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American Museum of Natural History.
DEPAKEMENT OF. VERTEBRATE
PALAZONTOLOGY.
Volume II.
ARTICLES COLLECTED FROM THE AMERICAN
MUSEUM BULLETINS OF THE YEARS
16958-1903.
BY
HENRY FAIRFIELD OSBORN, J. W. GIDLEY,
iui. WORTMAN, F, W. LOOMIS,
W. D. MATTHEW, BARNUM BROWN,
Or, P. HAY, Re oh eis,
WALTER GRANGER, W. K. GREGORY.-
SMITHSON AY
APR 2 1 1987
SRARIE?
Collected and issued for purposes of sale anc
New York, December, 1903.
§
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= ri THE New ERA PRINTING COMPANY,
Pm 7 - - LANCASTER, PA.
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EMAC EES.
PerLORATIONS, AND: RESEARCHES
OF THE
Department of Vertebrate Paleontology.
PREFACE ‘TO’ VOLUME. I.
BY HENRY FAIRFIELD OSBORN, CURATOR.
In the preface of Volume I| a brief outline was given
of the foundation of the Department in May, 1891,
together with the successive appointments of the cur-
ator and different members of the staff, of the purposes
of the Department, of the twenty-one expeditions sent
out between 1891 and 1897, and of the acquisition of
the Cope collection of fossil mammals; also a sum-
mary of the collections, embracing in 1897 the remains
of 3,000 animals ; and of the paleontological, geologi-
cal and stratigraphical work accomplished ; and a re-
sumé of the chief scientific results contained in the
twenty-one Bulletins.
It appears appropriate to introduce Volume II with
a similar summary of the work accomplished in the six
years between 1897 and 1903.
Staff.— The preface of this second volume must
first refer with regret to the resignation of Dr. J. L.
Wortman in March, 1899, to take charge of vertebrate
palazontology in the newly established Carnegie Mus-
eum. Dr. Wortman’s services to this Department
both in the field and in scientific writing were of the
highest order. Under his able and energetic direction
in the field seventeen of the twenty-one expeditions
were conducted, laying the foundations especially for
111
IV Preface.
the remarkably representative Eocene and Oliogocene
collections of mammals and the Jurassic collection of
fossils. Dr. W. D. Matthew, now Associate Curator,
was appointed in Dr. Wortman’s place. In January,
1900, Dr. O. P. Hay, now Associate Curator of Chelo-
nia, joined the staff in connection with the cataloguing
and arrangement of the Cope Collection of fossil rep-
tiles, amphibians and fishes. Mr. Walter Granger con-
tinued in charge especially of the Jurassic field work
until the season of 1903, when he reéntered the mam-
malian fossil field in the Bridger basin. Mr. J. W.
Gidley joined the staff in 1899, with reterence to
strengthening our collection of Miocene, Pliocene and
Pleistocene mammals, and in 1900 he took charge of
the exploration for fossil horses under the Whitney
Fund. Mr. Barnum Brown joined the staff in 1896 in
connection with the Jurassic explorations, and in 1900
was put in charge of the exploration of the Creta-
ceous.
We are indebted to Mr. F. B. Loomis, of Amherst
College, for codperation in the expeditions of 1900,
1901 and 1902; and to Professor R. S. Lull, of the
Amherst Massachusetts Agricultural College, for
codperation in the Cretaceous and Jurassic expeditions
of 1899 and 1902. In 1903 Mr. Peter Kaison, who
formerly served as field and Museum assistant, took
charge of the continued excavation of the Bone Cabin
quarry.
Mr. Adam Hermann has continued as head prepar-
tor, and has been constantly perfecting his methods of
mounting. Mr. W.K. Gregory joined the staff in 1900
as an assistant to the curator and in charge of the
Department Library. Mr. A. E. Anderson has con-
Preface. V
tinued in charge of the photographic and art depart-
ment. Among the artists who have illustrated our
publications should be mentioned especially : Messrs.
Weber, Christman, Yoshiwara ; Mrs. Stirling and Miss
Box.
Total Summary of Collections. —The collec-
tions made by our field expeditions during the four-
teen years embrace the remains of over 5,000 fossil
animals.
Our collections have also been strengthened by
friendly exchange with American and foreign museums,
and by purchases, the most important being that of
the second portion of the famous collection brought
together by Professor Edward D. Cope, known as the
Cope Collection of Fossil Fishes, Amphibians, Reptiles
and Birds. This was generously presented to the
Museum by President Morris K. Jesup in 1902. - At
the same time the Pampean Collection of Ameghino,
Larroqgue and Brachet, originally exhibited at the Paris
Exposition in 1881, and there purchased by Professor
Cope, was secured for this Museum by the gift of
Messrs. H. O. Havemeyer, D. Willis James, Adrian
Iselyn, Henry F. Osborn and the late James M. Con-
stable and William E. Dodge, Trustees of the Museum.
Among the other purchases are those from Charles H.
Sternberg from his explorations in the Kansas chalk.
Altogether the purchases have added 1,800 speci-
mens.
The chief exchanges have been with the museums
of London, Paris, Munich, St. Petersburg, Berlin,
Stuttgart, Lyons and Buenos Aires.
The collections as a whole now include fourteen
thousand four hundred specimens, representing :
vi Preface.
PossilemarnmalSe.,csc ces. scapes neiesw soe camer sed nen sienoinieinisriems 9,373
Fossil reptiles. ............csssesceseeeceeseseneecates sceeeeeuees 2,694
ThtaAS SH Lobes by se sacisneoceotinekseobbea0 Huodestindobstieceo soon. coudabodn 57
Fossil amphibians,.,.......2..cseeeeesseseeseerceeteneee receeeees 593
TASCrIlN AIS) VES A Aen aaciace erasbononcoade ago, neoprcaseaqcse Sscagcaoge 1,202
ARS TEI sec agnnoonenePeonodsdd: cg to Speadasoucnt rscecaboceen 14,420
Beginning in 1891, the horizons have been explored in the following
order :
Permian, Texas, 1902.
Triassic, North Carolina, 1894, 1895.
Upper Jurassic (Como Beds), Wyoming, 1897-1903 ; Montana, 1903.
Upper Cretaceous (Niobrara), Kansas, 1897.
Upper Cretaceous (Fort Pierre), South Dakota, 1903.
Upper Cretaceous (Laramie), Wyoming, 1892, 1900 ; Montana, 1902.
Basal Eocene (Puerco), New Mexico, 1892.
Basal Eocene (Torrejon), New Mexico, 1892, 1896.
Lower Eocene (Wasatch), Wyoming, 1891, 1896 ; New Mexico, 1896.
Middle Eocene (Wind River), Wyoming, 1891, 1896; . Colorado,
1897.
Middle Eocene (Bridger), Wyoming, 1893, 1895, 1903; Colorado,
1897.
Upper Eocene (Uinta), Utah, 1894, 1895.
Oligocene (White River), South Dakota, 1892, 1894, 1901, 1902.
Oligocene (White River), Colorado, 1898, 1901; Montana, 1902.
Middle Miocene, Colorado, 1898, 1901, Igo2.
Upper Miocene (Loup Fork), South Dakota, 1894, 1902, 1903.
Upper Miocene, Texas, 1899, 1900, 1901 ; Montana, 1902.
Pliocene (Blanco), Texas, 1900, 1go!.
Pleistocene (Sheridan), Nebraska, 1893, 1897; Texas, 1899, Igo0o,
1gOl.
Pleistocene Arkansas, 1903.
General Exploration. — Two series of expeditions,
for mammals and for reptiles, conducted by from two
to four parties, have been sent out each year since 1897
in the systematic search for fossil reptiles and mammals
in different formations.
These expeditions during the past five years, namely
between 1898 and 1903, have had a considerable meas-
ure of success. They have added some 2,500 speci-
Preface. vii
mens to the collections, including some thirty more or
less complete skeletons, eight of which have already
been mounted and placed on exhibition, as follows :
Equus scotte. Merycodus osbornt.
Oruztholestes hermann. Glyptotherium texanum.
Flypohippus equinus. Dinictis squalidens.
Neohtpparion whitneyt. Cynodictis grevarius.
So Po)
RECENT EXPLORATIONS:
Exploration for Dinosaurs. — A large part of
the field work of the Department since 1897 has
been directed to securing the remains of Dinosaurs,
especially from the Upper Jurassic (Como Beds) of
Wyoming and Colorado, and from the Cretaceous.
A rich and extensive deposit, the ‘Bone Cabin
Quarry,” was opened up in Wyoming in 1898, and
has been worked with good results for six years.
Evolution of the Horse. — The generosity of
one of the Trustees of the Museum, Mr. William C.
Whitney, enabled the Curator to send out in 1899
and subsequent years a series of expeditions into
various Tertiary formations of which the chief object
was to obtain materials to illustrate the evolution of
the horse.
GEOLOGICAL RESUETS.
The field parties beginning in 1900 have with one
or two exceptions made very exact stratigraphical
field records. The result is the accumulation of im-
portant data concerning the distribution of faunas and
especially concerning the subdivision of horizons which
were formerly considered single or incorrectly identi-
fied. We may summarize these results since 1900 as
follows :
vill Preface.
1. Uinta (Upper Eocene) subdivided by Peterson
into Telmatotherium and Diplacodon Horizons.
2. Puerco (Basal Eocene) subdivided by Wortman
and Matthew into Puerco and Torrejon.
3. Two other Eocene horizons determined and corre-
lated in the Huerfano Basin of Colorado by Osborn and
Matthew.
4. White River (Oligocene) subdivided by Wort-
man and Matthew by the addition of two upper di-
visions, Protoceras beds and Leptauchenia beds.
5. The three divisions of the White River, first made
by Hayden and Leidy, further defined and correctly
-correlated with the stratigraphy, and the existence
shown of two contemporary faunal phases in each
division (Wortman and Matthew).
6. The various JZzocene formations which have been
grouped under the general names of Loup Fork and
Deep River, distinguished and correctly correlated with
their respective Miocene horizons, and the so-called
Palo Duro horizon eliminated (Matthew and Gidley).
7. The ALolian Theory. — Our field researches
have resulted in replacing the lake basin theory by the
fluviatile and colian theory, especially under the ob-
servations of Dr. W. D. Matthew, the arguments de-
rived from which are fully summarized in his memoir.
This demonstration has been nearly simultaneous
with that of Messrs. Hatcher, Davis, and Gidley, who
have reached and published similar results.
8. Stratigraphical Succession of Horizons. —
The most complete paper on stratigraphical succession
is that by Matthew, No. 28, entitled ‘A Provisional
Classification of the fresh water Tertiaries of the West,”
summarizing results obtained by our field parties by
Preface. 1X
comparison with those obtained previously by Leidy,
Cope, Marsh, and others. Of the same nature are the
exact researches by Mr. F. B. Loomis, on the Jurassic
stratigraphy of Wyoming (Nos. 37, 48.)
PAE AZSONTOLOGICAL -RESULYS.
Dinosaurs and other Reptiles. Bulletins. —
The first paper, No. 27, is an erroneous identification
by Osborn of Camarasaurus Cope with Brontosaurus
Marsh, it having since been found that Camarasaurus
is more nearly allied to Morosaurus. In this paper
the vertebral structure of Sauropoda is discussed. The
fore and hind limbs of carnivorous and herbivorous
Dinosaurs are discussed by Osborn in later papers.
The latest contributions are an account of the skull of
Creosaurus by Osborn, No. 61, and of the skull of
Triceratops by Lull, No. 60.
Attention should be called here to a series of import-
ant memoirs on fossil reptiles, namely: A complete
Mosasaur Skeleton, by Osborn; A Skeleton of Diplo-
docus, by Osborn; Ox the Reptilian Subclasses Diap-
sida and Synapsida and the early history of the Diapto-
sauria, by Osborn,
Evolutionary Series. — One of the principal ob-
jects of our expeditions has been the completion of
the evolutionary or phyletic series so far as possible,
with a view to working out the development of differ-
ent orders and families from their first appearance to
their extinction.
To this subject the following papers have been espe-
cially devoted: No. 7, Azcestors of the Tapir, by
Wortman and Earle; No. 19, on the Ganxodonta and
L:dentata, by Wortman ; No. 24, on the Camelida, by
x Ie, refa BE
Wortman; No. 26, on the Amblypoda, by Osborn ;
No. 29, on the Canide, Viverride and Procyonide,
by Wortman and Matthew; No. 33, on the AAznocer-
oses of Europe, by Osborn ; No. 35, on the Creodonta, by
Matthew; No. 40, on the 77tanotheres, by Osborn ; No.
43, on the Eocene Primates, by Osborn.
Systematic Revisions. — We have also taken up
one group after another, and have succeeded in giving
a more or less complete and final revision of certain of
the groups, incidentally determining questions as to
priority, synonymy and the location of type specimens.
Among the papers of this character have been the fol-
lowing: No. 2, Spectes of Coryphodon, Earle; No. 8,
Upper Cretaceous Mammals, Osborn; No. 12, Puerco
Mammals, Osborn and Earle; No. 10, Species of -
Flyracothertum, \Nortman ; No. 21, Puerco Mammals,
Matthew; No. 26, Pantolambda and Coryphodon, Os-
born ; No. 35, Cveodonta, Matthew ; No. 36, American
Species of Equus, Gidley; No. 43, Eocene Primates,
Osborn ; No. 46, the yfertragulide, Matthew ; 49,
Cretaceous Actinopterous Fishes Hay; No. 53, Me.
Lebanon Fishes, Hay.
Morphological Series.— The chief morpholog-
ical or anatomical papers are the following: No. 3,
Protoceras, Osborn and Wortman; No. 5, Acerathe-
rium tridactylum, Osborn; No. 9, Patriofelis, Wort-
man; No. 14, dgrtocherus, Wortman; No. 18, Psz¢fa-
cothertum, \Wortman ; No. 22, Teleoceras, Osborn ; No.
23, Coryphodon, Osborn; No. 25, Phenacodus, Osborn:
No. 30, Oxyena, Wortman; No. 32, Lguus scorn:
Gidley: No. 34, Oxyena and Patriofelis, Osborn ; No.
41, Dinocyon, Matthew; No. 42, Bunelurus, Mat-
thew ; No. 45, Ceratogaulus, Matthew ; No. 46, Hypr-
Preface. X1
sodus, Matthew; No. 54, Orzztholestes, Osborn; No.
55, Neohipparion, Gidley ; No. 57, Glyptotherium, Os-
born ; No. 58, Paramylodon, Brown ; No. 60, 772cera-
tops, Lull; No. 61, Cveosaurus, Osborn.
Papers especially relating to phzlosophical anatomy
are those on the teeth and feet by Osborn and Wort-
man and Gidley and especially No. 39, Doltchocephaly
and Lrachycephaly, by Osborn and No. 52, Zhe Fle-
phants’ Skull, by Gregory.
Faunal Lists of Different Horizons. — We
have also attempted to give complete faunal lists of
different horizons. The most comprehensive paper of
this character is No. 28, by Matthew, including faunal
lists of ‘the entire Areshwater Tertiaries of the West.
Other papers are No. 12, Puerco Mammals, Osborn
and Earle; No. 21, Puerco Mammals, by Matthew ;
No. 47, Pleistocene Fauna from Hay Springs, by Mat-
thew ; No. 50, Lower Oligocene Fauna of the Pipestone
Springs, by Matthew; No. 59, Azocene Fauna of
Northwestern Texas, by Gidley.
Memoirs on Fossil Mammals. — These include
The Extinct Rhinoceroses, Part!, by Osborn, and Fosse
Mammals of the Tertiary of Northeastern Colorado, by
Matthew. The memoir on the Rhinoceroses will be
continued and the memoir on the Fvolution of the
fforse is 1n preparation by Osborn with the codépera-
tion of Mr. J. W. Gidley.
PoAOTOGR AP HY.
Field Photographs. —In connection with the var-
ious expeditions enumerated on page iii a series of
580 field photographs have been taken, the negatives of
which are filed in the department for use in connection
xii Preface.
with our publications as well as for lecture and illus-
trative purposes.
Museum Photographs. — These include photo-
graphs of mounted skeletons of skulls, of feet and
other portions of fossil mammals and reptiles number-
ing altogether 590 negatives, also photographs of the
restorations of fossil mammals and reptiles by Charles
R. Knight.
RESTORATIONS, AND MODEES:
The work of the department would be incomplete
without reference to the very interesting series of res-
torations of fossil mammals and reptiles executed by
the artist Mr. Charles R. Knight, chiefly under the
direction of Professor Osborn, a complete list of which
is given at the close of this volume and in a new cata-
logue which will be issued in 1904.
December, 1903.
Preface. Xill
PUBLICATIONS FROM: THE DEPARTMENT OF VERTEBRATE
PALASONTOLOGY.1
MEMOIRS.
1. The Extinct Rhinoceroses. Part I. By Henry Fairfield Osborn.
Vol. I, Part III, pp. 75-164, pll. xiia-xx, and 49 text figures.
April 22, 1898. Price, $4.20.
. A Complete Mosasaur Skeleton. By Henry Fairfield Osborn. Vol.
I, Part 1V, pp. 165-188, pll. xxi-xxili, and 15 text figures. Oc-
tober 25, 1899.
3. A Skeleton of Diplodocus. By Henry Fairfield Osborn. Vol. I,
Part V, pp. 189-214, pll. xxiv-xxvili, and 15 text figures. Octo-
N
ber 25, 1899. Price of Parts IV and V, issued under one cover,
$2,00.
4. Fossil Mammats of the Tertiary of Northeastern Colorado. By W.
D. Matthew. Vol. I, Part VII, pp. 353-446, pll. xxxvil-xxxix,
and 34 text figures. November, 1901. Price, $2.00.
5. Lhe Reptilian Subclasses Diapsida and Synapsida and the Early
Flistory of the Diaptosauria. By Henry Fairfield Osborn. Vol.
I, Part VIII, pp. 449-507, pl. xl, and 28 text figures. Novem-
ber, 1903, (Price, 42.60.
BULLETINS.
Contents of Volume I.
1892
1. Fossil Mammats of the Wahsatch and Wind River Beds. Collec-
tion of 1891. By Henry Fairfield Osborn and J. L. Wortman.
Vol-1V, No: 1, Article xi, pp. 81-147, pl..1v, and 19 text figures.
October 20, 1892. Price, 80 cents.
Revision of the Species of Coryphodon. By Charles Earle. Vol.
IV, No. 1, Article xii, pp. 149-166, 2 text figures. October 18,
Kag2) ‘Pree, 15 cents:
3. Characters of Protoceras (Marsh), the New Artiodactyl from the
Lower Miocene. By Henry Fairfield Osborn and J. L. Wort-
man, Vol Vj, No. 1, Article xvii; pp. 351-371, 6 text figures.
December 30, 1892... Price, 25 cents.
Nv
1893
4. Artionyx, a New Genus of Ancylopoda. By Henry Fairfield
Osborn and Jacob L. Wortman. Vol. V, Article i, pp. 1-18, 5
text figures. February, 1893. Price, 20 cents.
1 Separates of most of these papers can be obtained of the Librarian of the
Museum at the prices indicated, or in exchange.
XIV
Preface.
135
Aceratherium tridactylum from the Lower Miocene of Dakota. By
Henry Fairfield Osborn. Vol. V, Article vii, pp. 85-86. April
29, 1893. Price, 15 cents.
On the Divisions of the White River or Lower Miocene of Dakota.
By J. L. Wortman, M.D. Vol. V, Article ix, pp, 95-105. June
27, 093. ‘Pace, 15 cents.
Ancestors of the Tapir from the Lower Miocene of Dakota. By J.
L. Wortman and Charles Earle. Vol. V, Article xi, pp. 159-
180, 7 text figures. August 18, 1893. Price, 25 cents.
Fossil Mammals of the Upper Cretaceous Beds. By Henry Fair-
field Osborn. Vol. V, Article xvii, pp. 311-330, pll. vil and
vill, and 4 text figures. December 15, 1893. Price, 40 cents.
1894
Osteology of Patriofelis, a Middle Eocene Creodont. By J. L.
Wortman, M.D. Vol. VI, Article, v. pp. 129-164, pl. 1 and 5
text figures. May 24, 1894. Price, 40 cents.
Fossil Mammats of the Lower Miocene White River Beds. Col-
lection of 1892. By Henry Fairfield Osborn and J. L. Wort-
man. Vol. VI, Article vi, pp. 199-228, pll. 11 and 11, and 8
text figures. July 28, 1894. Price, 50 cents.
On the Affinities of Leptarctus primus of Leidy. By J. L. Wort-
man. Vol. VI,- Article, vii, pp.. 2290-231. July 30, 1928
HCE; 15, cents:
1895
Fossil Mammats of the Puerco Beds. Collection of 1892. By
Henry Fairfield Osborn and Charles Earle. Vol. VII, Article
i, pp. I-70, 21 text figures. March 8, 1895. Price, 80 cents.
Fossil Mammals of the Uinta Basin. Expedition of 1894. By
Henry Fairfield Osborn. Vol. VII, Article ii, pp. 71-105, 17
text figures. May 18, 1895. Price, 50 cents.
On the Osteology of Agriocherus. By J. L. Wortman. Vol. VII,
Article iv, pp. 145-178, pl. 1, and 24 text figures. June 17,
1895. Price, 60 cents.
ertssodactyls of the Lower Miocene White River Beds. By
Henry Fairfield Osborn and J. L. Wortman. Vol. VII, Article
xii, pp. 343-375, pll. viii to x1, and 12 text figures. December
23, 1895. Price, 80 cents.
1896
Species of Hyracotherium and Allied Perissodactyls from the Wah-
satch and Wind River Beds of North America. By J. L. Wort-
man. Vol. VIII, Article vi, pp. 81-110, pl. m1, and 18 text
figures. May 12, 1896. Price, 50 cents.
Preface. XV
17. Zhe Cranial Evolution of Titanothertum. By Henry Fairfield
Osborn. Vol. VIII, Article ix, pp. 157-197, pll. 111 and Iv, and
13 text figures. July 31, 1896. Price, 75 cents.
18. Psittacotherium, a Member of a New and Primitive Suborder of the
Edentata. By Dr. J. L. Wortman. Vol. VIII, Article xvi, pp.
259-262. November 30, 1896. Price, 15 cents.
1897
19. Zhe Ganodonta and thetr Relationship to the Edentata. By J. L.
Wortman. Vol. IX, Article vi, pp. 59-110, and 36 text figures.
March 22, 1897. Price, 85 cents.
20. The Huerfano Lake Basin, Southern Colorado, and its Wind
kiver and Bridger Fauna. By Henry Fairfield Osborn. Vol.
IX, Article xxi, pp. 247-258. October 20, 1897. Price, 15
cents.
21. A Revision of the Puerco Fauna. By W. D. Matthew. Vol. IX,
Article xxii, pp. 259-323, and 20 text figures. November 16,
1897... Price, 75, cents.
List of Casts, Models, Photographs of Skeletons and of Restorations,
issued March 15, 1898. Price, 40 cents.
BULLETINS.
Contents of Volume II.
1898
. A Complete Skeleton of Teleoceras fossiger. Notes upon the Growth
and Sexual Characters of this Species. By Henry Fairfield
Osborn. Vol. X, Article iv, pp. 51-59, pll. iv, va. March 18,
1898. Price, 25 cents.
23. A Complete Skeleton of Coryphodon radians. Notes upon the Loco-
motion of this Animal. By Henry Fairfield Osborn. Vol. X,
Article vi, pp. 81-91, pl. x, and 2 text figures. April 4, 1898.
Price, 20 cents.
24. The Extinct Camelide of North America and some Associated Forms.
By J. L. Wortman, M.D. Vol. X, Article vii, pp. 93-142, pl.
XI, and 23 text figures. April g, 1898. Price, 70 cents.
25. Remounted Skeleton of Phenacodus primevus. Comparison with
Euprotogonia. By Henry Fairfield Osborn. Vol. X, Article ix,
pp. 159-164, pl. x11, and 4 text figures. May 6, 1898. Price,
20 cents.
26. Evolution of the Amblyfoda. PartI. Taligrada and Pantodonta.
By Henry Fairfield Osborn. Vol. X, Article xi, pp. 169-218,
and 29 text figures. June 3, 1898. Price, 80 cents.
iS)
i.)
30.
oie
33:
34.
36.
Preface.
Additional Characters of the Great Herbivorous Dinosaur Camara-
saurus. By Henry Fairfield Osborn. Vol. X, Article xii, pp.
219-233, 13 text figures. June 4, 1898. Price, 35 cents.
1899
A Provisional Classification of the Fresh Water Tertiary of the
West. By W. D. Matthew. Vol. XII, Article iii, pp. 19-77.
March 31, 1899. Price, 50 cents.
The Ancestry of Certain Members of the Canidae, the Viverride and
Procyonide. By J. L. Wortman and W. D. Matthew. Vol.
XII, Article vi, pp. 109-139, pl. v1, and Io text figures. June
21, 1699, Price, 40 cents.
Restoration of Oxyena lupina Cope, with Descriptions of Certain
New Species of Eocene Creodonts. By J. L. Wortman. Vol.
XII, Article vii, pp. 139-148, pl. vir, and 3 text figures. June
Zt eisgo. Price, 20 cents,
Fore and Hind Limbs of Carnivorous and Ferbivorous Dinosaurs
from the Jurassic of Wyoming. Dinosaur Contributions, No.
3. By Henry Fairfield Osborn. Vol. XII, Article x1, pp. 161-
172, 8 text figures. October 30, 1899. Price, 25 cents.
1900
. A New Species of Pleistocene Horse from the Staked Plains of Texas.
By J. W. Gidley. Vol. XIII, Article xiii, pp. 111-116, and 5
text figures. August 18, 1900. Price, 15 cents.
Phylogeny of the Rhinoceroses of Europe. Rhinoceros Contribu-
tions No. 5. By Henry Fairfield Osborn. Vol. XIII, Article
xIx, pp. 229-267, and 16 text figures. December 11, 1900.
Price, 50 cents.
Oxyena and Patriofelis Re-studied as Terrestrial Creodonts. By
Henry Fairfield Osborn. Vol. XIII, Article xx, pp. 269-281,
pll. xvi and xix, and 4 text figures. December 21, 1900.
Price, 35 cents.
Igor
- Additional Observations on the Creodonta. By W. D. Matthew.
Vol. XIV, Article i, pp. 1-38, and 17 text figures. January 31,
1901. Price, 50 cents.
Tooth Characters and Revision of the North American Species Oat
the Genus Equus. By J. W. Gidley. Vol. XIV, Article ix, pp.
91-142, pll. XvilI-xx1, and 27 text figures. May 31, Igo1
Price, $1.20.
37:
38.
Ao.
4I.
43.
44.
45.
46.
47.
48.
Preface. XV
On Jurassic Stratigraphy in Southeastern Wyoming. By F. B.
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A Complete Skeleton of Teleoceras fosst-
ger. Notes upon the Growth and
Sexual Characters of this Species.
By Henry Fairrietp Oszorn.
PLATES’ ITV; anp [Va.
AUTHOR’S EDITION, extracted from BULLETIN
OF THE
American DUusewum of Datural History,
Vo. X, ARTICLE IV, pp. 51-59.
New York, March 18, 1898.
ae =<.
7
- :
e x
j
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- —
, i
= -:
ig
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a2 7
;
ry)
Article IV.—A COMPLETE SKELETON OF TELEO-
CERAS FOSSIGER. NOTES UPON THE GROWTH
AND SEXUAL CHARACTERS OF. THIS SPECIES.
By Henry FAIRFIELD OSBORN.
PLATES IV AND IVA.
The remarkable series of Rhinoceros skulls in the Cope and
American Museum Collections from the Upper Miocene or Loup
Fork Beds of Kansas and Nebraska, has finally been prepared
for exhibition and research purposes. Associated with them, and
of very great value, is a complete skeleton representing an aged
female of very large size, mounted from materials belonging to
several individuals secured by our excavations in Phillips Co.,
Kansas, under the direction of Dr. Wortman in the months of
September, October and November, 1894.
