Taxonomy'and Biostratigraphy of the Early Tertiary Taeniodonta

Taxonomy'and biostratigraphy of the early I Tertiary Taeniodonta (Mammalia: Eutheria)

R. M. SCHOCH Department of Geology and Geophysics and Peabody Museum of Natural History, Yale University. New Haven, Connecticut 0651 I

The Geological Society of America Bulletin, Pirt 11, v. 92, 1982-2267, 17 figs., 12 tables, 61 plates, December, 1981, Doc. no. M11205

Thus, the San Juan basin mammalian INTRODUCTION I faunas provide 'a sequence of faunas The Taeniodonta is an order of archaic which can provide the basis for inter-

mammals known exclusively from the lower basinal correlation. The Taeniodonta

Tertiary of western North America are animportant element of these'

(Schoch and Lucas, 1981a). Their re- faunas; many o# the type spccimeyts and

mains are found in Puercan (lowCr Paleo- important referred specimens of these

cene) to Uintan (upper Eocene) strata animals come from the San Juan basin.

of the Rocky Mountain intermontane However, except for Pa?terson's (1949b)

sedimentary basins (Figs. 13, 14; brief review, a comprehensive study of

Table 12). In the San Juan basin of this order has not been published for

Hpw Mexico and Colorado, the Puercan to more than 80 years (Wortman, 1897b).

Wasacchian (lower EL-cene) sedimentary Before a group of animals can be used,

sequence includes the type localities bibstratigraphic correlation, its

for the Puercan,.. Torrejonian (middle species- and genus-level taxonomy, and Paleoc'ene), and Tiffanian (late Paleo- its known stratigraphic occurrences

cene) land mammal - "ages" (Wood and must be well established. .In this paper,

others, 1941) as well as a classic I revise the species'level taxonomy of

Wasatchian fauna (Lucas and others, 1981). the Taeniodonta and document their 1982

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geographic and temporal (stratigraphic) specifically or generically indetermi-

distribution and I discuss their use- nate. In the figures and tables, I

fulness and implications for biostrati- illustrate and present dental measure-

graphic correlation in western North ments for each t-axon, both for use

America. as a practical guide to ideutifying

other taeniodont specimens and as a SCOPE AND METHODS source of data to serve as the basis

In this paper, I undertake a complete for further research. In the bibliog-

taxonomic revision at the genus- and raphy, I present references not only to

species-level of the Taeniodonta based works.mentioned in the present paper

on the majority of the known specimens. but also to other works? which discuss

The suprageneric classification and taeniodonts.

phylogeny of the taeniodonts is not ABBREVIATIONS USED considered here. Taxonomic judgments are

made solely on the basis of morphology; AC Amherst College, Amherst ,,Massa-

edtrinsic stratigraphic and geographic chuse t t s

data, although recorded and considered AMNH American Museum of Natural History,

important, are not uskd to define taxa. New York

Referred specimens listed for each BAWSM ,Bayerischen Akademie der Wis-

taxon indicate the distribution (strati- senschaften Sammlung Mhchen,

graphic/geographic) of the taxon. How- Mun-ich

ever, these lists are not meant to be BNM Basel Naturhistorischen Museum,

comprehensive,.> only representative. Basel Several other taeniodont specimens exist CM Carnegie Museum, Pittsburgh 4 which are not specifically mentioned or DNHM Dinosaur Natural History Museum,

illustrated in this work; however, many Vernal, Utah

of these remaining specimens are FMNH Field Museum of Natural History,

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Chicago L Length

IVPP Institute of Vertebrate Paleon- W Width

tology and Paleoanthropology, Tr Trigohid

Beij ing Td Talonid

MCZ Museum of Comparative Zoology, * Asterisks indicate approximate

Harvard University , Cambridge, measurements of damaged 0-rworn

Massachusetts teeth or measurements or alveoli.

MPM Milwaukee Public Museum, Milwaukee All dental measqements are given in mm.

RU Princ; ton University , Princeton Tooth nomenclaturq fol'lows Szalay (1969,

Texas Memorial Museum, Austin p. 198Y03, Table 1, Fig. 1). UALP University of Arizona Laboratory SYSTEWTIC PALEONTOLOGY of Paleontology, Tucson

UCMP University of California Museum Order Taeniodonta Cope, 1876a of Paleontology, Berkeley EquaZs or incZudes:

UK University of Kansas, Lawrence Stylinodontidae ,Marsh, 1875b, p. 221.

UM University of Minnesota Taeniodonta Cope, 187Ga, p. 39.

Minneapolis Ectoganidae Cope, 1876a, p. 39.

UNM University of New Mexico Calamondonfidae Cope, 1876, p. 39.

Albuquerque Hemiganidae Cope, 1888d, p. 310.

USGS U.S. Geological Survey, Paleon- Ganodonta Wortman, 1896a, p. 259.

tology and Stratigraphy Branch, Conoryctidae Wortman, 1896a, p. 260.

Denver Stylinodontia Marsh, 1897, p. 137.

USNM National Museum of Natural Conoryctinae Schlossei, 1911, p. 414.

History, Washington, D.C. Stylinodontinae Schlosser, 1911, p. 414.

uw University of Wyoming, Laramie Onychodectini Winge, 191.7, 'p. 105.

YPM Peabody Museum of Natural History, Conoryctini Winge, 1917, p.. 105.

Yale University, New Haven Stylinodontini Winge, 1917, p. 106.

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Psittacotherinae Matthew, 1937, p. 237. trigonids and talorids of all molars

Taeniodontidae Szalay, 1977, p. 368. subequal in size (length and width);

Inc Zuded genera: Onychodectes Cope, trigonids bear subequal protoconids

1888d; ConorycteZZa Gazin, 1939; Conorycte and metaconids; molars decrease in size

Cope, 1881a (= llexoion Cope, 1884a); posteriorly, hypoconulid/talonid not

Huerfanodon Schoch and Lucas, 1981b; expanded on Mg. Taeniodonts can also

Vortmania Hay, 1899; Psittacotheriwn Cope, be distinguished by a tendency toward

1882b (= Hemiganus Cope, 1882~); Ectoganus hypsodbnt cheek teeth. In relatively

Cope, 1874 (= CaZamodon Cope, 1874 = "primitive" ffxms (Onychodectes,

Dryp,todon Marsh, l876b = Coiiicodon Cope, Conoryctella, Conoryctes, Huerfanodon

1894 = Lampaddphoms Patterson, 1949a) ; and Nortmania) this take.s the form of

and StgZinodoh Marsh, 1874. "tooth-base" or crown" huposodonty

Distribution: Pucrcan (early Paleo- (White, 1959), characterized by the

cene) to Uintan (late Eocenej of western labial extension of the enamel on.

North America. the lower cheek teeth and the lingual

Discussion. I recognize \$theorder extension of the enamel on the upper

Taeniodonta as a monophyletic taxon whose cheek teeth. In more "advanced"

members share the following derived taeniodonts (Psittacotheriwn, Ectoganus,

characters: relatively narrow upper and SqjIZnodon) , this forh of hypsodonty

molars, with protocones, protoconules, and is combined with "root" hypsodonty

metaconules small and placed far lingually, (White, 1959), in which the roots of

paracones and metacones moderate-sized, the cheek teeth fuse and become ever-

punctate and placed far labiafly with re-. growing. Taeniodonts are also charac-

duced stylar shelves; pre- and postcingula terized by the possession of a

lacking on upper molars; hypocone absent leptictimorph astragalocalcaneal

or developed by a splitting off from the morphology (Szalay, 1977).

protocone; lower molars lack cingulids; Jacob Wortman (1896a, 1897b) first

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recgonized-theTaeniodonta in its modem Insectivora) composed of three tribes:

sense. Up until the work of Wortman, Onychodectini (Onychodectes),

different workers included varipus genera Conoryctini (Conoryctes), and'

in. the Taeniodonta. Several genera now Stylinodonitni ("Hemiganus" [=

considered taeniodonts, such as Onyclzo-' Nortmania] , Psittc zotherim, "Ca Zamodon"

gectes , Conoryc5es, Wortmania, and = Ectoganrisl , and Sty Zinodon) . Matthew 8 Psit$acotheriwn, often were placed in (1937) recognized the family Stylino-

the Tillodontia, Creodonta, or Insecti- dontinae (sole family of the order

vora (for example, Cope, 1882b, 1888d). Taeniodonta) composed of the Onychodec-

Wortman placed 'the taeniodonts in their tinae (dnychodectes)2 tonoryctinac

own suborder, Ganodonta, within the orpcr (Conoqctes), Psittacotherinae

Edentata, and included two families iA-the (Nortrnanic., Psittacotheriwn, and

"ganodonts": the Conoryctidae, composed "Ca lamodon" [= Ectogams 1 , and.-,S tyfino-

of Onychodectes and Conoryctek-, and the dontinae (StgZinodoiz). Simpson (1945)

Stylinodontidae, composed of "llemiganzis" : adopted Matthew s (1937) classj fication

(= Najotmaxia) , Fsitt-acJtheriwn "Ca Zamodon" of the Taeniodonta in its broad out-

(= k$oganiis), and StgZinodon. Wortman lines, but reduced the number of sub- .c. viewed the conoryctids and stylinodontids families to two. Thus, Simpson (1945)

as two evolving phyla forming a graded recognized one family of the order

ancestor-descendant series from one genus Taeniodonta, Stylinodontidac, composed

to the next., of the subfamilies Conoryctinae

Schlosser (1911) recognized the (Ongchodectes, ConocicteZ Za, and

Ganodonta.[sic] as a family of the Conomjctes) and Stylinodontin,ae (Nor&-,

Edentata and reduced Wortman's maina, Psittacotheriwn Ectoganus arid

Conoryctidae and Stylinodontidae to StyZinodon). The most recent student

subfamilies. Winge (1917, 1923) recog- of the order, Patterson (1949b),

nized the family Stylinodontidae (order adopted Simpson's (1945.) claS'sification

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of the Taeniodonta, adding his new genus Cope, 1888d (= Onychodectes rm

"Lampadophoms" (= Ectogcntus) to the Osborn and Earle, 1895).

Stylinodontinae. Included species. Only the type

It is not my purpose here to address species.

the spprageneric taxonomy of the Dist~butfon. Puercan of New

Taeniodonta; only the species- and Mexico and Utah.

genus-level taxonomy and the temporal Revised diagnosis. Small taeniodonts;

and geographic distribution of the teeth moderately hypsodont (relatively 4 taeniodonts are analyzed. In this less hypsodont than ConozycteZZa); P

paper, the taeniodont genera are listed nonmolariform with a well-developed

in order of increasing size, from protocone., paracone and incipient u smallest -to largest. A suprageneric metacone, parastyle, stylocone, and 4 analysis of the phylogeny and clas- metastyle; P meta-stylocone small

sification of the Taeniodonta will be to moderately well developed; lower

presented elsewhere. The term molar trigonids hear large, subequal,

"conozyctine" informally refers to the and sharply punctate protoconids and

relatiyFly small and generalized metaconids with only slightly'smaller

taeniodonts, Onychodectes , Conm'ycte Z la, lingually placed pazaconids; lower

Conoiyctes, and Huerfanodon; likewise, molar'talonids bear high and punctate

'Is t y linodon t inel' inf orma 11y refers to hypoconids, slightly smaller and

the larger and more specialized taenio- punctate entoconids, and small hypoconu-

donts, ldortmmin Psittncotheriwn, 1.1ds.

Ectoganus , and StyZinodon. Onychodectes tisonensis Cope, 1888d

Onychodectes Cope, 1, ?8d Onychodectes tisonensis Cope, 1a88d,

Onychodectes Cope, 1888d, p. 317. p. 318.

Type species. Onychodectes tisonensis (see synonymies' under the subspecies)

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Type subspecies. Oncyhodectes tiso- Qpe Specimen. AMNH 3405, right 43 nensis tisonensis Cope, 1888d. and Left maxillae with P -M , left

Included subspecies. The type sub- dentary with M2 and associated right

species and Onychodectes tisonensfs rmus astragalus (Pl. 4, Figs, 1-5; il-

Osborn and Earle, 1895. lustrated by Cope, 1888d; P1. 5, Figs.

Eagnosis. Same as *that for the genus. 8, 9). Several'different individuals

of 0. t. tisonensis have been cata- Onychodectes tisonensis tisonensis Cope, 1888d logued under.AMNH 3405; any of these

(Tables 1; Figs. 1, 2; Pls. 1, 2, 3, specimens that can be demonstrated tb

4, Figs. 1-5, 9, 10; P1. 5, Figs. 1- be from the same individualas Cope's

4, 6-;9, 13-15; PI.. 6, Figs. 7-9; Pls. illustrated specimens may be >regarded

7, 8, Figs. 3-8, 11, 12; P1. 9, Figs. as part of-the type specimen (see

11, 12, 15-18, 21-24) Cope, 1888d, p. 318).

Onychodectes tisonensis Cope, 1888d. p . 318. Horizon and ZocaZity of the tgpe.

Onphodectes tisonensis (Zapsiis ca Zami) : Collected by David Baldwin in 1885

Osborn and'Earle, 1895, p. 40. from presumably Puercan-strata of the

hzychodectes tisonensis (Zqsus caZami): Nacimiento'Formation, San Juan basin,

Wortman, 1897b, p. 97. New Mexico.

S. tisonensis lapsits cahi: Wortman, Referred speicmens. AW 902a, 72 1897b, p. 97. upper molar (M' ); AMNH 3405 (not the

Onychodectes tisonensis: Matthew, 1937, type specimen, see above), left maxilla 42 .p. 239. wik P -M (Pl. 5, Fig. 3); ANNH 3407, Onychodectes n. sp.?: Robison and Lucas, right and left dentary fragments with

1980, p. 302. left P1-2, right and left P3-M1, right

M2- 3 9 roots of left C1, right Ple2,

alveoli for left M2-3; ILW 3408,

left dentary fragment with P2(?), M1-3;

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P4 M2 M1 M3 p1 p2 L v L w &L .w 1 W 1 W 1 W AMNH 785 4.8* 5.9* 5.3 7.0 4.7 5.4 786 812 822a 902a 5.3 5.6 824' 3040a' 6.9 8.1 '3405a 5.2 5.1 6.5 6.8 6,O 6.9 4.3 6.4 3405 4.5 5.0 6.0 6.6 5.6 6.5 3406 5.6 7.4 3407 2.7* 1.9" 4.2* 2.6* 3408 3.7 2.5* 3409 3411 4.5* 5.2 5.7* 7.0 5.1* 6.7 3576a 16405 5.7 6.3 ,7.1 8.4 6.2 8.1 4.3 3.2-' 16406-.. 4.4 '2.6 16407 16408 16409 3f.t 16410 3.9 2.9 16411 5.0 5.5 16528 5.0 6.0 6.0 61.9 5.3 6.7 4.1 5.1 2.4 2.1 4.J' 2.8 -7. 23090 5.0 5.1 7.1 7.8 6.3 7.6 27608 5.8 7.0 27678 27679 6.1* 7.1 6.2* 6.9 36070 6.6 7.5 36071 5.0 5.1

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p3 p4 M1 M2 M3 L W L W L W W L W W 1 W W Tri Td Tri Td Tri Td

5.1 3.4 5.5 4.0 6.7 5.4 5.0 7.4 5.5 4!6 6~05.7 3.8 6.4 5.1 4.9' 6.4 4.9 4.9 5.4 3.3 7.8* 5.1 6.4 5.2 4.7 6.5 4.7

7.8 4.6 4.7 6.9 5.0 5.0

6.5 4.7 4.G

.. 4..9 3.3 5.4 3.9 6.1 4.6 4.6 4.9 6.0 4.5 3.7 6.7 4.6 6.1 4.8 5.9. 4.4 5.2* ..3.1, 6.1 4.8 4.4 2.9 5.1*- 3.3 6.5 4.6 4.6 5.9 4.8 4.8 5.7 4.1 3.7

5.4 3.1 'r . 5.5 4.6 4.7 5.4 4.3 3.7 . ,.. . '.. 5.1 3.8 :6.0 4.9 7t247.15.5, 5.9 6.7 5.5 5.5 5.9 4.6 4.1 - ..c . 4.9 3.3 5.5* 3.9 7.8 'if0 5.5 3.3 - 5.8 3.7 6.4* 4.9" 6.6 4.6* 4'.7 6.4 5.0 4.6

' 5.9* 4.3 6j5 4.8 5.8 5.2 3.1 5.5 3.7 6.8. 5.1 '4.6 4.8 2.8 6.0 4.5 4.1 5;2 3.9 4.7 3.1 5.2 3.2 5,9 4.6 4.3 5.4 4.3 4.1 5.4 3.5 3.2

5.4 3.3 '7.8* 5.1 6.4 5.2 4.7

6.6 4.6 4.7 .-6.3 4.8 4.7

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58059a 58172 UCMP no number 5.1* 5.9.* 1.1 8.0 5.9* 8.1* 31293 5.6* 5.4* 6.1* .6.9* 31817 4.1 3.0 31819 365U 4.8* 5.6 6.3 6.6 5;5 6..7' 4.5 5.6 68668 68687 5.2 5.3 74792 .5.2 7.2 89695 5.9 7.0 92156 UK 8114 6.5 7.5 8116 5.1 5.5 9416 941;

12711 13000 USNM 15534 15535 15536. 4.9 .5.4 5.9* 6.8

N 15 15 . 11 11 18 18 4 4 7' 7 Mean 5.03 5.59 6.35 7.17 5.84 7.19 4.40 5.63 4.iO 2.80 Standard Dev. .33 .39 .52 .61 .53 .53 .26 .56 .24 .25 Coefficient of Variation 6.6 7.1 8.2 ,8.5 9.1 7.4 5.9 9.9 5.9 8.9

a Type specimen of Onychodectes t. tisonesis.

bType specimen of 2. t. rarus.

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6.7 4.9 4.3 5.7 4.4 3.9. 5.8* 4.2* 6.8 4.9 4.6 6.7 5.0 4.7 6.2 4.2 3.5*

5.0 3.3 5.8 &. 5.3 h.6 6.1 5.3 5.8 5.1

5.2 3.6

6.1 5 .o 4.5 6.1 L4 3.7

6.5: 4.7 4.6 5.9 4.9 4.8 d 6.7 5.2 4.9

6.4 4.6* 4.5 6,O 4.8 4.4 6.1 4.0 3.4* 6.1 4.1 7.9 5.2 5.2 4.9 3.3 5.4 3.8 6.3 5.1 4.6 5.7* 4.3" 5.0 3.1 6.7 5.2 4.6 6.3 5.2, 4.4 5.9 4.6* 3.5*

12 12 19 18 24 24 16 23 24 20 15 13 13 4.96 3.23 5.52 3.72.. 6.80 4.92 4.78 6.29 4.93 4.63 5.80 4.31 3.68 .27 .26 .29 .48 .59 .29 .37 .45- .28 .32 .29 .31 .24 5.4 8.0 5.3 .12.9 8.7 5.9 7.7 7.2 5.7 6.9 5.0 7.2 6.5

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AMNH 3409, left dentary fragment with two isolated lower molars; UCMP 92156, 4 P4-M1; AMNH 3411, lef; maxilla with P - right dentary fragment with M2-3 and 2 M (P1.'5, Fig. 1) and right dentary roots of M1; UCMP no number, left 32 fragment with P3-M3 (P1. 5, Figs. 14,. 16, maxilla with P -M , left dentary frag-

17) and postcranial fragments; UCMP ment with roots of P 3-A and complete 36514, right maxilla fragment with M1-3 .and edentulous right dentary 43 P -M (Pl. 4, Figs. 9, 19); all from fragment; UK 8114, left maxilla with 2 4 presumably Puercan strata of the M 1 UK 8116, left P (PI. 5, Figj 7);

Nacimiento Formation, San Juan basin 7 UK 9417, left MI; UK 12711, left New Mexico. dentary fragment with M1-3; USNM 15535,

AMNH 785, skull with left.P4, M~-~, right dentary' fragments with P 3' Mi-3; 41 a,"d &ght Pi2-3, alveoli €or right and USNM 15536, right maxilla with P -M

left 13, C1, right P4, right and (Pl. 5, Fig. 4): all from Puercan 1 left M , left dentary fragment with strata of the Nacimiento Formation,

M2-3, and right dentary fragment with De-na-zin wash, San Juan baqin, New

roots of P2 and complete P3-M3 (Pls. Mexico:

3; 6, Figs.. 7, 8); AMNH 786, right Ml 16406, left dimtary fragment

dentary 'fragment with M1-2, roots of with P2-M1, alveoli for\P1, M2-3;

Cl1 P2-4y alveolus for P1; AMNH 812, AMNH 16408, right dentary fragment with

riiht dentary fragment with M,, alveoli M1-2 and alveoli for C1-P4 and M3' L I for P4-M1; AMNH 822a, right dentary (Pl. 6, Fig. 9); AMNH 16409, left

fragment with M2; AMNH 27678, left dentary fragment with P4-M1, M3;

dentary fragment with P4-M2; UCMF' 31293, AMNH 16410, left dentary fragment with '4 1 3 left maxilla with P,-M and partial P , P2-M1, alveoli for C1-P, , both humeri, 2 M ; UCMP 31817, left dentary with P2-4; proximal portion of Left uha, sacrum,

UW31819, right dentary with R4-M2 and radius, and other postcranial frag- 4' alveolus for M3; UCNP 68687, left P and ments (Pls. 5, Fig. 11; 7, Figs. 2-5;

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8, Figs. 11; 12; 9, Figs. 21-24); USMN basin, New Mexico.

15534, left dentary fragments with P3-4, AMNH 36070, left M2 (Pl. 5, Fig7 0; 4 and roots of P1-2 (Pl. 5, Figs. 15, AMNH 36071, left P : both from Puercan M2- 3 ' -. 18, 19): all from Puercan strata of the strata of the North Horn Formation,

Nacimiento Formation, Alamo wash, San Wagonroad local fauna, Emery County,

Juan basin, New Mexico. Utah.

AMNH 16411, isolated teeth and tooth Revised diagnosis. Subspecies of

fragments including two incisors, ,right Onychodectes tisonensis with relatively 4 4 ..- P , right dP (?), left P3, right and simpler crowned premolars and molars

left M2, right M and postcranial than in 0. rarus; anterior internal 3 t. fragment9 (Pl. 5, Figs. 8, 9); AMNH accessory cusp absent on P4; external

16528, skull and lower jaws wi,th right accessory cusp between trigonid and .. 12(?), left Cl-Pl, right and left P2-M3, talonid lobes of M1-2 absent.

roots of right and left I2 and C1, right D Otiychodectes tisonens-% rarm and left P1, left P2, fight and left Osborn and Earle, 1895

P3-M3, right manus, left pes, and (Table 1: Pls. 14, Figs. 6-8, 11, miscelfaneous vertebrae (Pls- 1; 2; 7, 12; P1. 5, Figs. 5, 10-12; P1. 6, Figs.. 6, 7; 9, Figs,. 11, 12, 15-18); 1 Figs. 1-6; P1. 8, Figs. 9, 10; P1. 9, AMNH 58059a;right M2(?): all from Figs. 1-8, 13, 14, 19, 20) P.uercan strata of the Nacimiento Forma-

tion, Kimbeto wash, San Juan basin, Onychodectes Tarus Osborn and Earle,

New Mexico. 1895, p. 42.

AMNH 27608, (?)left M2; AMNH 58172, Onijchodectes rhrus: Wortman, 1897b, p. 97. 2(?). right M3(?); U.CMP 74792, left M , hychodectes rarus: Matthew, 1937, p. 249.

UCMF' 89695, left M2(?): all from Type specimen. AMNH 824, left dentary

PuerGan S'trata of the Nacimiento Forma- fragment with Pi1- (p1. 4,.Figs. 6-8).

tion, Betonni'e Tsosie wash, San Juan Horizopi and locality, of the type.

