2001. J. Linn. SOC..58: 147-158
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
. . . . . . . . .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
147
147
148
149
151
152
154
156
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 n o general scheme comparable with Simpson’s has
been proposed that might incorporate these advances.
ADVANTAGES O F THE CATEGORY “SUPERORDER”
Since the Superorder is not universally employed as a taxonomic category, it
would seem quite proper t o 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
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 t o 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 dwell 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,
1839-1864 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
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 t o 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 (1779-1797),
who used Linnaeus’ name “Bruta”. A separate mammalian subclass for the
edentates has been advocated by de Blainville ( 1 83 9), Gervais ( 1 8 5 8 ) , 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 t o 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 t o 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
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 t o 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,
1811, 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 t o 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 & OettingenSpielberg (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 t o 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 (1896, 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 o r 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.
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 t o 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 h e r e . . .” Doubting the strict monophyly of his
assemblage, Simpson added, “ I t 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,
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).
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 POLYPROTODONTA)
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. 1091 ).
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: 21. 24;
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
E. C. MINKOFF
154
*Superorder AMBLYPODA Cope 1875: 24, 28 sensu Romer 1966:
385; new as superorder.
*Order PANTODONTA
* Order DINOCERATA
*Order PYROTHERIA
*Order XENUNGULATA
(?)*Order DESMOSTY LIA
Superorder MESAXONIA Marsh 1884: 9, 177.
Order PERISSODACTYLA
Superorder PARAXONIA Marsh 1884: 9. 177.
Order ARTIODACTYLA
REFERENCES
BLAINVILLE, H. M. de,
Sci. SOC. Philom. Paris
BLAINVILLE, H. M. de,
not seen.]
BLAINVILLE, H. M. de,
1816.Prodrome d’une nouvelle distribution systimatique du rkgne animal. Bull.
(sbr. 31,3: 1 1 3-24.[ Pp. 1 1 3-20are incorrectly numbered 105-121.
1834. “Cours de la facult6 des sciences, 1834.” [Cited by Palmer (1904:780);
1839. Nouvelle classification des mammifkres. Ann. fr. btrang. anat. physiol. 2:
268.
BLAINVILLE, H. M. de, 1839-1864.Ostebgraphie ou description iconographique comparbe du squelene
e t du systkme den taire des mammif6res. 4 vok.
BLUMENBACH, J. F., 1779-1797.Handbuch der Naturgeschichte, 2 vols. Gottingen: Johann Christian
Dietrich.
COPE, E. D., 1875. Systematic catalogue of Vertebrata of the Eocene of New Mexico collected in 1874.
Rept. Engineer Dept., U.S. Army, Geogr. Explor. Surv. (Lt. George M. Wheeler), 37 pp.
DEHM, R. & OETTINGEN-SPIELBERG, T., 1958. 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: 1-54.
EMRY, R. J., 1970. A North American Oligocene pangolin and other additions to the Pholidota. Bull.
A m . Mus. nut. Hist., 142: 455-510.
FLOWER, W. H. & LYDEKKER, R., 1891.A n introduction to the study o f mammals living and extinct,
763 pp. London: A. & C. Black.
GAZIN, C. L., 1953. The Tillodontia: an early Tertiary order of mammals. Smithson. misc. Collns.,
121(10): 1-110.
GERVAIS, F. L. P., 1858. Histoire Naturelle des MammifGres, 2: 344 pp. Paris: H. Curmer.
GILL, T., 1870.On the relations of t h e orders of mammals. Proc. A m . Ass. Advmt. Sci. 19: 267 70.
GILL, T., 1872. Arrangement of t h e families of mammals with analytical tables. Smithson. misc. Collns.,
11(1):1-98.
GREGORY, W. K., 1910.The orders of mammals. Bull.Am. Mus. nut. Hist., 27: 1-524.
HOPSON, J . A., 1970.The classification of nontherian mammals. J. Mammal., 51: 1-9.
HUXLEY, T. H., 1872. A manual of the anatomy of vertebrated animals, 431 pp. New York: D.
Appleton.
HUXLEY, T. H., 1880. On t h e application of t h e laws of evolution to the arrangement of the Vertebrata
and more particularly of the Mammalia. Proc. zool. SOC.Lond.: 649-62.
ILLIGER, C., 181 1. Prodromus systematir mammalium e t avium additis terminis roographicus utriudque
classis. 301 pp. Berlin: C. Salfeld.
KERMACK, K. A. & MUSSETT, F., 1958.The jaw articulation of the Docodonta and the classification of
Mesozoic mammals. Proc. R . SOC.Lond. (Ser. B), 149: 204-15.
LINNEAUS, C., 1758. Systema naturae per regna tria naturae . . ., 1: 824 pp. Editio decima, reformata.
Stockholm: Laurentii Salvii.
LINNAEUS, C., 1766. Systema naturae per regna tria naturae.. ., 1: 532 pp. Editio duodecima,
reformata. Stockholm: Laurentii Salvii.
McKENNA, M. C.,1961.A note o n t h e origin of rodents. A m . Mus. Novit., 2037: 1-5.
McKENNA, M. C., 1966.Paleontology and the origin of the primates. Folia Primat., 4 : 1-25.
MARSH, 0. C., 1880. Notice of Jurassic mammals representing two new orders. A m . J . Sci. (Ser. 3),20:
235-9.
MARSH, 0. C., 1884 [?18851. Dinocerata. A monograph of an extinct order of gigantic mammals.
Monogr. U.S.Geol. Surv., 10: 1-237.
MAMMALIAN SUPERORDERS
155
MATTHEW, W. D., 1918. 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. A m . Mus. nut.
Hist.. 38: 565-657.
MOODY, P. A,. COCHRAN, V. A. & DRUGG, H., 1949. Serological evidence on lagomorph relationships.