By the comparison of the 16 skulls and 13 jaws, representing
both sexes and all stages of growth, we are enabled for the first
time to positively define the animal long known as Afphelops
fossiger, to distinguish it both from Rhinoceros and Aceratherium,
and point out its important sexual and individual variations. The
writer’s attention was first drawn to the largely disregarded sexual
and age characters of fossil Ungulates in studying the group of
Titanotheres ;' the extinct Rhinoceroses conform to the laws
which were observed in that group, and which are familiar enough
among living types, namely: males, of larger size with more
robust and rugose skulls; horns, if present, more prominent ;
canines largely developed; incisors and anterior premolars dis-
_ appearing in adults.
We owe to Hatcher® the valuable demonstration that Aphelops
fossiger bore a terminal horn upon the nasals, although he as-
signed this character to a type which he supposed represented a
PP. 157-107-. i ; :
* American Geologist, March, 1804, pp. 149-150.
[51]
52 Bulletin American Museum of Natural History. [Vol. X,
major proves to be a middle-aged male of A. /ossiger, and _ his
distinction of Zeleoceras as a genus supersedes Aphelops Cope,’
because Cope’ originally applied the term Apfhelops to A. megalodus,
defining it as an Acerathere with only three premolar teeth in the
lower jaw. This is true of the type species (4. megalodus), but
this species should, so far as we know at present, be referred to
the genus Aceratherium, in which the lower premolars vary from
four to three in number according to age and individual variation,
as in the living Rhinoceros.
Technically, however, Hatcher’s definition did not clearly dis-
tinguish Zeleoceras from Rhinoceros, as he himself stated (of. c77.,
page 245). Our abundant material proves not only that Ze/eoceras
is a Rhinoceros with a median horn on the tips of the nasals, but
that it is fully distinguished from the genus AAznoceros as follows :
Horns. Lower Premolars. Digits.
Genus RAznoceros.. Upon anterior portion of nasals. .4 in young, 3 in aged
individuals........ 3-3
Genus 7¢e/eoceras...Upon tips of nasals........... 3 in young, 2 in aged
individuals. .......3-3
The reduction of the lower grinders to 5 in 7. fossiger (as com-
pared with 6 in RAznoceros) is a very important and distinctive
character, as it absolutely excludes Zeleoceras fossiger from the
ancestry of any of the modern Rhinoceroses, and shows it to have
represented a distinct side phylum, as Scott and Osborn had
already determined from its skeletal characters.
EXCAVATION OF THE SKELETON.
The Phillips County Quarry,* near Long Island, Kansas, was
discovered in 1883 by Mr. Charles Sternberg, who collected for
the University of Kansas and for the Harvard University Museum.
From the latter collection Scott and Osborn procured materials
1 American Naturalist, March, 1894, pp. 241-246.
2*On Some New Extinct Mammalia from the Tertiary of the Plains.’ Palason, Bull. No.
14, Proc. Am. Phil. Society, July 25, 1873.
8 See Williston, ‘ Restoration of Acerather*um fossiger Cope,’ Kansas University Quarterly,
Vol. II, No. 4, April, 1894, pp. 280-290. 3
1898. | Osborn, Skeleton of Teleoceras fossiger. 53
for the restoration which they published in 1890." Subsequent
collections were made by Sternberg and Hatcher for the United
States Geological Survey, between 1884 and 1886. Later Professor
Cragin collected here, and in 1891 Mr. E. P. West of the Univer-
sity of Kansas, aided by Mr. T. R. Overton, began the extensive
collections which led to the preparation of the skeleton for the
University under the direction of Professor Williston.* This skele-
ton, as mounted in the Kansas Museum and described by Williston,
gives a much more accurate idea of this animal than the pre-
vious restoration by Scott and Osborn, in which the chest is
represented far too shallow.
Its principal dimensions are as follows: Length, not including
fail, go it. > height, 4 ft:; greatest girth, 9 ft., 4 in.
The measurements of the American Museum skeleton as mounted
are: Length, 1o ft. 2 in. to bend of tail; height at withers, 4 ft.
I in.; greatest girth, g ft. 2 in.
From the above accounts, and especially from our own observa-
tions, it is seen that this quarry represents an old bone-bed, probably
the deposit of some stream or small river along which the rhinoce-
roses herded in great numbers. The materials collected by the
American Museum party are extremely numerous, especially in
the skeletal parts, the figures running somewhat as follows: Skulls,
Meastapilce-e7r vettewice, £59 ; humeri, 13 > radii, 207; ulnz, 10’;
carpals, 90; metacarpals, 38; pelves, 5; femora, 8; tibiz, 20;
astragali, 22; calcanea, 18.
In this typical bone-bed are mingled individuals of both sexes
and of all sizes, and the proximity of one specimen to another is
not a certain guide. There are certain spots, however, where
considerable portions of individual skeletons have drifted to-
gether. We associate the skull and pelvis in our mounted speci-
- men, for they are of similar age and were found within about six
feet of each other, the skull being that of a fully adult female,
and the pelvis indicating a corresponding age, because the ilia
are united above the sacrum; with the pelvis moreover was found
a part of the jaw belonging to the skull; also with this pelvis
_ 1 ‘Preliminary Account of the Fossil Mammals from the White River Formation contained
in the Museum of Comparative Zodlogy.’ Bull. Mus. Comp. Zodl., Vol. XIII, No. 5, p. 92,
1890.
2 OP. cit.
54 Bulletin American Museum of Natural History. (Vol. X,
belong a femur, tibia and fibula, astragalus, calcaneum and cuboid
of one side, several metacarpals and metatarsals and two cervical
vertebra. The selection of the other limb and foot-bones was made
from these as a guide.
Similarly about 300 feet distant were found the principal ribs
which have been selected for this mount, characterized by the
very rugose appearance and oblique lines for the insertion of the
abdominal muscles (sacro-lumbalts, longissimus dorst). Near these
ribs were large jaw and limb-bones corresponding in size with
those placed in the mounted skeleton. Apart from these proba-
ble associations, the main principle of selection adopted through-
out has been that of the age and size standard, after a careful
comparison of all the elements. In each region the largest and
oldest bones were chosen. Upon this principle the ribs are shown
to be of very great length; the chest girth exceeds that indi-
cated in the Scott-Osborn restoration, and equals that in the
mount in the Kansas Museum, which has heretofore appeared
extreme. In additional support of this correlation of material
belonging to different individuals, this bone-bed gives evidence
of the existence of only one species of Rhinoceros, namely,
T. fossiger. All the differences observed are due to growth,
individual and sexual variations, as set forth below.
The following description is supplementary to the very full
statements of the skeletal characters of Z. fossiger made by
Osborn in 1890.
DESCRIPTION OF MOUNTED SKELETON.
(Museum Catalogue Number, 2604.)
Mounting.—The composite skeleton shown in ihe accompany-
ing plate (Plate IV) has been mounted with remarkable skill by
Mr, Adam Hermann, preparator. All the bones are traversed by
small steel rods, rendering them firm and solid and the limbs self-
supporting. As shown in the photograph, the only visible parts
of the metal framework are the two uprights for the shoulder and
skull, and pelvis, ‘The bones are in a beautiful state of preserva-
1898. | Osborn, Skeleton of Teleoceras fossiger. 55
tion, and except in the case of the artificial elongation of a few
of the ribs and completion of the upper border of the scapule
(from complete scapulz of smaller size) no plaster was necessary.
Skull—Nasals smooth, expanding into a laterally-compressed
beak anteriorly, probably characteristic of adult females, with
sharp sides and a lateral notch. Temporal ridges forming a
sessile sagittal crest ; premaxillaries with triple infraorbital fora-
mina ; lachrymals with well-defined, knoblike projection ; zygo-
mata very deep, with extensive attachment for masseter muscle
inserted on a well-defined ridge on angle of jaw. Occiput broad
and low. Jaw with a single mental foramen below the second or
third premolar, and a marked median depression between the
canines upon front surface of the chin,
Dentition. — Formula: I, C$, P3, M3. Lower and median
upper incisors vestigial; lower canines worn, enamel measuring
4o mm. (1% inches). Adult condition indicated by very slight
wear of crown of third superior molar. Upon outer surfaces of
upper grinders parastyle nearly obsolete. First upper molar with
enamel crown of same length as that of premolars. Second
upper molar with sudden elongation or hypsodontism, with enamel
crown measuring 7omm. Thus m3 and m% elongate or hyp-
sodont, and of great service as reserve teeth for old age. Molars
with secondary folds characteristic of the species.
Vertebre.—Atlas much narrower than in A&. zzdicus, with verte-
braterial canal directly traversing the transverse process ; axis
with a very low spine; cervicals 1-3, with transverse process
restored ; characters of inferior lamellz somewhat conjectural.
Supposed Vertebral Formula: C.7, D.19, L.3, S.5.
The above formula is purely conjectural. It is made to con-
form to that of the living 2. wuicornis and R. sumatrensis.’ Sev-
enteen of the ribs are provided with both capitular and tubercular
facets. The ribs are extremely long and powerful, not very widely
arched ; lower line of chest nearly reaching the ground, as in
1 The definitely ascertained formula of the Oligocene A. ¢7/dactylum is D, 19; L, 5; S, 3.
This animal gives us no clue to A. /ossiger, because it belongs to the Dicerathere series.
56 Bulletin American Museum of Natural History. \|Vol. X,
the Hippopotamus ; girth (9’ 2’’) exceeding that of R. wnicornis
(8’ 9’’) ; chest section deep and heavy, rather than rounded as in
R. unicornts.
Scapula very characteristic, triangular in contour, with pointed
upper border ; narrow supraspinatus and very broad, triangular
infraspinatus fossa ; acromion placed midway on spine, reflected
backwards, so that attachment of deltoid and trapezius muscles is
elevated. Coracoid process forming a prominent rugosity for
short head of biceps ; bicipital tendons passing through a double
osseous groove upon front of humerus (as in A. umicornis, R. bi-
cornis and R. simus)"* and inserted distally in a prominent rugosity
upon front of radius. Humerus exhibiting prominent rugosity
for subscapularis muscle interior to inner tendon of biceps ;
greater tuberosity consisting of a large incurved hook for supra-
spinatus muscle, and a separate and distinct knob for infraspinatus
muscle ; powerful deltoid ridge, everted but not hooked ; distally
a large external condyle for extensor muscles, which exhibit
rugose insertion areas in the proximal ends of metapodials.
Pelvis with ilia arching over and coalescing above the five sacral
vertebral spines, affording a stout area of origin for /atssimus
dorst, erector spine and gluteus maximus muscles, correlated with
support and propulsion of the enormous abdomen; a foramen
piercing the posterior superior border of the ilium. Femur with
rugose but not very prominent greater trochanter; lesser trochan-
ter for insertion of gluteus maximus muscle, much less promi-
nent than in 2. wacornis, and not hooked ; third trochanter not
prominent. Pubis and ischium more slender than in &. waccornis.
Tibia with a characteristic fissure in the cnemial crest dividing it
into two distinct tuberosities for patellar tendons (unlike single
crest of R. unicornis and A. malacorhinus). Fibula fused with
tibia in aged individuals.
? Busk‘ On the Ancient or Quaternary Fauna of Gibraltar,’ Trans. Zo6l. Soc., 1877, p. 97.
2 De Blainville, ‘ Ostéographie,’ Atlas 3, Gen, Rhinoceros, PI. iv.
1898. | Osborn, Skeleton of Teleoceras fossiger. 57
COMPARATIVE MEASUREMENTS.
Teleoceras Rhinocerus
Sossiger. indicus.
Feet. | Meters.| Feet. | Meters.
Motaiglenathetoy bend Ofitailie anes ces. =o. o'-- 10. 2 | 3.10 | 10.8 3.05
Height, Stl eA er aceers ante 6 toe PARES Cerne Ax Ty | es
WILE GG Hye operat cei eh ayar chek 8) drac.s fohcy 0) se Sool! Spiel som 5.64 | 1.69
Breadth, across pelvis. . Sas Sane Sere 2.10 | 0.87 2.8 0.82
oS SRLS ors Graig wri Ie eae aoe 2.11 | 0.89
Skull, fengthucondiyles toys jeriets ste) stale) eel I. 11} 0.59 2.1% | 0.65
width across arches, dental series....... Ee 3)0538
Vert. column, total excluding Gardals 0s 3st wnat: FeO 2530 8.2 2.48
Cervicals, including intervert. pyacee SA Oot O. Oe I. 64) 0.47
Dorsals, Bs A oa anes 4. 64] 1.39
Lumbars, SS “a RMON See Ncoags sate ee oe “5 5| Ose
CANGRAIIS Spee eS AROS Oho ee, eR ECR ee on On] (eg 0.21
Rib (? sth)—length (Groundtthetcunve)e neo s|\02- a 0.80
(Pao) ET dee anasto n so Seep eae eno cee 3. 34\) 1-08
Girth of chest at 11th rib (estimated).......... O), 2 |) BAKO 2.9 2.66
Fore limb, total flexed (ball of hum. vert. to
LOU) eect crers ates ce aly AICHE EER Pae Rees 2. 44| 0.72 3.5 1.04
GA Pla tems wins aerstaiss Sichala'at siete laid a a FOUuol| toca dal pOLste!
NSAI CLUS eteralieys fel hore clepeaa ray sietete eis stam velave ts Seapevete< Net ets OTL 1.4 0.40
IReeUC ULI Sue peeeee Peer Red estat cate casey ersitnis tov ate “ei /oy et ei|t Tal wie ate 0.24 a3 0,38
WOlitrveampeereetetets co ete ttcae ty cae tepe es hits alae elo leldeaielats $06 406 0.34
IWC yvewere, sre cihe cracks celeste icisre oo avale a S65 ||" 4doah al Chios 8 0.20
Hind limb, total flexed (ball of femur vent. to
ground) RS OIA ia SRO e Rea aan LS TEOr 2. 74| 0.80 3.84 | 1.13
ELE TU tgueeerensten weve ed ty oepotet se artont ate etoile, ous ofa: oa" St imiasis Sele nests | LOUAT 1.8 0.52
TUITE by Selenite nice DO Sno ae etna ae error) OF23 1.24 | 0.36
INE S006 Goto Ae Os oe oe ENERO ELI Rete ee a ae om O.LOO4l 77s Bell Ong
Lower jaw, total length, condyle to tip of
GRINNING omactehd of Hee LH aH EE PRIA te ep rocices OSH!
Wentition toltiprolscanine:.5-.-2-.--. .-- betel rouse 0.35
(Cian? SEIntES Ao Cbd ce 6 + coe eee ee! (eee 0.26
Piltenothicaudalsi(alonesctinve) a. +. 22.6522 -6\-- 350 bon 5] (Opts
From the above measurements it appears that from head to tail
T. fossiger is only six inches shorter than 2. wnicornzs, while the
_ back is eighteen inches (.580 mm.) nearer the ground. This
remarkable lowering of the trunk is chiefly caused by the great
reduction of the fore arm, fore leg and metapodials. The hume-
rus and femur are respectively only g0 and 110 mm. shorter than
in 2. untcornis, while the radius and tibia (typically shorter ele-
ments) are respectively 140 and 130 mm. shorter, and the meta-
carpals and metatarsals are respectively go and g50 mm. shorter.
This limb reduction is very striking. At thesame time the abdom-
58 Bulletin American Museum of Natural History. \|Vol. X,
inal girth exceeds that of 2. unicorns, justifying Cope’s conclusion
that this animal had rather the proportions of the Hippopotamus
than of the Rhinoceros. It will be recalled’ that R. wnicornis has
a lower abdominal line than A. sondaicus or R. sumatrensis, or than
either of the African Rhinoceroses. 7. fossiger, therefore, had a
totally different external appearance from any existing form.
JUVENILE, SENILE AND SEXUAL CHARACTERS.
There are conspicuous differences in the dentition of different
specimens, all of which may be explained as due to influences of
growth or sex.
In the young calf jaw (No. 2608) the milk cutting teeth are as
follows: diz, dcz, dpy. In the young A. sondaicus (R. javanicus),
according to de Blainville, we similarly observe two milk incisors.
A somewhat older calf of 7. fossiger shows diz, dc;z, dpz.
Even in older jaws there is evidence in one case (No. 8391) of
two lower incisors upon one side, the formula being: 13-4, Cq, Pz,
mz, as indicated by the incisor alveoli. The outer incisors (iz) in
the lower jaw tend to drop out ‘at an early age, leaving only the
alveoli ; but the vestigial upper incisors (i2) are remarkably tena-
cious, although entirely useless.
The canines vary strikingly in the sexes. In the females (Nos.
2604-6, 2610-11, 2623), as shown in the photographs, they are of
moderate size. In certain males (Phillips Co., Kansas, No. 2612 ;
Republican River, Nebraska, Nos. 8391-2) they exceed in size any
that have been recorded in other Rhinoceroses living or extinct,
as shown in Pl. IVa.
No jaw shows any vestige of pz. Ps is present in young jaws,
and invariably absent in very aged jaws. Its dehiscence is cor-
related with the coming into use of m3. The upper molars,
especially the second and third, are extremely hypsodont, the un-
worn enamel of the crown measuring, respectively : m2,=m3=.
They are reserved for middle and old age.
The size of the skull differs considerably in the two sexes, the
female skulls (Phillips Co., Kansas, Nos. 2604, 2607, 2622-3; De-
catur Co., Kansas, No. 8388; Republican River, Nebraska, No.
_ 1 See Sclater ‘On the Rhinoceroses now or lately living in the Society’s Menagerie,’ Trans,
Zo6l. Soc., 1875, pp. 645-651.
1808. | Osborn, Skeleton of Teleoceras fossiger. 59
8393) being smaller and less rugose, with less prominent sagittal
crests, and decidedly smaller nasals, as shown in Pl. IVa. In old
females the nasals acquire a slightly rugose surface, and probably
bore a small horn. In the males (Decatur Co., Kansas, Nos.
8385, 8396; Republican River, Nebraska, No. 8420) the nasals
become greatly thickened at the extremities (Pl. IVa), forming a
vertically compressed plate, which undoubtedly bore a consider-
able horn. Differences in size are observed in skulls from various
localities, those from Decatur Co., Kansas, and from Nebraska,
being larger than those from the Phillips Co. quarry, which are
probably due to differences of geological level, the species run-
ning into a larger and more robust type before its extermination.
Growth-changes in the limbs are especially observed in the
close fusion of the fibula with the tibia, and of the remarkable
arching over of the sacrum by the superior borders of the ilium ;
this whole area above the sacrum forming a solid plate.
CONCLUSIONS.
T. fossiger may be briefly characterized as a brachycephalic,
extremely short-limbed Rhinoceros, partly aquatic in its habits,
with a very large brain and no diploé of the skull.’ It parallels
the African Rhinoceroses #. s¢mus and &. bicornis, in the form of
the humerus, femur and atlas, and in the terminal position of the
nasal horn. The occiput, however, is widely different from that
of the African Rhinoceroses, as well as of AR. sumatrensis, resem-
bling rather that of &. waicornis, although less pitched forward,
The limbs are much shorter than in any living type, and,
as pointed out by Pavlow,’ at once recall those of &. érachy-
pus and R. aurelianensis. A further comparison of 7. fossiger
strengthens the resemblance to the latter form. The proportions
of the skull, limbs and metapodials are very similar. In both the
cnemial crest of the tibia is double; the secondary folds of the
superior molars are similar, as well as the general form of the
skull. Further details will be given in the writer’s forthcoming
Memoir on the Extinct Rhinoceroses.
1 See Scott and Osborn, of c7t.. 1890, p. 93.
2+ Les Rhinoceridz de la Russie et le développement des Rhinoceridz en général,’ Bull. d.
la Soc. d. Nat. d. Moscou, 1802.
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ArticLE VI, pp. 81-91. _
‘Article VI-—A COMPLETE SKELETON OF CORY-
PHODON RADIANS. NOTES UPON THE LOCO-
MOTION OF THIS ANIMAL.
By HENRY FAIRFIELD OSBORN.
PLATE X.
The chief object of the writer in planning the American
Museum Expedition of 1896 was to complete materials for the
investigation of the evolution of the Amblypoda, and extend our
knowledge of Cvrypiedon. ‘The observations of Cope, Marsh,
Osborn, Wortman and Earle have been principally upon scattered
and imperfect material, and it seemed of the utmost importance
to secure materials sufficient to determine the relations of this
animal to its ancestral form, Pantolambda, and to its successive
form, Uzntathertum ; also the proportions of the body, the posi-
tions of its limbs and the number of its vertebre. Accordingly
our party, led by Dr. Wortman, aided by Mr. Granger of the
Museum, and Messrs. Brown and Riggs of the University of Kan-
sas, spent the months of April and May in northwestern New
Mexico, revisiting the locality where Cope’s most complete Covy-
phodon, C. elephantopus, had been found. The search here in the
‘Coryphodon’ or ‘Wasatch Beds’ was entirely unsuccessful, but
fortunately the underlying ‘ Torrejon Beds’ yielded a remarkably
complete series of Pantolambda. The party moved to the north
in June, and devoted July and August to a most energetic explo-
ration of the Big Horn Basin, especially of the exposures on the
south side of the Gray Bull River from Brown’s Ranch towards
the Big Horn River below Otto.
The party soon discovered two skulls, both in the sandstone.
The first (No. 2867), upon level A, with four vertebre and some
fragments of limb bones associated with it, the teeth being badly
| March, 1898.| [81] 6
82 Bulletin American Museum of Natural History. (Vol. X,
weathered, has been mounted in our skeleton. ‘The second (No.
2963) was found upon level B, 4o feet higher; it exhibits the
form of the top and back of the skull, together with the complete
teeth and palate. Shortly afterwards, upon the same level A as
No. 2867, the skeleton (No. 2865) was discovered with fifteen ribs
and vertebrz, the humerus, ulna, radius and two or three of the
sternal bones. This was followed by the discovery on level A of
the hind limb (No. 2869), several pairs of jaws, and finally, upon
level B, the part of a crushed skeleton (No. 2829), including the
skull, jaws, all the ribs and vertebrae in position. This, as Dr.
Wortman wrote, made the party absolutely certain of a mount.
The four skulls; Nos. 2827, 2867, 2963, 2865, with the associated
skeletons, were thus found substantially upon the same geological
level, they are in the same stage of evolution, and are found to
belong to the classic species C. (Bathmodon) radians Cope, the
first known in America.
The party then moved to the overlying Wind River Beds, and
discovered a unique skull (No. 2977) of a distinct species, which
reverses the natural order of evolution, since the sagittal crest is
a little broader than one’s finger. This exhibits the desired tran-
sition between Pantolambda and Coryphodon, and represents, in all
probability, a persistent primitive type.
Altogether parts of 18 individuals were found in the Wasatch
Beds (supplementing the 30 individuals found by Dr. Wortman in
1891), and 7 individuals in the Wind River Beds. ‘The selection
of nine individuals for mounting was done with great care as fol-
lows: The mounted skull, No. 2867, agrees exactly in size, and is
specifically identical with the skull and jaws of No. 5829. ‘The lat-
ter (No. 2829), while laterally crushed, had associated with it the
right scapula and complete forelimb, left scapula and parts of left
limb which were used in mounting ; also all the vertebrze as far
back as the pelvis; these vertebra, while too much crushed to
mount, enabled us to determine the formula and select, from series
Nos. 2865 and 2863, vertebree which exhibit the same characters.
The latter individual (No. 2863) included the pelvis and hind
limb, thus determining Aosztevely the correct proportions of the entire
animal. ‘Vhe mounting was done with great skill and care by Mr,
Hermann,
1808. | Osborn, Skeleton of Coryphodon radians. 83
DESCRIPTION OF SKELETON.
Composite of nine individuals, all from the Big Horn Beds, as follows: No.
2867 ; skull, left humerus, left mtc. I and ectocuneiform ; caudals 1-3, 10-11.
No. 2867, lower jaw. No. 2829; right scapula and fore-limb complete ; left
scapula, magnum, trapezium, trapezoid, metacarpals I-V and phalanges. No.
2865 ; all cervical vertebre, dorsals and ribs I-g complete, 3 phalanges of right
pes. No. 2963 ; dorsals 10-15 and ribs complete, lumbars 1-5 complete, pel-
vis, left pes (excepting mts. I, III-V, and ectocuneiform). No. 2869 ; right
hind limb complete (excepting mts. IV—V and 3 phalanges. No. 4320, left
ulna, femur and tibia. No. 258; caudals 4, 5, 8. Sternals complete from No.
2825. Restored - left radius, left fibula, right mts. IV—V, left mts. III-V, cau-
dals, 6, 7, 9, and 12-22.
In general one is struck by the very large size of the head,
formidable front teeth, the shortness of the ribs, the heavy char-
acter of the girdles, the heavy limbs, and the semiplantigrade or
subdigitigrade condition of the feet. It is probable, as already
shown by the writer, that in the hind foot the calcaneum nearly
touched the ground in the forward step.
The sku// presents a very peculiar appearance with its power-
ful and spreading upper and lower canines, and widely spaced
incisors, slender zygomatic arch and broad, flattened cranium.
The following characters distinguish this specific type: Premaxil-
laries short, not reaching nasals; free portion of nasals short ;
nasals projecting between maxillaries, and then spreading upon
inner side of frontal tuberosities; naso-frontal suture disap-
pearing posteriorly ; maxillaries occupying a broad area and bulg-
ing Out opposite the canine alveoli and concave behind; lachry-
mals not clearly defined ; frontals above the orbits supporting two
prominent knobs or convexities, confluent with parietals poster-
iorly ; parietals expanding above middle portion of temporal
-fossa, and lateral parietal crest thickening, as indicative of a rudi-
mentary parietal horn; mid-parietal region depressed and
extremely rugose for muscular attachment (occipito-frontalis) ;
malars extending to the front of orbit, exhibiting a downward
masseteric process slightly behind the orbit, expanding widely but
slender in section ; postglenoid processes for squamosal rather
slender; external auditory meatus widely open, and paroccipital
and posttympanic region compressed into a narrow ridge; occiput
84 Bulletin American Museum of Natural History. [ Vol; xe
low and broad; lower jaws with condyle facing upwards and
backwards; prominent depression posteriorly; a mental foramen
below first premolar, a second mental foramen below lower
canine.
Dentition.—The superior teeth, wanting in this specimen, are
perfectly preserved in No. 2829, and agree with the fragmentary
type specimen of Coryphodon (B.) radians Cope.
Vertebral formula: C.7, D.15, L.5, 5.4.
This remarkably low vertebral formula is ascertained from
No. 2829, in which all the dorsals and lumbars are retained in a
single block and can be counted with considerable certainty.
The formula is very low, but not more so than in 7Z?¢anotherium,
in which D. L.=20. ‘lhe chief characteristic of the vertebral
column is the series of low, undifferentiated neural spines, which
are ill correlated with the heavy skull.
Cervicals—Atlas moderately broad with a slightly expanded
transverse process, perforated slightly above the base by the ver-
tebrarterial canal. ‘The spine of the axis extends equally forwards
and backwards. The remaining cervicals exhibit a gradual
development of the inferior lamella, which is well marked in C.6,
but lacks the strength exhibited in the larger Perissodactyla.
The cervical centra are very short, and the vertebral centra grad-
ually increase in length and depth toward the lumbar region,
The most striking feature of the dorsal vertebra, which is
shared by Phenacodus, is the great prominence of the transverse
process supporting the tubercle of the ribs. This projects widely
out from the side of the vertebra in D.1, and gradually recedes to
D.11, which is apparently the last vertebra in which the rib tuber-
cle articulates. In all the dorsals the head is placed directly be-
tween the adjacent vertebre from D.1 to D.15. The low spines
characteristic of the cervical region extend back as a feature of
the dorsals, the vertebrae exhibiting terminal tuberosities for the
fascia of the hgamentum nuche. ‘The dorsal vertebral spines
gradually thicken in antero-posterior diameter; as they pass
backwards they decrease in height. In the lumbar region they
are cleft on the dorsal line. |The lumbars terminate inferiorly in
compressed keels.
mn
1898. | Osborn, Skeleton of Coryphodon radians. 8
MEASUREMENTS OF SKELETON.
|U FEET AND. INCHES. METERS.