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Puercan strata of the Nacimiento Forma- 11; 12; 6, Figs. 1-6); AMNI1 16407,

tion, De-na-zin wash, San Juan basin, left dentary fragment with, P3+ and

New Mexico. anterior portion of M1, roots of P2; 42 Referred specimens. AMNH 3040a, AMNH 23090, left maxilla with P -M : 2 (?)right maxilla with M and partial all from Puercan strata of the Naci- 1 3 roots of M and M ; AMNH 3576a, isolated miento Foyation, Alamo wash, San Juan

righf'and left M3, right dentary fragments basin, New Mexico.

with P4,* M2, scapula, vertebrae, and Revised diagnosis. Subspecies of

other postcranial fragments (Pls. 8, Onychodectes tisoneTsis with the

Figs. 9, lo; 9, Figs. 1-8, 13, 14, 19, following characters moderately to

20): both from presumably Puercan well developed (as compared to 0. t. 43 strata of the Nacimiento Formation, rams): P -M slightly broader and

San Juan basin, New Mexico. with better developed ectocingula; lower I. AMXH 27679, right maxilla wi$ M1-' premolars slightly more molariform with

and left dentary fragment with M 1-2; UCMp better developed talonids on P 3-4 and 68668, right dentary fragment with P4; an anterior internal accessory cusp

UK 9416, right M1, left M2 (Pl. 5, Figs. on P4, M1-2 with external accessory

10-12), and dentary fragments; UK 13000, cusp or CUSDS between the trigonid and

left dentary fragment with P4-M1, alveoli talonid lobes.

for P2-3: all from Puercan strata of ?Onychodectes sp. the Nacimiento Formation, De-na-zin (Pls. 8, Figs. 1, 2; P1. 9, Figs. wash, San Juan basin, New Mekico. 9, 10) 32 AMNH 16405,-left maxilla with P -M 3 and roots of M , right dentary with Referred specimen. AMNH 3404, (?)

canine stub, parts of P1-2 and complete right illium (Pl. 8, Figs.. 1, 2), lumbar

P4-M3, left dentary with canine, root vertebra (PI. 9, Figs. 9, 101, and

of P1, and complete P2-M3 (Pls. 4, Figs. other bone fragments from presumably

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Puercan strata of the Nacimiento Forma- C -M (Pls. 4, Figs. 11, 12; 6, Figs. 13 eion, San Juan basin, New Mexico. 1-6) referred to Onychodectes "rarus"

Discussion. This. specimen was col- by Matthew (1937, p. 249), bears this

lected by David Baldwin for E. D. Cope external cusp on M1-2, but not on M3,

in 1885 from the San Juan basin, New and also possesses a small anterior 42 Mexico. According to the AMNH label, internal accessory cusp on P4. P -M

it orfj-~lnallymay have .included an upper of AMNH 16405 are somewhat broader and ,- jaw fragment with molars identifiable have better developed ectocingula than

as Onychodectes; however, I have not some specimens of 0. tisonemis (Matthew,

bFen able to locate this maxilla. While 1937). However, these are all rather

several individuals (some nontaeniodont) minor and vari'ablc features in

are catalogued under AMNH 3404, a (?)right Onychodectes; wien a number of specimens

ilium and lumbar vertebra are in the arc lined up side by side there is a

size range of Onphodectes and may be continuous gradation from one extreme

referable to this genus. to the other. Thus, most specimens (for

example, USNM 15534, AMNH 3411, AMNH Description and Discussion of Onphodectes 16410; P1. 5, Figs. 13-19) lack an

I consider Onychodectes ttrarus'' external cusp on the lower molars and

Osborn and Earle, 1891, to be a junior have at most an incipient anterolingual

subjective synonym of Onychodectes cusp on P4. AMNH 27679 possesses a very

6sonensis Cope, 1888d, at the specific small, incipient external cusp on M1;

level. Ongchodectes "rams': is based UK 9416 (Pl. 5, Figs. 10-12) .possesses

on AMNH 824, the diagnostic feature adsmall external cusp on M.l; AMNH 3576a

being an external cusp between the and UK 13000 both have a small external

trigonid and talonid lobes of M1-2. cusp on the talonid of P4 and a small 3 AMNH 16405, a left maxilla with.P -M2 anterolingual accessory cusp on P 4; and both lower dentaries with incomplete and AMNH 3576a has two small externsl

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cusps between,f he trionid and talonid although there is the possibility that lobes of PI2. Other specimens which bear only two upper incisors.were present

the characters attributed to 0. ''rams'' (compare Matthew, 1937, p. 240). The

by Matthew (1937) in varying degrees of incisors are of moderate size with I 3

development are: AMNH 3040a, AMNH 16407, and I2 slightly enlarged. . AMNH 23090, AMNH 27579, and UCMF' 68668. The canines of Oqchodectes are

Likewise, the size of both the upper sharply pointed and of moderate size. 1 and lower molars of specimens of P are small, simple, pointed teeth and I 1 2 Onyclwdectes is somewhat variabl; are single-rooted. P is two-rooted and

(Table l), but there are no clehr gaps oval jn cross section, being compressed

and the majority of coefficients of vari- transversely and elongated anteroposteri- 3 ation range from approximately five to orly. P is triangular in cross section

nine. Much of this size variation may be and bears a large paracone labially and

hue to the extremely worn. condition of a moderate protocone lingually. 4 these teeth and the consequent diffi- P (Pl. 4, Figs. 7, 8) is roughly

culty in measuring homologous points on triangular in cross section and labially

diffbrent specimens. Although more speci- bears a large oval paracone elongated

mens in a better state of preservation anteroposteriorly with at most an

might demonstrate that Onychodectes is incipient mctacone on its posterior 4 composed of more than one species, it is face. Labially, P bears a very slight

most reasbnable to assign all specimens ectocingulum,aminute parastyle with a

to one species at the present time. The slight stylocone, and small metastyle

variable, but recognizable, morphological with a small metastylocone. kingually, 4 differences are here relegated to sub- P bears a moderate-sized protocone

specific status. flanked by pre- and postejngula.

The probable dental formula of The,precingulum connects the anterior 3143 Onychodectes is I3 C1 P4 M3 (Fig. 11, base of the protocone to the parastyle

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, I I cm

Figure 1. Restoration of the skull, mandible and dentition

bf Onychodectes tisonensis tisonensis, based primarily on

MlNH 16528 and MlNH 785. -(a) Left lateral view of skull.

(b) Left lateral view of mandible. (c) Occlusal view of upper d left dentiFion. (d) Occlusal view of lower right dentition.

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and bears and bears a small pericone.,r\The Anteriorly, P2-4 bear large, high,

postcingulum connects the poster- base anteropostcriorly elongated, and

of the protocone to the metastyle and laterally compressed protoconids w6ich '.. bears a small ''metaconule" flanked by a are sharp and slikhtly recurved

smaller "hypocone" labially (the homolo- .posteriorly. Posteriorly, P2-4 bear

gies of these cusps are not certain). small talonid heels which increase in

M1-3 (Pls. 4, Figs. 1, 9-12; 5, Figs. relative size from P2 to P4.

1-6) decrease in size posteriorly, but M1-3 (Pls. 2; 5, Figs. 10-19; 6) de-

otherwise are of similar morpholoy. These crease in size posteriorly, but other-

teeth bear moderate-sized, sdequal, wise are of similar morphology. The

conical, punctate, labially appressed, and trigonids and talonids are subequal in

lingually inclined paracones and meta- size, subcircular in cross section and

cones. Labially, M1-3 bear well-defined not particularly Compressed anteroposteri-

and minutely cuspidate parastyles, ecto- orly. The trigonids bear large, sub-

cingula, and metastyles. Their sfylar equal, and sharply punctate protonids 0 shelves are at most extremely narrow. and metaconids with o~lyslightly smaller

The trigon basins are shallow. M1-3 bear paraconids. The talonids bear high

moderately small and poorly defined and punctate hypoconids, slightly

protocones on the lingual edge of the smaller and punctate entoconids, and

teeth that are slightly recurved labially. smaller hypoconulids. Minute ecto-

Small protoconules (paraconules) and conulids and/or mesoconids may be

metaconule's occur relatively labially variably present; one or two small to

along the far anterior and posterior ihnute external cuspids are .sometimes 1- 3 edges of M . present on the anterior portion of the

(Pls. 2; 5, Figs. 13-19) are talonid lobe. "2-4 all simple, nonmolariform, double-rooted All of the cheek teeth are moderately

teeth which ipcrease in size posteriorly. hypsodont and show the characteristic

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taeniodont trait of "rolling eruption'' hypsodont (the enamel extends far down

(Patterson, 1949b) with enameLextending the lingpal face of the tooth) but is

far lingually on the upper cheek teeth very shallow labially.

and far labially on the lower cheek teeth. The skull of Ongchodectes (Fig. 1;

Associated with AMNH 16411 is a Pls. 1; 3; 7, Fig. 1) is known from

molariform tooth (Pl. 5, Fig. 9) which MlNH .785 and AMNH 16528 and has been

because of its enamel color (it is described and figured by .Matthew (1937,

slightly lighter in color than the othkr p. 241-243; Fig.'58, PI. 58, Figs. 3, 4)

teeth associated with it, which are and Wortman (1897b, p. 97-99, Figs. 31-

.presumably of the same individual) and 33). The skull is long and harrow, with

its dimensions (relatively small &nd relatively Gxge premaxillae, long

transversely widened; length is 4.7 mm, nasals expanded posteriorly, and reduced

width is 6.5 mm) appears to be a,right and slender zygomatic arches. It lacks 4 dP of Onychodectes. This tooth bears a postorbital processes and possesses a

large conical paracone, a slightly weak sagittal crest. The lower jaw

smaller metacone positioned well is long and of moderate depth with a

labially, a narrow stylar shelf, slight long, weak, and shallow symphysis. The

ectocingulum, medium-sized and broad ascending ramus is broad and flat with

(but broken) parastyle, small meso- the last molar forward of its anterior

style, and a shallow ectofltxus. The border. The condyle is at or slightly

trigon basin is relatively shallow and above the level of the tooth row.

bears a poorly defined protocone lingual- The postcranial skeleton of

ly, and incipient paraconule, pre- and Onycltodectes (Fig. 2) is known from

postparaconule wings, an incipient meta- portions of the forelimbs and hindlimbs

conule, and pre- and postmetaconule wings. and some vertebrae (Pls. 7, Figs. 2-7;

This tooth lacks any cingula, as do other 8; 9) which have been described in detail

teeth of taeniodonts. It is very by Matthew (1937, p. 243-248, Figs. 59, 60)

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Figure 2. Reconstructed skeleton and life restoration of Onychodectes.tisonensis

based on specimens described and illustrated in the text and plates. Top: left

lateral view of the reconstructed skeleton. Middle: dorsal view of the recon-

structed skeleton. Bottom: life restoration.

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ConomjcteZZa Gazin, 1939 taeniodonts; P4 submolarifoh with a

CQnorycteZia Gazin, 1939, p. -276. single protoconid anteriorly and a

Conoryctes: Taylor, 1981, p. 250. moderately developed talonid posterior-

Type species. Coizomjcte Z la dragonen- ly; lower molar paraconids relatively

sb--%azin, 1939. large.

IncZuded species. The type species CongrzjctcZZa dragonensis Gazin, 1939 and Conorycte Z Za pattersoni Schoch and (Table 2: P1. 10, Figs. 5, 6) Lucas, 1981c.

Distribution. Torrejonian (including ConoryteZZa dragonensis Gazin 1939, p. 276.

the "Dragonian") of New Mexico and Utah. Conoryetella dragonensis: Gazin, 1941,

Revised diagnosis. Small taeniodonts, p. 15 (in part).

larger than Onychodectes but smaller Conorycte Z Za dragonensis : Schoch and

than Conoryctesi; teeth-more hypsodont Lucas, 1981c.

than Onychodectes but less hypsodont Type and onZy -known specimen. USW '4 42 than Conoryctes; P nonmolariform with 15704, left maxilla with damaged P -2.1

wcll-developed pro2ocone and paracone, and part of P3 alveolus (PI,. 10, Figs.

but only an'-incipient metacone; stylocone 5, 6).

and parastyle absent on P4 or only Horizon and ZocaZity of the type.

slightly developed (in contrast to Dragon local fauna, Torrejonian strata

Onychodectes and Conoryctes, in which of the upper part of the North Horn

they usually are moderately well Formation, NW1/4, sec. 8, T. 19 S., R.

develop'gd) ; symphysis of .lower jaw 6 E., Emery County, Utah.

unfused; lower canine relatively large, Revised diagnosis. Largest known

heavily invested with enamel, tri- species of ConorycteZZa; differs from

angular in cross section, transversely C. pattersoni in the following

compressed, and tending toward the features: P4 with sligh& better

yooCkss condition seen in advanced developed metacone and inicipient

USNM AMNH UNM USNM USNM USNM UK LTK UK UALP

15704a 3412 B-1258b 15722 16173 18358 7807 7888 9562 i1661 1 c -L 9.8* c1 w 5.6* P4 L 7.4 7 &%L. 5.8 P4 w 10.1 8.1” 7.9 M1 L 8.1 8.3* 7.2 7.4 M1 W 11.1 9.8* 8.6 9.2, M2 L 7.4 6.8 6.0 M2 W 8.9 8.0 7.7 M3 L 5.7 M3 W 6.4 5.5* 4.9 c1 5.2 p3 3.7 p3 6.9 p4 4.9 p4 El1 L 8.7 8.0* 7.7 5.8 5.6* 6.4 5.3. M1 8.0 7.8 7.2 7.4 M2 5.3 5.7 5.3 5.2 M2 6.9 6.2* M3 4.5 4.8 M3

a Type specimen of Conoryctella dragonensis.

bme specimen of c. pattersoni:’

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~ ~

stylocone; M1-2 relatively wide with Canyon, San Juan basin, New Mexico.

reduced stylar shelves, slight ecto- Refer~edspecimens. USNM 15722,

flexi, and small mesos tyles. right dentary fragment with M1 and

base and roots of P4; USNM 16173, ConorycteZZa pattersorti Sdhoch and Lucas, 1981c fragmentary and isolated MI and M2;

ConoryctsZZa drayonensis : Gazin, 1941, USNM 18538, right dentarv fragment

p. 15 (in part). with M3: all from Torrejonian strata

conoryctine, n. gen. and sp.: R. W. of the upper part of the North Horn

Wilson, 1956, p. 82. Formation, Dragon local fauna, NW1/4

n. gen. et sp.: Russell, 1967, p. 68. sec. 8, T. 19 S., R. 6E., Emery

Conoryctella: Schoch and Lucas, 1981a, County, Utah.

p. 225. UK 7807, right dentary fragment with

n. gen. and n. sp.: Taylar, 1981, M2 and roots of M3; lJK 7888, right

p. 259. dentary fragment with M2 and partial

n. gen. et sp.: Tscntas, 1981, p. 272. alveolus of M UK 9562, left maxilla 1; 2 Conorgctes coma: Taylor, 1981, p. 250. with partial M1, complete M , and

ConorycteZla pattersoni Schoch and partial M3 and right dentary fragment

Lucas, 1981 c. with complete M and roots of C1 and 1 lijpe specimen. UNM B-1258, palate P1-4; UALP 11661, left MI: all from 43 \yith right and left P -M and roots of Torrejonian strata of the Nacimiento

right and left P2-3, right dentary Formation, Jutz Canyon, San Juan basin,

with C1, P3-M3, and roots of I 1-3' New Mexico. incomplete right and left ulnae (Pl. Diagnosis. Smallest species of

10, Figs. 1-4, 7-9). Conorgetella; differs from C.

Horizon and locality of the type. dragonensis in the following features: 4 Torrejonian strata of the Nacimiento P lacks a parastyle, stylocone, and 4 Formation, UNM locality B-1096, Kutz metacone; P postprotocristid only

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~~~~

slightly developed; upper molars be further described or discussed here.

relatively narrow. Conoryctes Cqpe, 1881a

?Conoryetella cf. . C. dragonensis Conoryctes Cope, 1881a, p. 829.

Conoryctella cf.' C. dragonemis: Hexodon Cope, 1884a, p. 794.

Schoch and Lucas, 1981c. non Hexodon Olivier, 17b9, p. 1.

RefeweJ specinierz. MlNH 3412, a Type species. Conoryctes coma 4 poorly preserved left maxilla with P - Cope, 1881a (= Hexodon moZestus Cope, 3 M largely encrusted with ironstone 1884a).

concretion and a left upper canine: IncZuded species. Only the type

from Paleocene strata of uncertain species.

age, Nacimiento Formation, San Juan Distribution. Torrejonian of New

basin, New Mexico. Mexico.

Discussion. This specimen may Revised diagnosis. Medium-sized

represent Conorgctel la, but the teeth taeniodonts, about the size of

are so heavily encursted with an im- Huerfanodon; teeth relatively hypsodont

pregnable ironstone concretion that (crown hypsodonty); canines lack 3 their morphology is largely obscured and internal grooves; P nonmolarifom,

definitive taxonomic assignment thus is bears a simple large paracone and a

rendered difficult. It is described minute to small metacone afllingual 4 and illustrated in Schoch and Lucas cingulum; P molariform with a large

(1981~). protocone, smaller paracone, still

smaller metacone, and small parastyle, Description and Discussion of Conoiycte Z Za stylocone, metastyle, and metastylocone;

ConomjcteZZa (Fig. 3) fs thoroughly mesostyle absent on P4; d-3bear large

described, illustrated, aqd revised in protocones and smaller, conical,

Schoch and Lucas (1981~)and need not labially placed paracones and metacones;

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I 1 c.m

Figure 3. Restoration of the skull, mandible, and dentition

of ConorycteZZa patterson;, based primarily on UNM B-1258. The

skull is hypothetical. (a) Left lateral view of skull. (b) Left

lateral view of mandible. (c) Occlusal view of known upper dentition 43 3 (left P -M and alveolus for P ). (d) Occlusal view of lower right

dentition.

M1-3 mesos tyles variable : absent to Fig. 6).

moderately well developed; P sub- 4 Ilorizon and locality of the t;ype. ,molarifom, trigonid bears a single Collected by David Baldwin for E. D. Cope

large prqtoconid and simple talonid; from presumably 'Torrejonian strata of

lower molars with snall paraconids. the Nacimiento Fornation, San Juan

basin, New Mexico. Conoryctes coma Cope, 1881a Referred specimens. AMNH 3396, 3 (Table 3: Figs. 4: Pls. 11; palate with left C1, right P , Eight 43 14; 15, Figs; 14-17; 16, Figs. 1-6) qnd left P -2.I , mandible with left C;,

Conoryctes coma Cope, 1881a, p. 829. P4-M3, right C1, P3, M?, and Ma, roots

Conomjctes coma: Cope, 1881b, p. 486. of left 11-3, P3, and right 12,

Ilexodon molestus Cope, 1884a, p. 794. right proximal three-quarters of the

Conoryctes corn: Cope, 1884c, p. 198. humerus, and right distal end of the

Conoryctes mmma- (= Hexodon molestus) : radius (type specimen of "Hsxodon

Cope, 188d, p. ,317. molestus"; P1. 14, Figs. 1-6, 8, 10)-

Conoryctes coma: Wortman, 1897b, p. 101. AMNH 3397, left dentary frapmght with

Conoryctes coma: Matthew, 19.37, p. 250. P4-M?? AMNH 3398, parcial skull and

Conoryctes coma: R. W.Wilson,1956,p.82 mandible with four miscellaneous in- 1 non Conoryctes coma: Van Valen, 1978, cisors, left C , right P3-M3, frag-

p. 58. mentary right P1, left P4, M2-3: all-

F non Conoryctes coma: Taylor, 1981, p. collecte&& David Baldwin from pre-

250. sumably TersttQnian strata of the

Conoryctes coma: Schoch and Lucas, 1981b Nacimiento Formation, San Juan basin, L

Qpe spec.i.meji. "QW 3395, left New Mexico.

dentary fragment with P4-M2, alveolus for AMNH 16029, right dentary fragment

P2, roots of P3 and b13, isolated lower with M1-2 (Pl. 14, Fig. 9); UNM B-890, 32 1- 2 right canine (Pls. 15, Figs. 14-17; 16, palate with right P -M and left M ,

C1 P3 P4 M2 LW LW LW LW L w L-w AMNH 339sa

3396b 8.8 7.5 6.3 6.'8 7.5 11.5 7.9 12.1 7.2 11.1 5.5 8.2

3397

3398 8.8 7.5 7.3 7.4 8.3 11.7 7.8 11.6 7.9 12.5 7.1* 8.9

16029

UK 8033

UNM B-890 7.6 7.5, 8.2 12.0 9.1 13.0 7.6 11.7

USNM 22484 7.1 8.3 8.4 13.9 7.9 16.9* 6.6 11.1

N 44 4 433 4 4 3,3

Mean 7.08 7.50 8.10 12.28 8.27 12.23 7.65 13.05 6.40 9.40

Standard Deviation .56 .62 .41 1.10 .72 .71 .33 2.63 -82 1.51

Coefficient of 7.9 8.3 5.1 9.0 8.7 5.8 4.3 20.2 12.8 16.1 Variation

a Type specimen of Conoryctes comma.

bType specimen of "Hexodon molestus".

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p3 p4 32 M3 L WL W L L L ,w W L W W L W W Tri Td Tri Td Tri Td 10.1 7.9 10.7 7.6 9.4 8.1 8.0 7.7

9.5 7.5 7.2 6.4 9.9 7.1 8.3 7.9 8.7 7.0 6.6 7.8 5.4 5.0

10.1 8.4 8.1 9.2 7.9 7.3 7.9 6..3 5.5

10.3 7.9 7.9 9.2 7.5 6.9

9.0 6.6 9.8* 7.9 7.5

8.8 6.5 10.8 7.4

7.7 8.3 10.8* 7.9 11.5 9.2 10.8 8.2

33 4 4 5 6 5 4 5 3

8.33 6.50 10.55 7.50 10.22 8.dO 7.88 9.48 7.66 6.93

.99 .10 .44 .34 .79 .49 .23 .91 .45 .35

11.9 1.5 4.2 4.5 7.7 5.9 2.9 '9.6 5.9 5.1

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alveoli for right M3 and left P3-4, Flatthew (1937, p. 249-254). Cope 3 M , loose incisor, left P right P4, 3’ (1884a) originally proposed a separate and fragments of cranium (Pl. 16, Fig. taxon, “llexodon mo2estus ,” for AMNH

1); USNM 22484, skull and jaws with 3396 (Pl. 14, Figs. 1-6, 8, lo), 3 2 right and left P3-4, M , and left M , claiming that it dsffered from

.right C1, P2, P4-M2 (Pl. 11); USNM 22483, Conoryctes comma (Pls. 15, Figs. 14-

partial left manus (Pl. 14, Fig. 7) 17; 16, Fig. 6) by having four lower

catalogued with a skull of Triisodon premolars instead ot three. However,

quivirensis may actually belong to the in his revision of Conoryctes, Cope

same individual as the skull of USNM (1888d) correctly pointed out that

22484 (see discussion below): all P is extremely small adsingle- 1 from Torrejonian strata of the Naci- rooted. According to Cope, its ab-

miento Formation, Torreon wash, San sence in AMNH 3395, the type specimen. Juan basin, New Mexico. of c. comma, is a preservational arti- UK 8033, ieft dentary fragment with fact. Otherwise, C. comma and ”H.

P3-M1: , from UK New Mexico locality molestu!! are indistinguishable and

15, Torrejonian strata of the Naci- therefcre he synonymized the two

miento’Formation, SW1/4 sec. 20, T. 22 N., (Cope, 1888d, p. 317). Van Valen

R. 6 W., Sandoval County, San‘Juan (!978) has recently assigned MlNH

basin, New Mexico. 3224 (Pls. 15, Figs. 10, 11; 16,

Diagnosis. Same as that for the genus. Fig. 8), the type specimen of

“Tm:

Conomjctes (Fig. 4) i.s a nonotypic bears a large paraconid diagnostic

genus and has been thoroughly described of Huerfanodon and is tentatively

and revised by Cope (1888d, p. 316- assignable to that genus, .although

317), Wortman (1897b, p. 101-102), and indeterminate at the species

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Figure 4. Restoration of the skull, mandible, and

dentition of Conoryctes coma. Skull based primarily G on USNEl 22484 and AMNH 15939 (undetermined conoryctine

skull). Dentition based primarily on AMNH 3395, AMNH

3396, and UNM B-890. (a) Left lateral view of skull.

