [Perissodactyls] Superorder Paraxonia Order Artiodactyla Class Mammalia Subclass Prototheria Order Monotremata *Subclass Allotheria *Order Multituberculata *Subclass uncertain *Order Triconodonta Subclass Theria * Infraclass Pantotheria *Order Pantotheria 11. Simpson (1945)

12 158 E. C. MINKOFF *Order Symmetrodonta Infraclass Metatheria Order Marsupialia Infraclass Eutheria Cohort Unguiculata Order Insectivora Order Dermoptera Order Chiroptera Order Primates *Order Tillodontia *Order Taeniodonta Order Edentata Order Pholidota Cohort Glires Order Lagomorpha Order Rodentia Cohort Mutica Order Cetacea Cohort Ferungulata [ Ungulata plus Carnivora] Superorder Ferae Order Carnivora Superorder Protungulata *Order Condylarthra *Order Litopterna *Order Notoungulata *Order Astrapotheria Order Tubulidentata Superorder Paenungulata *Order Pantodonta *Order Dinocerata *Order Pyrotheria Order Proboscidea *Order Embrithopoda Order Hyracoidea Order Sirenia Superorder Mesaxonia Order Perissodactyla Superorder Paraxonia Order Artiodactyla