Length incisors to perpendicular of | |
Si tad ta Hee ene ga | 7 9g e235
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Fore Limb :
SGApUl ave yas l= ORE BAS Ty iSte 43
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IEC UIGHS Ria tera pip soi oa actn ene 10" ae Ol
Miamiisstotalane cena sects oh 7 \ .82
Hind Limb : yf
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ISM UIrseran Soraya ate fo t 644 47 ) 2’ rrilg'
ALT OY Ee cy ROUGE UN See ae ae ne Tale .28 or
es htotall Sir rene coc eeere 4 che 6" 15 \ .gO
These measurements show that the tibia is only an inch longer than the radius, while
the femur is three inches longer than the humerus.
The anterior pair of 77s is extremely short, the succeeding
ribs increase in length and decrease in diameter, passing from a
flattened into a trihedral form in D.6 and 7, and finally into an
oval form in D.10-15. A characteristic feature of the ribs of D.
6-11 is a pit upon the upper surface just external to the
tubercle.
The zygapophyses have horizontal faces as far back as D.15.
In D.14 and 15 they turn obliquely upwards, the faces being
vertically flattened. In L.1 to L..5 the zygapophyses are sharply
concave and nearly vertical in position. The characters of the
caudals are not certainly known, the few centra being restored
from a number of different specimens. We have not ventured to
give the tail the remarkably flattened character already described
in a specimen found in 1892.
Fore Limb.—The scapula is vertically elongate, terminating ina
point superiorly, and distinguished from that of U7zntathertum by
nearly subequal supraspinatus and infraspinatus fosse. It is
partly restored in the mount, but perfectly preserved in No. 2873.
The spine rises near the superior border, is slightly thickened and
eflected, and passes down into the acromion process, which turns
sharply forwards and overhangs the great tuberosity of the
86 Bulletin American Museum of Natural History. |Vol. X,
humerus. The humerus is distinguished by the prominent
greater tuberosity, which enters inferiorly the prominent and
recurved deltoid ridge, extending far down upon the anterior
surface, two-thirds the length of the shaft. The characters of
this bone are best seen in No. 2780, an animal of the same size.
The entepicondyle is a rugose tuberosity, the ectepicondyle is
more elevated on the shaft, and is marked by an anteverted
ridge. The forearm is perfectly preserved upon the right side.
The radius covers the front face of the humerus, and the distal
faces of the ulna and radius are placed obliquely te the transverse
axis of the body, facing upwards and throwing the fore feet and
toes outwards rather than forwards. In this specimen the cunel-
form does not articulate with the fifth metapodial, as observed in
certain other specimens. The manus, as above described, is sub-
digitigrade, the lower surfaces of the proximal ends of the meta-
podials being slightly raised above the ground. The position of
the metapodials in the forward step is, however, much more
oblique than in the manus of the Elephant, the lower surfaces
being nearer the ground.
The felvis is partly restored in this mount. Its characters are
better shown in No. 258. The iliaexpand widely, but the antero-
inferior border is not extended very far down. The ischia and
pubes are strongly developed, and enclose. a wide obturator
notch.
A very characteristic feature of the skeleton is the long and
rather slender form of the femur and the disproportion between
the femur and the tibia, which is much greater than that which
exists between the humerus and the radius. The great trochan-
ter does not rise to the level of the head. ‘The third trochanter
is a long rather low crest, much less strongly marked than in
Pantolambda, placed on the upper third of the inner side of the
shaft. The lesser trochanter is very prominent, and lies slightly
below the middle of the shaft. The patella is a very character-
istic bone, but there is some doubt as to its position; it appears
probable that the slender pointed spine of the patella faces up-
wards, as the long patella facet is thus made to correspond with
the long facet on the inner side of the front face of the trochlea of
the femur. The tibia is a very stout bone with a rather low
1898. | Osborn, Skeleton of Coryphodon radians. 87
cnemial crest, and rests by a slightly concave distal face upon the
broad flat astragalus. The position of the pes is probably fairly
represented in the left right foot, and although it is possible for
the astragalus to be brought still nearer to the ground in the long
forward step, it appears that in this type plantigradism is not so
marked as has been stated by Osborn; probably the different
species varied in this respect. As in the fore feet, the median
digit faces outward. ‘The astragalus has a well-marked astraga-
lar foramen.
GENERAL APPEARANCE OF CORYPHODON.
The most accurate forecast of the appearance of the animal
was that made by Cope’ in 1874 :
“The general appearance of the Coryphodons, as determined
by the skeleton, probably resembled the Bears more than any liv-
ing animals, with the important exceptions that in their feet they
were much like the Elephant. Tothe general proportions of the
Bears must be added a tail of medium length. Whether they
were covered with hair or not is, of course, uncertain; of their
nearest living allies, the Elephants, some were hairy and others
naked. ‘The top of the head was doubtless naked posteriorly,
and in old animals may have been only covered by a thin epider-
mis, as in the Crocodiles, thus presenting a rough, impenetrable
front to antagonists.
“The movements of the Coryphodons, doubtless, resembled
those of the Elephant in its shuffling and ambling gait, and may
have been even more awkward, from the inflexibility of the ankle.
But, in compensation for the probable lack of speed, these ani-
mals were most formidably armed with tusks. These weapons,
particularly those of the upper jaw, are more robust than those of
the Carnivora, and generally more elongate, and attrition pre-
served rather than diminished their acuteness. ‘The size of the
species varied from that of a Tapir to that of an Ox.”
Osborn’ in 1892 wrote as follows :
“The fact is, the position of the fore and hind feet of Cory-
phodon is absolutely different. he fore foot was digitigrade,
like that of the Elephant, the A7nd foot was plantigrade, \ike that
of the Bear. In other words, the carpus was entirely raised from
the ground and the manus rested upon the distal ends of the meta-
eats and upon the spreading phalanges; while the caleaneum
1 ifeatenite Palsontllcsy. V ‘ih IV, Ww heeler eee P- 203 :
2 Fossil Mammals, of the Wasatch and Wind River Beds, Collection of 1891, Osborn &
Wortman, Bull. Am. Mus. Nat. Hist., Sept., 1892, p. r2t
88 Bulletin American Museum of Natural History. |Vol. X,
and tarsus rested directly on the ground together with the entire
plantar surface of the foot. This substantial difference between
the advanced state of evolution of the fore foot and retarded
evolution of the hind foot, is of great interest. It is clearly
shown in the accompanying figures.”
—@
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Fig. 1. Coryphodon hamatus, as restored by Marsh, 1893. One-twentieth natural size.
In 1893, Marsh,’ in his description and restoration, presented
quite a different conception of the animal as wnguligrade. In
regard to these matters he makes the following statement :
“This restoration is given one-twelfth natural size. The posi-
tion shown was. chosen after careful consideration, and is believed
to represent fairly one naturally assumed by the animal in life
when standing at rest. The figure represents a fully adult indi-
vidual, and one of the largest species of the genus which, when
alive, was nearly six feet in length and about three feet in height.
The basis of this restoration is the type specimen of Coryphodon
hamatus. This was supplemented by other remains, which
appeared to be superficially identical. A large number of such
specimens were available, some of them in excellent preservation.
For parts of the skeleton where such remains were wanting speci-
mens from nearly allied forms were used, but no serious error
can thus result.-..The fore feet presented 4n the ‘present
restoration are constructed mainly from the same specimen (that
first figured and described by the writer), and the position given
in the original figure has been essentially retained....The posi-
tion first given to the figure is retained in the restoration after
a careful investigation of the whole posterior limbs in a number
of well-preserved specimens. In Denoceras the terminal
1 * Restoration of Coryphodon,’ Amer. Journ. Science, Oct., 1893, p. 324.
Osborn, Skeleton of Coryphodon radians.
1898. |
so that they
ly free,
thus bore a greater weight, the digit being undoubted
phalanges are much larger than in the Elephant,
although a pad may have helped to support the feet.
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phodon the digits were still more elongate and the termi
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were covered with hoofs that supported the feet. This would
agree with the position given them in the restoration, which coin-
cides with the anatomical structure of the entire hind limb.”
This clear statement of Professor Marsh’s as to the position of
these limbs is consistent with his restoration, in which not only
the metapodials but the phalanges are raised from the ground,
and the animal is represented as walking upon the tips of its toes,
or terminal phalanges, the latter being supported by hoofs. The
morphological importance of this restoration is very great. If
correct it places Coryphodon among the Unguligrada, widely
removed from the unquestionably plantigrade Puntolambda.
Contrary evidence that Coryphodon, so far from being unguligrade,
was transitional between complete plantiyradism and digitigradism,
is given below. Many other important morphological characters
are involved in Marsh’s restoration, and are now found to be
incorrect. The scapula is given a trihedral form, with a very
broad angular infraspinatus fossa; the pelvis is extraordinarily
reduced ; the limbs are elongate and, together with the above-
mentioned unguligrade action of the digits, elevated the body
very much from the ground. In proportion to the scapula, the
humerus, the ulna and the radius are of very great length, and
similarly the tibia is only slightly shorter than the femur. The
most important character, however, is that assigned to the
vertebral column, there being 1g dorsals and, as far as can be
determined from the drawing, 6 lumbars, or D. L.=25, a formula
exceeding that of the Rhinoceros. The net result of these obser-
vations is to give the animal the general appearance and charac-
ters of a modern Perissodactyl,’ with the single important excep-
tion of the five digits preserved in the fore and hind feet.
It appears from our more complete material that the difference
between the feet was exaggerated by Osborn, as already observed
by Marsh. There is no doubt, however, that as seen in the
mounted specimen, in the forward step the calcaneum rested very
near the ground, being separated merely by a thick plantar pad.
The digits of the fore and hind feet have nearly the same relations
to the ground. Both feet are in a somewhat similar stage of tran-
sttion between plantigradism and digitigradism. Pantolambda has
a long tuber-calcis and pes like that of the Bear. Uzntatherium
1808. | Osborn, Skeleton of Coryphodons radians. gi
has a very short tuber-calcis and bore the pes slightly more planti-
grade than the Elephant. Coryphodon has a tuber-calcis inter-
mediate in length ; in the astragaius the upper facet for the tibia
and lower facet for the navicular presents an oblique angle, the
astragalus thinning out to a sharp edge in front (whereas in U7nta-
thertum these facets are more nearly parallel, and the astragalus
is truncate in front). The angles between the tibial and navicu-
lar facets of the astragalus, as shown in sections in Fig. 2,
afford the most decisive evidence that the pes of Coryphodon was
intermediate between the nearly plantigrade Pantolambda and the
sub-digitigrade Uzntatherium.
Coryphodon had a very short back and short, spreading limbs,
with a very clumsy, shuffling gait.
The rudimentary horn observed for the first time in the parie-
tals is prophetic of the great parietal horn of U?ntatherium.
Many other characters of the skull and skeleton are also prophetic,
but there is little tendency displayed to reduce the upper incisors
or lower canines into the Uintathere type.
1 The Perissodactyl affinities of the animal were dwelt upon in Professor Marsh’s earlier
papers (Am. Jour. Sc., 1876, page 428; of. c7t., 1877, page 84), but were abandoned subse-
quently (*‘ Dinocerata,’ 1884, page 177), in which CoryAhodon was correctly associated with
Uintatherium in the Amblydactyla (Amblypoda).
Butcetin A. M. N. H. Woe, Doky eau s Nabe D6 ¢
MouNtTED SKELETON OF Coryphodon radians,
Slightly exceeding one-fifteenth natural size.
rth A
Associated Forms.”
<
Worray, M.D :
3 one wreye yi
AND. TY-THREE T
Br aN i a¥e i ; Eee oe ii ‘ = i eS ie s
THOR’S EDITION, extracted from BULLETIN —
wsenm of Watuval History, —
w York, April 9, 1898.
2
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“
Say
Article VII.—THE EXTINCT CAMELIDA OF NORTH
AMERICA AND SOME ASSOCIATED FORMS.
By J. L. Wortman, M.D.
PLATE XI, AND TWENTY-THREE TEXT CUTS.
Among the living selenodont Artiodactyla the Camels and
Llamas of the Old and New World represent a very aberrant
group. Even if one were not familiar with the wonderful record
of their past history as revealed in the Tertiary deposits of this
country he would be quite justified, from the number of anatom-
ical peculiarities which they exhibit, in placing their origin far
back in the Tertiary, at a time when the primitive divergence of
the various lines of the Selenodonts was taking place.
The evidence is not yet sufficiently complete to trace the
phylum with absolute certainty below the upper Eocone or Uinta
stage, but from this point on to the present time there is very
little to be desired, in the way of intermediate species, to form a
compact and closely connected series, reaching to the modern
types.
More or less elaborate studies of this group have been made
by Cope’ and Scott,’ to whom we are especially indebted for
much knowledge concerning the extinct forms, and while it would
be difficult to add anything to their statements from the speci-
mens known to them, yet the acquisition of a large amount of
new material bearing upon this subject by the various Museum
expeditions within the past few years has rendered it especially
desirable to review the whole subject, with a view to defining, if
possible, the exact limits of the various genera and species of the
extinct North American representatives. The object of the
present paper, therefore, is: (1) A review of the genera and
species of the North American Tylopoda, with descriptions of
1 ‘Phylogeny of the Camelidz,’ Amer. Nat., 1886, p. 611. :
2 On the Osteology of Poébrotherium,’ Journ. of Morph., 1891. ‘The Mammalia of the
Uinta Formation,’ Trans. Amer. Phil. Soc., Aug. 20, 1889.
L93]
94 Bulletin American Museum of Natural History. |Vol. X,
additional materials of known types, as well as of new allied
forms ; (2) a careful consideration of the various steps in their
evolution ; and (3) a study of certain osteological characters of
the higher selenodont Artiodactyla in its direct bearing upon the
transmission of acquired characters.
The oldest members of Tylopoda which we can determine with
certainty are found in the Upper Eocene deposits of the Uinta
Basin, which strata, it may be noted, contain the first remains of
true selenodont Artiodactyla in this country, a fact originally
pointed out by Marsh, who was the first to explore this region for
fossil remains. In the preceding deposits of the Bridger Basin
the remains of Artiodactyla are found, but are very rare, no truly
selenodont types being known. It is trué that in several genera,
such as Homacodon and Helohyus of Marsh, we have a distinct
foreshadowing of the selenodont molar, yet it is not until the
Uinta is reached that the true Selenodonts appear.
Mr. O. A. Peterson, to whom we are largely indebted for the
beautiful collection now in the Museum from this horizon, divides
these beds into three stages,’ which he designates as the lower,
middle, and upper Uinta Beds, or, to use his own field designation,
Horizons 4, &,and C. A large part, if not the entire lot, of these
remains were obtained from the lower part of Horizon C, or the
upper part of Horizon #4, so that their stratigraphical position
would be correctly stated to be at least 300 to 400 feet below the
top, and at least 800 feet from the bottom of the Uinta formation.
No fossils are known from the upper levels of Horizon C, nor do
we know any Artiodactyla from the Brown Sandstones, 800 feet
in thickness, constituting Horizon A. If, therefore, close con-
nections between these Uinta forms and the preceding Bridger
species, on the one hand, and the succeeding White River, Oligo-
cene species, upon the other, are not shown to exist, the fact is,
in all probability, due to our lack of knowledge of the species
which lived during the time of deposit of these intermediate
strata.
The identification of these Uinta Cameloids is attended with
more or less difficulty, owing in part to the fragmentary condition
wre Osborn’s * Fossil Mammalia of the Uinta Basin,’ Bull. Am. Mus. Nat. Hist., Vol.
> P- 74-
1808. | Wortman, Extinct Camelide of North America. 95
of some of the material, and in part to other lines which resemble
them in certain points of skull and limb structure. These
resemblances are, no doubt, due to the close proximity to the
point at which the respective phyla began to diverge.
For the purpose of bringing into stronger relief the characters
of the Cameloids of this horizon, it is necessary to compare them
accurately with the cotemporary Selenodonts, and, since several
of them apparently represent new genera, they are herewith
described.
Leptoreodon marshi,' gen. et sp. nov.
This genus and species is represented in the collection by an
almost perfect skull in good state of preservation, a number of
vertebra, and a few fragments of the limbs (No. 2064), which I use
as the type. There are several other specimens of a more frag-
mentary character which are probably to be referred to the same
Fig. x. Side view of skull of Leptoreodon marshi.
genus and species, but they contribute little additional informa-
tion to the knowledge of the skeleton. The genus differs from
all the Oreodonts hitherto described in the possession of a short
diastema in front of, and a longer diastema behind, the first
superior premolar, together with a considerable diastema between
the first and second premolar in the lower jaw. ‘The incisors are
1 This species is dedicated to Prof. O. C. Marsh, in recognition of his numerous contribu-
tions to American paleontology.
96 Bulletin American Museum of Natural History. |Nol. X,
present in full in both jaws; the inferior canine is small and
incisiform ; the superior canine is large, with the characteristic
D-pattern of the Oreodonts on cross section, and the first inferior
premolar is enlarged and caniniform as in the Oreodonts. The
first superior premolar is two-rooted with a high, compressed
cutting crown, the second is similar but somewhat larger, the third
has a principal broad, lunate external cusp and a faint internal
cingular ledge. The fourth premolar crown is composed of a
single external and internal cusp, much as in the Oreodonts.
The superior molars closely resemble those of Pvroforeodon
(Zomeryx) in the composition of the crown, so far as can be
determined in their advanced stage of wear in the type specimen.
It is impossible to say whether or not there were anterior inter-
mediate cusps present, but judging from certain appearances in
this region of the crown, I am inclined to think that less worn
teeth would show them. The mesostyle consists of a vertical
pillar as in the Oreodonts generally, and not of a wide open loup
as in Agriocherus.
In the lower jaw the incisors and canines are of the typical
oreodont pattern, but they are unusually procumbent in position.
The first premolar is enlarged and caniniform, the second simple,
the third with a small internal cusp and posterior heel, and the
fourth similar in pattern, except that the internal cusp is smaller
and the heel more pronounced. ‘The lower molars are almost
identical in structure with those of the early Oreodonts.
The whole skull differs from that of the Oreodonts in its more
slender proportions. ‘This is particularly noticeable in the lower
jaws, which are relatively long and shallow, especially in the
region of the symphysis, in marked contrast with the deep and
abrupt chin of the Oreodonts in general. There does not appear
to have been a preorbital pit present, and the orbit was not
enclosed by bone posteriorly. ‘The present genus may be distin-
guished from its contemporaries in the following dental characters,
viz.: from Proloreodon (Lomeryx) in the possession of diastema
in both jaws and the full number of incisors in the upper jaw.’
1 In all of our material [ have not yet seen a specimen among the Oreodonts other than
Leptoreodon that has a full set of incisors in the upper jaw. Marsh figures the type of
Eomeryx pumilis with but two superior incisors, and if Protoxeodon has the full complement
as believed by Scott, then the two genera are certainly distinct. In two specimens in the
Musuem collection which correspond closely with Protorcodon parvus, as described by Scott
there is but a single incisor on each side above, and the premaxilla are widely separated from
each other in the median line,
1808. | Wortman, Extinct Camelide of North America. 97
From AHyomeryx it is readily distinguished by. the full number
of superior incisors and by the diastema, although it resembles
this latter genus, which is described by Marsh’ as having more
slender jaws than Protoreodon (Homeryx). From the cameloid,
Leptotragulus, it is easily separated by the numerous oreodont
characters which the skull exhibits, although the symphyseal region
is strikingly similar in the two genera.
Of the hind foot, the cuboid, navicular and the head of the
third metatarsal are sufficiently preserved to afford characters for
identification. These bones indicate an animal with far more
slender limbs and feet than any of the Oreodonts with which I
am familiar. The navicular has an inconspicuous posterior hook
unlike that of the Oreodonts, and, judging from the much reduced
facet on the cuboid, the fifth digit was considerably diminished
in size if not entirely rudimental. The limb-bones are not well
enough preserved to confirm or negative this conclusion of the
slender and delicate proportions of the animal, but, upon the
whole, I think it may be safely concluded, from the evidence at
hand, that Zepforeodon held the same position with reference to
the American Oreodontide that Xvphodon did to the European
Anoplotheriide.
The second genus to be described in this connection contains
species somewhat smaller in size and less perfectly selenodont.
Bunomeryx montanus, gen. et sp. nov.
There are two specimens in the collection which I classify
under this head, viz. : an anterior portion of a cranium somewhat
crushed, containing the maxillary dentition complete upon one
side, together with the greater part of the left mandibular ramus
of the left side having all the true molars and the last premolar
in good preservation (No. 2071). ‘The second specimen consists
of a portion of a lower jaw with a few teeth, the posterior part of
the cranium, a nearly complete fore foot, portions of the hind
limbs and other parts of the skeleton (No. 2070). The first of
these specimens may be taken as the type, but there can be very
1‘ Descriptions of Tertiary Artiodactyles,’ Amer. Jour. Sci., Vol. XLVIII, Sept., 1894,
p. 268.
[April, 1898.] vis
98 Bulletin American Museum of Natural History. |Vol. X,
little doubt that the second specimen is identical with the first and
can be regarded as a collateral type.
This genus most nearly resembles HYomacodon Marsh, from the
Bridger Beds, although it presents some dental characters similar
to Dichobune of the European Eocene. The more important
generic characters may be stated as follows :
Dentition, 13, C1, Pm4, M3. Superior molars, having well-defined cres-
centic outer cusps and a distinct mesostyle and parastyle ; first molar provided
with two conic internal cusps (protocone and hypocone), with anterior and poste-
rior subcrescentic intermediates; second molar having anterior subcrescentic inter-
mediate, subconic protocone, a posterior subcrescentic intermediate and no
hypocone ; third molar similar to second. ‘The superior premolars are present
in full number ; the two anterior have simple cutting crowns, while the crowns
of the third and fourth are made up of single external and internal conic cusps
well developed. In the lower jaw the structure of the molars is intermediate
between the bunodont and selenodont pattern ; there are only three premolars,
the anterior two of which have simple compressed crowns, while the last or
fourth of which is provided, in addition to the principal cusp, with anterior and
internal cusps, together with a well-defined heel.
As compared with Homacodon, Bunomeryx is readily distin-
guished (1) by the possession of three premolars in the lower
jaw ; (2) by the crescentic character of the external cusps of the
superior molars ; (3) by the presence of a well-developed para-
style and mesostyle ; (4) by the absence of the hypocone on the
second superior molar, and (5) by thesubcrescentic character of
the intermediates. (6) The internal cusp of the third superior
premolar is, moreover, better developed in Bunomeryx than in
Hlomacodon, and (7) the fourth inferior premolar is much more
advanced in structure. The structure of the inferior molars is
much more selenodont in Bunomeryx than in Homacodon.
From Dichobune the present genus is readily distinguished by
the absence of all traces of the anterior cusp of the trigon in the
lower molars as well as the more crescentic character of the outer
cusps of the superior molars, and the possession of well-defined
mesostyle and parastyle. The complete adult dentition of Décho-
dune is apparently not known, but there can be but little doubt
that it had the full number, forty-four teeth, in which case Buno-
meryx would be sharply distinguished by the inferior premolar
formula. I cannot at present say in what manner Bunomeryx
1898. | Wortman, Extinct Camelide of North America. 99
differs from the European Dezlothertum, Spaniotherium, Metrio-
theritum, Moutllacithertum and Oxacron of Filhol, which are
placed by Zittel in the subfamily Dichobuninz. On account of
the very imperfect knowledge we have of these forms, no com-
parisons are at present possible.
In specimen No. 2071, the upper and posterior portion of the
cranium is sufficiently preserved to indicate a relatively high over-
hanging occipital and a strong sagittal crest, the latter dividing
into two well-marked lateral postorbital branches. In advance of
the point of division of these two branches a strong ridge is
continued forward upon the frontals in the median line as in
many of the lower forms of the Selenodonts. The postorbital
process is well developed, but it does not join the molar, so that
the orbit is not enclosed by bone posteriorly. There is no evi-
dence of the presence of any long horn-cores.
Of the fore limb, the distal ends of the ulna and radius are pre-
served, but they are considerably crushed. ‘There is apparently
little or no tendency to codssification of the bones, although the
shafts are closely applied to each other in the lower third of
their extent. The articular end of the radius shows distinct
facets for scaphoid and lunar, but does not touch the cuneiform.
The distal end of the ulna articulates solely with the cuneiform.
The carpus is of the typical artiodactyl pattern, and especially
resembles that of the earlier Selenodonts. In the proximal row
the cuneiform rests exclusively upon the unciform, the lunar
about equally upon magnum and unciform, while the scaphoid is
supported below by magnum, trapezoid and trapezium. In the
distal row the unciform articulates distally with Mt. III, and to a
slight extent with Mt. II. In the modern Suillines, the Cameloids
and the later Oreodonts, the second metacarpal has lost all con-
nection with the magnum, but in the early Oreodonts Mt. IT still
retains a contact between these two bones. In /Pvofoceras of the
Oligocene a very minute contact is observable.
The trapezoid is free, and shows no tendency to unite with the
magnum as in Leptomeryx, the later Tragulines and Pecora. The
trapezium is not preserved in the specimen, but judging from the
well-marked facets upon the scaphoid, trapezoid and Mt. II, it is
quite certain that it was not only present and of good size, but
100 Bulletin American Museum of Natural History. |Vol. X,
that it supported a very considerable vestige of the first digit. It
would not indeed be a matter of surprise to find this digit com-
plete in more perfect specimens very much as in Oreodon.
There are four metacarpals preserved of which the median
ones, Mt. III and IV, are the largest and subequal in size. Mt. II
is slightly larger and longer than Mt. V, and in all of them the
distal. keels are confined to the palmar surfaces, as in all primitive
Ungulates. The phalanges of the fore feet are not known.
Of the hind limb the materials are not so complete as of the
fore limb, but enough is preserved to make out its more important
characters. The fibula was much reduced, and probably incom-
plete in the middle part of the shaft. The distal end of the
tibia displays no usual form of the more generalized Selenodonts,
as do the tarsal bones. The cuboid and navicular were not
coossified, and there is evidence of four complete metapodials,
the lateral ones, however, being unusually slender and delicate.
The first two phalanges resemble those of the early Cameloids,
Protoceros and Leptomeryx in their form, as do likewise the
unguals in being relatively high-pointed and _ flattened upon their
opposed surfaces.
Bunomeryx elegans, sp. nov.
A second species of this genus is indicated in the collection by
a portion of a cranium containing the last three premolars and
the molars, in excellent preservation, together with both mandibu-
lar rami bearing all of the teeth with
the exception of the incisors and
canines.
The most important difference be-
tween the two species is seen in the
presence of a short diastema between
premolars two and three of the lower
Fig. 2. Crown view of upper
and lower teeth of PBunomeryx jaw in B. elegans. In f53, Mmontanus
elegans. hy., hypocone; fr., pro- . : :
tacone - $2; protoconale, this diastema is absent, and the teeth
of the lower jaw were apparently in
a continuous series or closed row. &. edegans is smaller and more
delicate than B. montanus, a fact that is not only indicated by all
the teeth but particularly emphasized by the fourth inferior pre-
1898. | Wortman, Extinct Camelide of North America, Ol
molar, which is, considerably narrower and has a less development
of the internal cusp. Another important distinction between the
two species is found in the greater development of the vestigial
hypocone of the second superior molar of B. elegans. In
4. montanus this cusp has almost entirely disappeared, the only
evidence of its presence being indicated by a cingulum in this
portion of the crown.
One fact of great interest in connection with this genus is the
probable light which it throws upon the homologies of certain
cusps of the molar crown in the higher selenodont Artiodactyla.
It is here that we witness the actual passage from the bunodont
to the selenodont type of molar in this important group. If we
can trust. the evidence before us, Bunomeryx is a direct lineal
descendant of the Bridger Homacodon, and it is a matter of the
utmost moment to note that in the latter genus there are six fully-
developed cusps upon the crowns of the first and second superior
true molars ; in the third there are only five cusps present. In
Bunomeryx, as already indicated, the full six cusps are found on
the first superior molar only, while in the second molar there is
‘but a vestige of the postero-internal cusp or hypocone. ‘The
evidence appears to be conclusive, therefore, that the true homo-
logical hypocone is in process of retrogressive disappearance, and
in proportion as this cusp is reduced, the posterior intermediate
is pushed out to take its place. As a further evidence of the
truth of this proposition it may be stated that the true hypocone
of both the first and second molars of Homacodon, as well as the
first molar of Sunomeryx, exhibits no tendency whatever to
develop a selenodont structure, while the posterior intermediates
especially in Bunomeryx, exhibit very decided advances in this
direction. The very position of this cusp, moreover, precludes
‘any possibility of its entering into the formation of the single
posterior internal crescent of the more perfectly developed seleno-
dont molar of the higher types.