(b) Left lateral view of mandible. (c) Occlusal view

'of upper left dentition. (d) Occlusal view of lower

right dentition.

Figure 4 appears on the following frame.

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Figure 4.

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level (Schoch and Lucas, 1981b; also smaller, conical, labially placed

see Huerfmzodon below). paracones and metacones. The stylar

The den.tal formula of Conoryctes shelves of M1-3 are extremely narrow 133 cononc probably is I: or 3 C1 P4 M3 and the ectoflexi are weakly developed

(compare Matthew, 1937, p. 251). 11-2 are or absent. Labially, M1-3 bear

slightly enlarged arid the I2 i’s enlarged minutely cuspidate ectocingula; meso-

relative to the I1. The canines are styles either are absent (UNM B-890)

relatively large, projecting and deeply or moderately_ well developed (AMNH

rooted. Apparently, P1 is absent. P2 3396).

is small, possibly single-rooted, and P1-2 are extremely reduced, single-

unicuspid. rooted, and unicuspid. They are sub- .. .. P’ (Pl. 16, Fig. 1) is three-rooted circular in cross section. P3 (Pls.

and triangular to subcircular in cross 14, Figs. 2, 8; 16, Fig.\6) is single-

section. It bears a large paracone rooted, oriented slightly obliquely

and a minute to small metacone and in the jaw, and anteriorly bears a

lingual cingulum. large, anteroposteriorly elongated 4 P (Pl. 16, Fig. 1) is molariform and laterally compressed protoconid.

and bears a large protocone lingually A small, posterointernal talonid

and a smaller, conical paracone and heel may be variably developed on P3.

metacone, both placed far labially. The P4 of Conoryctes (Pl. 16, 4 P also bears a small parastyle, Fig. 6) is submolariform and bears a

stylocone, metastyle, and metastylocone. large, simple protoconid anteriorly.

It lacks a mesostyle or ectocingulum. The talonid is comma-shaped in

M1-3 (Pls. i4, Fig. 10; 16, Fig. 1) cross section (hence the name

decrease in size posteriorly, but @noryctes coma Cope, 1881a) and

otherwise are of’ similar morphology. bears a large hypoconid and.smaller

M1-3 bear large protocone$ lingually and hypoconulid and entbconid.

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of Conomjctes(P1s. 14, Fig. 9; "1- 3 Schoch and Lucas, 1981b). In general, 16. Fig. 6) decrease in size posteriorlv, the skull of Conomjctes is larger,

but otherwise are of similar morphology. heavier, and more robust than that of

The trigonids are slightly compressed Onychodectes(Fig. 1; Pls. 1, 3) and has

anteroposteriorly and bear large, a relatively short face.

conical protoconids and metaconids. The postcrania of Conomjctes are

Extremely small paraconids and para- poorly known. A partial humerus and

cristids are present. The talonids distal end of a radius are preserved

are subcircular in cross section and with AMNH 3396 (Pl. 14, Figs. 5, 6).

bear moderate-sized hypoconids, smaller They are not notably different from those

entoconids and hypoconulids, and still of Onychodectes. Robert T. Bakker

smaller mesoconids, entoconulids, (Johns Hopkins University) has identi-

and metastylids. Two smaller cusps fied the artial left manus of USNM

are present on the postcristids on 22483 (Pl. 14, Fig. 7), catalogued

either side of the hypoconulids. under the same number as a skull of

The cheek teeth of Conomjctes Triisodon quivirensis, as the foot

are hypsodont (crown hypsodonty) and of Conorgctes. It-may belong to the

show taeniodont pattern of "rolling same individual as USNM 22484, a

eruption'' (Patterosn, 1949b) in its skull and mandible of Conoryctes (Pl. 11)

extreme form. which was collected together with USNM

Wortman (189ib, p. 101-102, Fig. 36) 22483. This foot' is larger, but

and Matthew (1937, p. 250-251, Fig. 61; otherwise similar to the foot of

P1. 58, Figs. 1, 2) described and hzychodectes (PI. 7, Fig. 6), and will

illustrated the skull and mandible of be described in detail by R. T. Bakker

Conoryctes(Fig. 4; P1. 11). It and the author at a later date.

closely resembles the skull of Huerfanodon Schoch and Lucas, 1981b

Huerfanodon (Fig. 5; P1. 15, Figs. 1-9; Husrfanodon Schoch and Lucas, 1981b.

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Qpe species. herfanodon torrejonius 16, Figs. 2, 3, 5, 7)

Schoch and Lucas, 1981b. iluerfanodon torrejonizcs Schoch ,and Lucas ,

IncZuded species. The type species, 1981b.

Huerfanodon polecatensis Schoch and Qpe specimen. USNM 15412, a partial

Lucas , 1981b, and "Triisodon hei Zprixianuz skull with right P3, right M1-3, left M1-s! Cope, 1982c, nomeiz dubiwn. , associated fragments of the

Kstribution. Torrejonian of New central uppel; incisors, partial root of

Mexico and Wyoming right P2, alveoli for right P4 and

Kagnosis. Medium-sized taeniodonts, left P2-4, plus associated right 1- 4 approximately the size of Conoryctes: dentary fragments bearing C1, P , 3 3 P submolariform with a large paracone M , and right C1, plus an associated

and small but distinct metacone, proto- right dentary fragment with alveolus

cone, and hypocone; M1-2 with well for P4, roots of M1-2, and complete

developed and protruding mesod?!yles M (Pl. 16, Figs. 3, 7): from Tor- 3 connected to the metacones by slightly rejonian strata of the Nacimiento

cuspidate premetacristae, distinct Formation, 2.5 km northwest of

and cuspidate parastyles, ectocingula Nageezi, San Juan basin, New Mexico.

and metastyles, and broad trigon basins Dictgnosis . Species of Iluerfanodon

with shallow valleys and cuspidate with a submolarifor& P4: the trigonid

lingual margins bearing moderate-sized bears a large protoconid with a

J "protocones," Ilparaconules," and "meta- slightly developed cingulid on its

conules'l; lower canine with internal anterolingual aspect but lacks the

groove; lower molars with promineLC large and distinct metaconid seen

paraconids and paracristids. in H. polecatensis; slightly smaller

dentally than I!. po leeatensis. Huerfanodon tc@?qjb?;ius Schoch and Lucas, 1981b

(Table 4: Fig 5: Pls. 15, Figs. 1-9;

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-H. torrejonius USNM 15412 7.9 8.0 8.5 10.2 8.1 9.3 5.4 8.1 8.4 7.0

MCZ 20181 8.1 9.1 7.8 8.6 -8. polecatensis PU 14178 ?g."heilpinianus" AMNH 3224

p3 p4 M1 M2 M3 L WtL LWWLWWLWW Tti Td Tri Td Tri Td

11.3 8.2 10.1 8.5 8.5 8.7 6.9 6.7

MCZ 20181 8.5 6.0 5.3 H. polecatensis PU 14178 8.5 6.2 11.6 8.5 11.1 9.4 8.7 9.5 8.6 7.5 1H. "heilpinianus" 3224 AMNH 10.3 6.4

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Huerfanodon polecatensis Schoch and Lucas C'onoryctes coma: Van Valcn, $378, 1981b p. 58 (in part). (Table 4: Fig. 5: Pls. 15, Figs. 12, ?Ifuerfcmodon "hei Zprinianus": Schoch 13; 16, Fig. 4) and Lucas, 1981b., Huerfanodon po lecatensis Schoch and Lucas, Type specimen. AMNH 3224, left 1981b. CI dentary fragment with N2 (Pls. 15, Type and only knm specimen. PU 1471t Figs. 10, 11; 16, Fig. 8;. right dentary fragment with P3-M2, root Horizon and locality of the type. of P2, alveoli for C1, P1, and M3 (Pls. Paleocene strata of uncertain age, 15, Figs. 12, 13; 16, Fig. 4). Nacimiento Formation, San Juan basin, Horizon and locality of the tpe. New Mexico. Torrejonian strata of the Polecat Bench Discrcssion. The M2 of AMNH 3224 Formation, Rock Bency Quarry, SCC. 36, bears a large paraconid as in T. 57 N., R. 99 W., Bighorn Basin, Huerfanodon; however, without the P4 it Wyoming. is not possible to assign it to a Diagnosis. Species of Iluerfmdon species and I here consider ?Huerfanodon with a molariform P4, bearing a large "Iyilprinianus" a nomen dubiwn (see and distinct metaconid which approaches Schoch and Lucas, 1981b, for a full the protoconid in size; slightly larger discussion of this specimen). dentally than H. torrejonius.

cf . ?Huerfanodon sp . ?Huerfanodon "hei lprinianits" (Cope, 1882~)

I Conoryctes comma: Simpson, 1937, p. 169. (Table 4:. Pls. 15, Figs. 10, 11; Referred specimen. USh- 9678, left 16, Fig. 8) 3 P : from Torrejonian strata of ,the Triisodon hei lprinianus Cop,?, 1882c, p. 193. Lebo Formation ("Fort Union"), Silberling

Eoconodon heilprinianus : Matthew and Quarry, Crazy Mountain Field, Sweet-

Granger, 1921, p. 6 (in part). grass County, Montana.

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3 D-iscussion. This.specimen, a left P , generically indeterminate due to the

is in the size range of fhei*fanodon and incompleteness of the material.

bears a small, but distinct, metacone Conoryctine genus indeterminate B and protocone; thus, it may pertain to

Huerfanodon, but without more material Conoryctes coma: Simpson, 1937, p. 169.

it is impossible to make a definite Referred specimens. USNM 9597, left

assignment. upper molar: from Torrejonian strata

of the Lebo Formation ("Fort Union"), Description md Discussion of herfanodon Gidley Quarry, Crazy Mountain Field,

Huerfanodon (Fig. 5) is thoroughly Sweetgrass County, Montana.

described and discussed in Schoch and USNM 9816, two upper molars: from

Lucas (1981b) and need not be further Torrejonian(?) strata of the Lebo

described or discussed here. Formation, Simpson's Locality 6, Crazy

Mountain Field, Sweetgrass County, Conoryctine genus indeterminate A Montana.

Referred specimen. AMNH 832, right USNM 9826, cheek tooth fragments:

dentary fragment with roots of C1, from late Torrejonian(?) strata 6he p2(?) , of complete P3(?> , three upper molar Melville Formation ("Fort Union"),

fragments, right P3 of a second individu- .Simpson's Locality 28, Crazy Mountain

al(?) and other tooth fragments: from Field, Sweetgrass County, Montana.

Torrejonian strata of the Nacimiento Discussion. The above specimen'c

Formation, Escavada wash, San Juan basin, document the presence of a conoryctine

New Mexico. in the size range of Conoryctesand

Discussion. The above specimen docu- lluerfanodon at these localities, but

ments the presence of a conoryctine are generically indeterminate due to

in the size range of Conoryctes and the incompleteness of the material.

Huerfanodon in Escavada wash, but is

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Figure 5. Restoration of the skull, mandible, and

dentition of Huerfaxodon. The skull and mandible is

hypothetical and based on MCZ 20181, USNM 15412,

USNM 22484 (Conomjctes coma), and AMNH 15239

(undetermined conoryetine). Upper dentition based

on IISNM 15412 (Huerfanodon torrejonius). Lower den-

tition based on PU.14178 (top Pg- 4) and alveoli for

: H. poZecatensis) USNM 15412, and 3224 p1-2 , 2 (?H. "heiZprinianus"). (a) Left lateral view of

skull. (b) Left lateral view of mandible. (c) Occlusal

view of upper left dentition. (d) Occlusal vl.ew of

lower right dentition.

Figure 5 appears on the following frame.

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Figure 5.

Downloaded from http://pubs.geoscienceworld.org/gsa/gsabulletin/article-pdf/92/12_Part_II/1982/3418970/i0016-7606-92-12-1982.pdf by guest on 01 October 2021 2021 I Conoryctine genus &determinate C obliquely set in the jaw and anteriorly bears a large protoconid as in (Pls. 12, 13) Conoryctes and Wuerfanodon. Posteriorly

Conoryctes comma: Matthew, 1937, p. 250 there is a well-developed second conid

(in part). in the position where a posterior heel

Referred specimen. AMNH 15939, skull is often variably developed in

and mandible with roots or alveoli for Conoryctes. In the type specimen of 1 right and left 12-3, C and complete Huerfmodon poZecntensis, there is

P 33-M (the left P3-4 and right P4 fiave also a slight posterior heel inLthis , position 16, Fig. 4). However, come out..of the skull), alveoli for (Pl.

right and left 12-3, C1, M3 and left this conid is better developed in hMNH

complete left P3, right and left 15939 than in any other conoryctine p1-2 P4-M2 (Pls. 12, 13): from Torrejonian specimen known to me.

strata of the Nacimiento Formation, I Nortmania !lay, 1899 Torrejon wash, San Juan basin, New

Meki.co. Hemigmus: Wortman, 1897b, p. 67.

Uiscusszon. This specimen was de- Wortmania Hay, 1899, p. 593.

scribed and figured by Matthew (1937,

p. 252-254, Fig. 61) as Conoryctss 1 Cope, 1885a. I .TncZuded species. Only the type However, the teeth of Awn 15939

are extremely worn, such that, irhercas species.

it represents a large conoryctine An the Distribution. Puercan of New Mexico.

size range of Conoqctes and T!uei>fanodon, Revised diagnosis. . Medium-sized

it.is generically indeterminate. The taeniodonts (smaller than Psittacotherim

left:P3 of ANYH 15939 is the only tooth but larger than Conoryctes) with rela-

in this specimen which preserves any tdvely low-crowned teeth; all teeth

details of the crown morphslvgy. It is relatively shallow-rooted; upper and

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lower ca'nines enlarged and oval in cross for left M2 and Tight M3; associated

section; skull and mandible rbbust; lower left half of atlas, central and right

premolars bear a single anteroexternal portion of axis, three cervical verte-

conid and a well-developed posterinternal brae, (?)right second metacarpal,

cingulid ("talonid") ; lower molars with un'gual phalanx of manus, (?)left

both trigonids and talonids compressed lunar, left tibia, proximal half of , anteroposteriorly and expanded trans- left femur, proximal half of left

versely, with wear forming two blunt, ulna, portion of the right ulna (in-

transverse lophs; trigonids wider than cluding the semilunar notch) and left

talonids. radius (Pls. 17, Figs. 1-3; 18, Figs. 1-

3; 19, Figs. 2-5; 20). wortmania otttrzidens (Cope; 1885a) Horizon and Zocality of the type.

(Table 5: Fig. 6: Pls. 17-20) Collected by David Baldwin in l$85 from

Hemiganus otariidens Cope, 1885a, p. 492. presumably Puercan strataohthe'Naci-

Hemiganus otariidens: Cope, 188d, p. 311. miento Formation, San Juan basin, New

Hemiganus otariidens: Worfman, 1897b, p. 311 Mexico.

Wortmania otariidens: Matthew, 1937, p.271 Referred specimens. AMNH 755, left

non Nortmmia otariidens: Rigby and C 1 and bone fragrnents-(Pl. 18, Fig. 5); Lucas, 1977, p. 56 (based on a very worn UCMP 36528, left dentary with complete

maxilla fragment of Ectoconzts ditrigonics ; P1-2, roots of 11, C1, P3-4: both

S. Lucas, .19$1, personal commun.). from presumably Puercan strata of the

Qpe specimerr. AMNH 3394, partial Nacimiento Formation, San Juan basin,

skull with damaged and fragmenta'ry New Mexico.

right C1, P3-4(?.) and left 13(?), C1, UCMP 89280, isolated and well worn 3(?) P2-M1(?); lower mandible with right P2 or P3; USNM 17655, left P

(Pl. 17, Fig. 5): both from Puercan and left 13, C1-, P3r4*,.right M2, left M1, roots of righf'M1 and alveoli strata of the Nacimiento Formation,

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TABLE 5. DENTAL MEASUBEMENTS OF WORTMANIA OTARIIDENS

c1 P2 P3 L IJ 1 W 1 W 1 W

AMEM 755

3394a 7.6* 6.4" 14.5* 11.7" 5.4* 6.6* 6.2* 8.3*

16342

UK 12998

USNM 15429

15654

15655 6.7* 9.7*

UOlP 31819

89280

a Type specimen of Wortmania otariidens.'

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TABLE 5. (COnhh~ed)

P4 M1 M2 - I3 c1 p1 L W L W L W L W 1 W 1 W 13.7 10.0

6.7* 9.4* 8.5* 8.1* 4.0* 14.0* 11.0* 5.5 8.8

7.7* 12.5* 5.7 8.3

5.5 7.2

7.8 10.6 7.7 11.1

8.3 11.7

10.2* 5.7" 13.7* 13.6* 6.9 9.8

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TABLE 5. (CO&uLed)

L W L W L W L W W L W W Tri Td Tri Td

-I . .. 6.1 9.1 "6.7 @JJ 6.7* 8.5" a.5* 8.1* 6.9* a.i* 8.0 6.8*

6.2 8.9 * - 6.5 8.4 8.3 6.9* 5.5

6.7 9.9

5.4 8.9 * \

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Betonnie Tsosie wash, San Juan basin, (1937, p. 270-277).

New Mexico. The teeth of Nortmania are very

UK 12998, five (?)lower premolars poorly known (Fig. 6). The teeth of

(right and left P and unde- 1-2(?) the palate of the type specimen termined fragment) and a right M2; (Pl. 17, Figs. 1-3) are heavily worn

USNM 17654, right P 4(?) (Pl. 17, Fig. and poorly preserved. The skull

6): both from Puercan strata of the has been plastered together since

Nacimiento Formation, Kimbeto wash, Matthew (1937) wrote on Vortmania,

San Juan basin, New Mexico. and the left upper cheek teeth are

UShW 15428, right dentary with C1 only set in plaster and might not be

(Pls. 18, Fig 4; 19, Fig. 1); USNM in their original positions. At the 4 15429, left P -M I(?) (Pl. 17, Fig. 7): posterior portion of the left maxilla

both from Puercan strata of the Naci- behind the tooth here interpreted as

miento Formation, De-na-zin wash, San M1(?) is a tooth set in the plaster

Juan basin, New Mexico. which is so incomplete that it cannot

AMNH 16342, left dentary fragment with be determined if it belongs to

roots of C1, P3-4 and an associ- Nortmania. Likewise, the two halves

ated maxilla fragment with M2 and of the lower mandible (Pl. 18, Figs. 1- 1 3 alveoli for M and M (Pls. 17, Fig. 4; 3) have been plastered together, and

18, Fig. 6): from Puercan strata of the the incisors,. canines, and two anterior

Nacimiento Formation,- Alamo wash, San premolars are only set in plaster.

Juan basin, New Mexico. The upper incisors apparently are

Diagnosis. Same as that for the genus. roughly circular in cross section and

moderately large. The lower incisors Deswipt ion- and Discussion of Nortmania are smaller than the uppers and

Wortmania is an extremely rare elongated anteroposteriorly. The

monotypic genus last reviewed by Matthew upper and lower canines are large and

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Figure 6. Restoration of the skull, mandible, and

detition of Wortmania otariidens, based primarily on

AMNH 3394, AMNH 16342, USNM 15429, USNM 17654, and

USh 17655. (a) Left lateral view of skull.

(b) Left lateral view of mandible. (c).Gcclusal

view of upper left dention. (d) Occlusal view of

lower right dentition.

Figure 6 appears on the following frame.

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Figure $.

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stout (Pl. 18, Fig. 5). internal cingulids (or "talonids") . 1- 2 The poorly preserved P apparently P1-4 are slightly claw-like and

are short anteroposteriorly and rela- lingually recurved.

tively wide transversely (labio-lingual- M1-., (Pl. 18, Fig. 3) bear

ly). They bear large and high antero- moderate-sized protoconids and meta-

external cusps; o'therwise they are not conids and smaller paraconids. All

distinctive. detiils of the talonids on AMNH 3394

P3-4' (Pl. 17; Fig's; 5-7) bear large, and UK 12998 have been removed by wear.

centrally placed paracones labially Both the tigohids and talonids are

and smaller protocorics lirigually, slightly compressed anteroposteriorly

slight ectocingula, parastyles, meta- and expanded labio-linpually to farm

styles, and para- and metacristae. two blunt transverse lophs. The

of Wortmania (Pl. 17, Figs. 4, trigonids are wider than the talonids.

7) bear moderate-sized protocones, large The cheek teeth of F!ortmania show

protocones displaced far labially, and the typical early taeniodont character-

slightly smaller metacones partially istic of hypsodonty in the form of

fused with the paracones (USml 15429). a small degree of lingual expansion

M1-2 also bear slight parastyles, ecto- of the enamel on the upper cheek

cingula, metastyles, and may possibly teeth and labial expansion of the enamel

bear small 'lprotoconules'l and "meta- on the lower check teeth.

conulesl' (although wear on the teeth The degree of crown hypsodonty of

obscures these features and homologies Nortmania is less than that of the

are: uncertain). smaller genus Onychodectes. The teeth

PI-4 (Pl. 18, Figs. 3, 6) are of Ir'ortmania appear to have been

transversely oriented in the tlentary shallow to moderately rooted. The

with large and high anteroexternal roots of the type specimen (AMNH 3394)

conids ("protoconids") and postero- cannot be seen, but the referred (?)lower

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premolars of UK 12998 each bear one 277, Figs. 68, 69; Pls. 61, Fig. 1;

Stout root. The molar of UK 12998 bears 62, Figs. 1-4; 63) have thoroughly

two roots, one underlying the trigonid described and illustrated the skull

and the other underlying the talonid. (Fig. 6) and postcrania of the type

The upper molars of USXM 15429, USNM specimen of krortmania(AMM1. 3394; PIS.

17654, and USNM 17655 are all three- 17, Figs. 1-3; 18, Figs. 1-3; 19, Figs.

rooted, having two medium-sized roots 2-5; 20) and they need not be described

labially and one large root underlying here. In general, th.e skeleton of

the protocone lingually. Vortmania shares many of the character-

Matthew (1937) considered the prob- istics of later and more advanced

able dental formula of Vortmania to be taeniodonts. The skull is short and 1 1 3. 3- I1 C1 P4 M3, whereas Patterson (1949b) deep with a massive mandible. The neck

coniidered it tQ- be 1: C: P; PI:. From is short and stout. The ulna bears a

the known material, however. it can be relatively large olecranon process.

stated with certainty only that The.manus carries large, laterally

Wortmania had at least one incisor compressed, and recurved claws. The

above and below (but note that Wortman, limb bones arc relatively stout and

1897b, believed that Vortmania had 'two robust, foreshadowing the skeleton of

pairs of upper incisors, which does later advanced taeniodonts.

not seem unreasonable); one canine Stylinodontine genus indeterminate above and below; at least three upper

premolars, four lower premolars; an un- (Pl. 27, Figs. 1, 2)

determined number of upper molars; undetermined stylinodontine genus: Robi-

and three lower molars. son and Lucas, 1980, p. 302.