Evolution, 3: 25-33.
OSBORN, H. F., 1893. On t h e rise of the Marnmalia in North America. Stud. Biol. Lab. Columbia CON.
( Z O O L ) , l(2): 145.
PALMER, T. S., 1904. Index generum mammalium: a list of the genera and families of mammals. N. A m .
Fauna, 23: 1-984.
PARKER, T. J. & HASWELL, W. A.. 1879.A textbook o f z o o l o g y , 2: 683 pp. London: Macmillan.
PATTERSON, B., 1949. Rates of evolution in taeniodonts. In G. L. Jepsen, E. Mayr & G. G. Simpson
(Eds.), Genetics, paleontology, and evolution: 243-78.Princeton, N.J.: Princeton Univ. Press.
PAULA COUTO, C. DE, 1952. Fossil mammals from the beginning of the Cenozoic in Brazil.
Condylarthra, Litopterna, Xenungulata, and Astrapotheria. Bull. A m . Mus. nat. Hist. 99: 355-94.
ROMER, A. S., 1966.Vertebrate paleontology, 3rd ed., 468 pp. Chicago: Univ. of Chicago Press.
SCHLOSSER, M., 1923. Mammalia. In K. A. v. Zittel, F. Broili & M. Schlosser, Grundzuge der
Palaontologie (Palaozoologie). vierte Auflage, zweite Abteilung, Vertebrata: pp. 402-689. Munich &
Berlin: R. Oldenbourg.
SCOTT, W. B., 1903 [-19041.Mammalia of the Santa Cruz Beds. Part 1. Edentata. R e p . Princeton Univ.
Exped. Patagonia, 5(1): 1-364.
SHRIVASTAVA, R. K., 1963. Deltoid musculature of the Edentata, Pholidota, and Tubulidentata.
Okajimas Folia anat. jap., 3 8 : 25-38.
SIMPSON, G. G., 1931.Meracheiromys and t h e Edentata. Bull. A m . Mus. nut. Hist., 59: 295-381.
SIMPSON, G. G., 1945. The principles of classification and a classification of mammals. Bull. A m . Mus.
nut. Hbt.. 85: 1-350.
SYCH, L., 1966.Czy przodkami zajaca byly ssaki kopytne? Were the ungulated mammals ancestors of the
hare? Przegl. zool.. 10: 65-71.
SZALAY, F. S. & McKENNA, M. C., 1971. Beginning of the age of mammals in Asia: the late Paleocene
Gashato fauna, Mongolia. Bull. A m . M w . nut. Hist., 144: 274-317.
THOMAS, O.,1887.On t h e 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., 1 8 8 7 8 :
443-62.
VALENTINE, J . W., 1973.Coelomate superphyla. Systematic Zool.. 22: 97-102.
VAN VALEN, L., 1963. The origin and status of the mammalian order Tillodontia. J. Mammal., 44:
364-73.
VAN VALEN, L., 1964.A possible ongin for rabbits. Evolution, 18: 484-91.
VAN VALEN, L., 1966.Deltatheridia, a new order of mammals. Bull. A m . Mus. nat. H b t . 132: 1-126.
VAN VALEN, L., 1971.Adaptive zones and the orders of mammals. Evolution, 25: 420-8.
WEBER, MAX, 1891. Beitrige zur Anatomie und Entwicklung des genus Man&. Zoologische Ergebnisre
einer Reise in Niederlandisch Ost-Indien, 2: 1-116.Leiden: E.J. Brill.
WEBER. MAX, 1904. Die Saugetiere. Einfuhrung in die Anatomie und Systematik der recenten und
fossilen Mammalia, 866 pp. Jena: G. Fischer.
WEBER. MAX, 1927-1928.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., 1961.Revision o f the uintatheres. Bull. Peabody Mus. nut. Hbr., 14: 1-93.
WOOD, A. E., 1957.What, if anything, is a rabbit? Evolution, 11: 417-25.
WOOD, A. E., 1962. The early Tertiary rodents of the family Paramyidae. 7 h n s A m . Phil. Soc., 52:
3-261.
WORTMAN, J . L., 1896. Psittacotherium. a member of a new and primitive suborder of the Edentata.
Bull. A m . Mus. nat. Hist.. 8 : 259-62.
WORTMAN, J . L., 1897. The Ganodonta and their relationship to the Edentata. Bull. A m . Mus. nut.
Hisr., 9: 59-110.
ZITTEL, K. A. v., 1839.Handbuch der Palaontologie. Abteilung I. Palaozoologie. Band I V, Vertebrara
(Mammalia): 799 pp. Munich: R. Oldenbourg.
11
E . C. MINKOFF
156
APPENDIX
Synopsis of four influential classifications
(Extinct taxa are indicated by an asterisk)
Blainville (1839-1864)
Monodelphes [Placental mammals]
Maldentb [Edentatal
Bien dentis
Primatis [Primates]
Secundatks [Carnivores and insectivores]
Tertiatks [Rodents]
Quaternatks [Ungulates]
Gravigrades [Proboscidea and Sirenia]
On logrades
f d o i g t s impairs [Perissodactyla and Hyracoideal
A doigts pairs [Artiodactyla]
Gill (1870)
Subclass Ornithodelphia
Order Monotremata
Subclass 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
Gregory ( 1910)
Class Mammalia
*?Subclass Promammalia
Subclass Prototheria
Order Monotremata
MAMMALIAN SUPERORDERS
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
Simpson (1945)
Class Mammalia
Subclass Prototheria
Order Monotremata
*Subclass Allotheria
*Order Multituberculata
*Subclass uncertain
*Order Triconodonta
Subclass Theria
* Infraclass Pantotheria
*Order Pantotheria
11.
157
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