I believe therefore that the history of the formation of the four
crescents of the superior molar crowns of the Selenodonts has
been as follows, tracing it from the five-cusped Pantolestes’ of the
1 From this genus I exclude the type of Paztolestes etsagicus Cope as belonging to a distinct
genus ancestral to and leading directly up to the bunodont Artiodactyla. It is very probably
synonymous with Zohyus distans of Marsh, who properly placed it among the Bunodonts.
102 Bulletin American Museum of Natural History. |Vol. X,
Wasatch. The primitive condition of this oldest type of the
Artiodactyla was two buniform external cusps, two buniform
intermediates, together with one large more or less lunate internal
cusp, flanked by a rudimental postero-internal cusp which is
clearly shown to be an outgrowth from the cingulum. There is
no evidence that this postero-internal cusp was ever developed on
the last molar, because in all the forms from Pavtolestes to Buno-
meryx it is persistently absent. The next step consisted in the reduc-
tion of the large lunate internal cusp and the full development
of a well-marked postero-internal cusp, or hypocone, on the first
and second molars. ‘This condition is seen in, and is character-
istic of, Homacodon. The third step consisted in the disappear-
ance of the true hypocone and the gradual usurpation of its place
and function by the posterior intermediate in the crown of the
second true molar, a condition seen in Aunomeryx. As a fourth
step in this development one can readily imagine this process
extended to the first true molar, when it would be complete.
This hypothesis may be objected to on the ground that Awno-
meryx cannot stand as the direct ancestor of any of the Seleno-
donts at present known, on account of its reduced premolar
dentition in the lower jaw, but if we are to regard the type of
superior molar exhibited by either Homacodon, Dichobune or Helo-
hyus as the one which preceded, and from which was derived the
tetraselenodont or four-crescented crown, then this hypothesis
must be accepted as true.
The only case so far known wherein the true hypocone has
been preserved and has become crescentic, is in Cenothertum and
Plestomeryx,and here we have three well-developed crescents upon
the posterior moiety of the crown, of which the inner one repre-
sents the hypocone and the middle one the posterior interme-
diate. It is possible that the cusps of the two anterior superior
molars of X7phodon are to be interpreted in the same way, and
that the posterior innercrescent is composed solely of the posterior
intermediate, the true hypocone having come to occupy a more
anterior and median position. In this case the anterior internal
crescent would be made up_of protocone and the anterior inter-
mediate. Future discovery will no doubt reveal considerable
variety in the formation of the internal crescents in the various
1898. | Wortman, Extinct Camelide of North America. 103
phyla of the Selenodonts, but it appears to me certain that the
hypothesis herein advanced is the correct one for the formation
of the tetraselenodont superior molars of the Cameloids, Pecora,
Tragulines, and probably the Oreodonts and Anthracotheres.
Parameryx (Leptotragulus) proavus 5S. & O.
This genus was first described by Marsh’ and later by Scott
and Osborn,’ who considered that it to belongs to the Traguline
division of the order. Later Scott gave a fuller account of it’
and placed it in the Tylopoda’ immediately ancestral to Poébro-
thertum of the White River Oligocene. The materials in the
Museum Collection do not add very materially to the knowledge
of this form ; however, there are some important points to be
made out from it. There are four specimens which I refer to this
species, the most important of which are a fragmentary skeleton
containing a fairly good hind foot, together with the posterior
part of the last lower molar (No. 2509). The other specimens
pertain exclusively to the lower jaw (Nos. 1803, 1805 and 1808).
In the lower jaw there is one diagnostic character by means of
which the last lower molar can be recognized, and that is the
presence of an extra cusp upon the inner border of the heel near
its point of junction with the postero-internal cusp. It is by
means of this character alone that I associate the fragmentary
skeleton with this species. The lower molars are of the typical
selenodont pattern, and the cusps more elongated than in any of
the cotemporary Selenodonts.
The inferior premolars are three in number, the fourth being pro-
vided with a well-developed internal cusp and heel. The second
and third are simple and without accessory tubercles. In advance
of the second premolar there is a considerable diastema, in front
of which is the large procumbent alveolus for the canine. The
incisors are not preserved, and this region of the jaw is so much
broken as not to reveal their alveoli.
1 “Introduction and Succession of Vertebrate Life in America,’ 1877. _ ; ‘
2‘ Preliminary Report on the Vertebrate Fossils of the Uinta Formation,’ Proc. Am. Philos.
Soc., 1877, pp. 255, 264. , i a : i
3 ‘Mammalia of the Uinta Formation,’ Trans. Am. Philos. Soc., N.S., Vol. XVI, Part 1ii,
Aug. 20, 1896, pp. 479-486. f
4 Marsh had, however, clearly recognized the affinities of this genus with the Tylopoda ten
years previously, since we find in the address above quoted the following statement : * A most
interesting line, that leading to the Camels and Llamas, separates from the primitive seleno-
dont branch in the Eocene, probably through the genus Parameryx.
104 Bulletin American Museum of Natural History. [Vol. X,
Of the bones of the hind foot, the entire tarsus is preserved
with the exception of the cuneiform. ‘These parts of the skele-
ton present a most striking resemblance to those of Poébrothe-
rium in all the details of their structure, the only difference
discoverable being that of size. The third metatarsal is present
but unfortunately a small part of the shaft is missing so as not
to exhibit its full length ; there is enough, however, to indicate
that it was unusually long and slender, much flattened upon the sur-
face which it offered to the second metatarsal, and that the form of
the shaft, moreover, had that peculiar squarish outline upon cross
section, a feature so highly characteristic of the Oligocene
Cameloids. Another distinctive cameloid feature is seen in the
increased size of the medullary cavity. The lateral or fifth meta-
podial was reduced to a mere splint, as is indicated by the much-
reduced facet upon the cuboid ; this facet is relatively as small
as it is in the cuboid of Poébrotherium. The phalanges have
about the same proportions and shape as the corresponding
bones of the White River species.
That Parameryx (Leptotragulus) was a member of the Tylo-
poda, as has already been pointed out by Marsh and Scott, there
can be very little doubt, but at the same time the evidence is
equally conclusive that it does not stand in direct ancestral line
with the succeeding Poébrotheres. The evidence against such a
conclusion is to be found in the fact that Parameryx (Leplotraga-
/us) has only three premolars in the lower jaw, an enlarged canini-
form canine and relatively short, thick inferior premolars, the
last of which, or fourth, has a considerable development of the
internal cusp. It may therefore be looked upon as a precociously
specialized side branch which died out at the close of the Eocene
and left no modified descendants.
Protylopus petersoni,’ ven. et spec. nov.
This genus and species is primarily founded upon the anterior
portion of a skull from which the left ramus is missing. The
specimen is broken obliquely in such a manner as to show upon
the right side all of the facial portion, including the orbit and the
1 This species is named in honor of Mr. O. A. Peterson, whose explorations of the Uinta
Beds have been attended with such marked success.
u
:
i ce ct aly
1808. | Wortman, Extinct Camelide of North America. 105
anterior root of the zygomatic arch, while upon the left side the
greater part of the orbit is missing. Fortunately the skull
contains the dentition nearly complete. In association with it
were found the greater part of an ulna and radius of the same
individual, A second specimen which I refer to this genus and
species includes a large part of both hind legs, together with a
large number of vertebrz, ribs and other parts of the skeleton. A
Ty ray
4 Mii -
‘
Fig. 3. Side view of skull of Protylopus petersoni.
third specimen contains hind limbs and vertebrae, while a fourth
includes the greater part of a hind foot.
The more important generic characters may be stated as
follows :
Molars tetraselenodont without intermediate cusps. Teeth of the typical
number, forty-four, arranged in a continuous series. Canines of both upper
and lower jaws small and incisiform, the first inferior premolar not caniniform.
The first three superior premolars elongated from before backwards, secant and
without accessory cusps, the fourth with single external and internal crescentic
cusps. The inferior premolars elongated and cutting, the fourth without
internal cusps. Hind feet provided with but two functional digits, the outer
ones, second and fifth, reduced to mere vestiges. | Lumbar vertebral formula 7.
Ulna and radius, at least in old individuals, coéssified in the middle part of
their shafts but free at their proximal ends.
The skull is crushed laterally so as not to reveal the exact form
of the face, but it can be safely stated that the muzzle had
106 Bulletin American Museum of Natural History. |Vol. X,
moderate length, with slightly overhanging nasals, much as in
Poébrotherium wilsont. The premaxillz are relatively broad and
extend upwards and backwards to articulate with the nasals.
The orbit is not enclosed by bone posteriorly, but exhibits a
marked tendency towards that peculiar roofing so highly charac-
teristic of Pocbrotherium and the later Tylopoda. In advance of
this bony shelf is seen a faint though distinct indication of the
supraorbital notch, so constant a feature of the cameloid skull.
The lower jaws may be described as long and slender, with a
considerably elongated symphysis.
The superior incisors are relatively small, of a more or less
conical form, and directed downwards. ‘The premaxillae were
apparently not in contact in the median line. The superior
canine is but little larger than the outer incisor, of a more or less
hook-shaped appearance, and provided with a distinct sulcus upon
the outer portion of the crown as in Poébrotherium wilsont, The
first premolar follows after a very short interval and, like the
second, is a simple two-rooted cutting tooth. The third premo-
lar has a faint internal cingular ledge, while the fourth, as already
mentioned, is provided with single external and internal crescents.
The molars are much worn, and do not show clearly whether or
not intermediate tubercles were present, but I think it may be
safely assumed, from the general appearance of the crown, that
they were absent. In thesecond and third molars, between the
internal crescents, is to be seen a small styliform cingular cusp
which is entirely absent, so far as I can determine, in Poébro-
thertum.
In the lower jaw the incisors are of a more spatulate form and
more procumbent in position. As in the upper jaw, the canine is
slightly larger than the
outer incisor, but of a
very marked incisiform
pattern. After a very short
interval or diastema, is
placed the first premolar,
a two-rooted tooth whose crown closely resembles that of the
canine, the two teeth being about equal in size. The second and
Fig. 4. Crown view of lower teeth of Protylopus
petersont.
third premolars have elongated secant crowns like the Tragulines.
1898. | Wortman, Extinct Camelide of North America. 107
The fourth has a well-marked heel and anterior basal cusp,
but there is apparently no internal cusp present. The molars
are so much worn that their structure is not
very apparent. There can be very little
doubt however that they had the usual
structure. In the heel of the last molar a
prominent accessory cusp is seen upon the
border of the inner side near the point
where it joins the lower posterior inter-
nal cusp. In Poébrotherium this cusp is
clearly present, but it has fused with the
postero-internal, producing a prominent
angle at this portion of the crown. In per-
fectly unworn teeth of Poébrothertum wit-
soni, it can be readily demonstrated to be
an independent cuspule.
Of the vertebra, unfortunately, no cervi-
cals are known, consequently it is impossi-
ble to say whether they exhibit the peculiar
features of the more typical Camelidz or
not. The dorsals are well represented in
specimen No. 2564, the whole series being
present, with the exception of the first
three or four, together with all the lumbars
locked in position. The vertebra resemble
those of the modern Llamas closely in their
general proportions. The bodies of the
anterior dorsals are but moderately keeled,
and towards the posterior end of the series
strongly keeled; they increase gradually
in size from before backward. ‘The neural
spine of the fifth is long and recurved,
those of the succeeding dorsals decreasing
in length posteriorly. ‘The neural spines of
the last two are considerably shorter and
broader, having an almost vertical direc-
tion. The rib facets in the anterior region
have their usual relations and positions, the
ribs articulating with the vertebra by two
Vertebre of Protylopus petersoni (?)
108 Bulletin American Museum of Natural History. (Vol. X,
distinct facets, but in the last two the capitular and tubercular
facets appear to be fused together as in these dorsals of the Llama.
The lumbars are seven in number, the constant formula for the
Tylopoda; they resemble closely the corresponding bones of
Poébrotherium and the later Cameloids. The sacrum is composed
of only four vertebrze, but it is highly probable that another one
or two was added from the caudal region as age advanced, just as
in Poébrotherium and the modern Llamas. The three anterior
vertebrae of the sacrum have very reduced neural spines, while in
the fourth the spine is well developed. ‘The ribs do not display
any characters of especial importance.
The pelvis is in a very fragmentary condition, but it may be
stated that the ilium is well expanded, and, so far as one can
judge, the whole bone would correspond closely with that of
Poébrotherium. Yhe femur is present in its entire length with
both ends in a good state of preservation, although the shaft is
somewhat crushed. The proximal end has practically the same
relations and arrangement of the different parts as that of Poébro-
therium and other members of the group. ‘The distal end thus
early gives slight though conclusive evidence of the peculiar and
characteristic appearances which this part of the bone assumes in
the later Camelidze. This is especially seen in the great extension
of the condyles backwards behind the median line of the shaft
as well as the forward projection of the borders of the rotular
groove, which serve to increase the antero-posterior diameter of
this part of the bone. Although not clearly indicated on account
of crushing, yet there seem to be distinct traces of the beginning
of that peculiar depression at the proximal end of the rotular
groove so highly characteristic of the later Tylopoda. In a like
manner the patella has begun to assume the distinctively cameloid
form by the great elongation of its lower border into a long,
pointed process.
The tibia, which about equals the femur in length, shows a
great resemblance to that of Poébrothertum. ‘The cnemial crest
is unusually well developed, and extends quite one-third of the
way down the shaft. The fibula is much reduced, and although
the specimen does not show whether or not the shaft was com-
plete, the probabilities are that it consisted of a distal portion
—
1898. | Wortman, Extinct Camelide of North America. 109g
only. That part of the shaft which is preserved is very slender
and closely applied to the shaft of the tibia. In the hind foot the
tarsal bones have nearly the same relations as in Poébrotherium.
The tuber of the calcaneum is somewhat shorter proportionately
than in the White River genus, but otherwise both the caleaneum
and astragalus are strikingly alike in the two genera. The cuboid
of Protylopus is slightly narrower in proportion to its height than
the corresponding bone in
Poébrotherium, and the navi-
cular is provided with a
much better developed pos-
terior hook. As in Poébro-
therium, there are two cunei-
forms present, the inner of
which is a vestigial nodule
of bone only.
There are but two func-
tional metapodials, the third
and fourth, the second and
fifth being reduced to mere
vestiges. Upon one side the
vestige of the second meta-
podial is preserved in place,
and it is seen to articulate
. : Fig. 6. Left hind foot, femur, tibia and fibula
by a peculiar ledge-like facet, , of Protolypus petersoni (?). cad., calcaneum ; as.,
a: . _astragalus; cé., cuboid; za., navicular; cu®., ex-
upon the principal cunel-_ ternal cunciform; /., head of femur; 7., greater
3 ; trochanter; ¢7?., lesser trochanter; ¢., tibia; /@.,
form. Upon its posterior fibula. ;
surface is a distinct facet
by which it articulates with the small cuneiform. The remnant
of the fifth is not preserved, but the facet by which it articulates
with the cuboid is very small, and there can be no doubt that it
was as much reduced as the second.
The functional metapodials are relatively much shorter than
in Poébrotherium, and of a considerably more primitive form.
They are well flattened upon their opposed surfaces in the upper
half of the extent of their shafts. Below this the inner surfaces
of the two bones are well rounded. Unlike the metapodials of
Poébrotherium, they lack that characteristic four-sided appear-
TIO. Bulletin American Museum of Natural History. |Vol. X,
ance of the later Camels, but on the contrary, are more or less
triangular upon cross section, especially in the proximal half of
their shafts. The metapodials as well as the long bones show
their cameloid affinities in the unusually large size of the medul-
lary cavities. The phalanges exhibit comparatively few differ-
ences from those of Poébrothertum, the unguals being flattened
upon their opposed surfaces. The fore foot is entirely unknown,
but itis highly probable that it will be found to possess four
complete functional toes.
It may transpire that the association of this skeleton with the
above-described skull is incorrect, and that these bones belong to
separate and distinct species; however, they agree so well in the
matter of proportionate sizes of the different parts, and both are
so distinctly cameloid, that I am persuaded to believe that they
refer to one and the same species. It may be noted here, how-
ever, that in one of the specimens referred to above (No. 2067),
there is evidence that at least one of the lateral metapodials of
the hind foot was complete though very slender, and should prob-
ably be referred to another species on this account. The bones
are, moreover, somewhat more slender and delicate than the one
here described. At all events, whatever form of skull belongs
with these skeletal parts it is nevertheless certain that the skull of
Protylopus, above described, is just such a type as is required to
satisfy all the necessary conditions in order to occupy a position
in direct ancestral relation with Poébrotherium. ‘The true Tylo-
pod phylum is therefore traceable directly to it. Beyond this,
there is at present no satisfactory evidence to establish, with any
degree of certainty, the identity of the true Camel pedigree.
Poebrotherium /ezdy.
With a consideration of this genus we pass from the Eocene
to the Oligocene representatives of the group. It was estab-
lished by Leidy as early as 1847 upon an imperfect skull presen-
ted to the Philadelphia Academy by Mr. Alexander Culbertson
of Chambersburg, Pa., who was at the time engaged in the
western fur trade. It was among the first of the mammalian
fossils from the remarkable Bad Lands of the Cheyenne River
1898. | Wortman, Extinct Camelide of North America. 11
region, whose treasures were destined in later years to play such
an important part in the development of American palzontology.
Leidy at first’ regarded the skull as pertaining to a genus nearly
allied to the Musk Deer, but later pointed out its true position
among the Camelidez.
The generic differences between Poébrothertum and Protylopus
are not great, and indeed it would appear at first sight that they
are insignificant. It is more than probable, however, as stated
Fig. 7. Side view of skull of Poébrothertum wilson?.
above, that Protylopus will be found to have four complete and
functional digits in the fore hmb. So far as our knowledge
extends at the present, the chief distinctions are as follows: In
Poébrothertum the molars are much more selenoid and the crowns
more lengthened than in Protylopus ; the third superior incisor
is larger than the superior canine; the ulna and radius are firmly
coossified, even before the epiphyses of the bones are joined to
the shaft and the shaft of the fibula has completely disappeared.
Poebrotherium wilsoni Ze/dy.
This species, although very abundant in the White River Beds
of the Cheyenne River region, has not been very fully described.
All of the specimens in the Museum collection have been found
in the Lower Oreodon level, and it is doubtful if the vertical
range of the species extends much above this point. It differs
very markedly from its successor, P. Zabéatum, in the practical
1*Ancient Fauna of Nebraska,’ Dec., 1852, p.
2* Extinct Mammalian Fauna of Dakota Piel Nebrasiad? 1860, p. 141.
Ll2 Bulletin American Museum of Natural History. \Vol. X,
absence of diastemata in the lower jaw. The canines of this
series are, moreover, broad and incisiform, being separated from
the first premolars by very short diastemata. In the same man-
ner the second premolars follow after a very short interval. In
Fig. 8. Fore and hind foot of Poébrothertum wilsont.
Fig. 9. Ulna and radius and humerus of Poébrothertum wilsont.
Fig. ro. Femur and tibia of PoébrotherZum wilsonz.
P. labiatum, on the other hand, the lower canines and outer
incisors are almost in contact, the canines are subcaniniform in
shape, and there is a short diastema in front of, and a long dias-
tema behind, the first inferior premolar. The bones of the
limbs and other parts of the skeleton are, as far as can be deter-
mined, very much alike in the two species. As in /. /abiatum,
there is a considerable range in size in the various specimens
referred to this species.
1808. | Wortman, Extinct Camelide of North America. 113
Poebrotherium labiatum Cv/e.
The type of this species consists of the larger part of a skele-
ton of a single individual from the White River Beds of north-
eastern Colorado (No. 6520). Associated with this specimen
are two almost complete lower jaws from the same locality (Nos.
6517, 6518) showing the characteristic diastemata of P. /abia/um,
but considerably smaller. These specimens were erroneously
referred by Cope to P. wz/sont. I have not been able to correlate
with certainty the level from which these specimens were taken, with
that in which similar remains in the Cheyenne River region occur,
but judging from Cope’s unpublished sketch of the section of the
bed, there can be httle doubt that it corresponds closely with the
upper part of the Oreodon horizon. ‘This surmise is strength-
ened by the fact that there is one specimen in the collection
(No. 638), from the extreme upper part of the Oreodon Bed,
which agrees in every way with the type of P. /abiatum, except
that it is a little larger. Another specimen from the Cheyenne
River Bad Lands includes a lower jaw and a good part of the
skeleton. The lower jaw exhibits the characteristic diastemata of
P. labiatum, but is much smaller than the type, and of the same
size as the two jaws mentioned above. Unfortunately the exact
level of this specimen is not known, but it has every appearance
of having come from the upper part of* the Oreodon stratum.
Whether or not these smaller specimens are to be referred to a
species distinct from P. Zabéatum is a matter which requires a
greater amount of material than we at present possess in order to
decide correctly. So far as one can determine at present, the
only distinction between the two is one of size, and this is not
great. I have thought best to regard them as belonging to the
same species until other differences are shown to exist. Taken
as a whole, P. /aééatum, as exemplified by the larger individuals,
was considerably larger than /. qw¢/sonz, and in the possession of
diastemata in the lower jaw, as well as the more caniniform
shape of the lower canines, makes a distinctive approach to the
species from the John Day Beds. In this connection it is proper
to observe that no remains of Camels are known from the Proto-
ceras level of the White River Beds. When such are found they
| April, rS98.\ S
114 Bulletin American Museum of Natural History. [Vols
will probably establish a complete transition between /. dabiatum
and the John Day species.
Gomphotherium Co/e.
It is especially to Cope that we are indebted for the discovery
of Camels in the John Day Beds. The first remains secured by
him from this horizon were referred to Poébrotherium, but later
he established the genus Gomphotherium' for their reception,
ey.
ET)
inp
| Mh
MMM
7 | Wi Ming AY),
(\
Fig. rr. Side view of skull of Gomphotherium sternbergi.
which he distinguished from Poébrotherium by the more simpli-
fied character of the crown, and the one-rooted condition of the
first superior premolar. As this distinction was founded largely
upon error, | am now able to give the more important and true
characters which serve to separate the two genera in a satisfactory
manner.
In Poébrotherium the inferior canine is either in contact with
the outer incisor, or is separated from it by a very short dias-
tema, and the form of the canine is either like that of an incisor
or very imperfectly caniniform. In Gomphotherium, on the other
hand, the inferior canine is either separated from the outer
incisor by a very distinct diastema or the diastema is absent,
and the shape of the canine is strongly pointed and recurved, as
1° The Phylogeny of the Camelida,’ Amer. Nat., 1886, p. 618.
1898. | Wortman, Extinct Camelide of North America. UTS
in many of the later Camelide. In Poébrotherium again, the
orbit is not inclosed by bone posteriorly, whereas in Gomp/ho-
thertum the posterior boundary of the orbit is complete. Another
important distinction is seen in the character of the articular
facets of the third and fourth metapodials of the fore foot. In
Poétbrotheritum these bones give evidence of having been more
widely separated in the living animal, and capable of consider-
able independent movement, the facets being relatively large
and the opposed surfaces comparatively smooth. In Gompho-
thertum these facets are much reduced, the metapodials closely
applied to each other and their contiguous surfaces much rough-
ened, clearly foreshadowing the codssification of these elements
into a cannon bone.
Gomphotherium sternbergi Copc.
The type of this species consists of the greater part of the
skeleton of a single individual in good preservation from the
lower beds of the John Day Valley, Oregon. Other specimens
from the same horizon include more or less perfect foot-bones,
fragments of jaws, and other parts of the skeleton. The form of
the skull presents a striking resemblance to that of the modern
Camels in its general make up. The nasal bones are, however,
proportionately longer, the bony roof of the orbits not so broad,
and the muzzle apparently more laterally constricted in front of
the infraorbital foramen. As compared with Poébrotherium and
the Llama the face is less bent down on the basicranial axis, in
this respect resembling more the skull of the Camel. ‘The verti-
cal depth of the face immediately in front of the orbit is rela-
tively greater than in Poébrotherium, and the opening of the
posterior nares has a more forward position.
A very interesting transition from the relatively low, much-
swollen otic bulle of Poébrotherium, to the high, little-swollen
condition of these parts in the living species, is observable. The
otic bull of all the Camels are highly characteristic ; they con-
sist of an inner, longitudinally-directed swollen part, together
with an outer vertical buttress, which joins the inner part at an
angle, and at the upper limit of which ts placed the external
116 Bulletin American Museum of Natural History. |Vol. X,
auditory meatus. Immediately behind the point of junction of
these two parts is seen the deep recess where the hyoid arch is
articulated to the skull. In Poébrotherium the inner portion of
the bulla is much the larger, and the recess for the tympanohyal
is inconsiderable. In Gomphotherium the two parts are about
equal in size, and the tympanohyal recess much more pro-
nounced. In the living genera, Came/us and Auchenia, the nner
Fig. r2. Humerus, ulna and radiaus of Gomphothertum sternbergi.
part of the bulla is much reduced and the tympanohyal recess
is converted into a deep circular pit surrounded by bone.
In the skeleton of the limbs the lower end of the femur is
peculiar in the unusual size and development of the areas of
attachment of the outer and inner heads of the gastrocnemius.
This same peculiarity is seen in the femora of old individuals of
both Poébrotherium labiatum and Procamelus occidental’s, although
to a somewhat less extent, and is doubtless a result of age. The
head of the humerus shows the first distinctive change leading to
the development of the double bicipital groove, a feature so char-
1808. | Wortman, Extinct Camelide of North America. 1g)
acteristic of the later Camelidz. In ne individual in the collec-
tion is there evidence, even in those of the most advanced age,
of any traces of bony union of the metapodials. In size
G. sternbergt exceeded P. labiatum by at least one-third.
Fig. 13 Fig. 14.
Fig. 13. Femur, tibia and fibula of Gomphothertum sternbergt.
Fig, 14. Hind foot of Gomphotheriun sternbergi.
Gompotherium cameloides, sp. nov.
This species 1s represented in the collection by an almost com-
plete mandibular ramus from the uppermost levels of the John
Day deposits (No. 8179). To this same species I also refer an
upper-dentition (No. 7915), an almost complete fore limb (No.
7912), as well as several other fragments. The chief distinctions
118 Bulletin American Museum of Natural History. |Vol. X,
Fig. 15. Lower jaw of GomphotherZum cameloides.
between this species and the older G. sternbergi are seen in the
increased size and the absence of a diastema between the lower
canine and the outer incisor in G, cameloides. The comparative
measurements display these differences in size at a glance ; they
are as follows:
G. sternbergt. G. cameloides.
MM. MM.
Length of sup. ms. and three posterior pms. .. 60 83
a “inf. ms. and three posterior pms. .. 65 97
eb MRCHtiLe nme OnmCentt toni —me-lenee 110 170
os PeAntehiOnmmetapocdialSa-wec create. 180 228
Fig. 16. Upper teeth of Gomphotherium cameloides.
It will therefore be seen that G. cameloides shows
the same increase in size over G. sternbergi as G.
sternbergt does over Poébrothertum labiatum. Of
the bones of the anterior limb, no differences are
observable between them and the corresponding
Fig 17, Humer- parts of G. sternbergi, except in the matter -of size
us of Gomphothe- -
rium PASSE De already noted above.
1898. | Wortman, Extinct Camelide of North America. 11g
The exact stratigraphical position of this species is several hun-
dred feet above that of G. sternbergz, and there can be no doubt
whatever that G. cameloides is not only the direct lineal descend-
ant of the older species, but is, at the same time, the progeni-
tor of the succeeding Loup Fork species. This conclusion is
somewhat at variance with the view expressed by Scott,’ in
which he says: “ The Camels of the John Day formation do not
present any important modifi-
cations of the dentition; in
some of them the first upper
premolar has but a single fang,
and others are decidedly re-
duced in size; the former Cope fo
has erected into a separate
genus, Gomphotherium. It seems
probable that these forms are
not in the direct line of the
’
cameline descent.’ A careful
examination of Cope’s type of
Gomphotherium sternbergt, the
only species, by the way, with
the exception of the one above
named, which has so far been
described from these beds, re-
weals’ *the tact “that <the- first
superior premolar, instead of
being a single-rooted tooth, is
strongly two-rooted ; and that
the succeeding premolars ex-
hibit a most interesting and in-
structive transitional stage be-
Fig. 19. Fore foot
of Gomphothertum
cameloides.