Cope (1888d, p. 311-316, Pls. 4; 5, Refelired specimen. AMNH no number,

Figs. 1-7), Wortman (1897b, p. 67-71, cheek tooth (Pl. 27, Figs. 1, 2): from

Figs. 2, 3), and Matthew (1937, p. 270- Puercan strata of the North Horn

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3 Formation, Wagonroad local fauna, Emery external groove; I enlarged, deeply

County, Utah. rooted, and caninifom; lower incisors

Discussion. A1t hougbmast of the of.moderate size and rooted; cheek

crown is missing, the fused roots and’ teeth moderately hypsodont with

labial and lingual extensions of the re- relatively shallow roots (as compared

maining enamel indicate that this tooth to Ectoganus); posterior cheek teeth

-\ belongs to an advanced taeniodont in the double-rooted or single-rooted with

size range of Wortmnn7:a or Ps7:ttacotliem:iun traces of the fused roots;-upper pre-

molars subowl in cross section and Psittacotheriiun Cope, 1882b transversely elongated, bearing a large

Psittac9tkeriwn Cope, 1882b, p. 156. protocc;ne and paracone connected by

Henriganus Cope, 1882e, p. 831. low to incipient transverse anterior

Qpe species. Psittacotheriwn multi- and posterior crests; upper molars

fragwn Cope, 1882b (= Psittacotheriwn tritubercular, suboval in cross section

aspasiae Cope 1882c = Herriganus and transversely.elongated, bearing

vultuosus Cope, 1882e = Psittacotheriiun small paracones and metacones placed

megalodus Cope, 1887b). far labially and anteriorly, and large

Included species. On! y the type species protocones lingually; minutely cuspidate

Distribution. Torrejonian of Wyoming postmetaconule wings well developed

and New Mehco; Torrejonian-Tif fanian (especially on M2-3) and extending

of Montana and Texas. posterolabially to a point posterior

Revised diagnosis. Medium-sized and just lingual of the metacone;

taeniodonts with greatly enlarged, P1 reduced-nd single-rooted; p2 rooted, subgliriform canines Gith enamel single-rooted and placed transversely

limited to the anterior face of the in the lower jaw, bearing a large

tooth and with both crown and root external (labial) conid and a smaller

greatly elongated; upper canine with internal conid; Pq submolarifom with

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a transversely widened trigonid and Psit tacotherim mu 2tifragwn: Mat thew,

small, lingually placed talonid; Mi-3 1937, p. 255.

decrease in size posteriorly; M1-3 Psittacotheriza aspasiae: Matthew,

moderately bilophodont with broader 1937, p. 269.

trigonids than talonids, talonids Psi t tacotherim mu Ztifragwn: Simpson,

placed lingually. 1937, p. 169.

Psittacotheriwn rnuZtifragwn: R. W. Wilson, Psittacotheriwn rnuztifragwn Cope, 1882b 1956, p. 82. (Table.6: Fig. 7: Pls. 21-26; 27, Psittacotherim muZtifragwn?: Simpson, Figs. 4-14; 28-31) 1959, p. 6. 'Psittacotheriwn muZtifragm Cope, 1882b, Psittacotherim ci. P. mu2tifragwn: p. 156. J. A. Wilson, 1967, p. 159, 161. Psittacotheriwn muZtifragwn: Cope, i882~, Psittacotheriwn Cope, 1822 (lapsus ca&) p. 191. or Lampadophoms Patterson, 1949: Psittacotheriwn aspasiae Cope, 1882c, p. Schiebout, 1974, p. 19. 192. Psittacotherium multifragwn: Rigby, 1980, Hemiganus vuZtuosus Cope, 1882e, p. 831. p. 98. Psittacotherim multifragwn: Cope, 1884c, Psittacotherim mu2tifragwn: Schoch, p. 196. 1981a, p. 180. Psittacotheriwn aspasiae: Cope, 1884c,

p,. 196. Type specimen. AMNH 3413, mandible

Semiganus uu Ztuosus : Cope, 1884c, P1. 32c with left 13, C1, right Pg, M1, M2,

Figs. 7-12. roots of right 13, C1 and alveoli for

Psittacotheriwn megaZodus Cope, 1887b, left M2-3 and right P1, M3 (Pl. 21,

p. 469. Figs. 1-4).

Psittacotheriwn mu Zti fragum: Wortman, Horizon and 2ocaZity of the type.

1897b, p. 71. From presumably Torrejonian strata of CuZwnodon?: Douglass, 1'908, p. 22 the Nacimiento Formation, near Huerfnno

1 'I C1 P -__-. P2 . P3 P4 L W L W 1 W 1 W 1 W 1 W

AMNH 754 17.0" 10.7 8.1 11.8

756

75 7

2453 18..6 .9;7 9.4 15.7 JO.9 15.9 10.5 18.0"

3390' 25.2 16.4

3391 8.9 16.1 9.7 16.6

3392

3393

3413a

3416b

3418d

15938 8.0 15.9 11.3" 16#.5*

16660

16661 21.5 15.1 9.3 13.9 8.6'

16731 19.1 11.6 10.9 15.1 10.1 16.6

88383

100563 9.8 15.0

CM 1674 9.3 14.7

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M1 M2 P I3 'cl p2 3 L W L W L W L W L W L W L W

26.6 19.1

12.0 17.5 10.5 13.1 9.4 13.9

10.2 15.9 13.5 6.9 28.9 18.2

28.7 17.0

28.5 18.2

7.1 13.0

27.8 16.0 9.8 16.4

11.5* 16.5" 10.0" 15.5" 1l.t 13.1

27.0 18.3

25.4 15.9

8:8 12.9

31.2 19.5

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L W L W L W LW

11.5* 14.5*

10.3 15.4

13.5* 14.5 11.1 10;s *

11.4 10.8

10.4 10.1 9.3 9.0

9.2 9.0 9.3 8.3

13.6 16.1

11.9 12.9 11.9 11.7 10.4 10.0 12.9

12.5* 15.5" 14.0* 1l.O*.' 15.5* 11.0"

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"MM 40147-3 10.2 14.7

40148-2

40535-86

40536-119 9.5 13.4

40537-26

40537-33

40537-61

UK 8035 11.5" 17.0* 13.0* 18.4 10.0* 17.5*

UNM NP-220

USNM 6162

15410

15411 18.8 13.0 9.3 10.2 9.3 13.2 8.4 13.7 8.7 12.9

N 3 3 3 3 7 7 3 39 8

Mean 18.23 10.66 21.83 14.83 9.61 15.27 10.77 16.0 9.78 15.98

Standard 1.09 .95 3.21 1.72 1.20 1.32 2.30 2.35 .85 1.67 Deviation

Coeff4cient 6.0 8.9 14.7 11.6 12.5 8.6 21.4 14.9 8.7 10.5 of: Variation

a Type specimen of Psittacotherium multifragum.

bType specimen of g. "aspasiae". C Type specimen 06 "Hemiganus vultuosusQ1.

dType specimen of P. "megalodus".

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12.7 7.0

10.2 14.6

10.5 14.5

10.5 14.8

8.9 l5Sl

26.9 16.7

28.7 18.6

8.9 14.1 8.$-?12.9.. 7.6* 12.0*

4- 4 5 5. 10 10 3 3 10.65 16.0 9.44 13.24 27.97 17.75 8.77 14.43

1.39 1.43 1.51 1.13 1.61 1.27 1.46 1;76

13.1 8.9 16.0 8.5 5.6 7.2 16.6 12.2

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TABLE 6. (CO~HU~)

10.6 10.8 9.1 9.5

12.0 11.9

8 8 6 5 3 3 3 3

12.09 13.95 11.82 10.82 9.20 9.17 11.73 9.77

1.10 1.89 1.41 .60 .10 .29 3.31 1.37

9.1 13.5 11.9 5.5 1.1 3.2 28.2 14.0

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Peak, San Juan basin, New Mexico. and left C1, left P4, M3, roots of

Referred specimens. AMNH 754, partial left M1, alveoli for right and left

skull and mandible with right and left 139 '1-3, right P4-M3, partial left 31 11 I -C , fragmentary right P -M and premaxilla and maxilla with roots of 31 partial right 13, left P1, right and I -C , tooth, cranial, and vertebral

left P4 (Pl. 24); AMNH 757, (?)lower right fragments (Pls. 26, Figs, 3, 4; 29,

canine; AMNH 3416, left dentary fragment Figs. 5-8): all from presumably

with-partially erupted M3,alveoli for Torrejonian strata of the Nacimiento

M1-2 and crushed M2 cemented to the Formation, San juan basin, New Mexico.

outside of the jaw (type of P. "aspasiae"; UK 7749, upper cheek tooth fragments;

P1. 21, Figs. 9, 10); AMNH 3418, right UK 9564, canine, vertebra, and hind-

dentary with C1 root and alveoli for limb fragments; UK 9565, canine frag-

P1-M2 and isolated right P2 (type of ment; UK 9566, lower incisor; UK 9567,

P. %egaZodusl'; P1. 21, Figs. 5-8) ; canine fragment; UNM "-220, dentary 1 AMNH 3390, left C1, right 13, right C fragments with right and left M1, right

all from Torrejonian strata of and upper cheek tooth (type of M2: "Herriganus vuZtuosusll; P1. 21, Figs. the Nacimiento Formation, Kutz Canyon,

11-18); AMhW 3391, right P4(?), left San Juan basin, New Mexico'. 2? P4(?) undetermined cheek tooth (P 1, AMNH 15938, (?)right upper incisor

canine tip and vertebra (Pl. 29, Figs. tip, right P2(?); r.ight and left P4(?), left M1-2, right M 3 , right 3, 4); AMNH 3393, broken upper cheek

tooth and right C1; AMNH 3414, left and left P4, left distal tibia (Pl. 30,

premaxilla and maxilla with 13, C1; Figs. 8, 9); AMNH 16661, upper left 1 AMNH 3417, left dentary fragment with C , right and left P2, P4(?), unde-

alveoli for M1-3; AMNH 3419, right termined upper cheek tooth fragment,

dentary with broken C1 and crushed left C1, right P4, right and left M1,

molar; AMNH 88383, mandible with right right M3 (Pl. 22, Figs. 24-31); LJNM

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2 1 3 B-850, right P1 or P ; USNM 15413, right maxilla with C and P roots, alveoli

and left dentary fragments with unerupted for 13, P1-2, P4, right dentary frag-

right and left M3, deciduous C 1' ment with alveoli for M2-3; UK 8006, deciduous P1-2(?) (Pl. 27, Figs. 4-14): edentulous mandibular symphysis

all from Torrejonian strata of the with alveoli for right and left C1-P4;

Nacimiento Formation, Torreon wash, USNM 15410, left dentary with C1 (Pl. 23,

San Juan basin, New Mexico. Figs. 5, 6); USNM 15411, palate and 1 UK 8035, right side of skull with partial skull with right and left C , 3 1 4 ~3,M , roots of p2, p4, alveoli for right P , right and left P2-3, left P , 1 2 13, C1, mandible with alveoli right and left M , left M and right 3 for right and left.C1-M3, tooth and M (Pls. 23, Figs. 1-4; 31, Figs. 1, 2;

bone fragments (Pls. 25; 26, Figs. 1, USNM 15410 and USNM 15411 may represent

2): from Torrejonian strata of the a ,single individual, both were collected

Nacimiento Formation, UK New Mexico at the same time, place and bear the

Locality 15, SW1/4, sec. 20, T. 22 N., same original field number): all from

R. 6 W., Sandoval County, San Juan basin, Torrejonian strata of the Nacimiento

New Mexico. Formation, Kimbeto wash, San Juan basin,

AMNH 3392, lower canine: from New Jexico.

Torrejonian strata of the Nacimiento AMNH 756, dentary, canine and

Formation, Gallegos Canyon, San Juan other tooth fragments, including a right

New (Pl. 22, Figs. 22, 23); AMNH 2453, basin, Mexico. p4 (? AMNH 16560, ulna and radius, left mandible, upper and lower cheek teeth 3 femur, left proximal fibula(?) and and fragments including left I , left 21 3 partial left pes (Pls. 30, Figs. 207; P -M , M , right P2, left P4, right

31,, Figs. 3-5); AMNH 16660, mandible M1, right unlna, radius and manus

with right C1, alveoli for left C1, (Pls. 22, Figs. 1-14; 30, Fig. 1); 4 right and left P1-M3; AMNH 16662, left AMNH 16731, heft 13, right P2, left P ,

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3 left M , right M (Pl. 22, Figs. Wyoming. I(?) 15-21): all from Torrejonian strata 01 1674, right P (?) from Tiff anian

of the Nacimiento Formation, Escavada strata of the Melville Formation ("Fort

wash, San Juan basin, New Mexico. Union"), Douglass Quarry, northeast

AMNH 36000, skull in concretion with df Melville, Sweetgrass County, Montana. 1 3 roots of right and left C , right I , TMM 40147-3,- fragment of incisor, ..-- manibular symphysis with roots of right fragment of canine, right P 3(?).,

and left C1, alveoli for left P1-3, TMM 40147-7; edentulous right mandible

canine and bone fragments: from fra'gment; TMM 40148~2,left M I(?).., I, Torrejonian strata of the Nacimiento TMM 40535-86, incisor fragment; TMM

Formation, Simpson's Locality 226, 40536-119, left-P2(?); TMY 40537-26,

I1northwest to westnorthwcst of south- left M1'(?); TMM 4-537-33, lkft M I(?). ,

eastern tip of Cuba Mesa on fourth TMM 40537-61, right M1(?); TMM 40537-91,

main spur projecting southward from the right dentary fragment; TMM 40537-140,

mesa, mainly or wholly in sec. 3, T. 20 skull fragments in concretion 'and part

N., R. 2 W." (Simpson, 1959, p. 5), San of right upper canine; TMM 40537-68,

Juan basin, New Mexico. claw; TMM 41366-1, maxillary fragments

USNM 6162, canine and ,cheek tooth with roots of premolars and molars;

fragments including a right lower canine TMM 41366-73, upper incisor roots

and left P4: from Torrejonian strata in jaw fragment; TMM 41364-1, parts

of the Lebo Formation ("Fort Union:) of clavicle, left femur, right

"fromthelevel of and near Silberling humerus, right and left tibiae, right

Quarry'' (Simpson, 1937, p. 169), astragalus, and other bone fragments

Sweetgrass County, Montana. (apparently the same individual as 4 AMNH 100563, left P : from TMM 41364-2, below: Pls. 28;' 29, %s.

Torrejonian strata of the Fort Union 1, 2); TMM 41364-2, skull fragment&

Formation, Swain Quarry, Carbon County, with bone canines and one incisor,

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sockets for six left cheek teeth and teeth of Psittacotherim are extremely

associated bone fragments: all from late worn or fragmentary, many measurements

Torrejonian (?) and Tif fanian strata are little more than estimates.

of the Black Peaks Formation, Big Bend Measurements taken on different teeth

National Park, Brewster County, Texas. are not necessarily taken between

Diagnosis. homologous points. 4. The hypsodont

teeth of Psittacotherium (and taenio- Description and Discussion of Psittucot2e/iiL(i donts in general) may change in size

Schoch (1981a) pointed out that with eruption throughout the life of

known dental remains of Psittacotheriwn an individual (that is, rolling

are highly variable in size (see Table eruption; compare Patterson, 1949b).

6). This variability may be due to Due to the virtual lack of complete,

several factors: 1. More than one or even partial, unworn teeth still

species of Psittacotheriwn may be in place in jaws, there is no way to

present, and the measurements of two judge how stereotyped or variable

or more species may grade into each the various teeth in the tooth row

other. Detailed crown and cusp of Psittacotlieriwn are. In view of

morphology that might distinguish these factors, and the fact that

several species of Psittacotherim of there are no clear gaps in size

similar size is obscured by the worn between the measurenents of teeth

condition of most dental remains of (Table 6), even though the coefficients

Psittacotheriwn. 2. Many specimens of variation are somewhat high for a

of Psittacotherim consist of isolated single species (Simpson and others,

teeth, and their exact position in the 1960), it appears most reasonable

tooth row is uncertain. Thus, several to assign all specimens of Psittacotherim

teeth might be presumed homologous to one species (Schoch, 1981a).

when they are not. 3. Because many The dental formula of Psittacotherim

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1143 probably is I1 C1 P4 M3 (Fig. 7; canines. The lower incisors (for

contra drtman, 1897b; Matthew, 1937). example, AMNH 3413; P1. 21, Figs. 1,

The number of incisors is uncertain. 2) are much smaller. At eruption the

AMNH 88383 includes a complete left crown of I3 was completely covered

premaxilla which bears the root of with enamel. Enamel is limited to

only one enlarged upper incisor, here the anterior face of the incisor 3 designated I . UK 8035 includes a after wear and, as in the canines,

I. complete right naxilla (Pl. 25) whose the anterior portion apparently

rats and alveoli show that Psittacotheriur formed a cuttinglshearing implement,

had an enlarged and rooted upper canine, whereas the posterior enamel-free

four shallowly rooted upper premolars, surfaces of the incisors and canines

and three upper molars. The lower were used for crushing. The root of

mandibles of A"% 754, MI 2453,. the upper incisor is grooved medially

AMNH 88383, and UK 8035 (Pls. 24, 26) and laterally.

demonstrate that Psittacotheriwn had The upper and lower canines of

one set of moderately enlarged lower Psittacotheriwn are enlarged and

incisors, here arbitrarily designated subgliriform. Both the crown and

I3 (Matthew, 1937, considered it to the root are greatly elongated. Enamel

be 12, hoinologous to the enlarged is limited to the anterior portions of

lower second incisor of Conoryctes); an the canines arrrhis striated parallel

enlarged and rooted lower canine; four to the length of the tooth before wear.

lower premolars and three lower molars. With wear and eruption the occlusal

The upper incisors (for example, surfaces of the canines change size

AMNH 16731, an upper incisor of and shape such that in an old individual

Psittacotheriwn; P1. 22, Fig. 15) are like AMNH 754 (Pl. 24) the upper

enlarged and subgliriform, forming canines consist of only large dentine

miniature counterparts of the enlarged stubs elongated anteroposteriorly with

Downloaded from http://pubs.geoscienceworld.org/gsa/gsabulletin/article-pdf/92/12_Part_II/1982/3418970/i0016-7606-92-12-1982.pdf by guest on 01 October 2021 Figure 7. Restoration of the skull, mandible, and

dentition of Psittacotheriwn multifragwn. Skull and

mandible is based primarily on AMhW 754,. AMNH 88383,

and UK 8035. Dentition is based primarily on AMNH 3413,

AMNH 2453, AMNH 16661, 16731, TMM 40148-2,

TED1 40537-26, 'I" 40537-33, TMM 40537-61, and USNN 15411.

(a) Left lateral view of skull. (b) Left lareral view of

mandible. (c) Occlusal view of upper left dentition.

(d) Occlusal view of lower right dentition.

Figure 7 appears on the following frame.

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Figure 7.

Downloaded from http://pubs.geoscienceworld.org/gsa/gsabulletin/article-pdf/92/12_Part_II/1982/3418970/i0016-7606-92-12-1982.pdf by guest on 01 October 2021 T ~~ .. 1 a small amount of anterior enamel posteriorly. Thus, with wear, P forms

remaining. The upper canines bear a a dentine peg with a thin strip of

lateral groove in the enamel of the enamel labially and lingually. 2 anteroexternal surface. P is similar to P', but slightly

The upper cheek of Psittacotheriwn larger (Table 6). Unlike P', the

are fairly well known from USNM 15411 enamel on P2 extends further lingually

I (Pls. 23; 31, Figs. -1-2), supplemented than labially. P 2 bears a 'large, P primarily by AMNH 2453, AMNH 15938, and labial paracone and a small lingual AMNH 16731 (Pl. 22). The largest protocone. A small posterior crest

upper cheek tooth probably was P4 or connects the protocone to the paracone. 3 4 M 1 . All of the upper cheek tyth P and P are of similaY size and

are hypsodont with the typical morphology. Boty bear large paracones

taeniodont pattern of greatly extended labially and variably sized, though at 32 enamel lingually. The teeth are most small, metacones. In P -M ,. the

transversely widened and slightly lingual protocones are connecte'd to

spaced, so that they do not contact the pdracones/metacones by minutely

interstitially. cuspidate anterior and posterior 3 4 P 1 (considered P3 by Matthew, 1937, transverse crests. Both P and P

p. 259) is a single-rooted, small bear two small, fused roots labially

bicuspid tooth set obliquely in the and a larger' lingual root which

jaw posterior to the outer angle of curves labially. Near the tooth

the canine. It bears a large paracone, crown all three roots are fused. 1 with an incipient metacone, and a .M Wars a small conical paracone.

smaller protocone. The enamel is A metacone at the labial edge of

greatly extended both labially and the tooth is closely appressed to the 1 lingually on PI, but limited to the paracone. M lacks a stylar shelf

top of the crown anteriorly and of ectocingulum and there is a small,

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minutely cuspidate parastule. M1 are less well known than the uppers

bears a minutely cuspidate postmeta- (gl. 22). P1-4 arc single-rooted,

conule wing which extends postero- whereas M1-3 are double-rooted, though

labially to a point posterior and just their roots are fused. P4 is the

lingual of the metacone. largest tooth. M1-3 are hypsodont 2 1 M is similar to M , but the meta- with enamel extending further

cone is smaller and less distinct than labially than lingually.

on M1 and there is only an incipient The PI of AMNH 754 (Pl. 24, Figs.

parastyle. The postmetaconule wing 4, 5) is relatively small and poorly

is well developed-. preserved. It is obliquely oriented

AMNH 15938 and AMNH 16731 include in che jaw and may have been bicu$pid 3 two M ‘s illustrated by Matthew (1937, like P2. The small and shallow P1

p. 260-261, Figs. 6k, 65; also alveoli of AMNH 754, AMNH 88383, and compare P1. 22, Fig. 21) that are LJK 8035 indicate that Psittacotherim

single-rooted due to fusion of the may have nearly lost PI. 3 original roots. M is ovoid in The P2 of the type specimen of

cross section, with the long axis P. multifraqwn, AMNH 3413, is

dcirected labiolingually. It bears unworn. It is transversely oriented,

a large and isolated labial paracone simple and bicuspid, bearing a large,

connected to the large and anteru- labial paracone and a smaller internal

lingually placed protocone. The protocone.

protocone is connected to the The P3 of Psittacotheriwn is not

paracone by cuspidate pre- and postproto- certainly known. However, squared

cristae. Posterolabial of the proto- alveoli preserve in the lower jaws

cone on the postprotocrista is a minute of AMNH 754 and AMNH 3418 indicate

hypocone. that like P4, P3 may have been

The lower cheek teeth of Psittacotheriw submolariform.

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AMNH 15938 and MlNH 16661 (Pl. 22, conids variably connected by minutely

Fig. 28) include moderately worn P4's cuspidate protocristids. There are

of Psittacotherirn. P4 is a sub- minute paraconids and minutely cuspidate

molariform and transversely bilophodont paracristids anterolingual of the

tooth. The anteroposteriorly compressed metaconids. The talonids bear small,

trigonid bears a large metaconid and relatively high hypoconids and lower,

protoconid forming a transverse crest. cuspidate postcristids and entocristids.

A small paraconid is sljghtly labial Associated with USNM 15413, right

of the center of the tooth. P4 also and left dentary fragments with un-

has a small, posterolingual, antero- erupted N3's, are what appear to be the

posteriorly compressed, talonid which dC1 and dP Psittacotheriwn multi- 1- 2 of bears a large entoconid connected to fragum4 (Pl. 27, figs. 4-10). When

the protoconid by a prominent post- compared to the C1, dC1 is sharply

cristid. pointed and re1atively.compressed

The lower mblars of Psittacotheriwn laterally with a thin, posterior,

are unworn in AMNH 3413,aMNH 3416, enamcl-free portion. DP1-2 are

AMNH 16661 (Pls. 21, 22), UNM NP-220, extremely similar to P2 (PI. 21,

and USm 15413 (Pl. 27, Figs. 11-14). Figs. 7, 8) in being simple, bicuspid

M1-3 are essentially identical but teeth with tall and thin labial

decrease in size posteriorly. M1-3 cusps that are slightly inclined

are shallowly rooted, transversely lingually and lingual cusps that are

bilophoaont teeth with anteroposterior- approximately half the height of

ly compressed trigonids and talonids. the labial cusps and are slightly

The talonids of MI-3 are narrower than inclined labially.

the trigonids and placed relatively Several skulls, mandibles, and

lingually. The trigonids bear sub- various postcrania of Psittacotherium

equal, !conical metaconids and proto- (Pls. 23-26, 28-31) were described

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and illustrated by Wortman (1897b, p. sec. 8, T. 19 S., R. 6 E., Emery

71-78, Figs. 4-21) and Matthew (1937, County, Utah.

p. 255-270, Figs. 62-67; PlS. 59, 60, Discussion. USNM 16204, a right

61, Figs. 2, 3; P1. 62, Fig. 5; P1. 64). 13, is morphologically identical to

Their descriptions need not be added the lower incisors of both AMNH 3413

to here, except to note that the (the type specimen of Psittncotlieriim

vertebrae and pelvis (AMNH 2455) mirltifz~agum)and AMNH 3394 (the type

d4cribed and iglustrated by Wortman specimen of Wortmania otariidem).