Fig. 18. Ulna and
radius of Gompho-
therium canteloides.
tween Poébrothertum and Pro-
tolabis, not only as regards their
form but their degree of re-
duction as well. I have been unable, after the most exhaus-
tive study, to find any evidence whatever tending to show that
1‘ The Osteology of Poébrotherium,’ 1801, p. 4o.
120 Bulletin American Museum of Natural History. |Vol. X,
these John Day species are not in the direct line of the tylopodean
ancestry.’
Protolabis Cofc.
The next step in the line of cameline ancestry is furnished by
the genus Pro/olabis, originally established upon the greater por-
tion of both premaxilla and maxille, together with the front of
the lower jaw upon one side. This specimen was found near
Pawnee Buttes, Colorado, and is from the typical Loup Fork
horizon.” Another specimen was found by the writer in the so-
called Loup Fork Beds of the upper John Day Valley of central
Oregon, which was described by Cope as a new species of the
same genus. It consists of the anterior portion of the cranium
bearing the complete upper dentition in good preservation.
As regards the type specimen upon which the genus was origi-
nally founded, it is necessary to remark that the only distinction
between it and Procamelus robustus is to be found in the posses-
sion of alveoli for the first and second incisors above, which
would indicate a complete superior incisor dentition. In Proca-
melus, of the adult stage at least, these incisors are wanting. A
careful examination of the specimen shows that the part of the
premaxilla in front of the third incisor, upon the right side, is
much shorter than that upon the left side, and instead of two
alveoli there are in reality three, whtch would make altogether
four incisors in the premaxilla upon this side. The correspond-
ing part of the premaxilla upon the opposite side is much longer,
and contains the two alveoli for the first and second incisors. In
every other detail the specimen agrees perfectly with Procamelus
robustus. Indeed, Cope has pointed out that these first and sec-
ond incisors of this series, in all the species of Procame/us, were
= ——— me re
1 In this connection it is proper to call attention to the fact that there are important faunal
differences between the upper and lower beds of the John Day deposits. ‘hese distinctions
are so marked as to entitle them to be regarded as sepatate aud distinct divisions of the
North American Miocene 3esides the species of cameloids above described, this upper
division contains two species of MWerycocherus among the Oreodonts. Mesohippus prestans
among the Horses, and Elotherium humerosum of the Elotheres, which have not been found
thus far in the lower division of the beds. There can be little doubt that other important
faunal distinctions will be discovered when careful collections are made with this particular
object in view. For the lower series the name Diceratherium Beds may be retained, but for
the upper series | herewith propose the name Merycochcerus Beds.
2 Proc. Philad. Acad., 1876, p. 145.
i)
foal
1898. | Wortman, Extinct Camelida of North America. I
retained in the jaw for a longer or shorter period, and in some
instances until the animal was almost if not quite adult. «In view
of these facts I am disposed to interpret this specimen as a case
of abnormal retention of the incisors of the rather abundant
species P. robustus. If this conclusion is correct, then the spe-
cies Protolabis heterodontus becomes a synonym of Procamelus
robustus. The second species regarded as belonging to this genus
from the Nebraska Loup Fork, Protolabis prehensilis Cope, was
founded upon the anterior portion of a lower jaw without teeth,
and as there is no indication whatever what the upper dentition
was like, the reference at most is mere guesswork. In fact, I
doubt very much if it even is specifically distinct from Pyroca-
melus robustus.
With the elimination of these forms from the Nebraska Loup
Fork, we have left the single species Protolabis transmontanus,
represented by the Oregon specimen above referred to. In this
specimen we have a genuine case of normal retention of the supe-
rior incisors, a fact which separates it at once from Procamelus,
but the distinctions between it and its predecessor, Gomphothe-
rium, are less clear. Cope assumed that it differed from this
latter genus, in the codssification of the metapodials into a can-
non bone, but of this there are no specimens at present known to
demonstrate the truth or falsity of such an assumption. It is true
that Camel remains have been found in the Deep River Beds of
Smith’s Valley, Montana, which deposits have been generally
looked upon as older than the Loup Fork of both Oregon and
Nebraska. These fossils have been uniformly referred to the
genus Profolabis by Cope and Scott, but it does not appear, from
any materials with which [| am acquainted, whether these remains
have been correctly determined or not. Neither skulls, jaws, nor
even teeth, are known from this horizon which would enable one
to say definitely whether the dentition was that of Profo/abis or
Procamelus. The metapodials of both fore and hind feet are
united into a cannon bone, and the size almost, if not quite,
equals that of Procame/us robusius, a circumstance which would
seem to indicate that if they belong to the genus /ofo/abis they
must represent a different and later species, since /rotoladis
transmontanus is much smaller, scarcely exceeding Gomphothertum
122 Bulletin American Museum of Natural History. |Vol. X,
cameloides in size. ‘The only valid distinction between P. ¢vans-
montanus and G. cameloides is seen in the marked reduction of the
second superior premolar in the former, and as this is in the
direct line of modification leading to the Loup Fork Procametus,
I have thought best to regard it as of generic rank, especially
until the question of the coéssification of the metapodials is defi-
nitely settled.
Protolabis transmontanus Cofe.
This species was somewhat smaller than the modern Llama,
and so far as one can judge from the imperfect remains, exhibits
very similar proportions in its general form. ‘The crowns of the
molars are notably shorter and have a more quadrate outline than
those of either the modern genus or Procamelus. ‘This, however,
may be due in a measure to wear. The superior canine Is smaller
than the third incisor ; the first premolar is two-rooted, and the
second is much reduced in size, as in Procamelus. ‘The first and
second incisors are of goodly size, implanted by strong roots and
with obliquely-directed, more or less spatula-shaped crowns.
The nasals are relatively longer than in tie Llama, and the
muzzle is moderately compressed laterally.
Procamelus Zeii/y.
In the widespread Loup Fork deposits of the plains region
remains of Camels are very numerous ; next to the Horses, they
are perhaps the most abundant fossils to be found in these beds.
A number of genera and species of these Camels have been
described, but, as has too frequently happened in the history of
the science in this country, unfortunately, misplaced zeal in the
finding and describing of new forms has been allowed to run riot,
and much of this work is apparently characterized by an utter
lack of the most ordinary display of judgment and discrimina-
tion. No less than four or five genera have been proposed, of
which not more than two are entitled to recognition. Of these
Procamelus 1s the more abundant, and is represented by three
well-marked and easily-distinguished species. ‘The most distinc-
tive features of the genus are the possession of a full complement
eee eee
ee a
1898. | Wortman, Extinct Camelide of North America. 123
of premolars in both jaws, the loss of the first and second pairs
of incisors in the upper jaw in the adult stage, and the at least
partial coéssification of the metapodials of both fore and hind
feet into cannon bones.
Procamelus robustus Zezdy.
I place as synonyms of the above those proposed by Cope,
Protolabis heterodontus and Protolabis prehensilis. The type
specimen, as figured by Leidy, consists of a lower jaw bearing all
the premolars and molars with the exception of the second,
together with the second and third molars of the upper jaw. The
jaw is broken just in advance of the first premolar so as not to
show the characters of the incisors, canine and chin. In Cope’s
type of P. heterodontus the lower jaw is preserved as far back as
the root of the last premolar, and this is associated with the
almost complete superior dentition. The type of P. prehensilis is
represented by the front of both rami of the lower jaw, associated
with a last lower molar. A careful comparison of these speci-
mens reveals a remarkable agreement in all details, and I do not
think there can be any question of their belonging to one and the
same species. The only difference worthy of note is seen in the
jaw fragment which constitutes the type of P. prehensilis ; in this
specimen the chin ts not so abrupt nor deep, and the lower border
of the symphysis is more nearly in line with the long axis of the
jaw. ‘The size is practically the same in all. I distinguish the
species by the following characters: Size, large ; lower molars
with much greater transverse diameter than in other species ;
symphysis not codssified ; chin abrupt and deep. In one speci-
men only is the upper incisor dentition known, and in this the
first and second incisors are retained. It may transpire that this
‘is not an abnormal case as expressed above, but that it is a further
character of the species. No other parts of the skeleton are
known with certainty.
Procamelus occidentalis Zezdy.
This species is by far the most abundant of the Camels in the
Loup Fork Beds. 1 regard the name proposed by Leidy, Homo-
124 Bulletin American Museum of Natural History. \|Vol. X,
camelus caninus, as synonymous with the above, together with
Procamelus angustidens given by Cope to various remains. It is
somewhat smaller than 7. robustus, and is altogether intermediate
in size between this latter species and P. gracilis. It can be
readily distinguished from P. vobustus by the coéssification of the
two rami of the lower jaw at the mandibular symphysis, and
especially by the narrow lower molars. The first and second
incisors disappear early in life and no traces of their alveoli are
seen in the adult skull. A large part of the skeleton is known,
and this is surprisingly like the modern genera. The two main
metapodials are firmly united into a cannon bone, and the ves-
tigial representatives of the two lateral metapodials are codssified
with them. No ungual phalanges, however, are known, and it is
a matter of uncertainty whether they were relatively high and
compressed like the early Camels, or whether they were broad
and depressed like those of the living came/us. 1t is probable that
when found they will show the intermediate conditions between
these two extremes. Most of the skeleton has been described
and figured by Cope,! and need not be repeated here.
Procamelus gracilis Lez.
The species, described by Cope under the name of P. jiss¢dens,
agrees in every particular, so far as the measurements are con-
cerned, with the above, and I therefore regard them as synony-
mous. P. gracilis is the smallest species of the genus, and its
remains are not uncommon in the Loup Fork Beds of Colorado
and Nebraska. It about equals the modern Llama in size, and is
considerably smaller than P. oce¢dentalis. Of the materials in the
collection referable to this species, there are three lower jaws.
One of these has the posterior part of the symphysis preserved,
and this agrees with P. rodustus and differs from P. occidentalis
in the lack of coéssification of the rami, although the well-worn
teeth indicate an old animal. The lower molars are of the nar-
row type, in this respect agreeing with /. occidentalis rather than
P. robustus. In two of the specimens the second premolar is
well developed and two-rooted, but in the third specimen this
tooth is much reduced and single-rooted, thus indicating a ten-
1 Surv. W. tooth M., p. 320.
1808. | Wortman, Extinct Camelide of North America. 125
Fig 20. Top and under view of skull of Procamelus gracilis, after Cope.
dency to disappear. This, it may be said, constitutes a decided
advance in the direction of the next genus, P/auchenta, in which
this tooth is permanently absent. I give herewith some compara-
tive measurements of the three known species in order that the
differences in size may be more readily understood :
126 Bulletin American Museum of Natural History. [|Vol. X,
P. robustus. P. occidentalis. P. gracilis.
MM. MM. MM.
Length of last three pms. and ms..... 152 126 etn
133
126
135
Length of last three pms. and / 106 go Gi
FITSCADWOMISH ee ene ie ee \ 85 76
88 76
92
Two incomplete metapodials in the collection, which are of
suitable size for those of this species, show lack of bony union,
notwithstanding the fact that the epiphyses are completely joined
to the shaft. It is not certain, but there is evidence, that these
metapodials belong to the fore foot, in which event it will not be
surprising to find in more complete and perfect specimens that
these bones are not codssified in this species, except in old indi-
viduals. In the hind foot the metapodials are completely united,
as is shown by several specimens. In one specimen, which corre-
sponds in size with P. occidentalis, the proximal part of the
cannon bone shows lack of bony union for a distance of nearly
two inches down the shaft.
Pliauchenia Co/fe.
The only character so far known in which this genus differs
from Procamelus is found in the absence of the second lower
premolar, and as this constitutes an important step in the dental
evolution of the Camels, the species displaying it are entitled to
rank as a separate and distinct genus. The exact stratigraphic
position of the group is not known, owing to the imperfect collect-
ing that has been done in the Loup Fork Beds. It is generally
stated that P//auchenia was contemporaneous with Procamelus m
the Loup Fork, and comes from the same horizon, but of this one
cannot be certain. It must be remembered that this deposit is
several hundred feet in thickness in places, and it would not be a
matter for surprise if it is found, upon more careful investigation,
that Procamelus comes from the lower levels and P/liauchenta
from the upper strata of the same bed.
Three species have been described by Cope, of which two are
from the Loup Fork of New Mexico and one from the later
1808. | Wortman, Extinct Camelide of North America. 127
Blanco Beds of Texas. One of these species, P. vulcanorum, was
founded upon a superior maxillary bone with most of the teeth in
place, but as the diagnosis of the genus rests exclusively upon the
number of premolars in the lower jaw, I do not see how this
species can be admitted until the lower dentition is determined.
It is therefore discarded.
Pliauchenia humphresiana Co/c.
The type of this species consists of the anterior portion of two
lower jaws from the Loup Fork of New Mexico. <A second
specimen which I refer to this species is from the Loup Fork Beds
of Long Island, Philips County, Kansas, collected by the Museum
party during their explorations there. The specimen consists of
the greater part of a lower jaw broken just in advance of the first
premolar and bearing all the molars and premolars with the
exception of the first and third. In size the specimen about
equals the smaller individuals of Pvocamelus occidentalis, which it
otherwise closely resembles in the structure of the teeth. The
molars are of the narrow and elongated type, and the premolars
have the same structure as that seen in the various species of
Procamelus.
Pliauchenia minima, sp. nov.
The smallest species of Cameloid yet known from the Loup
Fork deposits is represented in the collection by the greater part
of a lower jaw, from Decatur County, Kansas, which I provision-
ally refer to the above genus. The specimen is broken posterior
to the symphysis and does not display the incisors, canine nor
first premolar, if indeed this latter tooth were present. The
diastema in front of the third premolar is unusually long and the
ramus in this region remarkably slender; more so, in fact, than in
any known species of the Camelidz. The third and fourth
premolars are present but considerably reduced in size. The
molars are of the usual pattern. Some fragmentary remains of a
hind foot from this same region are probably to be referred to
this species. The phalanges and distal ends of the metapodials
are extremely cameloid, but unusually slender and delicate. ‘The
128 Bulletin American Museum of Natural History. |Vol. X,
metapodials are firmly codssified. The size of the species is
about equal to that of Poébrotherium labiatum of the White River.
The reference of this species to Pléauchenia is of ‘course only
provisional, and until the entire dentition is known the reference
is uncertain. If it should transpire that the first premolar is
absent it could not be placed in P/iauchenia, but would occupy a
position between this genus and Awchenia.
Pliauchenia spatula Cope.
‘This species is founded upon an unusually complete lower jaw
from the Blanco beds of Texas. According to Cope it is one of
the largest Camels yet found, and considerably exceeds the
modern Dromedary in size. A comparison of the measurements
of the three known species will exhibit the striking difference in
size ; they are as follows:
P. spatula. P. humphresiana. FP. minima.
MM. MM. MM.
Length of last two pms. and ms.... 185 110 70
Camelops Ze/ay.'
Camel remains in the Pleistocene, Equus Beds, are very
numerous; they have been found in nearly every State and
Territory west cf the Mississippi River, and are, in places,
exceedingly abundant. Unfortunately, however, in most
instances the fossils are so fragmentary that it is impossible to
determine the genus or species to which any given specimen is to be
referred. This fact has been taken advantage of, it appears to
me, to inordinately multiply the genera and species, upon no
better ground, frequently, than mere guesswork. ‘These remains
have been grouped at different times in no less than five distinct
genera and ten species. ‘The first of these genera, Came/ops, was
proposed by Leidy (Proc. Acad. Nat. Sci. Phila., 1854, p. 172),
upon the fragment of an upper jaw of a large Camel from the
gravel drifts of Kansas. The second genus, dZegalomeryx, was
also proposed by Leidy (Proc. Acad. Nat. Sci. Phila., 1858, p.
24), for the reception ot some large teeth supposed to belong to
an extinct Camel, from the Pleistocene of Nebraska. In 1872
1 Proc, Acad. Nat. Sci. Phila., 1854. p. 172.
1808. | Wortman, Extinct Camelide of North America. 129
Dr. Leidy again described some large Camel remains from Cali-
fornia, which he referred to the living genus Awchenia.' Follow-
ing this Cope’ described two new genera, Holomeniscus and
Eschatius, which he based upon fragmentary materials from
Oregon, Mexico and elsewhere. He distinguished these genera
from Pliauchenia, Auchenia and Camelus by the possession of a
single superior premolar, the fourth, and separated Lschatius from
Hlolomeniscus by the extreme reduction of this tooth to a simple
cone. The evidence upon which a knowledge of the superior
premolar dentition of these forms rests is furnished, so far at least
as I have been able to learn, by (1) a fragment of an upper jaw
of H. vitikerianus, containing the first and second molars, together
with the roots and alveoli of the premolar or premolars immedi-
ately in advance, as well as a portion of the free border of the
jaw ; (2) a much damaged fragment of a superior maxilla of //.
hesternus, in which no knowledge of the premolar dentition is
possible, since neither the teeth nor their alveoli are preserved ;
and (3) a portion of a superior maxilla of Eschatius conidens with-
out teeth but having nearly all the alveoli preserved.
It appears from a careful examination of this material that the
number of superior premolars in all these Pleistocene cameloids,
with the exception of Aschatius conidens, is uncertain. In the only
specimen in which it can be possibly made out, there are
undoubted traces of an alveolus for a third premolar. If there
were two premolars above, then the dental formula is the same
as in the living genus Awchenia, andis indistinguishable from it,
so far, at least, as the number of teeth is concerned. The
third superior premolar is very small in Awchenia, and it will
not be surprising to find, when a larger number of suitable
specimens of these North American Pleistocene species are
known, that in some cases a vestige of this tooth remained in
the jaw for a longer or shorter time during the life of the animal ;
at least this seems to be the rule in many cases wherein a tooth
is about to disappear from the series.
I therefore reject the definition given by Cope, but retain the
genus as distinct from Auchenia, upon an entirely different
1 Report U. S. Geolog. Surv. Territories, 1873, p. 225.
2 Proc. Amer. Philos. Soc., 1884, p. 16.
[April, 7898. | g
130) Bulletin American Museum of Natural History. [Vol. X,
ground. In the lower molars of Auchenia there is a very promi-
nent buttress, amounting almost to a lamina, developed at the
outer extremity of the antero-external crescent, especially in the
second and third molars. This structure is not present in the lower
molars of any of the North American species which I have seen,
although there are faint traces of it to be found in some of them.
As regards the generic name to be applied to these North
American species, I revive the one originally given by Leidy,
Camelops, notwithstanding the fact that no attempt was made by
him to give a generic definition. The extreme reduction of the
fourth premolar, together with the small size of the inferior
canine, satisfactorily distinguishes schatcus, which is undoubtedly
a distinct genus and is here regarded as such.
In the description of the numerous remaining species which
have been proposed, apparently very little latitude has been
allowed for individual and sexual variations, and because of the
very fragmentary condition of the material so far known any
accurate understanding of their limits is quite impossible. On
this account it is very difficult to determine what species are valid
and entitled to recognition. Until a larger number of specimens
in a less fragmentary state are obtained, I cannot distinguish more
than two, or at the utmost three species, and these determinations
rest solely upon size.
Camelops kansanus Zeid.
This species is by far the most abundant of all the North
American Pleistocene Cameloids, and of the widest geographical
distribution. I regard as synonymous of the above the following
specific names: JZegalomeryx niobrarensis Leidy,'| Auchenia hes-
terna Leidy, Holomeniscus hesternus’ Cope, Auchenia huerfanensis
Craigin, and Holomeniscus sulcatus Cope. Leidy’s type of
Camelops kansanus consists of a fragment of the anterior portion
only of the upper jaw, bearing the root of the incisor and a
portion of the alveolus of the canine, and does not reveal the
1 Proc. Acad. Nat. Sci. Phila., 1858, p. 24.
2 Rep. U.S. Geolog. Surv. Territories, 1873, p. 225.
3 Proc. Amer. Philos. Soc., 1884, p. 16.
4 Amer, Geologist, 1892, p. 257.
® Rep. Geol. Surv, Texas, 1893, p. 84.
1808. | Wortman, Extinct Camelide of North America. 131
number of premolars nor the character of the lower molars. It
agrees so well in size and character, however, with the corre-
sponding parts of other specimens from different localities in
which the dentition can be accurately determined, that there can
be very little doubt that they belong to one and the same species.
The type of Megalomeryx niobrarensts consists of lower molars
from the sand hill region of Nebraska; and since they do not
present any characters which will serve to distinguish them from
the corresponding teeth of C. kansanus they may be safely
regarded as belonging to the same species. In the same way the
teeth described by Leidy from California under the name of
Auchenia hesterna present no differences, either in size or
structure, that would lead one to consider them as belonging to a
distinct species. <Auchenia huerfanensis of Craigin comes appar-
ently in the same category and does not differ, so far as can be
made out from the description, from C. kansanus. Neither can
one detect any characters in the type of /olomentscus sulcatus of
Cope from the Pleistocene of Texas that separate it from the
above species. Cope described a sulcus upon the inner face of
the lower molars, which he thought distinguishes this species.
Upon careful comparison [ find this sulcus is quite as well devel-
oped in many other specimens, the reference of which to C.
kansanus there can be no reasonable doubt. I find it to bea
character which not only varies with the state of wear of the
tooth but is subject to a considerable degree of individual
variation,
The dentition of the species thus considered is as follows :
Ii, C1, Pm+?, M3. Im size the majority of the specimens about
equal the corresponding parts of the Dromedary, although some
are notably more robust, while others are decidedly more slender.
In fact, there appears to be a very great range, as far as size is
concerned, in certain bones, especially those of the feet. The
length does not seem to vary so much as the degree of robustness,
and this probably is due to differences in sex and age. I give here-
with the measurements of some of the principal limb bones of this
species from the Pleistocene sand beds near Hay Springs,
Nebraska, collected by the Museum expedition during last
summer. These bones, it is proper to remark, pertain to many
132 Bulletin American Museum of Natural History. |Vol. X,
individuals, and were found hopelessly mixed together. I also
give measurements of the corresponding bones of the Dromedary
for comparison :
C. kansanus. Dromedary.
MM. MM.
Length of posterior cannon bone............ 345 325
360
365
Length of anterior cannon bone...........-. 330 325
370
Wenothyoteulmaram Gara clits rteenneti-wetie settle 555 580
ILfesavendel Cin ibhiNSAS 4 & 5 smo saaaD oe ne sey ooe 375 420
LEM OF HAMA cos saxsce veaconaobceac 415 460
Length of phalanges, proximal row.......... 98 102
to
124
A specimen from an anterior cannon bone from the Silver Lake
locality in Oregon is considerably more robust than either the
Dromedary or those of C. kansanus from Hay Springs ; it meas-
ures 385 mm. There is yet another specimen in the collection
consisting of a complete posterior cannon bone from the Pleisto-
cene of Hitchcock County, Nebraska, which greatly exceeds in
length that of any known Camel. It is notably more slender than
the cannon bone of the Dromedary, has less distal spread of the
metapodials, but is nearly double the length; its exact measure-
ment is 555 mm. It is highly probable that this represents a
distinct species, but I refrain from creating another specific name
until we know more of the skeleton.
Camelops vitikerianus Cope.
A fragment of an upper jaw containing the first and second
molars, together with the roots and alveoli of the two superior
premolars, is the only specimen of this species so far known with
certainty. A complete lower jaw was described by Cope from
the Pleistocene of Texas under the name of Ho/omeniscus macro-
cephalus,’ which I strongly suspect belongs to this species. It
agrees, so far as one can judge, in size, but no exact comparison
is possible, since the inferior condition of C. vtikervanus is unknown.
Cope remarks inhis des¢ription: “ I observe here that it is not cer-
tain that the species now described is not an Auchenia, as the supe-
rior dentition is not known.” The lower molars, as figured, are
1 Rep. Geolog. Surv. Texas, 1893, p. 85.
1898. | Wortman, Extinct Camelide of North America. 133
certainly not those of any Awchenta with which I am familiar,
since they entirely lack the antero-external buttress of this genus
and agree with those of Camelops. I cannot see any reason
whatever to regard this jaw as belonging to a species different
from that represented by the upper molars above noticed. The
size of the species is near that of the living dAwchenca, and is hence
much smaller than C. ansanus, the only character to my knowl-
edge by which it can be distinguished.
Camelus americanus, sp. nov.
From the Pleistocene Beds of the Hay Springs locality were
obtained by the Museum expedition last year several specimens of
a Cameloid apparently different from anything hitherto described
from this country. One specimen consists of the greater part of
Fig. 2t. Lower jaw of Camelus americanus, side view.
both rami of the mandibles of a fully adult individual, in excellent
state of preservation, but broken in such a manner as to display
but little of the jaw posterior to the last molar. The second
specimen consists also of a mandibular ramus, but the individual
g, in which the milk molars had not yet been shed.
was young,
The interior dental formula of the adult specimen is as follows:
Iz; Cz, Pms, M3. This it will be noticed is the dentition of the
genus Camelus, from which I cannot distinguish it generically at
present. If this is a true Camelus, as I suspect, it is the first time
that it has been found in the western hemisphere.
The incisors display the usual spatulate cameloid pattern, and
are very procumbent in position. Almost immediately behind
the third incisor follows a rather weak incisiform canine. At the
posterior termination of a comparatively short diastema is placed
134 Bulletin American Museum of Natural History. |Vol. X,
the first premolar, a moderately well developed tooth, with a
compressed, internally grooved, pointed, recurved crown, very
similar to the lower canine of Awchenia. After a relatively long
interval follows the fourth premolar, which with the molars makes
a continuous series. The fourth premolar is relatively more robust
than that of the Llama or Dromedary, and gives evidence by this
fact of a less reduced condition. Its form is the same as in these
two genera. The molars display the same structure as those of
the later Cameloids in general, except the second and especially
the. third show the first indication of the external buttress so
characteristic of Auchenia. The two rami are firmly codéssified at
the symphysis. I distinguish the species from Camelus drome-
darius by the relatively small incisiform canine, as well as by the
less reduced fourth premolar and much smaller size. A compari-
son of the measurements of this species with C. dromedarius and
A. lama are herewith given :
A.lama. C. americanus. C. dromedarius.
MM. MM. MM.
Length from incisive border to end of
lastamolanye ti-.s Soe Saeko 165 205 205
Length of molars and fourth premo-
[Enos eereretote Med grso etal IIA eae 80 g2 132
Length from incisive border to fourth
PLCMOLAL mec i citer eee ec 85 114 233
Eschatius conidens Copc.
The last of the American Cameloids to be considered is this
species which, so far as we are now aware, was the most special-
ized of the Camelide. As already noted, the characters which
distinguish this group from all others is found in the extreme
reduction of the fourth superior premolar to a simple cone, as
originally pointed out by Cope; this is associated with a marked
reduction in size of the lower canine. ‘The specimens from which
1898. | Wortman, Extinct Camelide of North America. 135
the genus and species is known are, unfortunately, very fragmen-
tary, so that it is impossible to say much concerning it. It was
first found at the Silver Lake locality in southeastern Oregon, but
subsequently Cope identified remains referable to it from certain
localities in Mexico, showing that it was most widely distributed.
Two species have been described by Cope, &. contdens and E.
Jongtrostris, but it is very doubtful indeed whether the material at
c. ———*
Fig. 23. Crown view of upper teeth of Auchenia lama.
§- 23
present known warrants the recognition of more than a single
species. There are some trifling differences in the length of the
diastema of the lower jaw, it is true, but this is probably due to
individual variation. The size of the species about equals that of
the Dromedary.
THE EVOLUTION OF THE CAMELID&.
We come next to consider the evoiution of the Camelidz, and
although the principal facts of their development have been quite
fully set forth in the foregoing pages, yet it seems proper to
summarize them here. The earliest forms in which one can
detect true tylopodean peculiarities, as already pointed out, are
found in the upper Eocene or Uinta stage ; at the same time it is
quite certain that the phylum was detached from the main stem of
the Artiodactyla at a much earlier date. Both Cope and Scott
have expressed the opinion that the ancestry of the group is
traceable directly to the Wasatch genus Paztolestes, and Scott
believes that the connection is established by the Bridger genus
Homacodon. Whether or not these opinions will bear the test of
future discovery time alone will reveal. At present I am of the
opinion that the osteology of these genera is too imperfectly
136 Bulletin American Museum of Natural History. |Vol. X,
known to speak with any degree of confidence as to its correct-
ness.