(1897b, p. 82d87, Figs. 15-20), However, it is intermediate in size

ostensibly of Psittacotheriwn multifragm, between these two, and therefore I am probably belong to Pantolambda. cavikctus reluctant to definitely assign it to (Simons, 1960, p. 19-20). Matthew ei ther taxon.

(1937) also questioned Wortman's Ectoganus Cope, 1874 assignment of AMNH 2455 to Psittacotherim. Ectoganus Cope, 1874, p. 592. yet evidently used it in his recon- Calamodon Cope, 1874, p. 593. struction of the skeleton of P. Dryptodon harsh, 1876b, p. 401. multifrap (1937, P1.. 64). Conicodon Cope, 1894, p. 594.

?Psittacotherim sp. or Wortmania sp. non Calmodon Amaral, 1935, p. 203.

Lanpadophorus Patterson, 1969a, p. 41. (Pl. 27, Fig. 3) !@pe species. Ectoganus gZirifomnis

Stylinodont, near Psittacotherim: Cope, 1874 (= Calamodon simplex Cope,

Gazin, 1941, p. 17. 1874 = Calmodon arcmaenus Cope,

Referred specimen. USNM 16204, right 1874 = CaZmodon novomehicanus Cope,

I3 (Pl. 27, Fig. 3): from early Tor- 1874 = Dryptodon crassus Marsh, 1876b =

rejonian strata of the North Horn ?Psittacotheriwn 1obdell.i Simpson, 1929b =

Formation, Dragon local fauna, NW2, Lmpadophorus expectatus Patterson,

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1949a). Revised diagnosis. Largest species d IncZttded species. The type species of Ectoganus (Tab$ 7).

and Ectogmus cope; Schoch, 1981b. Ectogcmus g Zirifoks g Ziriformis

Distribution. Tiffanian-Wasatchian Cope 1874

of Colorado, Clarkforkian of/%tana, (Tables 7, 9, 10: Figs. 8, 9:

Clarkforkian-Wasatchian of Wywing, and Pls. 32, Figs. 2-10, 12-33; 35,

Wasatchian of New Mexico. Figs. 20-22; 36; 37, Figs. 1-8,

Revised diagnosis. Medium- to large- 15-21; 38, Figs. 5-22; 41, Figs.

sized taeniodonts; canines enlarged, 1-12; 42; 46, Figs. 1-4) 0 rootless, and compressed posteriorly Ectoganus g Zirifads Cope, 1874,

with enamel limited to the anterolabial p. 592.

aspect; cheek teeth moderately to CaZamodon s%Zex Cope, 1874, p.

extremely hypsodont; transversely 593.

bilophodont; P3-[, submolariform t'd Ca Zamodon arcmuenus Cope, 1874,

molarifom with talonids lingually p. 593.

placed and narrower than the trigonids; CaZamodon novomehicanus Cope, 1874,

M1- 3 transversely bilophodont with p. 594. subequal trigonids and talonids. Dryptodon crassus Marsh, 1876b,

p. 403. Ectoganus gZiriformis Ectoganus novomehicanus: Cope, 1877,

Ectoganus glirifornris Cope, 1874, p. 592. 159.

(see synonymies under the subspecies) Ectogmus glirifornris: Cope, .1877,

Bjpe subspecies. Ectoganu's gliri- p. 160.

formis gZiriformis Cope, 1874; CuZumodon arcmuenus: Cope, 1877,

IncZuded subspecies. The type sub- p. 163.

species and Ectoganus gZiriformis Cazam&don simpZex:: Cope, 1877, p. 189.

Zobdel Zi (Simpson, 1929b). CuZumpdon simplex: Cope, 1884c, p. 189.

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TABLE 7. DENTAL MEASUREMENTS OF ECTOGANUS GLIRIFORMIS

C1 Pl P2 P3 .P4 L W L W L W L W L W L W -E. g. pliriformis

AMN€l 4286 4287

16244 12-0* 18.5* 13.0 17.9

16245 16771 12.6 7.7 21.0 11.9 14.9 10.9 18.5* 17.P 12.5* 16.5* 10.5* 16.0*

48001

86859

CM 11497 16.7 9.5 34.0* 17.4 16.5" 17.5* 13.5 17.8 13.9 19.6 PU 13173 14.9 18.9

UM VP-6000

UNM B-970/971 12.0* 16.0* 11.3 15.7

USNM 1012b 14.2 15.7

1017' 1102d* 11.5* 17.5* 1137' (unmeasurable)

YPM lllOOe N 3 3 4 4 5 5 Mean 15.50 17.5 12.23 17.13 13.3 17.62

Standard Deviation 3.61 0 .93 1.26 1.71 1.73

Coefficient of 23.3 0 7.5 7.4 12.9) 9.8 Variation

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M1 M2 M3 I3 c1 p1 p2 L W L W L W L W L W L W L W

9.5" 7.6 29.2 16.1 17.6 12.9 19.1 16.7

16.5* 22.01s

12.8 15.4

14.8 16.3 15..2 14.5 11.1 11.9

12.2 16.5

8.1 6.3 26.5 15.0 14.2 12.0 14.5 16.5

13.3 14.7 12.9 15.9

6.7* 6.0 17.3 16.5* 11.5*

3 3 3 3h33

13.63 15.47 16.13 16.10 12.13 16.70 18.40

1.04 .80 1.15 1.73 .71 2.31 3.12

7.6 5.2 7.1 10.7 5.9 13.8 17.0

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p3 p4 M1 M2 M3 L W L W L w.L W L W

12.4 13.1 13.8 14.3 12.7 13.2 14.6 14.2 14.2 13.8

15.8 15.6 15.3; 14.7

14.4 13.0*

14.7 15.7 13.7

15.8 15.1 13.7 13.7

16.3 16.4

13.1 14.0 16.4 14.9 13.5" 15.8 14.0* 17.5*.13.4 14.7

14.6 14.2 14.2 14.1

15.1 13.0

16.0* 13.5* 16.0* 15.1 14.5" 1315".. 13.5* 14.5* 14.5* 15.0*

5 4 8 8 6 6 4 4 3 3

14.18 13.40 15.55 14.91 13..98.14-.17 14.30 14.80 14.03 14.50

1.32 .45 .89 .85 .90 .95 .70 1.91 .57 .62

9.3 3.4 5.7 5.7 6.4 6.7 4.9 12.9 4.1 4.3

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.. a.

-E. g. lobdelli: AMNH 22234f

22235 17.3 10.1

CM 11560 F"H PM 241

P 14906

P 15569 13.7 17.5

P 15575 14.1 17.9 P 26083g 12.0* 17.1

PU 18345 12.6 15.5 18954

18982 12.4 16.3

18994 13.8 16.5 20864

21499 11.5* 19.3" 13.1 18.2 13.7 18.9

N 4 4 4 4 Mean 12.8 17.28 13.55 17.20 Standard Deviation .75 .79 .66 1.50

Coefficient of 5.9 4.6 4.9 8.7 Variation

a Type specimen of Ectoganus gliriformis.

bType specimen of "Calamodon simplex". C Type specimen of "Calamodon arcamaanus":

%ype specimen of "Calamodon novomehicanus". e Type specimen of "Dryptodon crassus".

f"ype specimen of "?Psi t tacotherium" lobdelli.

%ype specimen of "Lampadophorus expec tatus".

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13.8 14.5

27.0 16.3 15.6 8.2

12.5 14.3

11.4 13.4

13.4 19.5 12.4 19.3 12.8 16.1 12:6 14.9

13.2 17.4

4 4

12.58 14.28

.98 .63

7.8 4.4

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14.6 15.5 14.9 15.5

13.1 12.1

14.7 14.9

16.4 14.5 14.3 12.6

14.9 15.8 14.3 12.7 13.7 12.9 13.5 11.7

16.3 15.6 16.4 16.0 14.4 14.7 14.6 13.8

5 5 4 4

15.46 15.34 14.03 13.03

.86 .63 .62 1.15

5.6 4.1 4.4 8.8

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TABLE 8. DENTAL MEASUREMENTS OF ECTOWNJS COPE1

C1 Pl P2 P3 P4 -E. 2. copei: L W L W L W L W L W L W

AMNH 15633

PU 14689

USGS 3838 14.3 6.7 24.0* 12.0* 13;5* 6.5 9.5 11.0* 11.5* 11.5"

USNM 12714a 10.9 6.7 19.0 10.6 10.7. 7.7 10.7 13.7 9.9 12.3

YPM 18618' 12.0 13.3

N 3 3

Mean 10.47 12.2

Standard Deviation 1.34 1.15

Coefficient of 12.7 9.4 Variation

-E. 2. bighornesie:

AMN€l 86852

PU 14678b 10.7 14.0

PU 18052 10.7 13.3

specimen of Ectoganus c. copdi.

bType specimen of E. 2. bighornensis.

C The identification of this tooth as a P3 is uncertain; it may be a 4 1 P orM.

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-~ ~~~ ~ ~~ ~ M1 M2 M3 I3 c1 p1 p2 L W L W L W L W L W L W L W

11.8 13.2 11.0 18.7

25.4 14.1 16.4 11.5 11.8* 18.9

11.0* 11.0* 22.9 15.0* 13.0" 14.5* 12.0" 17;0*

11.2 12.9 10.5 la2 10.8 18.8

3 3 4 4

11.33 12.36 11.40 18.35

.42 1.19 .59 .90

3.7 9.6 5.2 4.9

10.5 12.6 10.2 11.7 8.7 9.8

9.9 12.2

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p3 p4 M1 M2 M3 \ L W L W L W L W L W 10.9 -11.4

ll.O* 11.9

12.9 11.9* 11.8 12.0

11.8 11.0

11.5 11.1

11.5 11.3 11.7 11.3

4 dP3 dP dP4 L W L W L W

AMNH 16771a 10.9 12.3 10.9 17.6

USNM 1137b 10.3 14.1 9.8. 17.4 12.7 9.6

USNM 12714' 9.8 16.1 12.1 10.4

a Specimen referred to E. g. gliriformis.

bType specimen of g. g. glirifomis: 'Type specimen of E. c. copei.

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P3 M1 p2 M3 L W L W L W L W

Ectoganus Rlirifokmis:

4 N 8 8 4 4 5 5 10 10

Mean 12.56 17.20 13.53 15.95 i5p18.80 14.00 13.71

Standard Deviation .82 .98 :.a8 1.12 2.67 2.27 .$ .76 1.14 * '. Coefficient of 6.5 5.7 6.5 7.3 17.6 12.1 5.4 .a.3 Variation

-E. copei: N 5 5 5 5 4 4 3 3

Me,"" 10.56 12.78 10.88.12.38..l~.IR) 18m.35 11.97 11.43

Standard Deviation .95 1.16 . .72 .86 ' .59 .90 .81 .42 I Coefficient .of 9.0 9.1 616 6.9. 5.2 ,p.4w.9 6.8 3.7 7 Variation '..

All Ectoganus :

N 13 13 9 9 9 9 13 13

Mean 11.79 15.50 12.06 13.97 13.50 18.50 13..53 13,.18

Standard Deviation 1.31 -2.45 01.58 2.10 2.77 1.72 1.15 1.41 Y Coefficient of 11.1 15.8 13.1 15cf 20.5 9.3 8.5 10.7 Variation

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CaZmodon simplex: Wortman, 1897b, p.81 canine fraginents (type of “Calamodon

Calamodon arcarnnaeus (lapstis calami) : simplex”; P1. -32, Figs. 8, 13, 22,

Wortman, 1897b, p. 89. 23); USNM lOly, right and left dentary Ecotganus gliriform-s:3 Gazin, 1936, fragments, right M2 and caninel frag- p. 610 (in part). ments (type of “Cahodon arcamaenus”;

Ectoganus cf. simplex: Guthrie, P1. 32, Figs. 2, 12); USNM 1102, right 2 1967, p. 23. P (type of “Calpnodon novomehicanus”-

Ectoganrts simplex: Schankler, 1980, P1. 32,. Figs. 4, 14, 15): all from

b. 104. Wasatchian strata of the $an Jose

Type specimen. USNM 1137, right Formation, probably in Almagre Arroyo, 3 1 and left I and C fragments, (?)righ.t San Juan basin, New Nexico. dp3-4 ,- upper (?)deciduous incisor YPM 39805, left radius (Pl. 46,

fragment, right P2, left dP4, partial Figs. 3, 4): from Wasatchian strata

lower molar trigonid, partial lower of the San Jose Formation, San Juan

molar talonid, fragmentary upper molar, basin, New Mexico. 3(?) and associated bone and tooth frag- UNM B-970, left P ’(?)’ right P ,

ments (PI. 32, -Figs. 5-20, 16-21), iieht M2(?), left P4(?), right M1

Horizon andlocal& of the type. (Pl. 38, Figs. 5-10, 14-20); UNM B-971,

Collected by E. D. Cope in 1874 from right M (Pl. 38, Figs. 11-13);

Wasatchian strata of the San Jose UNEl B-973, canine fragments, ‘partial

Formation, probably in Almagre Arroyo, ulna, scapula and bone fragments (UNM

San Juan basin, New Mexiqo. B-970/971/973 were all collected

.Referred specZmens. USNM 1001, lqf t together and portain to a single

P2, molariform c6eek tooth, fragments indiadual) : all from Wasatchian‘ tl . of scapula, right humerus, ulna, radius, strata’of the San Jose Format$on,

magnum, ungual phalange, ana otner Go%ernador area, $an Juan basin, New 4 *bone.fragmefits; USNM 1012, left ‘P and Mexico.

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AMNH 86859, upper M3(?), right P4, Creek, Bighorn basin, Wyoming. 1 left M1 and canine and bone'fragments CM 11497, left 13, right C , 2-4. '(Pl. 35, Figs. 20-22): from Clark- (?)right P . from Wasatchian strata

Sorkian-early Wasatchian strata of of the Willwood Formatbon, 4.8 km S. W.

the "lower variegated beds" (see of the mouth of Elk Creek, Bighorn

McKenna; 1980a, p. 330), Togyotee Pass basin, Wyoming.

area, Purdy basin, northwestern UM VP6000, left dentary with C1-P2,

Wyoming. M1, and roots of right left T M3 and 3' AMNH 4286, mandible with right and left P3-4, M2: from Wasatchian

left 13, right C1, right and left P1, strata 'of the Willwood Formation, ap-

left P2-3, right and left P4-M3 (Pl. proximately halfway between Worland and

42, Figs. 1, 2) ; AhH *4287, ief t-' P2, Meeteetse (Fifteen Mile Creek drainage,

.right P4, right and left M1 (Pl. 37, Cottonwood Creek), Bighorn basin,

Figs. 19-21); AMNH 16771, right upper Wyoming.

(?)deciduous incisor, right and left. AC 2879, right dentarp fragment with 32 I -M , right dP3-4, right M3 and bone 13' C1' p1L.2, and M3: from Wasatchian

fragments (Pl. 36, "Figs. 1-12, 14-17) : strata (Lysite) of the Wind River

all from Wasatchian strata of the Formation, Wind River basin, Wyoming.

Willwood Formation, Bighorn basin: Revised diagnosis. Large Ectoganus

Wyoming. with all teeth extremely hypsodont;

PU 13173, (?)right P3-4 and left P4, incisors and P1-2 approach the totally 1- 2 A (Pl. 41, Figs. 1-12); USNM.no number, rootless condition of the canines.

.canine and molariform tooth fragments, Ectoganus glirifomis ZobdelZi (Simpson, partial left maxilla, partial left 1929b) '

femur and other bone fragments (Pl. 46, (Tables 7, 10: Fig. 8: Pls. 32,

.Fig+4/2) : both fkom early: Wasatchian Figs. 1, 11, 34, 35; 33; 34; Figs.

strata of the Willwood Formation, S. Elk 1-18; 39; 40)

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Psittacotheriwn sp. indet.: Simpson, FMNH P 26083, skull with right and 3 1929a, p. 121. left 2, P~-~,left M , alveoli for

?Psittacotheriwn Zobdeili Simpson, 1929b, right and left M1-2, right and.

p. 11. left distal humeri, (?)left scapula,

?Psittacotheriwn sp. : Patterson, 1936, rigk' and left ulnae, right and left

p. 397. femora, right and left tibiae, forefoot

Lampadophorus expectatus Patterson, 1949a, unguals and other bone fragments (type

p. 42. of ,"Lumpadophorus expectatus"; Pls. 33;

Lconpadophorus ZobdeZZC: Patterson, 34; 35, Figs. 1, 2); FMNH P 14906,

1949a, p. 42. right p3-4 (?) (P1: 35, Figs. 13-16); 3 Type specimen. AMNH 22234, right M FMNH P 14954, fragmentary lower left

(Pl. 32, Figs. 1, 11). molar (Pl. 35, Figs. 11, 12); FEN 4 Horizon and locality of the type. P 15575, left P (Pl. 35, Figs. 3, 4);

Clarkforkian strata of the Fort Union FMNi4 P 15008, fragmentary lower molar;

Formation, Eagle Coal Mine, BeAr FMNH 15569, right P3 (Pl. 35, Figs.

Creek, Montana. 5, 6); FMNH P 26106, right M l(?) (listed

Referred specimens. AMNH 22235, by Patterson, 1949a, as P 26093; PI.

left I3 (Pl. 32, Fig. 35); CM 11560, 35, Figs. 17, 18) and lower (?)left

right C1 (Pl. 32, Fig. *34): both from canine; FMNH 26090, left humerus (P1.

Clarkforkian strata of the Fort Union 38, Figs. 21, 22); FMNH P 26101,

Formation, Eagle Coal Mine, Bear lower molar talonid (Pl. 35, Figs. 7,

Creek, Montana. 8); FMNH PM 241, left I3 (Pl. 35, Figs. 3 PU 18345, right P3 and right M 9, 10) (the above FMNH specimens com-

crown: from a carbonaceous clay above pose the original hypodigm of

Coal #3, Clarkforkian strata of the "Lcmrpadophorus expectatus") : all from

Fort.Union Formation, Foster Mine, Bear late Tiffanian-Clarkforkian(?) strata

Creek, Montana. of the ldasatch ("DeBeque") Formation,

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3- 4 Plateau Valley local fauna, Mesa left M3, left P2(?), right and left P ,

County, Colorado. right MI, incisor, canine tips, bone

PU 18954, right M1 and twQ rooted and. tooth fragments (Pl. 40): from

incisors (deciduous?) ; PU 18982,. right Clarkforkian strata of the Polecat Bench

P3(?), right and left P4: both from Formation, NWk, sec. 27, T. 57. N., R.

Clarkforkian strata of the Polecat Bench 100 W., Bighorn basin, Wyoming.

Formation, west side of Polecat Bench, Revised dia*qnosis. Large Ectoganus 2 NWk,. sec. 21, T. 57 N., R. 100 W., Big- with P2 only slightly more hypsodont

horn Basin, Wyoming. than in Psittacotheriwn; posterior 33 PU 18994, right P2(?), right P 4(?) , cheek teeth (P -M moderately hypsodont 33 right P4, right and left M1 and mis- with relatively low, rounded, bulbous

cellaneous bone fragments (Pl. 39, crowns, and relatively shaTlbQ, com-

Figs. 16-24): from Clarkforkian strata pressed roots.

of the Polecat Bench Formation, west

t Ectoganus copei Schoch, 1981b side of Sand Coulee, Sa, sec. 14, T. 57 N., K. 100 W., Bighorn basin, Ectoganus cope; Schoch, 1981b, p. 000.

Wyoming. (see synonymies under the subspecies)

PU 20864, left M2(?), right and Ripe subspecies. Ecto&ms copei

left M 3(?) , le,ft P2(?), left P4-M2, cop& 2cboch, 1981b.

right and left M3 and miscellaneous IncZuded stlbspecies. The type

tooth and bone fragments (Pl. 39, Figs. subspecies and Ectoganus copei big-

1-15): from Clarkforkian strata of the hornen.& Schoch, 1981b.

Polecat Bench Formation, SFk, N~L, Dia.qnosis. Smal1es.t species of < see. 17, T. 54, *N., R. 96 W., Bighorn Ectoganus (Table 8).

basin, .Wyoming. Ectoganus copei cope{ Schoch, -1901b PU 21499, ,partial ?eft PZc?),

left P3, right and left P4, right M1, (Tables 8-10: . Figs. 8, 9: Pls. 38,

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Flgs. 1-4; 43; 44; 45, Figs. 1-4; 46, PU 14689, right and left dentary

Figs. 5-9) fragments with partial right and left

Ectoganus gZirifoks : Gazin, 1936, p. C1, right P1-2: from Wasatchian strata

597 (in part) of the Willwood Formation in the area

Ectoganus cope; copei Schoch , 1981b, of sections 16, 17, 20, and 21, T. 47.

p. 000. N., R. 93 W., 11.3 km northwest of

njpo specimen. USNM 12714, skull Worland, Bighorn basin, Wyoming.

and mandible with right and left I3 - USGS 3838, left maxilla and partial 14 2 M (P ‘s unerupted), left M , right right premaxilla dith roots of right 2 3 11 and left dP4, alveoli for right M and left I , left C -M and partial 3 and left M’, roots of right and left crown of left P , left dentary fragment

C1, right and ‘left P2, left P4 (un- with roots of C1-P2 and partial P3

erupted),. left dP4, right and left crown and fragments of the skeleton

M1, left M2, right and left M3, roots (Pl. 46’, Figs. 5-9) : from Wasatchian

of right P4, alveoli for right M1-2 strata in the SWk, ,S*, sec. 35, T.

(Pls. 43: 44; 45, Figs. 1-4). 48 N., R. 94 W. and in the NF&, NEk,

Horizon and Zocalittj of .the type. “5, sec. 2, T. 47 N., R. 94 W.,

Wasatchian strata of the Willwood Washakie County, Bighorn basin, Wyoming.

Formation, 13 km northwest of YPM 18618, right P 3(?) and other

Worland, Bighorn basin, Wyoming. tooth fragments el. 38, Figs. 1-4):

Referred specimens. AMNH 15633 (more from Wasatchian strata of the Willwood

than one individual), two right P2’s, .Formation, Sa, sec. 25, T. 49 N., R.

right and left P3, left M1’(?), and 97 W., Bighorn basin, Wyoming.

other tooth fragments (Pl. 37, Figs. 9- Diagnosis. Small Ectogirnus with

14): from Wasatchiah strata of the all teeth extremely hypsodont; incisors 1- 2 Willwood Formation, Fifteen bij le Creek, and P1-2 approach the totally rootless

Blghorn basin, Wyoming. condition of the canines; cusps on.

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upper premolars larger and better forkianearly Wasatchian strata of

developed than in E. c. bighornensis. the "lower variegated beds" (McKenna,

1980, p. 330), Togwotee Pass area, Ectogaxus cope; bihornensis Schoch, 1981b Purdy basin, northwestern Wyoming.

(Tables 8, 10: Fig. 8: P1. 35, Fig. Diapos&s. Small Ectoganus with

19; P1. 45, Figs. 5-19) relatively low crowned and shallow-

Ectoganus cope; bighornensis Schoch, rooted cheek teeth; cusps on upper

1981b. premolars smaller and not as well

Bzpe specimen. PU 14678, right and developed as in E. c. copei- 3 left P , right ,M1-2, left M3, right Taxonomic Distinctions in Ectoganm M1 and canine fragments (Pl. 45, Figs.

5-19). As here revised, llLurnpadophomsll

i!orizon, and ZocaZitrJ of the type. Patterson, 1949a, .is considered to be

Early Wasatchian strata of the lower congeneric with Ectoganus Cope, 1874.