The principal changes of structure in the evolution of the
group relate largely to the limbs and skull, although, as is
well known, the cervical vertebre are highly characteristic in the
whole family. Unfortunately the cervicals are entirely unknown
in the earliest representative, Protylopus, but in the succeeding
genus, Poébrothertum, they had already assumed the _ typical
cameloid peculiarities. If one is permitted to judge by analogy,
it is more than likely that Pyvotylopus possessed the peculiar
cameloid cervicals, and it seems highly probable that one of the
crucial tests of tylopodean affinity, in any genus older than this,
will be found in the incipient changes leading to this modifica-
tion. |
The changes in the limbs consisted mainly in (1) their gradual
elongation, (2) coéssification of certain bones, (3) modification of
the patella and loss of the greater part of the tibia, and (4) the
subtraction of digits, the change in the character of the feet and
modification of the phalanges, together with the codssification of
the podial elements into a cannon bone. In this connection
should also be mentioned the development of a double bicipital
groove upon the humerus and the modification of the carpal and
tarsal elements.
The question of the elongation of the limbs is closely asso-
ciated with the general increase in size, in which there is as
complete a gradation, from the little Pvotvlopus, scarcely larger
than a good-sized Jack Rabbit, to the more modern species,
larger than the Dromedary, as the most hypercritical opponent of
the Evolution Theory could possibly demand. In the matter of
the codssification of certain bones, the ulna and radius were the
first to be affected by this process. As we have already seen, it
had taken place in Protylopus in a very old individual, and then
only in the middle of the shaft, leaving the proximal ends entirely
ununited. In Poébrotherium the complete codssification of these
elements was accomplished in young individuals before the milk
dentition had been completely shed, and before the epiphyses had
united to the shafts of the long bones, as is demonstrated by many
specimens in the Museum collection. In all the later types these
bones are firmly united.
1898. | Wortman, Extinct Camelide of North America. 137
The patella of the later Camels is peculiar in that it is narrow
and of great vertical depth. The first evidence of this modifica-
tion is seen in the development of a long pointed process upon
the inferior border of the bone in Protylopus ; this is continued in
Poébrotherium and Gomphotherium, until in Procamelus the
modern condition is reached. The shaft of the fibula in all the
modern types has completely disappeared, and the bone is repre-
sented by a distal nodular element which lies under the end of the
tibia. In Protylopus a considerable part of the shaft of the bone
was present, but it was probably not complete. In Poébrotherium
the shaft is reduced to a very short bony spicule, and the distal
part is partially pushed under the end of the tibia; in Gompho-
therium the shaft has completely disappeared, and in Procamelus
the modern arrangement is attained.
In the matter of the loss of digits I cannot speak with any very
great degree of confidence, especially as regards the older types.
Our materials fail to reveal the number of toes in the fore feet of
Protylopus, but there is very little doubt that there were four. In
one specimen which I have referred to this genus, the lateral toes
of the hind feet are reduced to mere nodular splints, while in
another specimen in the collection there is evidence that the
lateral toes of the hind feet were complete but very slender. In
Poébrotherium the lateral toes are reduced to vestiges in both fore
and hind feet, while in the later forms even these vestiges
disappear.
The modifications in the character of the feet refer to what may
be properly termed a retrograde change, if we regard the subject
from the standpoint of the podial evolution of the Ungulates
generally. Protylopus, as well as Poébrotherium and Gompho-
therium, had apparently a fully developed unguligrade gait. The
ungual phalanges are relatively high, keeled upon their dorsal
surfaces, and more or less flattened upon their opposed sides.
The distal ends of the proximal phalanges do not have their
articular surfaces extended upon the dorsal side, while the distal
phalanges of the second row have this surface well extended upon
‘the dorsal portion of the bone. This arrangement is highly
characteristic of all the higher Artiodactyla, and is the strongest
possible evidence that can be adduced from the skeleton of the
unguligrade gait.
138 Bulletin American Museum of Natural History. |Vol. X,
In Procamelus, on the other hand, the distal ends of the
proximal phalanges, as well as those of the second row, have the
articular surfaces reaching well back upon the dorsal aspect, just
as in the modern Camel and Llama, and this we know to be
accompanied by the digitigrade gait and the peculiar cushioned
foot. There is likewise a marked change in the character of the
ungual phalanges, which are much reduced and flattened vertically
especially in Camelus.
The conclusion is obvious, therefore, that this peculiar tylopod
character is secondary, and was developed from the unguligrade
condition. Just what led to it is impossible to say. Zittell’s
explanation that it resulted as a consequence of the codssification
of the metapodials can hardly be the correct one, since the same
thing occurred in all of the Pecora in which the unguligrade gait
was retained.
The codssification of the metapodials and the consequent
formation of a cannon bone took place comparatively late in the
history of the phylum, much later in fact than the union of the
ulna and radius. In Protylopus and Poébrotherium the metapo-
dials were not only free but capable of considerable independent
movement, as indicated by the facets. "The opposed surfaces of
these bones, moreover, are relatively smooth and less flattened
than in the succeeding Gomphotherium. In this latter genus the
bones are more closely applied to each other, the surfaces rough-
ened, and the articular facets, by which they join one another,
much reduced. ‘The feet of Pvofolabis are wholly unknown. In
one species of Procamelus (P. gracilis), there is evidence that the
metapodials of the fore feet were not united until late in life, if
at all, while those of the hind foot were fully coéssified into a
cannon bone. Procamelus occidentalis had the posterior metapo-
dials fully united and those of the fore foot only partially coéssi-
fied, being free at their proximal ends for some distance.
In the Pleistocene species the metapodials were fully united
into a cannon bone early in life, and in the modern Camelidz bony
union of these elements takes place before birth, almost as soon
in fact as bony tissue is thrown down in the shafts.
It will thus be seen that the codssification of the metapodials,
as well as that of the ulna and radius, was a gradual process, and
1808. | Wortman, Extinct Camelide of North America. 139
is always found first in the old individuals ; its further advance
has consisted in reducing the time of its appearance to such an
extent that it is now altogether intra uterine. I leave for a future
paper the discussion of these important facts in their relation to the
question of the transmission of acquired characters. I may state
here, however, that they furnish very strong presumptive, if not
conclusive, evidence of the transmission of a pathological change.
The modifications of the skull include some minor changes in
the position of the orbit, its inclosure by a complete bony rim, a
broadening of the frontal region and a shortening of the nasal
bones. There have also occurred some modifications of the
tympanic bulle and a change in the position of the posterior
nares. The most important of all the modifications connected
with the skull, however, are found in the dentition, and here the
changes are quite as profound as have occurred in any other
group of mammals within the same length of time. In Protylopus
the structure of the molars is, to a large extent, intermediate
between the bunodont and selenodont pattern. ‘This is especially
seen in the lower molars, the internal cusps of which are rather
more conical than crescentic ; the outer cusps are not perfectly
crescentic, and the crowns of all the molars are very short. The
lower canines are small and incisiform, the inferior incisors are
sub-erect, and there is evidence of the fact that these teeth had a
more or less effective bite against the upper ones.
In Poébrotherium the crowns of the molars are much more
elongated and the crescents fully developed. The lower canine
is yet incisiform in the older species, 7. z//sonz, but has begun to
assume the caniniform shape in the later ?. /abiatum. ‘The lower
incisors have a very procumbent position, but while yet opposing
the superior incisors, the effectiveness of the bite is considerably
diminished on this account. The only important change in the
teeth of the John Day species is seen in the reduction in size of
the first upper premolar and the assumption of the caniniform
shape of the lower canine. The inferior dentition of Profoladis is
not known, but an important change has taken place in the second
superior premolar, in that it is much reduced in size.
If the specimen which i have referred to Procamelus robustus
is normal, it offers the first evidence of the diminution in size,
140 Bulletin American Museum of Natural History. |Vol. X,
leading to the final loss of the first and second pair of superior
incisors. In Procamelus occidentalis these teeth have completely
disappeared in the adult, but vestiges of them have been found in
the young, a fact which was demonstrated by Cope.
Pliauchenia furnishes us with the next step in which the second
premolar has disappeared from the lower jaw, leaving the first,
third and fourth of this series. The superior premolar dentition
of this genus has not yet been found, but itis highly probable that
some of the species will show a loss of the corresponding tooth in
the upper jaw. It is highly probable that at this stage, or the one
preceding, three diverging lines took origin, of which one con-
tinued into Awchenia, another into Came/us, and a third into
Camelops and Eschatius.
From Pliauchenia the transition is easy to Camelus, in which
there are three premolars in the upper and two in the lower jaw.
In Auchenia the premolars are still further reduced, there being
two above and only one below. ‘The first evidence of the peculiar
buttress found in the lower molars of this genus is seen in
Procamelus gractiis, and on this account I am of the opinion that
this species is the ancestor of the Llamas. It is highly probable
that the species migrated to South America at the close of the
Miocene, and that the intermediate links between it and the living
genus will be found in that country. Came/ops could have easily
been derived from Plauchenia spatula, requiring the loss of only
two premolars in the lower jaw and probably one above; in a like
manner /schatius follows Came/ops and carries dental reduction to
the extreme limit reached in the Camel group.
_ In this attempt to indicate the more exact specific evolution of
the group one is necessarily handicapped by want of knowledge
of the osteology of many species which are at present represented
by fragmentary remains only. I doubt not that it will be materi-
ally altered when we come to have a more perfect understanding
of these forms. I give herewith a table of distribution in time, of
the Cameloids treated of in the foregoing pages.
141
Wortman, Extinct Camelide of North America.
1808. |
PERIODS. ForMATIONS, THICKNESS. FAuNAL Divisions. EVoLuTION OF SPECIES.
Recent. Recent. Camelus. Auchenta.
2 Eschatius Bice, \
Pleistocene. Equus. 150 Equus. : Camelus americanus, )
Camelops kansanus. \ Camelops vitikerianus. |
. = - : r a ae as a
: Pliauchenia. ' \ /
Pliocene. Blanco. 100 ; ie ‘ Zz 1
Hippidium. Pliauchenta spatula. / i
a err ese oo
| Pliaucheniahumphresiana. P. minima. |
Loup Fork, 400 Procamelus. Procamelis occidentalis.“ ec
1 u Axe
7 ; : Procamelus robustus. P. gracilis,
Miscenes Deep Lone 200 Cyclopidius. Protolabis.
INTERVAL| OF MIGRATION, '
= oe
Merycocheerus. Gomphotherinm camelotdes.
John Day. 1000 ———_.
Diceratherium. Gomphotherium sternbergi.
pa ee +
Protoceras.
: 7
Oligocene. White River. 800 Oreodon. Poétbrotherium labiatum.
F ; { ; z
Titanotherium. Poebrotherium wilsont.
Diplacodon. Protylopus. Parameryx.
Uinta. 800 : badd =F z
Telmatotherium. Se Sea
Lae
Bridger. 2000 Uintatherium. ? Homacodon.
eo
Eocene, Wind River. 800 Bathyopsis. i
RRR ee z eae
Wasatch. 2000 Coryphodon. ? Pantolestes.
Torrejon. 300 Pantolambda.
Puerco. 5006 Polymastodon.
142 Bulletin American Museum of Natural History. \|Vol. X.|
BXPLANATION OF PLATE XI.
A, hind metapodials of Protylopus petersoni ; B, fore and hind metapodials
of Poébrotherium wilsont ; C, fore and hind metapodials and head of humerus
of Poébrotherium labiatum ; D, same species from higher level ; £, fore and
hind metapodials and head of humerus of Gomphotherium sternbergi ; F, fore
and hind metapodials of Gomphotherium cameloides ; G, fore and hind meta-
podials and head of humerus of Procamelus occidentalis ; H, head of humerus
of Auchenia lama.
Butietin A. M. N. H. WOT OX Am ON:
METAPODIALS OF EXTINCT CAMELOIDS.
— Remounted Skeleton of Phenacodus prime-
| vus. Comparison weth Euprotogonta.
y
By Henry FArrRrIELD Osporn.
~PLATE XII anp Four TExtT FIGURES.
See EDITION, extracted from BULLETIN
Amevican Museum of Natural History,
. (
\
VoL. x, ARTICLE IX; pp. 159-164.
New York, May 6, 7898.
Article IX.—REMOUNTED SKELETON OF PHENACO-
DUS-PRIMAIV US... COMPARISON WITH EUPRO-
TOGONIA.
By Henry FAIRFIELD OSBORN.
PLATE XII AND Four TEXT FIGURES.
This unique skeleton was transferred to the American Museum
with the remainder of the Cope Collection in 1893. It had been
mounted’ as found (Fig. 1), laterally crushed, a large portion of
the vertebrae and ribs concealed so that their number could not
Fig. 1.
tion of Professor Cope.
Skeleton of Phenacodus prima@vus as originally mounted and cast under the direc-
be definitely ascertained, and in such position as to convey a false
impression both of the proportions and mode of location of this
remarkably primitive Ungulate.
After very careful deliberation we decided to remove the skele-
ton entirely from the matrix, and remount it as nearly as possible
in the natural position. This removal cost many months of labor
1 See Cope, ‘ Tertiary Vertebrata,’ Plate LVII k.
[159]
160 Bulletin American Museum of Natural History. |Vol. X,
and two months more were occupied by Mr. Hermann, Prepara-
tor, in mounting the animal as represented in Plate XII. This
mount is now a model of its kind, since it not only displays the
real characters of the animal, but every bone upon one side of
the body or the other can be removed for purposes of detailed
study. Moreover, in course of removakof the stone and plaster
matrix, the two missing cervical vertebrae were found inserted
in the tail, and the number of ribs was definitely ascertained. to
be fifteen on each side, thus positively determining the dorsal
vertebral formula, a matter of very great importance. ‘These
results alone justify the labor involved.
As photographed in Plate XII, Phenacodus primevus strikes us
as a rather slenderly built, straight-limbed animal, digitigrade
like the Tapir, five-toed, but almost exclusively supported on
three toes both upon the fore and hind feet, with the median toe
considerably enlarged, well hoofed, and extended beyond the
others; therefore functionally of the tridactyl type. Flower’s
restoration, in his volume upon the Horse, is very nearly correct.
The upwardly-arched back, powerful lumbar vertebra, the long
hind-quarters (measuring 635 mm.), the long powerful tail, when
contrasted with the much shorter fore-quarters (measuring 460
mim.), the rather low withers and small head, are reminiscent of
Creodont ancestry.
Phenacodus was, in fact, a swift-footed, cursorial, small-brained,
microcephalic type (in distinction from Coryphodon). It was
largely propelled by its powerful hind limbs. ‘The skeleton is,
however, straight limbed at the elbow in contrast with the early
Amblypods, such as Pantolambda and Coryphodon.’ In_ this
respect it approaches that of the Perissodactyla. The terminal
phalanges of the three median toes are broad and spreading,
while the lateral phalanges have rather the narrow compressed
type seen in Luprotogonia.
The most striking features of the skull are the small size and
separation of the basicranial foramina, the simple primitive struc-
ture of the whole region around the ear at the base of the skull
(the auditory meatus being bounded posteriorly by the mastoid),
1 See Osborn, ‘A Complete Skeleton of Coryphodon,’ Bull. Am. Mus, Nat. Hist., Vol. X,
April 5, 1808, pp. 81-o1.
1898.| Osborn, Remounted Skeleton of Phenacodus prime@vus. 161
Fig. 2.
Spines, fifth lumbar vertebra and posterior caudal vertebre restored. One-fourth natural size.
I
CAUDALS.
Vertebral Column of Phenacodus primevus. s. f., spinous foramen; v. ¢ , vertebrarterial canal; c._, capitular facets ; 7, tubercular facets.
£1
[| April, 7898.\
162 Bulletin American Museum of Natural History. [Vol. X,
and the longitudinal grooves upon the superior intranareal sur-
faces of the maxillaries (also observed in Coryphodon). The
proportions and characters of the skull (exclusive of the teeth)
are identical with those in the ancient Amblypoda, such as Perz-
tychus and Pantolambda.
The distinctive feature of the vertebral column is the small
number of dorsal vertebra, namely 15, exactly as in Coryphodon
and 7ttanothertum, giving a dorso-lumbar formula of 20-21. In
Fig. 3. Skeleton of Phenacodus prim@vus placed in the standing position to show elevation
of the withers. One-twelfth natural size.
the writer’s opinion this number is characteristic of the primitive
Condylarthra or Protungulata. Marsh,’ on the other hand, has
assigned to the Holodactyla (Condylarthra) 23 dorso-lumbars,
and to the Protungulata 30 dorso-lumbars or more. The second
distinctive feature is found in the splitting of the transverse pro-
cesses on the posterior dorsals for articulation with the tubercles
of the ribs and for the support of the zygapophyses respectively,
as shown in Fig. 2, these processes being sharply separate.
1 *Dinocerata,’ pp. 171, 172.
1898.| Osborn, Remounted Skeleton of Phenacodus primevus.
163
TABLE OF MEASUREMENTS.
Phenacodus. Euprotogonia.
Feet. In. Metres. Metres.
Length, chin to perpendicular line of tail.... 4 434 1.340
Length, chin to ischiac symphysis 1.220 -590
Length skull, condyles to symphysis of pre-
nen exe ac uta ho PA Oe o eee eek eo -235
Height, dorsal spines at withers........... 550) .270
leicht .dorsalspines/at, hips.).o- . eee .585) .280
Limbs, total length of hind limb, outside
MEASUTEMEN ts Ae Leia ere cues ee tek eine 2 I .635 ) .310
Limbs, total length of fore limb, outside -
TH EASUMEME ME gayopey «cep sisselrersye tu shale evekensy erenelty ats I 6 .460 )
Euprotogonta puercensis thus appears to be about one-half the
size of Phenacodus primenus in all its measurements.
As shown
in Fig. 4, the lateral digits are considerably longer, reaching the
ground.
Fig. 4.
in outline from Phenacodus.
Skeleton of Exprotogonia puercensis as now mounted, with missing parts restored
One-twelfth natural size.
Same scale as P. primevus.
COMPARISON WITH EUPROTOGONIA.
Luprotogonia.
[TorrEjon.]
(1) Caudals 18 preserved, neural arches
complete in C 1-8. D. L. formula
unknown.
_ (2) Fore-limb. Posterior face of ulna
convex. (C.)
(3) Ectepicondylar ridge prominent. (C.)
(4) Magnum very small. (C.)
(5) ? Os centrale. (C.)
(6) Trapezoid very short. (C.)
(7) Terminal phalanges laterally com- |
pressed, intermediate between
hoofs and claws.
(8) Hind-limb. Femur with sharp and
prominent patellar trochlea.
Phenac
[WasatTcu. ]
odus.
Caudal formula indeterminate.
Neural arches complete anteri-
orly., Ws Es — 20-21.
The same, concave.
The same.
Magnum somewhat enlarged.
The same wanting.
The same somewhat enlarged.
Hoofs fully formed in P. prime-
UUs.
More compressed in ?. ( 77tsfous-
dylus) wortmani,
The same.
164 Bulletin American Museum of Natural History. |Vol. X.|
COMPARISON WITH EUPROTOGONIA.—Coxntinued.
Luprotogonta. Phenacodus.
[Torreon]. WASATCH.
(9) Tibial spines sharp and prominent; | The same. Cnemial crest less
cnemial crest very prominent and prominent, and shorter.
elongate. (C.)
(10) No fibulo-calcaneal facet. The same.
(11) Astragalo-cuboidal facet depressed. The same.
(12) An astragalar foramen ; tibial troch- | No astragalar foramen. Tibial
lea in front of foramen, z.e., sub- trochlea extended further back,
digitigrade. (C.) z.e., fully digitigrade.
(13) Meso-cuneiform very short. (C.) The same.
(14) Lateral digital reduction advanced: | Lateral digital reduction still more
ratio of Mts.V: Mts.III: 23mm. advanced : ratio of Mts.V : Mts.
37 mm., Or aS 2: 3. Ris 25 mim. 773 nimi. ior asere2)
(15) Metatarsals and phalanges slightly | The same nearly straight.
curved. (C.)
This strengthens the observation of Matthew,’ in his recent
comparison of these types, that the features in which P/enacodus
differs from Fuprotogonia are progressions from the Creodont
type, as indicated in the above column by the letter C.
A far larger proportion of Creodont characters are, however,
found in the Amblypod contemporary of Luwprofogonia, namely,
Pantolambda, which will be fully described by the writer in a forth-
coming paper.
! Revision of the Puerco Fauna,’ Bull. Am. Mus Nat. Hist., Vol. IX, 1897, pp. 305-308.
BULLETIN A. M. N. H. VoL. X, PLate XII.
MOUNTED SKELETON OF Phenacodus primevus.
One-twelfth natural size.
WSF
| New York, tae a , 1898.
t
TY,
ONE AP i
Article XI.— EVOLUTION OF THE AMBLYPODA.
PART I. TALIGRADA AND PANTODONTA.
By Henry FAIRFIELD OSBORN,
TWENTY-NINE TEXT FIGURES.
The Amblypoda constitute a sharply distinguished order of
Ungulates which probably sprang from the Creodonta during the
Cretaceous period and extended through the Eocene period in
three great stages of evolution known as the suborders Taligrada,
Pantodonta and Dinocerata, the latter entirely confined to North
America.
They are arrested or persistently archaic in structure through-
out, but especially in the brain, the triangular teeth, and the penta-
dactyl feet. The morphological problems involved in the skeleton
and teeth, and the phylogenetic problems involved in the suc-
cession and extinction of the main and collateral lines, form the
main subjects of this essay.
The American Museum has sent out a series of expeditions
after remains of the Amblypoda: First, into the Wasatch of
Wyoming, 1891 ; second, into the Bridger and Washakie Basins,
1893 and 1895; third, into the Torrejon, 1892 and 1896;
fourth, into the Wasatch of New Mexico, and Big Horn Moun-
tains of Wyoming, 1896. We have thus succeeded in bringing
together invaluable material for the history of this remarkable
group from the time it issued in the Creodont-like Pantolambda,
of the Torrejon Beds until it became extinct in the largest
Uintatheres of the Upper Washakie and Middle Uinta Beds.
Several very important results are obtained :
First.—The evidence can be clearly stated as to the succession
of the known types of Pantolambdide, Coryphodontide and
Uintatheride. Many prophetic or ordinal characters are now
observed in the earliest types ; certain species of Coryphodon are
found to show the rudimentary parietal horns and the incisiform
[169]
170) «Bulletin American Museum of Natural History. |Vol. X,
lower canines of the Uintatheres, but no known species leads
directly into Uintatherium.
Second.—It is now demonstrable that the Periptychide belong
in this group as a dunodont division, which probably arose and
diverged in the Cretaceous period.
Third.—The nearly complete skeleton of Pantolambda bath-
modon, the most archaic type of Ungulate known, places almost
upon the firm basis of fact Cope’s hypothesis, that the Ungulates
sprang from the Creodont division of the Unguiculates.
Fourth.—A fairly complete systematic revision of the entire
group is rendered possible.
The writer desires to express at the outset his indebtedness to
Dr. J. L. Wortman for his energetic and intelligent supervision in
the field of these various expeditions, assisted by Mr. Walter
Granger, Mr. Barnum Brown and others; to Dr. W. D. Matthew
for the skill and care with which the very fragmentary skeletons
of Pantolambda have been put together, and for critical assist-
ance in the diagnosis of specific types. Also to the various pre-
parators and draughtsmen of the department.
I.-—ORIGIN OF THE AMBLYPODA.
A. THE CRETACEOUS TRITUBERCULATE MOLAR.
Lypothetical Upper Cretaceous Ancestors.—In describing the
Laramie mammals in 1893 the writer’ directed attention to the
resemblance which certain isolated upper and lower molars bore
to the teeth of the Periptychide, especially to Actoconus and
flaploconus. ‘The figures of these teeth are here reproduced.
In C (Syaconodon), the crowns are laterally compressed, thus
bringing the primary proto-, para— and metacones, both of the
trigon and trigonid, very close together. This compression of
the primary cones is also true of / (Zctoconodon), in which type
the outer wall is reinforced as in Zctoconus by accessory tubercles.
1 Osborn, ‘ Upper Cretaceous Mammals,’ Bull. Am. Mus. Nat. Hist., Vol. V, pp. 325-329.
1898. | Osborn, Evolution of the Amblypoda. Part I. 1
This compression is an essential character of the Periptychide.
The upper and lower molars of Protolambda, H/, A and Ad, on
the other hand, resemble those of the Pantolambdide.
Fig. 1.
LARAMIE TRITUBERCULATES.
C. Synconodon sexicuspis. Upper and lower molar types. Nos. 2218, 2220. Am. Mus. Coll.
F. Ectoconodon petersonz. Upper molar types. Nos. 2224, 2223. Am. Mus. Coll.
A. Protolambda hatcheri. Upper molar types. Nos. 2201-3. Am. Mus. Coll.
Lower molars AZ, AZ (possibly correlated), Nos. 2241, 2230, 2231. Am. Mus. Coll.
These types may now be defined as follows :
Synconodon, gen. nov.
7ype.—Isolated upper and lower molars.
2220.
Am. Mus. Coll., Nos. 2218,
Definition.— Molars long and narrow. ‘Trigons laterally compressed,
primary cones approximated. ‘Valonid well developed.
S. sexicuspis, sp nov.
Definition.—Talonid of lower molars with three cusps, hypoconid, hypo-
conulid, entoconid. Type as above. Fig. 1, C.
Ectoconodon, gen. nov.
Type.—Isolated superior molars
Loc., Laramie.
2223 "220A
Am. Mus. Coll., Nos.
Definition.—Superior molars very broad.
Trigon laterally compressed.
Two external prominent cones (parastyle and metastyle), reinforcing the outer
wall of the crown.
E. petersoni, sp. nov.
Superior molars with conules and a rudimentary mesostyle.
above. Fig. 1, /.
Type as
172 Bulletin American Museum of Natural History. (|Vol. X,
Protolambda,' gen. nov.
7ype.—Four isolated upper molars. Loc., Laramie. Am. Mus, Coll., Nos.
2201-3.
Definition.—Superior molars with open trigon and elongate outer wall.
Paracone and metacone laterally compressed. Conules subcrescentic. Para-
style prominent.
P. hatcheri,’ sp. nov.
With rudimentary spur, metastyle on posterior external border.
Type, as above. Fig. 1, A.
The lower molars #/ and AJ, in Fig. 1, probably are associated with this
type.
mesostyle
_ Fig. 2. Pantolambda cavirictus. A, superior molars, No. 963; &,diagram of same; C,
inferior molars, crown view, No. 3961; D, lower jaw, type of P. cavzrictus. Am. Mus. Coll,
These types are quite distinct from the _ trituberculates
described by Marsh* from the Laramie, namely, D¢delphops,
Cimolestes, Pedomys, Telacodon, Batodon, so far as these are known.
These animals are defined and placed here hyfothetically
merely to show that zz the Upper Cretaceous are found types of
upper and lower teeth capable of giving origin to the teeth of the
Amblypoda.
1 Having a type of tooth antecedent to that of Paxtolamébda.
2 Dedicated to Mr. J. B. Hatcher, the well-known writer and explorer.
3 Discovery of Cretaceous Mammalia, Pts. I, II, I11, Am. Jour. Sci., July, 1889, to Mch., 1892.
1808. | Osborn, Evolution of the Amblypoda. Part I. ES
EVOLUTION OF THE MOLARS IN THE AMBLYPODA.
The upper and lower molars of the Periptychide are purely
bunodont, characterized by a compressed trigon, the development
of accessory external cusps, and crescentic internal cusps upon
the superior premolars. The latter recall the teeth of Panto-
lambda, Ectoconus is the only type in which the lower premolars
tend to become molariform.