Willwood Formation, southern tip of Patterson-Is (1949a, p. 42) original

Polecat Bench, T. 55 N., R. 100 W.., diagnosis of "Lampadoph wus" read3

Bighorn basin, Wyoming. as follows:

Referred specimens. PU 18052, right Canines rootless, enamel-free por- 3 1 P ;left M , left M2, partial left,M1 tions more compressed than in

and enamel fragments: from Wasatchian Psittacotherim. Cheek teeth with

strata of the lower Willwood Formation, cement at bases of crowns, roots with

southed tip of Polecat Bench, sec. lo1 vestiges of former.ldivisions; crowns

T. 55 N., R. 100 W., Bighorn basin, higher than in Psittacootherium,

Wyoming. 1&er than in Ectoganus; little or

Ecotganus cf . E. copei bighornensis : no tendency toward develqpment of

AMNH 86852, left M3, and canine frag- enamel-free bands on anterior and

ments (Pl. 35, Fig. 19): from Clark- posterior faces. P1-2 smaller than

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in Ectoganus,more crenulated than does not include the roots of any of 3 in Psittacotherim. M wider thar? the cheek teeth. However, they are

in Psittacotherim, hypocone less present on Patterson's referred speci-.

isolated than ih Ectoganrts. 3-4 mens- (P1. 35, Figs. 3-18) and on the

with independent talonid crests. type specimen of "L. ZobdeZZi ,'I and

None of these characteristics serve to do not differ from the roots of prbvi-

clearly distinguish "Lcmrpadophorus" ously accepted specimens of Ectoganus

from Ectogamts and each point is ad- (for example, USNM 1017). The "vestiges r dressed below. of former divisions" of the roots

Rootless canines with relatively mentioned by Patterson (see quote above)

compressed enamel-free portions are are variable, but seem to be no better

comman to both E'ctoganub and Stylinodon. 'developed in Patterson's "Lampadophorus"

Whether cement is preserved at the base than in previously acceptedfspecimens of.

of the enamel on the cheek teeth'ap- Ec'togunus (compare Pls. 32, 35).

pears to beean artifact of preservation, Crown height also appears to be ex-

Many specimens of-Ectoganus (senszc tremely variable in "Lmpadophomcs"- " Pattgpon) have 'cement at the base of Ectoganus. This may be partly due to

the enamel: USNM 1017 (PI. 32, Figs. the fact that crown height of the 2, .11,2),. the type specimen of "CaZomodon cheek teeth in Zctogcntus increases 1 zrcamadnus" Cope, 1874, as an example. anteriorly (that is, PI generally is

highest crowned and b13 lowest crowned) Furthermore, this, chargctek is not 3

preserved on either tlie type specimen and often it is difficult to position

of "L. expectatus" (FW P 26083, P1. isolated teeth in a tooth row accurately.

35, Figs. 1, 2) or the type specimen Thus, alleged crown height differences

of "L. Zobdelli" (ILhH 22234, P1. 32, can be largely an artifact of comparing

Figs. 1, .11). teeth 'which are from different positions

The type specimen of "L. expectatus" in the tooth roW. Furthermore, whereas

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the Crowns. of Patterson's type and not preserved in the type specimen or

referred specimens of "Lcmrpadophorus" among the referred specimens of. 1- 2 are higher than those of Psittacotheriwn, "L. expectatus." The alveoli for P

they are not demonstrably lower than on FMNH P 26083 are incomplete, dis-

all specimens .of Ectogmus (sensu Patter- torted, and unmeasurable. Further-

son;,compare Pls. 32, 36, 37, 41, 45). more, as P1-2 of Ectoganzis erupt, their

Anterior and posterior enamel-free dimensions change.

bands are best developed on the anteriot P3-4 of Ectoganus have independent 1- 2 cheek teeth of Ectogmus. (P,L2). talonid crests. The type specimen

Patterson, however, did not havelche of "LQniPadOphorus lobde Z 2i" (M ant%rior cheek teeth of '1LumpadopJzoms8' 22234) here is identified as a right

(see list of his referred specimens M3 (Patterson, 1949a, considered it to 4 f oro"Lurnpadophoms I' above) . The be a P ). The posterointernal cusp

posterior cheek teeth which he did hive ("hypocone'.') is less isolated on AMNH

show a definite "tendency towarb 22234 than the same cusp on the M3 of

development of enamel-free bands'' FMNH 26083, the type.specirnen of "L.

(compare P1. 35, Figs. '4, 6, 14, 18). expectatus". However, given the 3 Although enamel-free bands are better general variability of M Is (Gingerich,

developed (along with increased 1974; Gingerich and Schoeninger, 1979;

hypsodonty) in some specimens of Gingerich and Winkler, 19791, I would

Ectoganus , the develbpment of enamel- not judge this chafacter to be of much

free bands here is considered an ex- taxonomic value. The M3 of FMNH P

tremely variable character and worthy 26083 does not differ sZgnificantly

only of subspecific recognition. from the M3 of AMNH 16771 (Pl. 36,

It is impossible to say how Patterson Fig. 171, an upper dentition of

detehined that P1-2 of "Lampadophorzts" from the Wasatchian of the Bighorn

are smaller than in Ectogamcs; P1-2 are 1 basin, Wyoming.

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- 2 As Patterson already noted (1949b, M1 in

p. 251), the only other cheek.teeth The dP3-4 of USNM 1137 also are

preserved in the type specimen of indistinguishable from those of AMNB 3- 4 "Lampadophorus expectatus ," the P , 16771. However, the deciduous teeth

agree "almost cusp for cusp and groove of Ectoganus do not appear to be diag-

for groove" with the P3-4 qf Ectoganus nostic at the specific level. As 4 (AMNH 16771, P1. 36). Thus, MiP Gazin (1936) noted, the dP4 of USNM 1137

26083 and AMNH 16771 are morphologically also are indistinguishable from the 4 indistinguishable from one another and of USNM 12714 (see Table 9), the 4 *- must be placed in the same species. type specimen of Ectoganus copei (Pl.

The comparable parts of .AMNH 16771 45, Figs. 1-4). 'A lower molar trigonid

and USNM 1137 (Pl. 32, Figs. 5-10, 16-21, andValonid of USNM 1137 also are larger

24-33), the type specimen of Ectoganus than those of the lower molars of 1 gZiriformiS, dP3-4, a damaged M or E. copei . 2 M , and 'irncisor and canine fragments, Taxonomic problems within the genus

are indistinguishable. The upper molar Ectoganus are similar to those en-

of USNM 1137 is crushed and broken, countered with Psittucotheriwn(see

but is a bilophodont tooth bearing earlier discussion). Many specimens

cuspidate, transverse, anterior, and of Ectoganus are fragmentary, isolated,

posterior crests like those of AMNH and heavily worn teeth. Taxonomic

16771. The enamel distribution and decisions based on these specimens cannot

degree of hypsodonty appear to be! be made bith any degree of confidence. 1 identical in the left M of AMNH 16771 However, Ectoganus is better repre-

and the upper molar of USNM 1137. Con- sented by well-preserved specimens

sidering the crushed nature of USNM than is ?sittucotherium, and thus finer

1137, they also appear to be identical taxonomic distinctions can be made.

in size, and both are' larger than the Hcrc I formally recognize two species,

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each composed ot two subspecies, to 12) : and "Ca lamodon novomehicanus ," 2 encompass most known Ectoganus material. based on a rightP (USNM 1102: P1. 32,

The decision was made to recognize Figs. 4, 14, 15). When Gazin 11936)

subspecies on the basis of characters described USNM 12714 (Fig. 8; Pls. 43,

which are variable (forming a seemingly 44), a skull of Ectoganus, he noted

continuous grade) but recognizable that Cope's three types appear to belong.

in over 75% of the specimens (compare to a single species. Cope based his

Simpson, 1943, 1961, on subspecies). species on distinctions between teeth

At the specific level, I recognize which he believed to occupy the same

large (Ectoganus g ZiKgformis ) and positions in the tooth rdw. However,

small (Ectoganus cop&) species of by com'parison with the complere denti-

Ectoganus (compare Gazin. 1936, and tion of USEM 12714, Gazin recognized

Guehrie, 1967, p. 23, who-also recog- that these teeth o

nized a large and a smalf' species). t'ions in the tooth row. Gazin (h36)

As demonstrated above, the type considered the type of E. gZirifomis to

specimens of Ectoganus gZiKformis (US" represent the same, dentally small ,

11371, "?Psittacotheriiim Zobde Zli" species as USNM 12714. He also recog-

(AMNH 22234) and "krnpadophorus expectatus' nized that there might be a second,

(FMNH P 26083) all belong to the large larger species of Ectoganus which he

species of Ectogazus (Table 7). Cope provisionally called Ectogcmus shiplex

also described three other species .of (Gazin, 1936, p. 611). However, he

Ectoganus (= Ca Zamodon 1: "Ca Zamodon also tentatively subsumed all four of 4 simplex," based on a left P and canine Cope 's names under Ectoganus gliriformis

fragments (USNM 1012; P1. 12, Figs. 3, (Gazin,' 1936, p. 611).

13, 22, 23); "Calamodon arcamaenus ," As demonstrated above, USNN 1137, hhe

based on a right M2, canine and dentary extremely fragmentary type specimen of

fragments (USNM'1017: P1. 32, Figs. 2, Ectopanus gliriformzs also pertains to

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the large species of Ectogmus. Thus expected for a single species (compare

the correct name for the large ,species Simpson and others, 1960). In contrast,

is zctoganus gziriformis (= CaZmodon for E. gZiriformis alone they range from 5.4

sirriplet). USNN 12714 pertains to a to 8.3 and for E. copei alone they range 4 distinct small species of Ectoganus, from 3.7 to 9.1. From the frapentary

E. copei (Table 8), which at present material known, Ectoganus copei 'also I is distinguished from E. gliriform's appears to have had a relatively smaller

only by its smaller size. The type skull (and presumably body) than

specimen of "Dryptodon crassus" (YPM Ectoganrts qzirifo~::. Thus, USSM 12714 11100; P1..42, Figs. 3-5) is an (E. copei) and F"H P 26083 (E.

incomplete mandible with heavily worn gZirifoimis) arc both skulls of younp

teeth which falls in the size range individuals of comparable age, yet not

of E. gzirij'omis (Table 7). As Gazin only do they differ in the size of the

notcd (1936), it is indistingriishable teeth, but the size of FMSH P 26083 is

from Cope s Ectoganus and thus relatively,lar,qer; for example, the

"D. cras.=zts'' is a junior subjective length from the anterior fape of the

synonym of E. ?Zirifo>+s. . canine to posterior biz is 88 mm in USLW

Fctoaanus _oZiri,Com*sand Z. cocci .12714, and approximately 105 mm in FT.lhW

are easily recognizablc visually and P 26083. However, sexual dimorphism

when.coefficients of vaciation are cannot be ruled out, so I do not attach 1 \ calculated for P3, M , and (Table much taxonomic significance to these M 1 10, Fig. 8; the only teeth for which differences in size.

there is enough reliable information Within each species, I recognize

for both species: P2 is difficult to two subspecies: E. nliir:fopm:s is

I measure and chMs shape with erup- divided into E. g. qZi'y,.forn;is and I?. 9.

tion) they vary from 8.5 to 15.8,f~r. ZobdeZZZ; E. copci is divided into

the pooled data, which is higher than E'. c. cope; and I?. c. binhorwcnsis.

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I A

A A 0

9- 15-

14 -

8 13- 8 8 + 12 - P3 M’ 11 - 8 -L lo! L 9 10 11 12 13 14 9 io ii i2 i3 1’4 ii Y W all miawrements in mm 16i 0 Mi

13 0 -- A

Aa 8b +

11 + I I I I I a. isolaled tooth tl 12 13 14 15 16 t~ estimate

Figure 8. Rivariate plots of P3 width versus P3 length 1 (top left,):, bl width versus M1 length (top right), and M 1 wid.t) versus >f length (bottom) of specimens of Ectoganuc 1 listed in Tables 7 and 8. (Footnotes: a, measurements of

an isolated tooth that is questionably an.M b, approxi- 1’- mate measurements of a damaged tooth.)

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Subspecific dis.tinctions are made on and 40, Figs. 1, 2). In E. q. lobdeZZi 2 the basis of the variable characters the roots of P2 are.shorter and the of degree of hypsodonty, relative root internal and external bands of enamel- development, and over-all crown morphology are relatively wider anteroposteriorly

(compare Simpson, 1943, 1961, .on sub- and do not exfend as far up and down

1-2 ’ specific distinctions). Patterson the roots, In contrast, the P .. 1- 2 (1949b) recognized and tried to use of’E. g. gzififortrtis are deeply rooted,‘

differences in some of these Characters with long, ’thin internal and even

to make a case for the validity of longer, thin external enamel bands.

“Lumpadophoircs. ,The posterior cheek teeth’of E‘. g.

E. g. lobdelli differs from E. g. lobdelli ‘also are less hypsodont than

glififollnis in having relatively fcwer- those of E. g. glir4fomnis, with

hypsodont teeth, with the,exception of shallower, more compressed roots that

the canine which is eve,r+-growing in both sometime?form narrow points. The r‘ forms. The incisors of F:. g. glirifdmk posterio; cheek teeth of R. 5. pZi?Yi-

approach a totally rootless condition formis are gen’erally more hypsodont

and are relatively smaller in legnth than those of B. 9. lobdclli and have 3- 4 and width than those .of ‘R. g. Zobdelli, thicker, blunter roots, l’hc P

which arc less deeply rooted (compare crowns of E. g. lobdclfi are usually

Pls. 35, Figs. 9, 10; 36, Figs. 3, 4; simpler Jhan those of most E. 9. 2 and 32, Fig. 35). P2 in F:. 9. ZobdelZi gZii<,Comlc, lacking well-defined

(PI1 are unknown in B. p. Zobdelli), al- metacones and hypocones and haviny!

though much more hypsodont and more less cuspidate, or smaller cusps, on

deeply rooted than ;n Psittacothciliim, the transverse crests. The lower

are less so than in E. 9. plir7:fo.mis P3-M3 of E‘. g. 1d.BdelZi are relatively

(compare Pls. 32, Figs. 4, 14, 15; bulbous and globose compared to the

36, Figs. 7, 8; 39, Figs. 4-6, 16-18; corresponding teeth of E. 9. gliriformis

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and bear.smal1, but distinct paraconids. specimen of Ectoganus gziriformis, both

P of E. 9. glir”i,Comtis are less 33-M include a pair of upper incisors which 3 bulbous than those of E. a. lobdelli presumably are I .Is. In addition, both

and lack paraconids, or at best these specimens include a smaller (?)upper

cusps arc minute. incisor which may be inner incisors

E. c. t)i~?io~~ti~miisdirfers from (Pls. 32 and 36) comparabte to thc small

E. c. copci in having less hypsodont ‘pair of upper incisors o> Stylinodon (PI.

cheek teethOwhich are more shallowly 48, Fig. 3). PU 18954, here referred

rooted. ‘1he.cusps ‘of the upper pre- to E. glim’formis, also includes two

molars are larger and better developed relatively low-crowned and shallowly E in L‘. c. cop& than in E. c. rooted incisors. One incisor of PU 18954

b-ighorncnsfs (Pl. 45). is much larger than the other, but it

cannot be determined whether these

incisors are uppers or lowers. Thus,

The upper incisors of Bct,opanics (Fig., there is some evidence that Ectoganus

9) mimic the canines by having enamel glim:-forrrL? possessed two upper incisors

confined to an anterior band, and the on either side. Alternatively, the

posterior enamel-free portion is small incisors of USNM 1137 and MI

compressed lateral ly. USNM 12714, 16771 may be deciduous incisors (both

a skull of Ectopniis copei, add UShW 1137 and AMNH 16771 are the

USGS 3838, left and right premaxillae dentitions of young individuals that

and a left maxilla of B. c‘opei, both retain deciduous premolars), these

bear only one upper incisor (here incisors may be lower incisors, or they 3 designated I ) on either side and do may be teeth of a different individual. 2 not show any trace of I . USNN 1137, Both USNM 1137 and AMNH 16771 consist

tho holotype of Ectoganus glirifomnis , of only isolated, hut associatecl teeth

and AXNH 16771, a principal referred which are presumably from one individual.

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Figure 9. Restoration of the skull and mandible of

Ectogcnius copei and the dentition of Ectoganus glirifonnis.

Skull based. primarily on USM 12714; dentition based 4 primarily on AM" 4286,' ANhW L287, NlKW 15633, A@JH

16244, 1p345, AElNH 1.6771, and AElNH 48001. (a)

Left lateral vfew of skull. (b) Left lateral view of

mandible. (c) Occlusal view of upper left dentition.

(d) Occlusal view of lower right dentition.

Figured9 appears on the following frame.

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Figure 9.

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At present, I suggest that the smaller arc rootless and ever-growing.

incisors are deciduous incisors which Both species of Ectoganus have

may have been retained for some time seven cheek teeth on either side above

after the eruption of the permanent and below and the morphology of the

incisors. Both the larger incisors canines and*clieek teeth is very similar

and the stnailer incisor of AMNH 16671 in both species (compare Pls. 32,. 35-45). 1 show definite P are triangular in cross section signs of wear. The upper 1 incisors of USNN 12714 may have been and enamel is lirrited to labial and

rootless, whereas those of NlM1 22235 posterolingual bands. The crown 1 and PN 241 represent rooted incisors structure on is unknown. On PI FMNH c11. of Ecotganus. the labial enamel band is longer

The lower incisors of Ectoganus anteroposteriorly and extends further

gZirifor?nis are known with certainty to down the side of the tooth than the

consist of one peglike incisor on either posterolingual band. The anterolingunl

side placed lingual to the anterior enamel-free portion of P: complements

margin of the canines (MNH 4286: P1. and is labial to the posterolabial

42, Flg. 2). The lower incisors are not enamel-frcc compressed portion of the

known for 6. copci. canine. Thus, the labial enamel of 1 1 The upper and lower canines of C and P a continuous surface, 1 1 forms Ectopmts arc similar to those of broken only by the gap between the * PsittncotJi~~

canines of Ectogmus functioned in bears two cusps, a higher labial and

cutting, and posteriorly they functioned slightly anterior cusp, and a lower

in grinding. The canines of Fcto?an;ts lingual, slightly posterior cusp.

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~ - Except for around the unworn .ckown, of the protocone to the anterior base 2 enamel bands on P2 are limited to the of the paracone and the apex of the

labial and lingual sides of the teeth. hypcone to the base of the.metacone.

The labial band is slightly longer These transverse crests are highest

anteroposteriorly and extends down the lingually and decrease in height .

tooth further than the ltngual band. The labially. P3-4 are hypsodont with 2 distribution of enamel on P2 complements enamel extensions labially and 1 1 P just as the enamel on P1 complements 1 lingually and enamel-free portions C;. anteriorly and posteriorly. The'enamel

P3-4 are extremely si.milar to each extends further lingually than

other. They are subcqual in size, and labia 11 y .

it is nearly impossiblc to distinguish M1-3 are transvc'rsely bilophodont . .. between them on the basgs of isolated teeth. Each is approximately square

teeth. Both bear a large, high paracone in cross sectqon and anteriorly bears

hnteroln~allyand a variably dcvclloped a large transverse crest formed*by the *: ' tho&h small metaconc p 0st e ro1 a bia11 y . large protocone and smaller paracone _. ... Lingually, they bear small and' low, Connected by a minutely cuspidate

subdistinct to fused protocones and transverse ridge. The posterior crest

hypocbnes. On.the basis of their is slightly lower and is not as wide

position, these two cusps appear to be as the anterior crest; it is combosed

homologous to the single lingual of the large hypocone and smaller I protocone of the corresponding teeth metacone connected by a minutely cuspi-

in Psittncott2';7?-iim. Between the date transverse crest. Whereas the

protocone-hypocone and paracone-metaconc hypoconc is placed directly posterior'

is a deep valley bounded anteriorly of the protocone, the metacone is

and posteriorly by minutely cuspidate placed slightly posterolingual of the

transverse crests which connect the apex - paracone. Posteriorly, M1-3 decrease

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in size, in degree of hypsodonty, and, higher and the entoconidS.are ab.out

in the relative size of the posterior the same size as the hypoconids. "1- 3 crest. The enamel extends further are square in cross section.. Their

lingually than labially on the uppek trigonids are compressed antero-

. molars. posteriorly and bear large protoconids P P3-4 are of similar morphology, sub- and metaconids. which fuse to form

equal in size, and difficult to transverse lophs. In R. 9lij.i fomii::

I distinguish from onGanother. P3-4 thcrc oftcn are small protoconids at

are bilophodonk and bcar widc and high the anterolabjal bases of the metaconids.

trigonids and lower, narrower, The talonids of 14 arc also compressed 1- 3 lingually placed ta'lonids. The trigo- anterop'osteriorly. In E. cop& the

nids bear large, conical protoconids talonids bear subequal hypotonids and

and sllghtly smaller and lower meta- entoconids khich fuse to form posterior

conids connected to thc protoconids transverse lophs. Thc tal.onids of

by minutely cuspidate transverse crests. R. g~im:form~salso bear subequal

In E. gZiri,romis there are ogten small hypoconids and entoconids, and

paraconids at the anterolabial bases hypoconulids or cu,spidate transverse

of the metaconids. The talonids bear ridges occur between them to form

large hypocoflds and small, low posterior transverse lophs. Variably,

entoconids, both connected by minutely mesoconids and/or entoconulids

cuspidatc transverse crests. Pj-4 on the talonids are'present in E.

are hypsodont, with enamel extending qL7:iv:;Coirnis. M1-3 decrease in size

further down the teeth labially than and hypsodont? posteriorly, and enamel \ lingually. extends further labially than

M1-3 are transversely bilophodont lingually on the lower molars.

teeth of morphology similar to P3-4, Gazin (1936) thoroughly described

except that the talonids are widcr and and illustrated the skull of Ectop-aizzts i\

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~ ~-

(USNM 12714; Fig. 8, Pls. 43, 44). Cope E. cope< than in E. gli.fifods. How-

(1884c, p. 188-193, Pi. 24b, Fig. 1) and. ever, these skeletal differences may

Wortman' (1897b, p. 88-92, Figs. 22, 23) be due to sexual dimorphism.

described and illusrrated the mandible The skeleton of FMNH P 26083 is

of Ectogunrts (AM" 4286; P1. 42, Figs. poorly preserved; the bone has been

1, 2). Cope (1877, p. 166-169, P1. 43, heavily permeated with Ironstone, and

Figs. 1-12) also described a partial the ends of the long bones, in particu-

scapula, humerus, radius, ulna, maggum, lar, are missipg (perhaps reflecting a

and uiigual phalanx of F,'ctopzu:: (US" lack of epiphyseal fusion in a young

1001). individual). However, a left scapula,

The type specimen of "Lan;padophoms the distal ends of both humeri, both

cqectutz(s" (FMM1 P 26083; Pls. 33, 34.) ulnae, femora, and tibiae are present,

includes a crushed skull and partial together with a (?)metacarpal, and

skeleton of a young adult. The crushed proximal and ungual phal2ngcs of the

skull is similar to USNN 12714, but is forefoot. A complete left humerus

larger, relatively short anteroposterior- (FMN P 26090: P1. 38, Figs. 21, 22)

ly, and does not have a promirlent from the Plateau Valley lqcal fauna

sagittal crest. In view of the fact is identical to the humeri of FMNH

that both specimens are of young P 26083 and is referred to Ectogmus.

individuals of roughly comparable age, Overall, the skileton is rather small

these differences are especially in comparison to the size of the

important because they may represent skull.

further distihctions between the two The scapula is heavily encrusted

species, E'. glirifomtis and B. copci. with ironstone and by comparison with

Subjectively, it appears that in com- the scapula of StyZinodon (Pls. 53, r. parison to the size of the skull, the Fig, 5; 54, Fig. 31, it appears that

cheek teeth are relatively larger in most of the perimeter of the blade

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and the coracoid process have been lost. 26083, FMNH P 26090, and USNM 1001 and

This scapula evidently had a large are referable to Ectoganus.

acromion process. The ulna and radius of Ectogmlts

The humerus of derived taeniodonts is (Pls. 34, Figs. 3, 4; 46, Figs. 3, 4, 9:

highly distinctive (compare Cope, 1877; UShW 1001 in Cope, 1877) resemble those

Narsh, 1897; Patterson, 1949b; Wortman, of other taeniodonts. Both are short,

1897b; the humerus of Stylinodon, hcavy, and robust and the ulna has a

P1. 55, Figs. 1, 2). It is short and large olecranon process that apparently

robust with extremcly well-developed extends up to one-third the length of

muscular crests. The greater and lesser the bone as in Stylinodok(P1. 55,

tuberosities both are well developed Figs. 4-6). Thc radius evidently could

and thc deltopectoral crest extends bc rotated on the ulna.

over half thc length of the shaft. The The manus is similar to that of

teres eminence is large and centered Psittrrcotherim and StyZixodoon, bearing

mcdially at the mid-length of- the shaft largc, sharp, laterally compressed and

and points backward. Thc humcrus is recurved claws.

extremely broad distally with large, The hind limb of E'ctoplu.? is

well-developed internal and external relatively poorly known. The femur of

condyles, a well-developed supinator FMh?i P 26083 (which is crushed dorso

ridge, and a largc, subcircular ventrally) is short and stout with

entepicondylar foramen. The distal largc, well-developed greater

articular surface (capitellum and trochantcr. The proximal and distal

trochlea) is broad and smoothly curved. ends of the femur of FHhW P 26083 arc

The olecranon fossa is of moderate sizc. missing. A more complete femur of

YPM 27201, a left humerus, and UNM GE-097, tctoqmus is preserved with US" no

a right humerus (Fig. 17; Lucas, 1982) number (Pl. k6, Figs. 1, 2). The

are identical to the humeri of FMh31 P tibia of RfNH P 26083 is also crushed

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~ ~~~~

dorsoventrally and both ends.are incorporating the paraconid). This may

missing. It is relatively short and be a tooth of an aberrant individual,

stout. USGS 3838 includes a partial it may represent a distinct taxon, or

calcancum of Ectoganstc (Pl. 46, Figs. I may have wronbly'interpretcd it as

5, 6). a P4.