The Pantolambda superior molar (Fig.2 4) shows a crescentic
disposition of the three primary cones, which are, however, closely
approximated as in the Periptychide, the outer wall of the crown
extending widely into the parastyle. The lower molar shows a
lofty trigon distinguished by the marked elevation of the meta-
conid, and a talonid best shown in Fig. 2 Y. No true hypoconulid
is developed. From the entoconid a spur extends forwards and
inwards, to which the designation ‘entoconid 2’ may be given,
as seen in Fig. 3. This little cusp be-
comes a very important feature of the
crown in Bathyopsis and Uintatherium.
The Coryphodon superior molar (Fig. 4),
as homologized by Cope, Earle and the
writer, exhibits a protoloph and an ecto-
loph consisting of a greatly reduced para- Bathyopsts.
cone, a vestigial mesostyle and strongly pri met
crescentic metacone. In the last superior
molar these ectoloph elements are trans-
formed into a single oblique lophoid crest.
The inferior Coryphodon molar (Fig. 3)
shows a greatly elevated protoconid, an
enlarged metaconid and depressed para-
conid. ‘The talonid consists of a hypo-
conid, entoconid and a low spur, the
entoconid 2, prophesied in Pantolambda.
From this, the Bathyopsis and Uinta- Pantolambda.
therium lower molar is readily derived, as __,,Fig.3- Lower molars, Am-
: i 2 : blypoda. Diagrams showing
shown in Fig. Ze simply by the fission of evolution of crown. Not to
: A x scale. »
the metaconid into the metastylid, mld, —#r¢protoconid; pad para-
- = a ‘ conid ; me metaconid ; mid
and further reduction of the paraconid — metastylid; Zy¢ hypoconid ;
. aes 2 z end entoconid; e.2 ento-
and entoconid 2. ‘This fission is an ad- _ conid 2.
174 Bulletin American Museum of Natural History. (Vol. X,
ditional analogy with the Horse molar to those which the writer
has already pointed out.
The superior molar of Bathyopsis is unknown. That of Uznta-
therium probably repre-
sents, as Cope supposed,
metecone parastyle the ectoloph swung around
i dee so as to form with the
a protoloph a V opening out-
_metecone wards. Just internal to
the apex of the V the hypo-
cone is often developed.
F We thus observe a set
of profound changes re-
sulting finally in the
unique lophodont crown
of Uintatherium.
paracone
parastyle ae mesostyle /
BS t
(C7
Ap A)
ectolophS\\G Ys q
ei Pe
protocone protocone
Fig. 4. Upper molars of Coryphodon testis, show-
ing primary and secondary elements. Am. Mus. Coll.,
No. 274.
B. EVOLUTION OF THE UNGULATE FOOT FROM THE
CREODONT TYPE.
Cope’s famous generalization as to the serial character of the
primitive ungulate foot, supported and extended by Osborn, has
been recently disputed by Matthew (1897, p. 320) upon the very
strong ground that most of the earliest, z. ¢., basal Eocene, feet
are non-serial or displaced ; so far as known, both the Creodont
carpus and tarsus are certainly non-serial.. Cope’s generalization
Fig. 5. Creodont carpus (Déssacus carnifex), heavy lines showing hypothetical protun-
gulate displacement,
1898. | Osborn, Evolution of the Amblypoda. Part J. E75
therefore appears to be non-consistent with his own theory that
the Ungulates sprang from Creodonts.
The evidence as fo the earliest types stands thus :
Creodonta. Amblypoda. Condylarthra. Mentscothertum.
CanpuS seer Non-serial. Non-serial. Non-serial. | Non-serial.
ME ATSUS sie cuerey oo Non-serial. Non-serial. Serial. Serial.
Carpus.—-Matthew’s conclusion is important as concentrating
our attention upon the /xterlocking or alternating (lunar on
unciform, scapho-centrale on magnum) carpus of such a Creodont
type as Dissacus (Fig. 5), from which the carpus of the Amblypoda
and Condylarthra, and probably of all Ungulata, may be derived.
The manner in which this was probably effected is :
Amblypoda. Condylarthra.
Absorption or coalescence of Absorption or coalescence of
centrale. centrale.
Lunar remaining on unciform. Magnum greatly enlarged
Magnum somewhat enlarged. and spreading beneath
lunar, which is thus sepa-
rated from unciform.
The enlargement of the magnum, as observed in both Amblypoda
and Condylarthra, would tend to readjust the primitive Creodont
interlocking or displaced arrangement, and, especially where the
weight is concentrated on the median toes, give us the secondarily
serial Phenacodus type.
Fig. 6. Pes and Manus of Coryphodon testis, viewed from above, showing enlarged magnum,
Am. Mus. Coll., No. 258.
I 76 Bulletin American Museum of Natural History. (Vol. X,
Tarsus.—In the tarsus the case is different; while the non-
serial Amblypod (Pantolamébda) tarsus can be directly derived
from the Creodont, the serial Condylarth tarsus probably arose
from an unknown serial Creodont. Hence a double or parallel
derivation of the two-hoofed orders, as follows :
Persistent atypi- Carpus second- Carpus and
Protungulate ¢aj displacement In arily serial. Tar- tarsus typically
Creodonta. carpus and tarsus. sus persistently displaced.
serial.
Tarsus interlocking, t --+-Amblypoda.
interlocking. :
i oi ey tas t shar AM ere Condylarthra.....Diplarthra.
GiarSuS Seiial eermer tein:
Other characters of the primitive foot are given in the following
synopsis, also upon pages 184, 187, 188, 216 of this Bulletin :
2.—SYNOPSIS OF THE EVOLUTION OF THE AMBLY-
PODA.
The subjoined table brings out the three important laws of
phylogeny as applied to taxonomy :
First—TVhe persistent primitive characters (P.p.) are also the
ordinal characters, which serve to separate this phylum
from the Condylarthra, Proboscidia, Hyracoidea, Diplar-
thra and other ungulate orders.
Second.—The primitive (Pr.) progressive, (Pg.) and retrogres-
sive characters (Re.) constitute the subordinal characters.
Third.— Among the above characters are innumerable char-
acters both in skull, limb and foot structure in which the
Amblypoda farallel certain Perissodactyla, Artiodactyla
and Proboscidia. The latter are mainly approached in the
adaptations to great body weight, which first misled Cope
to place this group with the Proboscideans.
177
Primitive [Pr.], Persistent primitive [P.A ]. Progressive [Pz.] and Retrogressive [Re.]
Characters.
SKULL.
DENTI-
TION.
VERTE-
BRA.
Taligrada.
P.p. Brain small, ol-
factory lobes large,
hemispheres smooth,
cerebellum exposed.
Pr. No air cavities.
Sagittal crest. Fron-
toparietal sutures
open.
P.p. Anterior nares
terminal.
Pr. Premaxille sepa-
rate, reaching nas-
als.
Pry. Maxillaries
smooth.
Frontals smooth.
Py. Parietals smooth.
Pr. Nasals smooth,
extending between
orbits.
P.p. Mastoid (peri-
otic) widely exposed,
perforate, bordering
auditory meatus.
Pr. No alisphenoid
canal.
P.p. Zygomatic
arches slender.
Pr, Mandibular con-
dyle elevated, facing
upwards.
Pg. Molars triangu-
lar (tritubercular),
selenodont.
Pr. Paracone com-
plete.
Superior premolars
composed of two
crescents, unlike mo-
lars.
Pr. Superior incis-
ors present.
Py. Canines round,
normal.
Pr. Cervicals short,
dorsals and lumbars
with short spines.
[MWay, 7898.]
Pantodonta.
The same.
Pg, Extensive air
cavities. Cranium
flattened. Frontopa-
rietal sutures closed.
The same.
Premaxillee grooved
above, separate, not
reaching nasals.
The same.
Pe. Frontals with
supraorbital knobs.
Pg. Rudimentary
parietal horns.
The same, shortened,
extending between
orbits.
The same, much com-
pressed.
The same.
The same.
Condyle elevated, fac-
ing obliquely.
Molars triangular, se-
lenolophodont.
Pe. Paracone
duced.
The same.
re-
The same present.
Pe, Canines round,
trihedral, or com-
pressed, enlarged.
The same.
Dinocerata.
The same.
The same.
The same.
The same.
Re. Premaxillee di-
vergent, edentulous,
reaching nasals.
Pg. Maxillaries with
horns.
The same.
Large parietal horns.
Pr. Nasals with rudi-
mentary horns; re-
duced in length.
The same, widely ex-
posed and perforate,
bordering auditory
meatus,
Pg, An alisphenoid
canal.
‘The same.
4s. Condyle depress-
ed, facing back-
wards.
Molars triangular,
lophodont.
Re.
ing.
Paracone want-
Superior premolars
submolariform.
Ke. The same absent,
or vestigial.
Pe. Superior canines
lance-shaped. Infe-
rior, incisiform.
‘The same.
178 = Bulletin American Museum of Natural History. |Vol.
VERTE-
BRE.
RIBs.
ARCHES.
FORE-
Limp.
HUME-
RUS.
RADIUS
AND
ULNA.
CARPUS.
FEMUR.
TIBIA.
FIBULA.
TARSUS.
Talizrada.
Continued.
Pp. WD. ese:
Sacrals unknown.
Pr. Very short.
Pr, Scapula acumi-
nate, fossz subequal.
Py. \lium acuminate.
Pr. Bent outwards at
elbow : manus evert-
ed.
Pr. An entepicon-
dylar foramen.
Pr. Prominent del-
toid and ectepicon-
dylar (supinator)
crests.
Pr. Subequal.
Pr. Ulna, posterior
border convex.
Pry. Lunar resting
upon cuneiform.
P7. An os centrale.
Ye
Pr. Plantigrade.
Magnum small.
Pr. Prominent third
trochanter.
P.p. Rudimentary
spine and cnemial
crest.
Femoral facets, ap-
proximate.
Pr. Articulating with
calcaneum.
Pr. A tibiale.
Pr. Mesocuneiform
small.
Pr. Astragalus with
neck.
Pantodonta.
Continued.
27a) Sli
Sacrals unknown.
The same.
Pg. The same, infra-
spinatus fossae some-
what enlarged.
Peg. Ilium, border ex-
panded.
The same.
Pg. No entepicon-
dylar foramen.
Ps, Entepicondyle
somewhat reduced.
The same.
Poe whewsame, con=
cave.
The same.
Pg. Os centrale uni-
ted.
Pg. Magnum larger.
Pg. Sub-digitigrade.
Re. Reduced third
trochanter.
The same.
The same, approxi-
mate.
The same.
Pg. A tibiale, varia-
ble.
Poamluie samen ien=
larged.
Pe. Astragalus with-
out neck,
Dinocerata.
Continued.
ID IL ==?
Sacralsi—= 45
The same.
fe. Scapula triangu-
lar; infraspinatus fos-
sz greatly enlarged.
The same.
Pg. Straight, or ver-
tical at elbow.
The same.
The same.
The same.
The same, concave.
The same.
The same.
The same.
Digitigrade.
No third trochanter.
The same.
The same, confluent.
Pe. Not articulating
with calcaneum.
A tibiale, variable.
The same, enlarged.
The same.
1808. | Osborn, Evolution of the Amblypoda. Part J. 179
Taligrada, Pantodonta. Dinocerata.
Continued. Continued. Continued.
Taksus. | P.f. Astragalar fora- | The same, variable. The same, variable.
men.
Pr. An _ astragalo-| The same. The same.
cuboidal facet.
Pr. Astragalo-tibial| Pg. The same, ex-| Pe. The same, ex-
facet very limited| tended, horizontal, | tended, horizontal,
and facing obliquely | approaching astraga-| separated from astra-
outwards. lo-navicular facet] galo-navicular facet
anteriorly. anteriorly.
Pr. A calcaneo-fibu- | The same. Peg. No calcaneo-fibu-
lar facet. lar facet.
Pr. Mts. V. curved | Pe. Thesame straight, | The same, straight ;
with prominent ‘pe-| process reduced. process vestigial.
roneus_ brevis’ pro-
cess,
P;, Plantigrade. Pg. Subdigitigrade. | Pg. Digitigrade.
PEET. Py. Pentadactyle and | The same. The same.
Fig. 7. Pantolambda cavirictus. Cotype.
isodactyle.
Am. Mus. Coll., No. 963.
Lateral, inferior and superior views of cranium.
180 Bulletin American Museum of Natural History. [Vol.
3.—SYSTEMATIC REVISION.
Order AMBLYPODA Cope.
Most primitive order of Ungulates known. Avan very small, with smooth
Dentition
brachydont : Superior molars triangular (tritubercular, bunodont, selenodont,
cerebral hemispheres, large olfactory lobes and exposed cerebellum.
lophodont), rarely with functional hypocone. /ve¢ pentadactyl, isodactyl ; plan-
tigrade (as in Bear) to subdigitigrade (as in Elephant). sp/acement in carpus
(unlike Diplarthra) ; lunar resting on unciform only; in tarsus (like Diplar-
thra), astragalus resting on cuboid. A tibiale tarsi. Astragalus perforated or
grooved.
The three suborders are very widely separated.
SUBORDINAL CHARACTERS. |
Taligrada Cope.
Dentition.
Typical Eutherian form-
ula.
Molars tritubercular
selenodont.
Premolars simple,
two crescents.
with
Skull,
Nasals, frontals and
parietals smooth. A
sagittal crest. No alis-
phenoid canal; no air |
cavities.
Limbs.
Plantigrade. An entepi-
condylar foramen. A 3d
trochanter.
trale. Fibula articulat-
ing with calcaneum.
Astragalus with neck.
An os-cen- |
Pantodonta Cope.
| The same.
Lopho-selenodont,
Premolars two
crescents.
simple,
Rudimentary parietal
horns. Cranium flat-
tened. No alisphenoid
canal. Air cavities.
Semi-plantigrade. No en-
tepicondylar foramen,
A 3d trochanter. No
os-centrale. Fibula ar-
ticulating with calca-
neum. Astragalus
without neck.
| Sub-digitigrade.
Dinocerata Marsh.
No superior incisors.
Lophodont.
submolari-
Premolars
form.
Nasal, maxillary and pa-
rietal horns. Cranium
flattened. An alisphe-
noidcanal. Aircavities.
Ente-
picondylar foramen. A
3d trochanter. Os-cen-
trale wanting. Fibula
not articulating with
calcaneum. Astragalus
without neck.
1808. |
Osborn, Evolution of the Amblypoda. Part Tl. . 181
Suborder TALIGRADA Cope.
Family PERIPTYCHIDH Cofe. | Family PANTOLAMBDID&' Cofe.
Molars bunodont, primitive triangle | Molars selenodont, primitive triangle
compressed in superior molars; less compressed. Lower molars
secondary internal cusps developed without hypoconulid. No second-
(protostyle and hypocone). Lower ary internal cusps.
molars with hypoconulid. Third
and fourth upper and lower premo-
lars enlarged.
Family PERIPTYCHIDAL Cope.
In 1892 (op. cit., p. 47) Osborn pointed out that Periptychus
is distinguished from the order Condylarthra (in which it had
been placed by Cope) first, because the ¢arsus ts not serial, there
being a displacement of the astragalus upon the cuboid ; second,
Fig 8. Pantolambda bathmodon. Lateral view of skull, showing wide exposure of mas-
toid. Composition of Nos. 2550 and 2549, Am. Mus. Coll.
because it has the s¢récély trigonal molar of the Amblypoda. In
1897° Cope adopted this view and removed the Periptychidée to
the Amblypoda. The close comparison of Pantolambda and
Periptychus given below fully confirms this step. ‘The condition
of the upper premolars as composed of two crescents and (unlike
1 See Osborn and Earle, 182, p, 40.
' 2 fing Nat., 1897, p. 335. Matthew has, however, retained the Periptychide in the Condy-
arthra,
182 Bulletin American Museum of Natural History. \Vol. X,
the Condylarthra) with little or no tendency to acquire the molar
pattern, is essentially similar in the two families.
Subfamily ANISONCHINE O.&/. | Subfamily PERIPTYCHINE O.&L.
Smaller forms. ? Arboreal. | Larger forms. ? Terrestrial.
Superior molars with conules sup- Superior molars well developed.
pressed or wanting. Paraconid Paraconid strong.
reduced or wanting. Astragalus Astragalus with shallower trochlea.?
short and wide,! with deeper troch- |
lea.
The genera and species of Periptychidz, as recently revised by
Matthew,’ are given in the following geological table.
|
Creta-| Basal Lower :
ceous.| Eocene. Eocene. Middle Eocene.
Wasatch. i Bridger.
= 3
F : 2; % .
2 at etal coalesce Uae)
EUS ton lpr on |slias %| £ \£s
a ey lite mo | Oo | | oo] oo Te
See Wee lcs: Whey ee eee cess
HjalelA~l/el/ol/el/alere
Ofiixi yw. Viiei a iley ees coat somllborcor Meroe ore i>. <><
BATH VOPSIDIANE es, let oeychste oe ailleueenees epaae a Waeeedereet lke <llok
CORVEHODONDIDAE 2 see 5 Sotsllaara|loc-oleeGaeeGniles® less
PANTOLAMBDID.
Pantolambda: cavanictus. o.. sec aleereer Bell SS
2 bathmodoneere rere see elle
PERIPTYCHIDA,
HCHO OTS, GClnimeROeUS. 5 6c555a054l)550 20 Ds
Reriptychus thabdodon f=. ee| taser ror Kee
oF Catimidensm eneree ee see Hiatt.
i COATCTAtUS Hee eee ers eee SK
Conacodon entOconlsS.. eee ESS
oe COplater ay mesiar si sreys ete ><
Haploconuslineatus 3) eee ee foie uf x
MAYS COLMICUIA ISH ee eee eee an all Sx
Amisonchus Sectonusse sas se cceie leer Aiea aes
ee INET. coos coo So 55\|s000¢ x
Hemithleus kowalevskianus .....|..... x
INCERTA SEDIS.
Eictoconodonta casei aera oe Aia.<
SHUTGOMOTSYN Goo9 sadaescasoont ex
Rrotolambaarneeemereechin ene OS
1 Matthew, Bull. Am. Mus. Nat. Hist,, 1897, p. 297.
2 Op. cit., p. 265.
1898. |
Osborn, Evolution of the Amblypoda. Part 1. 183
Family PANTOLAMBDID® Cofe.
Genus Pantolambda Cope.
Dentition typical. First upper premolar one-rooted. Second, third and
fourth three-rooted, with internal cones. Canines rounded.
P. bathmodon. | TV GQULELGLUS.
|
Type: Mandibular ramus, No. 3956. | Type: Jaw, No. 3961.
Smaller size. Both dental series con-| Larger size. First lower premolar
tinuous. | close to canine and separated from
| second by a wide diastema. Pre-
molars reduced in size.
P. cavirictus' Cope.
The type lower jaw, described and figured by Cope (Am. Nat.,
Vol. XVIII, p. 1111) is peculiar in the close apposition of the
first lower premolar to the canine, and the wide diastema behind
it. This is the largest type known and the diastema is probably
prophetic of the diastema invariably observed in Corvphodon,
(Fig. 2).
The skull (Fig. 7) was mistakenly described by Osborn and
Earle in 1895 (1895, p. 43) as P. bathmodon. It differs in its
much greater size from P. bathmodon, and in the absence of dias-
temata from the P. caviricfus jaw. Unlike Corvphodon the upper
canines are mainly worn upon the inner posterior surface.
Fig. 9. Skeleton of Pantolambda bathmodon. Scapula wholly, pelvis partly restored.
Composition from several individuals. One-eighth natural size.
1 Am, Nat., Vol. XVII, 1883, p. 968.
2 Am, Nat., Vol. XVI, 1882, p, 418.
184 Bulletin American Museum of Natural History. [Vol. X
P. bathmodon' Cope.
The composition skeleton of P. bathmodon measures 2 feet 9
inches (830 mm.) from the premaxillaries to the back of the ischiac
symphysis, and 1 foot 134 inches at the withers. It is thus about
the size and proportions of a large Wolverene (Gu/o luscus).
Excepting in the selenodont teeth, it typifies the Aypothetical
Protungulate, being more primitive than either Awprofogonia or
Phenacodus. The step is that of the
Bear, the feet very broad and spread-
ing, the wrist and ankle being slightly
raised off the ground, and the phal-
anges terminating in hoofs.
The vertebrez preserved (Nos. 2549,
anca )indicate a short, neck (C,6—
tomm.) as in Periptychus, and a back
increasing in strength and power as we
pass towards the lumbar region, Thus
the dorsals are short anteriorly (D. 5
=15 mm, No, 2549) and indicate
less separation of the zygapophysial
and rib-tubercle facets than in P/ena-
codus. The lumbars (L. 4= 25 mm.,
ene, pie No. 2549) are longer; unlike most
Astragalus, metacarpal III and phal- Creodonts they present horizontal
anges. Am. Mus., Cope Coll., No. : a
3957: rather than vertical zygapophysial
facets. The tail is long and powerful.
PRIMITIVE OR PROTUNGULATE CHARACTERS.—Among the
persistent primitive or Creodont characters of Pantolambda are the
following :
Brain small, olfactory lobes large, hemispheres smooth. Skull with a sagit-
tal crest ; terminal anterior nares ; nasals very Jong and expanding posteriorly;
mastoid (periotic) widely exposed and forming lower posterior border of exter-
nal auditory meatus ; tympanic bones rudimentary ; zygomatic arches slender ;
no alisphenoid canal; basi-cranial foramina separate.
Dentition typical; no diastemata; molars tritubercular, incisors small,
cylindrical; canines rounded. Girdles: scapula unknown ; ilium acuminate
as in Phenacodus.
1 See Cope, Am. Nat., Vol. XVII, p. 406.
1898. | Osborn, Evolution of the Amblypoda, Part J. 185
Fore-limb strongly
bent outwards at el-
bow (as in Creodonta
and Carnivora), ma-
nus everted. Humer-
us with powerful del-
toid, pronator (ente-
picondylar) and supi-
nator (ectepicondylar)
crests; ulna with a
convex posterior bor-
der; carpus with an
os centrale, an ex-
tremely small mag-
num and short trape-
zoid, causing the me-
tacarpal IV to be in-
serted proximally be-
tween the trapezoid
and magnum (Fig.
2).
Hind-limb straight,
with three trochanters
upon the femur (Fig.
11). Tibia with a
rudimentary spine, a
very long cnemial crest
(Fig. 11) and femoral
facets approximate.
Tibia (Fig. 11) articu-
lating with calcaneum.
Probably an os-¢zbzace
(Fig. 12). Mesocu-
neiform short (analo-
gous to trapezoid in
the carpus), so that
metatarsal IV articu-
lates between ento—
and ectocuneiforms
(analogous to meta-
carpal IV). Articu-
lation between tibia
and astragalus slant-
ing obliquely inwards,
very limited in extent,
bounded. posteriorly
- i Ni 2
Fig. 11. Pantolambda bathmodon. Anterior view of fore
and hind limbs. showing powerful development of crests and tro-
chanters as in the Creodonta. Humerus, No 2549. Femur, No.
2523 (2551, 2549). Tibia, No. 2551. Ulna, Was 2550, 2547. Ra-
dius, No. 2547 (2546). Am. Mus. Coll.
186 Bulletin American Museum of Natural History. [Vol. X,
by astragalar foramen which issues posteriorly between the ectal and sustentacu-
lar facets (Fig. 12) (as in Creodonta). Astragalo-navicular head broad convex ;
two astragalo-calcaneal facets only. Fifth metatarsal curved with a prominent
external tuberosity for the peroneus brevis muscle, as in the Bear.
All these osteological characters are shared by Periptychus, so far
as the skeleton of the latter genus is known; (the carpus of
Periptychus is unknown). The skull, vertebra and proportions of
the limb bones in the two types are remarkably similar.
ORDINAL OR AMBLYPOD CHARACTERS. — Primitive ordinal
characters. It will be observed in the definition of the Ambly-
poda, p. 180, that many of these above-described primitive charac-
ters persist throughout the evolution of the order, and therefore
rank as ordinal.
MEASUREMENTS: SKULL, TEETH, SKELETON.
P. BATHMODON. P. CAVIRICTUS.
Mus. Nos. | Mm. |} Mus. Nos. Mm.
= se |
Skull, condyles to premaxillaries . . est. 2548 157 963 | est.272
Molar-premolar series, superior....... Wereecscre O55 ry 088
ime Ore en || 2550 060 3961 118
Lower jaw, symphysis to angle..... 5 2550 126 er est. 251
Fore-limb: Humerus, total length...)) 2549 124
oe Radius, ¥ ovolil Oxy 083
a Manus, oo ers = o81
Hind-limb : Femur, Me ete 2551 148
is Tibia, - es oy 11g
sey Pes, oe i 10g 3963 184
Ilium, total length, as restored........ 7 IIl
Sole “WALCithe sera eee scttectens tt Bee ell renee ee E 031
RESTORED SKELETON.
Skull to perpendicular of ischiac
SYMP MYSIS 5 save esata eve legekotet Mon svcenel sense etses ter eeeeqe as 830
Heioht -atjshoulder bladegen 50s serrl|l(eerei: <Hol) SEXO)
Se) Salami eos eee eee varcgere stereo ere lesen eves 332
SPECIALIZED ORDINAL CHARACTERS.—There are, however,
many structures in Pantolambda which amply affirm Cope’s view,
that Puntolambda is the ancestral type of the great lower and mid-
dle Eocene forms. ‘These are: the separation of the premaxillz
1808. | Osborn, Evolution of the Amblypoda. Part /. 187
in the median line (as in Coryphodon and Uintatherium), the widely
exposed mastoid (as in Uintatherium); the nasals extending far
back between orbits (as in Coryphodon) ; the molars triangular and
selenodont leading to the Coryphodon type. The scapula is
unknown, probably acuminate as in Periptychus. The most
striking likenesses to Coriphodon in the fore-limb are in the
muscular crests of the humerus, the outward flexure of the elbow,
the displacement of the lunar upon the unciform.
STRUCTURE OF THE FEET IN PANTOLAMBDA.
Both manus and pes are of exceptional interest. ‘The terminal
phalanges are hoof-bearing.
Fig. 12. Pantolambda bathmodon. Superior view of carpus No. 2546, partly restored, and
tarsus (composition No. 2551), showing essential Creodont structures. Coll. Am. Mus. Nat.
Hist.
Fore-foot.—As carefully put together by Dr. Matthew (see Fig.
12), the manus certainly possessed a separate centrale. Other
exceptional features are the curvature of Mtc.I] and the propor-
tions of the distal carpals.
Pantolambda | Magnum Trapezoid Trapezium
bathmodon. (| smallest. 4mm. intermediate. 9mm. large. 16mm.
Coryphodon | Magnum Trapezoid Trapezium
testis. ( large. 37mm. smallest. 30mm. largest. 4omm.
188 9 Bulletin American Museum of Natural History. \Vol. X,
These proportions show that the evolution of the carpus in the
Amblypoda marks an enlargement of the magnum and trapezium
and absorption or coalescence of the centrale.
Hind-foot—TVhe very large pes,
ze probably belonging to P. cavzrictus
4 (No. 3963), and small pes of P.
bathmodon (No. 2550) agree in the
many primitive characters enumer-
ated in the Table, p. 178. Analo-
gous to the manus we find a curved
first metapodial (Mts. I). As in
Periptychus and in Ursus the fifth
metapodial bears a prominent pro-
cess for the peroneus brevis abduc-
tors. As in most Creodonts the
cuboid is wedge-shaped proximally
for the calcaneum and astragalus,
and the astragalo-tibial facet faces
at pie ae obliquely inwards (instead of up-
Right pes. directly from above. No. wards as in Coryphodon) ; this facet
3963, Am. Mus. Coll.
is very narrow, bounded by the
astragalar foramen posteriorly and
a deep pit anteriorly. A very important feature in this species 1s
the thinning out of the inner side of the navicular, bringing the
ectocuneiform almost into contact with the astragalus.
Comparative Measurements.
P. bathmodon. P. cavirictus.
MM. MM.
Astragalus oneatest iam etetar mr eres sierra 20 44
Tarsalss transverse) measumemlem ts mya pats) r= tenelo = 33 68
Mighemichl I SBoonasess opsobsotcagno@onGouT 28 aire
a WO por ecers Ame ree oR recutne ero arc siorese 34 Est. 60
Of ¢ MMT eon tortie Sere tetioicio weet 40 61
‘a NY eRe MG OF cas. S000 OOS OF 30 60
ie WORE AS arate cpal si mot vette MeiensaodeTo shes ees 28 50
Suborder PANTODONTA Cope.