Ectogmus sp.

(Pl. 14, Figs. 13-27) (Fig. 17)

Referred spccirbz. UI: 1823, right E(:lognnu.s sp. : Lucas, 1982.

p4(?) , left p4(?), left Pi1(?), and Re fcrred specimens. uNE.1 GE-097,

canine fragments (PI. 41, 'Figs. 13-27): Fragmentary right humerus (Fig. 17):

from Wasatchian strata of the Willwood from Wasatchian strata of the lower part

Formation, Little Sand Coulee, S!; of of the Galisteo Formation, Cerrillos

sec. 26, T. 56 N., R. 102 W., Bighorn local fauna, S&, S&, sec. 16, T. 14 N.,

Ba s i n , \dy omi n g . R. 8E., Santa.Fe County, New Mekicp

e Lkkcusszon. The tccth of UW 1823 YPM 27201, left humerus: from

here interpreted as a right P4 and a Wasatchian strata of the Willwood Forma-

left M1 are indistinguishable from the tion, Bighorn basin, Wyoming.

corresponding teeth of I..'ctoganus gzi~- Discussion: See discussion of these I- Jro~m',r. However, the crown morphology specimens above.

of the tooth here interpreted as a left Ectoganus sp. or Sty?

centrolabially placed conids (the Referred specimeno. FMNH 1' 15602,

(?)protoconid and (?)metaconid) , a large tips of canincs; FMNH.PM 336, canine

cuspidate posterolingual "cingulid" fragment: both from Wasatchian strata .

(the (?)talonid), and a slightly smaller of the Wasatch Formation ("Rifle Member

anterolingual cingulid (perhaps of the DeBeque Format'ion") , Garfield

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County, Colorado. thin Strips of enamel labially and

Dismssion. Although generically lingually after moderate wear.

indeterminate, these apparently rootless

canines document the presente of an ad-

vanced stylinodontine. (Table 11: Figs, 10, 11: Pls. 47-

40; P1. 52, Figs. 3-13; PIS. 53-55; StyZinodon P1. 59)

Stylinodon'Marsh, 1874, p. 531. Stylinodon mims Marsh, 1874, p. 531.

CaZamodon: Cope, 1881~.p. 184. CaZmohn cyZindrifc~Cope, 1881c, p. 184.

Calwnodon: Cope, 1884c, p. 192 (in part). CaZmur;do?i azl~.pchi~c~~:Cope, 1884c, p. 192

Rjpc species. Sty2inodm mh.s Marsh, StlyZitiodon rm'rms: Marsh, 1897, p. 137.

1874 (= Ca lamodon cy Zindrz:feri Cope, Sty Zinodon q 1 indm'fer : Wo r t man, 18 9 7 b ,

1881~). p. 92.

Inclrtdcd species. The tfie species St!jZinodo,i mirus: Vortman, 1897b, p. 93.

and Sty Zinodon i?zexplicatics Schoch and .Sty Zinodon qrZi n&-re?? : \I% i t e , 195 2 ,

Lucas, 1981d. p. i93.

Di.s t PL bu t ion. Late IJa sa t c h i an o f St?yZ7:r?donsp. : Robinson, 1966, p. 44.

Colorado and Wyoming, Bridgerian of StyZinodo)? qiZii2drifer: Guthrie, 1971,

Colorado and Wyoming, Uinun of Utah, p. 66.

' and Bridgerim or Uintan of western Sty Ziqodon ?virus: Schoch and Lucas,

Texas. 1981d (in part).

Revised diagnosis. Taeniodonts with R!;x specinien. YPM IXl95, right 2143. the dental formula I C P and left dentary fragments -with partial 1143'M all of the teeth ever-growing and right P3-M1; left P and alveoli for 3 rootless; moderately worn teeth without right P2, N2-3 and*left P2, Pi-N1;

enamel entirely around their perime ter ; labial enamel'fragment of left P1

posterior premolars and molars with only (Pl. 47, Figs: 4-6).

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TABLE 11. DENTAL MEASUREMENTS OF STYLINODON

AMNH DMPl FMNH PU USNM YPM SPECIMEN $3 107954 V-25 P-12185 16102' 16664 - 1109Sa 2 I- L 10. o* 4.2 12.W 6 .O 3.6 L 13.5" 4.7 w 9.5" 4.5" c1 L 38.0" 36.0" 40.1 15 .o* c1 w 17.5" 17.5" 19.5" 9.0" Pl L 15.0* 18.0" Pl w 14.5" 15.5" P2 L 11.0" 10.5" P2 w 12.5" 12.5* P3 L 10.5" 10.5" 7.3 P3 w ll.O*( 12.0" 7.4 P4 L 11.0" 9.5" 6.2 P4 w 11 .o* 10.56 7.0 EptL 13. 5" 12.0" 7.O M1 W 12.5* 10. 0" 7.5 M2 L 13.5* 10.3 6.5 M2 W 13.5" 10.2 7.0 tl3 L 10.9 11.5* 9.7 5.2 M3 W 10.0 12.5* 9.2 5.7

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6.0 6.8* 7.7* I3 I3 w 6.1 6.2* 5.5" 33.8 33.0* 33.2 c1 20.1 15.4* 18.2 c1 28.1 28.0* 25.3 22.2+ p1 17.7 17.5* 17.0 p1 12.9 10.4* 10.5 p2 18.1 14.1* 17.O* p2 11.0 11.1* 10.5* p3 14.2 10.5" 12.0* p3 10.5 11.7* 11.2 p4 12.0 11.2* 12.3 p4 12. 10.6" 11.5 M1 5* 12.5* 10.7* 12.0 M1 12.0" 10.7 M2 12.0% 9.8 M2 13.7 11.8 M3 11.3 8.7 M3

a Type specimen of Styllnodon mirus

b"ype specimen of 2. "cyllndrlfer" CType specimen of 5. lnexplicatus.

Downloaded from http://pubs.geoscienceworld.org/gsa/gsabulletin/article-pdf/92/12_Part_II/1982/3418970/i0016-7606-92-12-1982.pdf by guest on 01 October 2021 Horizon and ZocnZihj of thz type. Rridger "A" beds, Bridger Tormation,

Collected by Sam Smith in 1874 from E'j, Sa, sec. 33, T. 22 N., R. 113

Bridgerian strata of 'the Bridger IJ., Lincoln County, Wyoming.

Format ion, near Pfi llersville, Bridger UShTl 16664. mandible with partial

basin, Wyoming. right and left C1-P1, partial right 3(?) Rcferrcd spec&cns. I\MNII 4810, M , P2-3, M3, alveoli for right and left

fragments of enamel from the upper 13, left P2-3, right and left P4-M1,

canine(?), and' fragments of the skull right M2, isolated cheek teeth, verte-

(Pl. 50, Figs. 11-22; the type of brae, accetabular portion of the

"Ca%modon cylindrf fer") ; AMNH 14743- pelvis, shaft of the femur, patella,

14745, canlne and tooth fragments: all ungual of the pes, and other bone

from late Wasatchian strata of tbe Wind fragments (Pls. 50, Figs. 1-10; 52,

River Formation, Wind River basin, Wyom- Figs. 3-13; 59, Figs. 10, 11): from

ing. Bridger "C" beds, Bridger Formation,

USNN 18440, cheek tooth: from late Sage Creek basin (Greater "Bridger

Idasatchian strata of the Wind River basin"), Uinta County, Wyoming.

Formation, east side of Big Horn River AMNH 107954, partial skull with left 1 2 113 and north side of Birdseye Creek, C , right M + roots of right C , M , M ,

Boysen Reservoir area, SId!t, sec. 5~ alveolibfor right and left 12-3, right

T. 39 N., R. 94 W., Wind River basin, P1-4, mandible with right and left

Wyoming. -P left M1,3, isolated t'heth, skull I 3 4' PlCZ 3477, canine tip: from late fragments, miscellaneous vertebrae and

Wasatchian strata of the Wind River ribs (Pls. 48; 59, Figs. 1-6, 8, 9, 12,

Formation, Muddy Creek, Wind River 13): from Bridger "D" beds, Bridger

basin, Wyoming. - Formation, SW:, NWk, sec. 20, T. 13 N., R.

UW 2270, edentulous complete skull, 113 U., Green River basin (Greater

mandible, and partial skeleton: from "Bridger basin"), Uinta County, Wyoming.

Downloaded from http://pubs.geoscienceworld.org/gsa/gsabulletin/article-pdf/92/12_Part_II/1982/3418970/i0016-7606-92-12-1982.pdf by guest on 01 October 2021 2088 .. MNH PM 3895, skull and mandible with Huerfano Formation, San Draw, 3.2 km

complete dentition and partial skeleton: northwest of Gardner, Huerfano basin,

from late Bridgerian strata, lower part Colorado.

of the Adobe l'own Member, lower part YPM 14616, (?)lower canine: from

of the Washakie Formation, along Manuel late h'asatchian or early Bridgerim

Road, south-southeast of Haystack strata of the Huerfano Formation, "south

Mountain, Sweetwater County, Wyoming of Ilausero's Ranch" (Robinson, 1966,

[Turnbull, 1972, 1978: this specimen p. 44), Huerfano basin, Colorado.

will be described hy William D. Turnbull FMNH P 12185, palate and partial

(FMNHII. skull with left C1, M2-3 and alveoli 1 11 YPM 11096, left dentary fragment for right C , right and left P -M ,

with alveoli for C P -M occiput of right (Pl. 47, Figs. 1-3): from 1' 3 3' skull, left scapula, humerus, ulna, Uintan strata, Horizon B, Wagonhound

radius and partial manus, seven cervical Plember, Uinta F .Jtion, Coyote basin,

vertebrae, first thoracic vertebra, Uinta County, Utah.

right and left first ribs and partial DN" V-25, skull with complete right 1 sternum (Pls. 53-55; 59, Fig. 7): from and left C , alveoli for right and left 13 late Bridgcrian(?) strata of the 12-3, P -M , lower jaw with complete

Washakie Formation, lower green,sand right and left M2, right and left

at Haystack Mountain, Washakie basin, roots of 13-E11 and Pi3 (Pl. 49): from

Wyoming. the center of sec. 24, T. 8 S., R. 24 E.,

AMNH 17525, molarifom tooth fragment: Uintan strata, Horizon B, Wagonhound

from late Wasatchian strata of the Member, Uinta Formation, Coyote basin,

lower Huerfano Formation, Garcia Canyon Uintan County, Utah.

region, Huerfano basin, Colorado. Reviscd diaposis. Largest species

AMNH 17451, (?)upper canine: from of Stylirodon; approximately twice

early Bridgerim strata of the upper as large as Stylinodon inexplicatus

Formation, Sublette‘ County, Wyoming. SQjlir?odon ZnercpZirntus Schoch and Lucas, 1981d TPlM 41444-3, TMM 41443-291, cheek

(Table 11; Fig. 12: P1. 51; P1. 52, tooth fragments: from late Bridgerian

Figs. 1, 2) or early Uintan strata of the Pruett

njpC a?id On i!y k?lOiJn Spec imP?l. Formation, basal Tertiary conglomerate

PU 16102, skull with complete and in the Aqua Fria area, Whistler Squat 3 unerupted right and left PI , roots of local fauna, Trans-Pecos, Texas (see 21 right and left I -C , left P1-3, right J: A. Wilson, 1974; Wood, 1973). 42 and left P -M , associated vertebrae, Eaton (1980) has reported Stylinodon

rib and indeterminate bone fragments sp. from Brldgerian-Uintan(?) strata

(Pls. 51; 52, Figs. 1, 2). of the Wiggins Fornation, Carter Mountain

Horiaou:m~d 2ocal;t;y of the t-ypc. area, Southeastern Absaroka Range,

Bridgerian strata (“Washakie All) of the Wyoming .

Washakie mmation, sec. 11, T..16 N., ~iscu,o.sion. Although specifically

R. 96 W., lJashakie basin, Wyoming. indeterminate, these rootless cheek

‘DicrLgnosio. Smallest known .species tooth fragments document the presence

of Stylinodon; approximately half as of Stylinodon in these areas.

large as StpZinodon miruc (Table 11). Undetermined stylinodont, cf. Stylinodon mirus StyZinodon sp. (Pls. 56-58)

Stylinodontine: West, 1973, p. 125. Undetermined stylinodont: Gazin, 1952,

Sty2;nodon miru.7 : Schoch and Lucas, p. 26.

1981d (in part). Referred specimens. USNM 18425,

Referred specincns. JXNH PM 15198- partial hind- and forelimbs and

15200, 15590, tooth and enamel frag-. miscellaneous vertebral fragments (Pls.

ments: from late Wasatchian-early 56-58): from late Wasatchian strata of

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the Wasatch Formation (upper "Knight j6-c' an undetermined stylinodont,

Formation"), 19.3 km north of Big Piney, Jbably referable to Skjlilzodon mir?~,

Qest side of US Highway 189, La Barge has been partially illustrated and fully

fauna, SWk, sec. 33, T. 32, N., R. 111 described by Gazin (1952, p. 26-32,

W., Sublette County, Wyoming. P1. 2,. pigs. 2-5, Pls. 3, 4).

CM 37474, partial hind foot: from As in h'ortnania, the cervical

Bridgerian strata of the Washakic Forma- vertebrae of Stylinoah ~~PZG(Fig. 11)

tion, east end of Haystack Mountain indicate that thc neck was short,

near the base, Swcetwater County, IJyoming. thick, and stout. The first ribs and

Discussion. See discussion bclow. manubrlum are large and massive. The

limb bones are short, massive, and Description and Discussion of Stij linodm .closely comparable to the limb bones

The dentition, cranial morphology, and of Ectoganits gZiri-Toi.mis. The

systematics of Stylino&?? (Figs. 10-12) metacarpals of the manus of Sty Zinodon

are discussed at length in Schoch and mims are short, robust and bear large,

Lucas (1981d) and need not be repeated laterally compressed, recurved claws

here. as in b'ortvaxia, Psittacotlierium, and

YPM 11096, a partial skull, mandible, Ectoganits (compare Patterson, 1949b).

cervical vertebral series, first ribs, The metatarsals are shorter in s. mirus

manubrium of the sternum and forelimb than in Psittacotheriwn (Pl. 31,'Fig. 3). of Stylinodon Wyus (Pls. .53-55; 5.9, All five digits of the pes of .%SjLinodon Fig. 7) has been described, illustrated, bcar large, robust claws which are less

and discussed by PlaTsh (1897, p. 137- compressed and recurved than the

144: Figs. 3-9), Wortman (1897b, p. claws of the manus.

93-95, Figs. 26-28), and Patterson OTHER SUPPOSED OCCURRENCES OF TAENIODONTS (1949b, p. 252-254, Figs. 3-9). USNM

18425, the partial limbs and feet (Pls. At present, taeniodonts are known only

dentition of Stylinodon mims, based primarily on Nlh

107954, DhqIl.1 V-25, and UIJ 2270. The teeth are shown

worn (unworn teeth of Stylincdon mirus are not knowri).

(a) Left lateral view of skull. (b) Left lateral view,

of thc mandible. (c) Occlusal view of upper left

dentition. (d) Occlusal view of lower right dentition.

Figure 10 appears on the following frame.

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Figure lo.

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Figure 11. Reconstructed skeleton and life restoration of

Stylinodm mfmk based on specimens described and illustrated 2 in thc text and plates. Top: right lateral view of the recon-

structed skeleton. Middle: dorsal view of the reconstructed

skeleton. Bottom: life restoration.

Figure 11 appears on the following frame.

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Z cm

Figure 12. Restoration of the skull of StyZinodon

i)zc.rpZica'ius based on PU 16102: left lateral view.

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from western North America (Schoch and 1958), a genus of Insectivora (sensic

Lucas, 1981a). Several supposed lato). Among other features which

taeniodonts have been reported from exclude I'Calmodon" crcropncics from being

outside of western h'orth America. These a tacniodont are: its relatively small

reporrs are: size; the distribution of enamel on its

1. Rijtimeyer (1890, 1891) reported tusk, which occurs primarily cn the

and described a new species of "Calanodon' labial and anteroexternal sides, rather

(= Ectoganus) , "C. europaeus , from than equally on the anterointernal and

the middle Eocene Egerkingen deposits of anteroexternal faces as in taeniodonts;

Switzerland. Rutimeyer thought his and the fact that the posterior enamel-

specimen, a lo!*er right incisor (Pl. 60, free portion of the tusk is not laterally

Figs. 6, 71, was a lower canine of compressed as in taeniodonts.

Ectoganus. Rutimeyer also reconstructed 2. Amgehino (1891) described

a left dentary fragment (Pl. 60, Figs. 4, ktocnsmus hetcrogenidens on the basis

5) to.,look like the mandible of EctogmiiL? of a premolar and incisor (Pl. 60,

(Pl. 42). Schlosser (1894) and Cope Figs. 1, 2) from the Tertiary of

(1894) were both aware of these specimens Patagonia and believed it to be a new

and believed that they did;extend the genus and species of "Ectoganidae. It

range of "Calmodon" *into Europe. However, Ameghino (1902) later

However, in his revision of the Eocene reassigned it to the Notoungulata,

faunas of Switzerland, Stehlin (1916) synonymizing It .En tocnsrnus iicterogenidens"

thoroughly redescribed and discussed with rJoto3iippus toxodoxtoides.

"Calmodon" europaeus , demonstrating 3. Chow (1963a) described a new

,beyond doubt that it only superficially genus and species, Chitngchicnirr

resembles a taeniodont. Stehlin (1916) sichuanica, from the upper Eocene of

thus referred "Calamodon" eitropcreus to southern Hcnan, China. This taxon was

"Anphichiromjs" (= Heterolzyus : Saban, based on what Chow claimed was a right

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mandibular ramus (P1. 60, Fig. 3) with seen in any known taeniodonts. The

a single cheek tooth in place, a partial enamel on the only known tooth of

alveolus for a second tooth behind, and Chungchienia is limited to the antero-

an internal, horizontally,oriented external face,-whereas in nll derived

alveolus for a second tooth behind, and taeniodonts, enamel is lost anteriorly

an internal; horizontally oriented alve- and posteriorly on the cheek teeth,

olus for a "chisel-like 'incisor' of but remains as thin bands both internally

rodent or taeniodont type" (Chow, 1963a, and externally. Therefore, I here.

p. 1891). Savage (1971) suggested that exclude Chungchicnin from the

Chungchieuia may represent a Chinese Taeniodonta, although the taxonomic

taeniodont. However, in his original pos it ion of C?iu?igchimia must remain

description, Chow (1963a) rejected the uncertain until more complete material

notion that Chmpchienia is a taeniodont is known.

and instead referred it to the 4. Chow (1963h) also described a

Xenarthra (Edentrata). I concur with gliriform tooth (Pl. 61, Figs. 9, 10)

Chow's judgment that Chungclzicnin which he originally referred to "Til-

probably is not a taeniodont, although lodontia, gen. indet. sp. 2" from the

I do not consider it to be an edentate, late Eocene of Shandong, China. Later,

due to the constricted band of thick Chow and others (1973) suggested that

enamel which the cheek tooth bears. this specimen is a lower ciinine ot a

Ciiungchienia shares with taeniodonts taeniodont and should be referred to

the characters of ever-growing cheek "?S@ Zinodon sp. I' However, Chow s

-teeth with restricted bands of enamel (1963b) original identification of this

and a large "tusk." However, specimen as the I2 of a tillodont appears

has a large diastema between the incisor to be correct. Tn'both stylinodontine

like todth and what is evidently the taeniodont canines and trogosine

first cheek tooth; this is a feature not tillodont second incisors the enamel

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is limited tfo the labial portion of pattern often seen in trogosine

the tooth as in IVPP V. 2766 (Patterson, tillodonts (cimpare P1. 61, Fig. 5;

1949b; Gazin, 1953). However, in an incisor fragment of T~ogosuc),and

taeniodont canines the enamel-free the preserved portion of enamel-free

portion is laterally compressed dentine is in the same plane as the

posteriorly (compare P1. 48), unlike enamel-covered portion. It does no€

the second incisors of tillodonts which show any evidence that the posterior

are not compressed (Gain, 1953). 111 enamel-free portion was laterally

IVPP V. 2766 the enamel-free portion compressed as in taeniodonts. There-

is not laterally compressed as in fore, it is possible that ME'M 30848

taeniodonts, but rather resembles the represents a tillodont rather thag a

Tiliodontia in having subparallel taeniodont, but the material is too

internal and external sides. There- incomplete to permit a definitive

fore, I refer IVPP V. 2766 to the identification. Trogosine ti1 lodmts

Tillodontia, genus indet. were large, herbivorous extinct mammals

5. West and others (1977), West (Gazin, 1953), and if MPM 30848 is a

(1978), and McKenna (1980b) have re- tillodont, it would still have the

ported a stylinodontine taeniodont from same paleoclimatic implications as

the upper portion of the Eocene Eureka would a taeniodont (compxe McKenna,

Sound Formation of Ellesmere Island, 1980b).

North-West Territories, Canada. This 6. Dehm and Oettingeh-Spielberg

occurrence is based on a single enamel (1958) described Bnsalixn basaZcnsis

fragment (MPM 30848, P1. 61, Figs. 6-8). as a new genus and species of

Examination of this fragment suggests stylinodontine taeniodonts from the

that it may be the enamel fragment of middle Eocene Kuldana Formation near

a tillodont lower incisor. The enamel Ganda Kas, Pakistan. Rasalina basnlcnsis

of MF'M 30848 shows the slightly beaded is based on a left dentary fragment with

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one preserved cheek tooth (Pl. 61, Figs. taeniodonts occur are, from north to

1-4). DnsaZi?za has recently been south: the Crazy Mountain Field

thoroughly reviewed and reinterpreted (south-central Montana), Bighorn

as a tillodont (Lucas and Schoch, 1981) basin (north-central Wyoming), Togwotee

in accord with earlier suggestions by Pass area (northwestern Wyoming), Wind

Gingerich and Gunnel1 (1979) and West River basin (central Wyoming), Green

(1980) that I?usnZinn is a tillodont. River-Bridger basih (southwestern

The preserved cheek tooth of Rmnlina, Wyoming), Washakie basin (south-central

originally described as an M1 (Dehm and Wyoming), Uinta basin (northeastern Utah),

Oettingen-Spielbcrg, 1958), appears Huerfano basin (south-central Colorado),

to be a molarifonn,’bunoselenodont P4 San Juan basin, (northwestern New Mexico

closely comparable to the P4 of Estlion!jx and southwestern Colorado), and Tornillo

and other tillodonts (Gazin, 1953). Flat. are (western Texas. Here, I place

It is unlike the hypsodont, transversely special emphasis on the San Juan basin

bilophodont cheek teeth of derived (Fig. 15), from which the early Puercan

taeniodonts. to Wa#;atchian taeniodonts arc best known.

The history of study and nomenclature THE GEOGRAPHIC AND BIOSTRATIGRAPHIC of the Tertiary strata of the San Juan DIST.RIBUTION OF THE TAENIODONTA basin (Fig. 16) has been discussed and.

Introduction reviewed in numerous papers, including

All unequivocally known taeniodonts Baltz and others (1966), Cnrdncr (1910),

come from the Rocky Plountain early Granger (1914), Kues and others (19771,

Tertiary intermontane sedimentary Lucas (1981), Matthew (1937), Reeside

basins of bestern North America (Figs. (1924), Simpson (1948, 1959, 1981),

13, 14; Table 12), Lhich were formed Sinclair and Granger (1914), Tsentas

or rejuvenated d.uring the Laramide (1981), and Wood and others (1941).

orogeny. The major areas where

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Figure 13. Localities at which taeniodonts have

been found. Numbers correspond to localities listed

in Tablc 12. For localities in the San Juan basin

(SJB), see Figure 15.

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-0nychodectes tisonensis rarus

Conoryctella dragonensis

- Conoryctella pattersoni

- Conoryctes comma

Huerlanodon torrejonius

0 Huerfanodon polecatensis

- Wortmania otariidens

-0- Psittacotherium multilragum .3

- -.3 - - -€doganus gliritormis lobdelli

-Ectoganus glirilormis glirifomis

Ectoganus copei cope8

.-. -- Stylinodon mirus . .3

Stylinodon inexplicatus

Figure 14. The biostratigraphic distribution of the

Taeniodonta.

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LOCALITY FORMATION AGE 1. Betonnie Tsosie Wash Nacimien to Puercan

2. Kimbeto Wash Nacimiento Puercan

3. De-na-zfn Wash Nacimiento Puercan

4. Alamo Wash Nacimiento Puercan

5. Wagonroad Ridge North ,Horn Puercan

6. Dragon Canyon North Horn Torre jonian

7. Kutz Canyon Nacimiento Torrejonian

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8. Kimbeto Wash Nacimiento Torrej onian

9. Torreon Wash Nacimiento Tor re j on 1 an

10. UK NM Locality 15 Nacimien to Torrej onian

11. Gallegos Canyon Nacimiento Torrejonian

12. Escavada Wash Nacimiento Torrejonian

13. Simpson's Locality 226 Nacimien to Torrejonian

14. Gidley Quarry Lebo Torrejonian

15. Silberling Quarry Lebo Torrej onian

16. Rock Bench Quarry Polecat Bench Torrej onian

17. Swain Quarry Fort Union Torrejonian

18. Douglass Quarry Melville Tiff anian

19. Tornillo Flat Black Peaks Tif fanian

20. Plateau Valley k'asatch ("DeBeque") ' Tiffanian-

Clarkforkian

21. Polecat Bench Polecat Bench Clarkforkian

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23. Togwotee Pass Area "Lower variegated beds" Clarkforkian?

24. Bighorn Basin Willwood Wasatchian

25. Ahagre brroyo Sen Jose Wasatchian

26. Gobernador Sen Jose Wasatchian

27. Cerrillos Galis teo Wasatchian

28. Wind River Basin Wind .Fiver Wasatchian

29.. Huerfano Basin Huerf ano Wasa tchian-

Br idgerian

30. Green River/Br idger Basin Bridger Bridger ian

31. Washakie Basin Washakie Bridgerian

32. Uinta Basin Uinta Uintan

33. Brewster County, Texas Pruett Bridgerian-Uintan

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TAXA REFERENCE

Onychodectes tisonensis Simpson, 1959; Sinclair and tisonesis Granger, 1914

Wortmania otariidens -0. 5. tisonensis Simpson, 1959; Sinclair and -W. otkriidens Granger, 1914 -0. 5. tisonensis Sinclair and Granger, 1914 -0. 5. rarus -W. otariidens -0. t. tisonensis Sinchir and Granger, 1914 -0. t. rarus -W. otariidens -0. t. tisonensis Robisoq and Lucas, 1980 stylinodontine indet.

Conoryctella dragonensis Gazin, 1941; Schoch and Lucas,

Conoryctella pattersoni 1981c

?Psittacotherium sp. or

?Wor tmania . sp .

-C. pattersoni Granger, 1917; Schoch and Lucas,

Psittacotherium multifragum 1981c; R. W. Wilson, 1956

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Huerfanodon torrejonius Schoch and Lucas, 1981b

-P. multifragum

Conoryctes comma Matthew, 1897; Tsentae, 1981

-P. multifragum 1. comma R. W. Wilson, 1956 -R mulqifragum

-P. \multi)€ragum Sinclair and Granger, 1914

conoryctinc indet. Kues and others, 1977

-P. multifragum

-P. multifragum Simpson, 1959

conorcytine indet. Simpson, 1937

?Huerfan?d= sp. Simpson, 1937

-P. multifragum

Huerfanodon polecatensie Gingerich and others, 1980

-P. multifragum Rigby, 1980

-P. multifragum Douglass, 1908; Simpson, 1937

-P. multifragum Schicbout, 1974; J. A. Wilson, 1967

Ectoganus gliriformis iobdeili Patterson, 1936, 1949a; Sloan

and others, 1980

-E. 8. lobdelli Gingerich and others, 1980

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-E. g. lobdelli Simpson, 1929a, b -E. g. pliriformis McKenna, 1972, 1980a -E. cf. E. copei bighornesis -E. g. pliriformis Brown, 1980; Schankler, 1980 -E. copel copei -E. 2. bighornesis

-E. g. pliriformis Granger, 1914 ; Simpson, 1948

-E. g. gliriformis Kues and others, 1977

Ectoganus sp. Lucas and Kues, 1979; Lucas,

1982

-E. g. gliriformis Granger, 1910; Guthrie, 1967

Stylinodon mirus 1971 --S. mirus Robinson, 1966

--S. mirus West, 1972 --S. mirus Turnbull, 1972, 1978 Stylinodon inexplicatus --S. mirus Black and Dawson, 1966 Stylinodon sp. J. A. Wilson, 1972, 1974

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I1zones, a lower Ectoconw "zone" Puercan-Torre j onian (also known as the kmith2aeus "zone":

Taeniodonts of Puercan and Torrejonian Van Valen, 1979) and an overlying C age occur in the Paleocene Nacimiento TaenioZabis "zone" (formerly known as

Formation at several localities in the the "Po lymns todon, 'I = Taenio2abis 'I zone" :

southwestern and south-central San Juan Lindsay and others, 1979; Matthew, 1937;

basin (Fig. 15; Table 12). The Osborn, 1929; Sinclair and Granger, B14).

Nacimiento Formation is composed of Localities in Betonnie Tsosie wash and

red and green, buff and gray clay shales Kimbeto wash are in the Ectoconu; "zone,"

and siltstones, black clay shales and whereas localities in De-na-zin wash and

lenticular arkosic and quartzose sand- Alamo wash include both "zones." These

stones (Baltz and others, 1966; Tsentas "zones" have been considered to repre-

and Lucas, 1980; Tsentas and others, sent superposed biostratigraphic units

1981). In the upper part of the well separated temporally. Alternatively,

Nacimiento Formation, a northern facies they have been tought to represent

of relatively high-energy fluvial deposits different facies or to reflect collecting

(with a greater over-all percentage of biases (Matthew, 1937). Previously,

sandstone) and a southern facies of Onychodcctes tisonensic has been thought

lower-energy fluvial and swamp deposits to occur throughout the Puercan in both

are recognizable (Tsentas and others, %ones, I' whereas 0. I'rarus'' and

1981). This distribution of facies Fortmnia ota,.iidens were restricted to

suggests a northern source area for the Taeniolabis l'zonctl (Russell, 1967).

milch of the tipper part of the Nacimiento Here, 0. "rams" is considered a

Formation (Baltz, 1967; Tsentas and junior subjective synouym of 0. tisonensis

others, 1981). at the specific level and N. otariidens

The Puercan strata of ,the Nacimiento is now known from the Ectoconus "zone"

Formation have been subdivided into two in Betonnie Tsosie and Kimbeto washes.

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4 N LOWER TERTIARY OF THE SAN JUAN BASIN, NEW MEXICO -20km 0 SAN JOSE FORMATION MIMIENTO F0fWAlK)N mANlNAs FoRwmcM. QOJO AM SA)(DSTONE

Figure 15. Geologic map of the lower Tertiary strata of the San Juan basin, New Mexico,

showing the major Paleocene localitites which have produced taeniodont fossjls. For lower Eocene Localities in the San Juan basin, see Figures 2 and 3 in Lucas and others

(1981) (see also Table 12; geology modified after Fassett and Hinds, 1971, P1. 1).

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YPRESIAN EOCENE

PALEOCENE

Lem CRETACEOUS

LEGEND I Figure 16. The stratigraphy of the lower Tertiary strata of the 0 ALLUVIUM San Juan basin. Lithologies are schematic. Stratigraphic units after @CONGLOMERATE

0 SANDSTONE Baltz (1967). Local faunas after Sinclair and Granger (1914), Matthew ANDESITE (1937), Simpson (1935a, 1935b, 1935c), and Lucus and others (1981). fl MUOSTONE North American land mamnial "ages" with European stages after Berggren fl COAL and others (1978). Abbreviations for faunas: A = Almagre local fauna, a UNCONFORMITV = G = = JUNFOSSILIFEROUS AW Alamo Wash local fauna, Gobernador local fauna, L Largo

P = = = I100M local fauna, Puerco fauna, T Torrejonian fauna, Ti Tiffany

ALMAGRE LOCAL fauna . IA FAUNA

In -LnOCn nel

Downloaded from http://pubs.geoscienceworld.org/gsa/gsabulletin/article-pdf/92/12_Part_II/1982/3418970/i0016-7606-92-12-1982.pdf by guest on 01 October 2021 However, as far as is known, 0. t. Torreon wash and University of Kansas

tisonensis does occur in both zones, New Mexico Locality 15 as well,as the

whereas 0. t . rants is known only from areas southeast of Kimbeto and just

the Taeiziolabis "zdne." Thus, better south of Cedar Hill (Rt W. Wilson, 1956;

knowledge of the Puercan taeniodonts re- Tsentas, 19.91). These "zones" also have

duces the distinctiveness of these been thought to represent superposed

"zones" and does not strongly support biostratigraphic units separated by a

1 the idea that they are separ3ted by a significant length of time (Lirfdsay

significant span of time. and others, 1979; Taylor and Bytier,

Both 0. tisonensis and N. otariidens 1980). Alternatively, it has been

are restricted to the Puercan. The suggested that the differences between

occurrence of 0. tisonensis in the the faunas .of the two "zoneso reflect

Wagonroad local fauna of the upper part facies differences and/or collecting

i of the North Horn Formation of east- biases (Matthew, 1937; Tsentas, 1981;

central Utah supports the correlation R. W. Wilson, 1956). Recently, a speci-

of this locality with the Puercan-aged men of Pantolambda was found in a

strata of the Nacimiento Formation Deltatherim "zone" horizon in Kutz

(Robison and Lucas, 1980). Canyon, supporting the idea of

The Torrejonian strata of the collecting biases (Lucas and O'Neill,

Nacimiento Formation also have been 1981). Conoryctes coma is known with

divided into two "zones" on the basis certainty only from the Pantozmda

of mammalian faunas (Lindsay-and others, "zone." The speclrnens reported by

1979; Matthew, 1937; Osborn, 1929). Taylor (1981) as Conoryctes Coma

The presumably lower De Ztntherium l'zonel' from a DeZtathefiwn "zone" horizdn 2 occurs in Kutz Canyon, Torreon wash, in Kutz Canyon here ar"c assigned to

apd Kimbeto wash, whereas the overlying Co?wrycteZZa (Schoch and Lucas, 1981~).

PmtoZambda "zone" is well known from However, because it is relatively rare,

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I do not stress the possible blostrati- Mexj.cn, Wyoming, and Montana suggests

graphic significance of the absence of that these strata all are approximately

C: coma in the Deltatherim "zone.'' the same age.

Huerfanodon torrcjonius is now Tomida (1981) recently has described

known in the San Juan basin only from a typical Torrej onian mamma 1ian faunal

the Deltatherim "zone" in Kimbeto wash assemblage from the San Juan basin in

(Schoch and Lucas, 1981b). Huerfanodon sediments that can be magnetostrati-

po-leeatensis is known from the Rock graphically correlated with the Dragon

Bench Quarry of the Polecat Bench local fauna'of the North Horn Formation,

Formation, Bighorn basin (Schoch and Utah momida and Butler, 1980), the

Lucas, 198% and ?Hiierfanodoii sp. type locality of the "r)ragonian" land

is known from Silberling Quauy, Lebo mammal "age" (Wood and others, 1941).

Formation, Montana. Both Rock Bench These sediments also are stratigraphically

and Silberling Quarries are considered below a Deltathcriwn "zone" horizon

to be in the upper Torrejonian and suggest the presence of a third,

(Pantohbch "zone") (Ginberich and earliest Torrejonian "zone. It

others, 1980). Although Huerfanodon Conorycte 2 la pattersoni occurs in

polecatensis may appear to be slightly Tomida's (1981) "Dragonian" zone in

more "advanced" (and perhaps the $an Juan basin, in the Kutz Canyon

younger?) than Iluerfanodon torrejonius, Deltatherim "zone" (R. W. Wilson,

it is unwise to try to hypothesize that 1956) and in the Dragon local fauna

one species is the ancestor of another of Utah (Schoch and Lucas, 1981~).

and then base the relative correlation Thus, these occurrences of Co)zoqetalla

and dating of the strata in different suggest that all three localities do

basins on such hypothetical lineages not significantly differ temporally.

(contra Gingerich, 1976). Rather, The Dragon local-fauna thus can be

the occurrence of Huerfanodon in New correlated with the "lower" Torrejoni&

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strata of the Nacimiento Formation. Pocket, is in the Animas Formation in

Psittacotherim is dar ranging southern Colorado, but has not produced

genus, both geographically and any taeniodonts (Barnes and others,

temporally. It occurs throughout the 1954; Simpson, 1935a, 1935b, 1935~).

Torrejonian strata (all three "zones") Psi t tacotherim mu Z tifragum does occur of the Nacimiento Formation. It may in lower Tiffanian strata of Montana

also occur in the Dragon local fauna, (Melville Formation; Simpson, 1937)

and, if this is substantiated, it would and Texas (Black Peaks Formation;

further support a Torrejonian age for Schiebout, 1974).

this fauna. Psiftncothcrium is prcsent "kmpadopliorus lobde 2 Zi" and

in the Torrejonian Swain Quarry, Fort L. erpectatus" are junior subjective

Union Formation, Wyoming, and in the upper synonyms of Bctogcnus 9 Ziriformis.

Torrcjonian-lower Tiffanian strata of Bctoganus g Zirifomis Zobde 2 Zi occurs

Montana (Gidley, Lebo, and Douglass in upper Tiffanian-Clarkforkian strata

Quarries) and Texas (Tornillo Flat, of Colorado and in the Clarkforkian

Black Peaks Formation. of the Bighorn basin, Wyoming. E.

g Ziri formis gliriforrrtiz occurs in the Tiffanian-Uintan possible Clarkforkian strata of

In the northeastern San Juan basin, the Togwotee Pass area, but otherwise

the Nacimiento Formation grades lateral- is restricted to the lower-middle

ly into the uppcr part of the Animas Wasatchian of Wyoming and New Mexico.

Formation and the Ojo Alama Sandstone Except for a single specimen of Ectopnus

grades into the lower Animas (FlcDermott cf. F. copei bighornensis from the

Member; Baltz, 1967; Barnes and others, possible Clarkforkian strata of the

1954; Rceside, 1924). The type Togwotee Pass area, both E. copei copsi

locality of the Tlffanian land mammal and E. copci bighopensis are restricted

"age" (Wood and others, 1941), Mason to the lower Wasatchian of the Bighorn

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basiri, Wyoming. Rose (1977) has used was collected from rocks stratigraphical-

"~adophoms"in part to support ly equivalent to the "Almagre beds"

the vaLYdity of the Clarkforkian land near Governador (UNM B-970/971/973;

mammal "age" ; the revised taxonomy of P1. -38, Figs. 5-20).

Ectoga72us eliminates "Lumpadopliorus" Baltz (1967) defined and mapped four

as a distinctive Clarkforkian, or even fbnnal members of Simpson's San Jose

latest Paleocene, genus. Formation. The Cuba Mesa Member is

The San Jose Formation "Wasatch" of the lowest and is essentially unfos-

early workers), a series of variegated siliferous (Lucas, 1977); the overlying

continental mudstones and sandstones Regina $fember includes the Almagre

(Simpson, 1948), unconformably overlies fauna and the lower part of the Largo

the Nacimfento 'Formation and is in the fauna; the Llaves Member is essentially

structural center of the San Juan unfossiliferous (Lucas, 1977) and

basin. Granger (1914) distinguished either overlies the Cuba Mesa Member

two Wasatchian facies In the $an Jose: or the Regina Member, or grades into

the lower variegated "Almagre beds" and intertongues with the Regina

exposed in Almagre and Blanco arroyos Member. The Tapicitos Member is the

near the present town of Regina, and youniest member and either overlies

the upper red "Largo beds" exposed or laterally grades into the Llaves

near Lindrith and Gavilan (see Member; it includes the majority

Simpson, 1948). Cope's specimens of the Largo fauna (Lucas, 1977).

of Ectogams and llCakmodon" as well The only taeniodont taxon known

as Marsh's "Dryptodon" almost surely from the San Jose Formation is

came from the Almagre fauna, as have Ectoganus gZirifonmk gZzXfoMnis.

most taenlodonts since then (Lucas, Although the vast majority of

1977; Lucas and others, 1981); Recent- specimens of Ectogm2us has come from

ly (1977) a specimen of Ectoganus the Almagre local fauna, E. gzirifohs

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glirifohs is also known from the Stylinodon n6ms in the Lostcabinian

Largo local fauna. AMNH 16245, a lower (Guthrie, 1967, p. 1971). Thus,

jaw fragment of E. g. glirifomis Ectoganus is not known from strata

(Pl. 36, Fig. 13), was collected by younger than Lysitean. The presence

Granger in 1912 from the "west branch of E. g. glirifomris in the

of Almagre Arroyo, upper beds." Almagre fauna thus suggdsts a

This locality corresponds to the lower Graybullian or Lysitean rather than

part of the.Largo of Granger (1914) and a Lostcabinian age for that fauna

is in the upper part of the Regina (contra Lucas, 1977) and the occurience

Member of the San Jose Formation. of E. 9. glirifonnis in the lower part

, The Wasatchian land mammal "age1' is of the Largo would also suggest that

usually divided into three "subages," that horizon of the Largo (Granger,

the 'Graybullian, Lysitean and Lost- 1914) in the ReSina Member of the San

cabinian, from oldest to youngest Jose Formation is of Graybullian or

(Granger, 1914; Wood and others, 1941). Lysitean age. Nevertheless, the

Ectoganus g Ziri formis 9 Ziriformi,? majority of the Largo fauna, which occurs

(= Ectoganus "sivp Zex"; Schankler, stratigraphically higher in the

1980, p. 104) occurs in Graybullian Tapicitos Member, may be of Lost-

and Lysitean strata of the Willwood cabinian age (Lucas, 1977).

Formation, Bighorn basin, Wyoming, but Southeast of the San Juan basin in

not in the Lostcabinian strata of the north-central New Plexico in the

Willwood. Likewise, in the Wind River Wasatchian Cerrillos local fauna of

Formation of the Wind River basin, the Galisteo Formation (Lucas and

Wyoming, the type area of the Lysitean Kues, 1979; Lucas, 19821, a humerus

and Lostcabinean, E. g. gzirifohs of Ectoganus sp. recently has been

occurs only in the Lysitean strata and found (Fig, 17: Lucas, 1982).

is superseded by the taeniodont supporting the assignment of a

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Cerrillos local. fauna. The species and subspecies of

The earliest occurrence of StyZinodon stylinodontines (k'ortn;cmia,

hrus is in the upper Wasatchian Psittucotheriwn, Ectoganito, Stylinodoon)

(Lostcabinian) strata of the Wind are either known only from a few speci-

River basin, woming. It also occurs nens or are so broadly distributed

in the upper Wasatchian-lower geographically and of such long durations

Bridgerian of Colorado, in the temporally that at present they are not

Bridgerian of the Green River-Bridger useful except for very general correla- -and Washakie basins of Wyoming, and in tions no more refined than the level of the middle Ulntan ("Horizon El') of the land mammal "ages. "

Uinta basin, Utah. Stylinodon inexpectatus ACKNOCILEDGMENTS is known from only one specimen of

Bridgerim age in the Washakie basin.. Ifthank Dqs. Malcolm C. McKenna and

Two specimens of StyZinodon sp. are Hicfixrd H. Tedford (MiNH); Drs. R. Dehm, I known fromupper Bridgerian or lower K. Heissig, and P. Wellenhofer (BAWSM);

Uintan-aged strata of the Pruett Dr. B. Engesser (BNN); Dr. Mary Dawson

Formation, Trans-Pecos, west Texas ((31); Mr. Alden Hamblin (DMM); Dr.

(Schoch and Lucas, 1981d). The Killiam D. Turnbull and Err. J. Clay

presence of SQjlinodon, therefore, Bruner (ET.iNH) ; Mr. Charles 'Sthaf f (HCZ) ;

indicates a Lostcabinian-Uintan age. Dr. Robert M. West (MPM); Dr. Donald

Baird (PU); Dr. John A. Wilson (TMM); Conclusions Dr. William Clemens, Jr. (UC?P); Drs.

In conclusion, the early to middle Robert I+.Wilson and Larry G. Martin (UK)

Paleocene conorystincs (Onychodectes , Dr. Robert Sloan (UM); Dr. Barry S. Kues

Conorycte 2 la, Conoryctes , Huerfanodon) (UNEI); Dr. Thomas Brown (USGS); Drs.

appear to have the most potential for Robert J. Emry and C. Lewis Gazin and

Downloaded from http://pubs.geoscienceworld.org/gsa/gsabulletin/article-pdf/92/12_Part_II/1982/3418970/i0016-7606-92-12-1982.pdf by guest on 01 October 2021 Plr. Robert Purdy (USNM); Dr. Jason A. Farelly. for draying P1. 7, Fig. 1; Lillegraven (LW);and Dr. John H. Ostrorn and Ruth Yanai for drawing P1. 61,

(=ti) for permission to examine specimens Fig. 2. I especially thank Spencer ., in their care. I thank R. M. West and G. Lucas for many helpful discussions,

J. H. Hutchison for finding, and kindly for reading the.entire manuscript in

allowing me to borrow, ANNH 107954, a several veqsiops, , Tor drafting

partial skeleton of Stylinodon. I Figures 9 and 13-16, and for providing

thank Robert T. Bakker for pointing out the photographs in Fig. 17 and Pls. 5,

USNN 22483 to me and for helpful dis- Fig. 6; 27, Figs. 1, 2; 30, Fig. 1;

cussion. I thank R. Holtzman, S. G. 60, Figs. 4-7; and 61, Figs. 1, 3, 4.

Lucas, C. Tsentas, and W. D. Turnbull All other photographs and drawings

for reviewing the manuscript and making in Pls. 60, Figs. 1-3 and 61, Figs. 9,

many helpful suggestions. I thank J. 10 are by the.author. I gratefully.

David Archibald; Earl Planning, John acknowledge support from the Nat'ional

H. Ostrom,andTing Su-Yin for many Science Foundation (NSF Graduate

useful discussions. I thank 3. D. , Fellowship) an'd facilities provided

Archibald, D. Edwards, B. Honey, C. by thc Department of Geology and

Kochan, Ii. Lack, P. Lewis, C. Luna, Geophysics and Peabody Museum of

L. Reichlin, L. Ruppert, M. C. Schoch, Natural History, Yale University. I

bl. R. Schoch, and C. Tsentas 'for thank Betsy Gorecki for help in typing

logistic support in visiting some of the manuscript and David Schindel for

the above institutions. I gratefully the use of photographic equipment.

thank Nienke Prins for drawing the

'i SELECTED BIBLIOGRAPHY dentitions in Figures 1, 3-8, and 10

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OCTOBER 1, 1981

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