The Pantodonta or Coryphodontia are distinguished by a very
great increase in size, the large development of the upper and
1898. | Osborn, Evolution of the Amblypoda, Part J, 189
lower canines, the lopho-selenodont molar teeth, and the broad-
topped skulls. ‘These and other advances upon the Pantolambda
type were effected in the interval between the deposition of the
Torrejon or Upper Puerco and the Wasatch Beds.
Fig. 14. Skull of Ceryphodon testis. Male specimen, No. 2867, as mounted in skeleton.
Lower jaw, No. 2872. Am. Mus. Coll. The back part of the skull is elevated by distortion.
REVISION AND CRITERIA OF SPECIES.
Twenty-one species were named by Cope, with as little regard
for the laws of individual variation as for the association of
skeletons with teeth or of jaws with skulls. It is @ prvorz improb-
able that such numerous species should have coexisted, consider-
ing that all the collections come from a few levels and a single
geographical region. Our knowledge of large living quadrupeds,
such as the African Rhinoceros, shows that rarely more than two
species of one genus coexist, and these have different local feeding
habits. The writer has found the same to be true of the Eocene
Titanotheres of Wyoming. FEarle’s revision of the species (1892),
therefore, marked a valuable advance but left much to be done,
owing to his lack of comparative material at the time.
190) Bulletin American Museum of Natural History. |Vol. X,
— = ————e ne —— = — <—__—______——
XIN
C. armatus
C. testis. )
Fig. 15. Typical upper molars. C. swbguadratus, type; C. cinctus, type; C. elephan-
topus, cotype; C. radians, type; C. testis, male, No. 274; C. anax, type; C. armatus;
C. testis, type, female. Allin the Am. Mus. Coll.
Altogether there are about thirteen distinct species known at
present, which are distributed as in the following ‘Table.
1898. |
Osborn, Evolution of the Amblypoda.
WASATCH.
WYOMING.
| Winp RIVER.
WYOMING.
| Evanston,
Black Buttes.
SERIES I.
Ge wontmaniiea.-
Series IT.
C, testis
““ repandus
marginatus .
* cinctus
semicinctus....
lobatus
einige
pachypus...:..
ae
cuspidatus
obliquus
latipes...
latidens
6c
oe
SERIES III.
C. armatus
‘* simus
INCERTA SEDIS.
C. radians ..
SSO AMS MB neo
subguadratus ..
“ee
ae
elephantopus...\..
curyicristts....|..
ae
singularis
Gray Bull, |
| Big Horn Mts.
Clark’s Fork.
Buffalo Basin,
Wind River
Mts.
| Big Horn Mts. |
XXX
-Morar Typrs.—It is essential, first, to clearly conceive the
correlated changes taking place in the upper and lower molar
teeth, as shown in the accompanying figures and diagrams.
In the upper molars :
ite
Pantolambda (Fig. 2) exhibits the triangular upper molar
ancestral to Corvphodon, ectoloph (with crescentic cones and meso-
style) at r7gAt angles to protoloph
192 Bulletin American Museum of Natural History. |Vol. X,
md FG + én?
Wns mek Gn me? m2
C. testis. 2
pr
_S
if z
me’ m3
C. simus, Carmatus.
Fig. 16. Typical lower molars. C. eocenus, cast of Owen’s
type. C. testis. C. cinctus, variety, with triangular heel, No.
4329. C. lobatus, typical lower molar, No. 4305. C. ventanus,
type, No. 2970. C. curvicristis, type, No. 4326. C. szmus,
No. 2563. C. armatus, type, No. 4316. Allin the Am. Mus.
Coll.
C. curvicristis.
2. C. radians pre-
sents an intermediate
stage, ectoloph (with
crescents and meso-
style disappearing)
oblique to protoloph
(Figs 15):
3. C.armatus pre-
stage,
oval crown, ectoloph
sents a final
(a slightly concave
crest) parallel to pro-
toloph (Fig. 15).
The second upper
molar slowly goes
through the
phases as the third,
and both approach
the Uintathertum
pattern.
same
In the ¢tizrd lower
molar various species
of Corvphodon ex-
hibit (Fig. 16):
1. Primitive stage,
a trilobed heel (hypo-
conid, entoconid, en-
2 i= CKEStS
heel with
cusps,
toconid
oblique,
three main
C. eocenus.
2. Intermediate
stage, a heel with two
main cusps (hypo-
conid, entoconid),
third cusp (entoconid 2) degenerate ; crests dess obligue, C. testis.
3. Final stage, a bilobed heel (hypoconid, entoconid), ento-
conid 2 absent, crests transverse, C. simus.
Tapir or Lophiodon type.
A tooth exactly of the
1898. | Osborn, Evolution of the Amblypoda. Part J. 193
Thus the crests of the upper and lower molars slowly become
transverse, and simultaneously also preserve their mutual inter-
locking shear; the stages, 1, 2 and 3, in each being probably
correlated.
It is, however, extremely difficult to determine the species by
these principles alone, because the hypolophid, or posterior crest,
of the third lower molar seems to be highly variable, not in its
obliquity of angle, but in the greater or less development of the
entoconid 2, as frequently seen upon opposite sides of the same
jaws (e. g., Nos. 2868 and 4321).
CurtinG Teeru.—The canines aid us: the primitive form is
round (persisting in C. wortmant) asin Pantolambda ; the inter-
mediate form is “7-
angular and antero-
posteriorly compressed
(C. testis, C. ventanus) ;
the final form is flat-
tened and laterally
compressed (C. arma-
tus), paralleling the
Uintatherium type, be-
cause the long axis of
the blade is longitudi-
nal.
SKULL.—When fully
known, the top of the
skull will prove highly
distinctive. At present
we know only the com-
paratively primitive
crested type (Fig. 18 By. Fig. 17. Upper and lower canines, lateral views and
sections. A, Coryphodon testis, 4. male upper and lower
and the flattened type canines, No. 274. ?, female lower canine. B&B, C. anax,
: : male lower canine, No. 4328. C, C. wentanus, male
(Fig, 18 G): which ap- upper canine, No. 2970. LD, C. armatus, upper canine
5 : and section, No. 4315. This very small canine is upon
proaches Cintathertun. double the scale of the other teeth.
[May, 1898.] 13
194 Bulletin American Museum of Natural History. \|Vol. X,
Fig. 18.
Comparative view of Crania of A, Pantolambda bathmodon,
small, of B, Coryphodon wortmani, intermediate, of C, Coryphodon testis,
large, showing flattening of the upper surface of the cranium, and appear-
ance of parietal horns, Pa.#7. All one-fifth natural size.
Disregarding synonyms and_ partially known
types, these characters, together with the form of
the canines, indicate three series, or lines of spe-
cies, typified as follows :
Levels. | Series LI.
Primitive. | Narrow
| crested skull.
| Canines rounded.
Wind River | C. wortmantz.
Wasatch... |
Series Lf,
Specialized. Broad
crested skulls.
Canines triangular.
Second lower incis-
ors enlarged; third
ditto reduced.
Lower molar crests
oblique to trans-
verse. M, trilo-
bate to bilobate.
C. armatus.
RTA NTL fe
Specialized. Medium
crested skulls.
Canines laterally
GO mMeEpEGensisieids,
grooved. Molar
crests nearly or
quite transverse.
M,, bilobate.
1898. | Osborn, Evolution of the Amblypoda. Part J. 195
The presence of the narrow crested C.
wortmant in the Wind River Beds, the
highest level in which the genus appears,
shows that the fwo serves evolved simultan-
eously, just as we observe among the flat-
tened and crested Eocene ‘Titanotheres.
In each the evolution of the molar crests
Bg els
in
fomiic)
oi
SoH
+, Co
8
ere:
3 :
SS :
S ES
Ss
Sy Q
= ZN
:
5 a
™ eg 3
JeimjeN jO WhasnyY uvooury ey} Ul UOJaTSys paluNoP
from oblique to transverse angles, with degeneration of the ento-
conid, loss of crescents, etc., was apparently parallel.
The sex characters, as clearly shown in numerous specimens
of C. ¢estis, are: Males larger, with powerful upper and lower
canines ; females smaller, with smaller upper and lower canines,
(See Fig. 17.)
196 Bulletin American Museum of Natural History. |Vol. X,
Among the specific or variable characters in the skeleton are
the presence or absence (compare Earle) of the tibiale facet, of
the astragalar foramen, and of the articulation of the cuneiform
with Mte. V, which variation is similarly observed in Ucntathe-
rium, as shown by Marsh. (‘ Dinocerata,’ p. 107.)
None of the generic characters assigned by Cope to Bathmodon,
Ectacodon, Metalophodon and Manteodon appear to the writer to
be valid, as they rest either upon errors in field collection or
upon individual variations.
Fig. 19. Coryphodon elephantopus, showing flat-topped skull. A, view of occiput; &
transverse section of cranium through region of olfactory lobes, showing air cells. Coll. U.S
,
Nat. Mus., No. rrr. ‘
Series I7.—LARGE CORYPHODONS WITH BROAD, FLarT-
TOPPED SKULLS, RUDIMENTARY PARIETAL Horns, TRIANGU-
LAR CANINES.
12. C. elephantopus Cope.
Type, U.S. Nat. Mus. Coll. Superior molar 3 ; inferior molar 3. Loc., New
Mexico. This type has been temporarily displaced.
Cotype, No. 111, U.S. Nat. Mus. Coll. Skull with upper dentition complete ;
inferior dentition and fragmentary jaw.
Definition.—Superior molars=158 6. Skull characters as in C, zestis, except-
ing m® more quadrate (cotype) with slightly convex mesostyle ; m* (type) with
hypolophid and entoconid 2 ; incisors equal sized.
Synonyms.
C. obliquus Cope. Type, U.S. Nat. Mus. Coll. A single fragmentary inferior
molar. Hypolophid oblique. Entoconid 2 reduced.
This is a smaller animal than C. ¢estis, but is found upon a
higher level.
Cope’s association of type and cotype is open to some ques-
tion. ‘The type has been temporarily lost, and unfortunately the
Er
Part 1.
Osborn, Evolution of the Amblypoda.
1898. |
TABLE SHOWING THE PROPOSED SPECIES wirH ORIGINAL Names, NATURE OF Type, ETC.
US NAME. Date, TYPE. FIGURE.
Nn |
1 Coryphodon eocenus Owen.......) Hist. Brit. Foss. Mamm., 1846, Pp. 299..../Inf. m. 2 and m, 3...|Figs. 103, 104.
2 ““ anthracoideus Bainville. |
3 = owent Aébert........ S|Amtlanscy Nat. Vile TOSO)sp.O 7. on. nee - Skull and teeth. |
4 |Bathmodon radians Cofe........ .|Proc. Am. Phil. Soc, wieby EO) 1872, D. 418, Sup. m. 2 and 3 with)
Skeletonmerte eer Menta \Ver emp Seediy fie
5 cal semicinctus Cofe...... * a ss SD 420s heeth:
6 Metalophodon armatus Cofe....... x o OP, (0b, GUIS co core Superior molars. =| Terk Ver., pl. 49.
7) Bathmodon latipes ae Beciogenee oie Me Me Ue Ey (Os Waa do ear bv Skeletons sity: Oo a tals
8 brevipes’. . --...+-.|Proc, Acad. Nat. Sci. Phila., 1873, p. 103,
9 we simus Cope. Pn og day ooo oils Wien, Io ING ile Wheel ler 1874, p. 120, Inferier molars...... Pal. of N.M., pl: 55.
IO s MOLEStUS| CapeHae mes s : Sf “p. 121d, Sup. and inf. molars, pls. 56-57.
iit a Tomas Cope se arena ne us Y uf SD L2e.| Intenoramolanses sires be ee iol, yates
12 ¥ elephantopus Cope. | nS ve a 3 “p. r21c, Last sup. and inf.m.3, ee “pls. 50-54.
13} bs. latidenss Capers semis oyse, Cat, Vert. Wa Mls Uti, jo, Ao) .|Jaw with teeth...... ot mS ““ 48-50
14 ag cuspidatus Cofe...... ; oe - ps 30. ...- |Inferior molars 3). es ve jails 416).
15 Coryphodon enna ILS WHT PSTES 30 46 Am. Jour. Sci. and Arts, 1876, p. 426.... Sup. and inf. molars, Mon. of Dinocerata,
MSs
16 e eisai Cope......,..|Pal, of New Mexico, W heeler, Sh 7/ 75 Dn AODy)o, Hs oo Wert. Palen .M., pl. 47.
17 as IkoORNIIS (Cop, Gccnacus Aon! UE 2 x ie 46.
18 Manteodon subquadratus Cofe.... . Proc. Amer. Phil.Soc., Dec.16,1881, p. 166, Last sup. m. with
incisors . soo) Weris Wetton, falls ale,
To) | Wetacodoncinctus) Cape.) nee or of ve ‘“_ p. 167, Superior molars... .. ™ Sa
20 |Coryphodon anax Cofe...........| - S 188r, fs HOS goad . Sup. and inf. molars, te pls. 44a-e.
21 * repandus Cope... .. .| id * (Op yl Repaecasote Inferior m. 2 and 3 ae pl. 44c.
22 a curvicristis Cope...... S a! PR W7Praadeo ‘iio, il, 2 voxel sonleyol,
Walch teehee) set oi TAGE
23 a marginatus ste sre i ss seas Ober ty): Nenrcren se: Superior molars 3 : - 44g.
24 Metalophodon testis Cofe.........| si Be Pea aah Grist eee ee core gs «Ada.
25 bBathmodon pachypus Cofe..... ...|Proc. Acad. Nat. Sci. Phila., 1882, p. 204, Skeletonteyes terre i pls. 44d-g.
26 Coryphodon ventanus Oséorz......)/ Bull. Amer. Mus. Nat. Hist.,X,1898, p. 210, Jaw and teeth....... + Op. Clty, pie aor
27 % wortmani Osborn.....| “f 1 3 p. 212, Skuli. Om Ot
28 = singularis Osborn ..... _ ‘- “ p. 2t4, Tarsus. We Sees seta ck
1 Name only.
liMolae
=
Ffistory.
198 Bulletin American Museum of Natural
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Osborn, Evolution of the Amblypoda,
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200 Bulletin American Museum of Natural History. |Vol. X,
characteristic last lower molar, m2 in the cotype, is wanting.
The species must therefore rest upon the characters of the cotype.
No lower teeth were found with the fine palate (No. 275, Am.
ai ROM
a TD Se
Fig. 20. Coryphodon elephantopus, Cotype. Lateral, superior and sectional views of
cranium. No. 11, U.S. Nat. Mus. Coll.
Mus.) rightly associated with the cotype by Earle. The cranial
and dental characters of the cotype (Fig. 20) are those of C.
testis and C. dobatus upon a smaller scale ; m® has a well-marked
postero-external elbow (Fig. 15), and the paracone is sharply dis-
1898. | Osborn, Evolution of the Ambiypoda. Part 1. 201
tinct from the metacone ; the ectoloph thus does not form a con-
tinuous crest as in the supposed type. ‘The parietal protuber-
ances or rudimentary horns Pa./7, are less pronounced but
equally rugose ; the premaxillary symphysial borders are extensive
although without contact ; the incisors are equal in size, 11 and
13 being fully as large as i2 in both jaws. This is therefore a
smaller and perhaps more primitive type than either C. /odatus
or C. ¢estis, although skull No. 275 is definitely recorded by
Wortman from the Buffalo Basin, the highest true Wasatch level.
Unfortunately the characteristic last lower molar is missing in
the cotype ; the series pm 1-m 2 measure 122 mm. ‘The lower
jaws of C. elephantopus are also represented either by Cope’s
C. obliquus or by his C. latidens (see below). ‘The former is more
probably the case for the following reason. According to the
ratio of upper and lower teeth established in the C. /estzs jaws
(see Table, p. 199, No. 3829), the lower grinders in C. elephantopus
should measure 167 mm. The type lower molar of C. od/iguus ap-
proximately agrees with this size (see Table, p. 199) and character.
The last lower molar of C. obdiguus agrees closely with that of the
supposed type of C. elephantopus. We may therefore consider the
greater or less development of the entoconid 2, which these
molars present, as variations similar to those which we have ob-
served in the other species of this series, namely, C. festis and
C. lobatus.
No complete jaw is nearer this size than No. 4321 (Am. Mus.,
Cope Coll.), in which the lower grinders measure 172 mm. ; this
specimen is also significant because the last inferior molar on the
right side agrees in form with C. cuspidatus (¢.e., entoconid 2, dis-
tinct), while the same tooth on the left side agrees with C. od/iqguus
(z.e., entoconid 2, obsolete). Another proof of the variability of
these cusps. This jaw, however, may belong to a small female of
C. testis.
INCERT# SEDIS.
21. Coryphodon repandus Co/e.
Type, No. 4309, Am. Mus., Cope Coll. Superior and inferior molars m*, m®,
Ms, Mz. Symphysis of lower jaws. Size=C. éestis, male. Loc., Big Horn,
Wyoming.
202 Bulletin American Museum of Natural History. |Vol. X,
This is an indeterminate type. It is distinguished by angula-
tion of ectoloph in m# (as in C. elephantopus, cotype) ; perhaps
also by the more transverse direction of hypolophid in mg ;
second incisors only slightly larger than first and third (as in C.
elephantopus, cotype). The nearest resemblance is therefore
to C. elephantopus, from which it is distinguished by larger size.
Superior molars No. 4366, from New Mexico, furnish a transition
in the angular form of the ectoloph of m# to the C-. ¢estis type.
Altogether C. repandus is of very doubtful validity.
19. C. cinctus Cope.
C. (£ctacodon) cinctus. Type: No. 4341, Am, Mus., Cope Coll. Superior
molars complete. A strong cusp appearing at postero-external angle of m®*.
Loc., Big Horn, Wyoming.
The distinctive feature of this type, viz., the quadrate form and
postero-external basal cusp of m® (Fig. 15), and to aless extent on
m2, are either individual variations or valid specific characters.
They are certainly not generic.
Lower teeth which may possibly be correlated (Nos. 4329, 4334,
266) have a triangular heel upon the last lower molar (Fig. 16),
with entoconid very distinct and extremely short and oblique
hypolophid.
24. C. testis Cope.
24. C. (Metalophodon) testis Cope. Type: No. 4317, Am. Mus., Cope Coll.
Superior molar series. Originally distinguished by reduction of posterior cres-
cent spur in m®.
Definition.—Sup. molars=169¢to 1826. Inf. molars=172¢to 192¢.
Third superior molar typically oval, with oblique posterior crest with primitive
paracone, mesostyle and meta-crescent more or less distinct. Third inferior
molar with oblique hypolophid, entoconid 2 reduced or vestigial. Second
incisor the largest.
This includes the most completely known Coryphodon. It has
been heretofore described by Earle and the writer as C. radians,
but is now found to be distinct. ‘The identification with Cope’s
type of C. desé’s is made by means of a careful comparison with
the superior molars in the female skull No. 2963. ‘The form and
measurements are identical. As this skull undoubtedly belongs
1898. | Osborn, Evolution of the Amblypoda. Part J. 203
to the same species as the male skull (Fig. 21) and skeleton, all
the characters of this fine type are now available.
This is the largest Coryphodon but one, and is very abundant
in the Middle Wasatch levels, being represented by a magnificent
series of skulls and skeletons in our collection. From these the
sexual characters are clearly made out. The large male skull is
used in the complete mounted skeleton, Fig. 18 4. The smaller
Fig. 21. Coryfhodon testts. Large male, showing rudimentatary parietal horns. Upper
canines partly restored. Skull No. 2867, lower jaw, No. 2872. Am. Mus. Coll.
female type of this species is represented in the skull No. 2963,
and jaws (Nos. 2868, 259) in contrast with the powerful male
skulls (Nos. 2829, 2867) and jaws (4322).
Variations in the last lower molar are considerable, from an
oblique to a bilobed (No. 259) or less oblique condition of the
posterior crest, with all the stages in reduction of the entoconid 2.
Exactly similar variations are found in the lower molars of the
larger and smaller members of Series Il. The development of
entoconid 2 also varies in the posterior molars upon opposite
sides of the same jaws of several specimens of C. lobatus.
The osteological characters have been fully described and fig-
ured by the writer (this Bulletin, 1898, pp. 81-91). Full charac-
ters of the vertebral column are shown in Fig. 23.
Certain specimens (skull, No. 2866) of the still larger C. Zobatus
have been found below it, and the much smaller C. edephantopus
204 Bulletin American Museum of Natural History. [Vol. X,
occurs in the higher levels of Buffalo Basin, Wyoming. Our
scanty evidence therefore appears to indicate a retrogression in
size in this series, but this is an inference by no means certainly
established.
Pao? “apai th pe m.1 ma
Fig. 22. Coryphodon testis. Superior molar series, male (Am. Mus Coll. No 274); inferior
molar series, female (Am. Mus. Coll. No. 2868).
17. C. lobatus Cope.
Type, Nat. Mus. Coll. Sup. molar 3; inf. molar 3; part of sup. and inf.
canine, indeterminate.
Definition.—Sup. molars=193 4. Inf. molars=1964. Dental characters
as in C. ¢estis, excepting elongation of protoloph and degeneration of posterior
metacone crescent in m®. Astragalus usually lacking astragalar foramen.
Cranium massive, with widened parietal horn rudiments.
Synonyms.
20. Coryphodon anax. Type: No. 4327, Am. Mus., Cope Coll. Superior
molar 3 ; inferior molars, premolars and incisors. Loc., Big Horn, Wyoming.
25. C. (Bathmodon) pachypus Cope. Type: No. 4335, Am. Mus., Cope
Coll. Astragalus, calcaneum, pelvis, femur, &c. Indeterminate type. Loc.,
Big Horn, Wyoming.
This is the largest Coryphodon known ; it surpasses C. fest¢/s in
size, the ratio being 50: 45, as indicated by the femora.
Unfortunately the name C. /obatus is prior to the more appro-
priate C. anax,and must supersede it. The lower molars defined
1808. | Osborn, Evolution of the Ambl
vpoda. Part J. 205
ye} TepNoseqny
*€z *S1q
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wood
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“uoya]axs pejanouw jo uurnyoo [erAqgei19 A
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z= 35)
~@ /> of its skull and molar teeth.
as C. lobatus Cope, which may be
considered as indeterminate types,
present exactly the C. avax measure-
ments (see Table, p. 198), and exhibit
an oblique hypolophid and depressed
entoconid 2 upon ms, as shown in
Fig. 16. ‘This may be considered the
typical C. Jobatus or C. anax third
lower molar. It is well shown in the
large jaw, No. 4333, and in the frag-
ment, No. 4305.
A variation, No. 266,in molars of
the C. Jobatus size is paralleled by a
variation, No. 4239, in molars of the
C. cinctus size, as represented in Fig.
16, in which the three cusps form a
posterior triangle, as also in C. cuspr-
datus (No. 4324). If these are not
variations they represent three dis-
tinct species, which is possible but
not probable.
Synonym.—The skeleton defined as
C. pachypus by Cope (No. 4335) un-
doubtedly belongs here. The astra-
galus of C. pachypus and of another
specimen (No. 2870) exhibit no astra-
galar foramina, and show a wider in-
terval on the front face of the astra-
galus between the tibial and navicular
facets than we find in C. éestis.
The massive male skull in our col-
lection (No. 2866) is most interesting
in its progressive development of the
parietal horn thickenings, parallel
with those of Uvdntatheritum. Its
geological level, however, according
to Wortman, is below that of C. éeszzs,
an observation very difficult to recon-
cile with the more advanced evolution
206 Bulletin American Museum ae Natural History. er x
14. C. cuspidatus Cope.
Type, Nat. Mus. Coll. Fragmentary inferior molars 2 and 3, and a portion
of the jaw. Originally distinguished by prominent entoconid 2.
Cotype, No. 276, Am. Mus. Coll. Complete lower dentition ; upper pm*—m®.
This specimen was referred to C. obfiguus by Earle (1892, p. 162).
Definition.—Inferior m.pm.=154. Superior m.pm.=150. Last superior
molar oval, antero-posteriorly compressed. Last inferior molar with oblique
hypolophid and more or less prominent entoconid 2. Metaconid with rudi-
mentary metastylid. Inferior incisors unequal in size.
This is a diminutive Coryphodon, of the size and very similar
in molar type both to C. eocenus Owen, from the London Clay,
and to C. owenit Hébert, from the Suessonian of France. The
characters of Cope’s type are very indefinite ; the complete upper
and) lower series of teeth, No 270.
therefore serve as a cotype to define
this species, the most diminutive of
the series. ‘lhe last lower molar of
the right side agrees in form and
measurement with Cope’s type, al-
though the entoconid 2 is less promin-
ent and isolated ; on the left side the
entoconid 2 is nearly obsolete, again
eRe att ee cr os demonstrating the variability of this
Coryphodon lobatus. Am. Mus. cusp. The very small lower canines
Cope Coll., No. 4335.
fr a ‘
(
%
indicate that the animal is a female.
The enlarged second incisors and
general form of m3 confirm its reference to Series II. A unique
feature is the reduplication of the metaconid in m2 and m3 into
a rudimentary metastylid, parallel with the large metastylid of
Uintatherium. Another example of this species is No. 4324.
13. C. latidens Cope.
Type, Nat. Mus. Coll, 4. Lower jaws and teeth, left premaxillary and in-
cisors, superior canine. Loc., New Mexico.
Definition.—Inferior m. and pm.=156. Inferior molars short and broad
with crests nearly or directly transverse (angle=e 85°); entoconid 2 vestigial or
wanting. Inferior incisors equal sized. Superior canines nearly straight,
antero-posteriorly compressed, subtriangular, with an external ridge.
1898. | Osborn, Evolution of the Amblypoda. Part J. 207
This imperfectly-known animal appears to represent a rather
small and specialized form in Series II.
It is distinguished from the type of C. elephantopus by the
straighter and more compressed superior canine observed in the
type, by the transverse position of the crests of the inferior
molars, and by the absence of entoconid 2. As shown in the
Table, p. 199, the measurements of Cope’s C. /atidens type are
identically the same as those of C. seus, although Cope speaks of
the latter as being much smaller than the former.
Cope has suggested the possible association of C. Zatédens with
the cotype skull of C. elephantopus. It appears to be distinguish-
ed, however, by the form and compressed section of the superior
canine. It is, however, certainly related to Series II by the sub-
triangular form of the canine and the characteristic swelling of
the jaw below m3. Unfortunately the types have been tempo-
rarily misplaced, and no determination of this question by direct
comparison can be made at present. If these jaws should prove
to belong to C. elephantopus, the species C. obliguus will have to
be revived.
It will be noted that both types come from New Mexico. In
New Mexico, also true Wasatch, we found in 1897 a lower jaw
(No. 2563, Fig. 16) of extremely small size, associated with
Mentscothertum, Ambloctonus and Didymictis, which may represent
a female of this species. Unfortunately the canines are not pre-
served. The total lower grinding series does not exceed 125 mm.,
so that this is the smallest Coryphodon jaw known; the last
lower molar measures only 30x 19 mm.; the posterior crest forms
an angle of 85° with the long angle of the jaw; a minute vestige
of the entoconid 2 can however be observed.
22. Coryphodon curvicristis Cope.
Type, No. 4326, Am. Mus., Cope Coll. Lower jaw fragments containing
pm.4 to m3; canine.
Definition.—Molar crests transverse. Posterior crest of my directly trans-
verse, crenulate, depressed. Superior incisors with sharply angulate anterior
faces. Canines as in C. Zestis.
The systematic position of this species (Fig. 16) is indetermi-
nate. It resembles C. /atdens in the transverse crest angulation
208 Bulletin American Museum of Natural History. [Nol. X,
and in the rather broad proportions of the molars, but exceeds
this species in size. The complete superior canine determines
the position of the animal in Series II]. The canine is powerful,
curved and antero-posteriorly compressed, partly as the result of
pressure. The inferior premolars are exceptionally short.
A fourth member of this series, C. vevtanus, is found in the
Wind River Beds. It appears to resemble C. /a/idens in the form
of the superior canines.
Sertes I[TT.—SMALLER CORYPHODONS. SPECIALIZED. RELA-
TIVELY NARROW, FLAT-TOPPED SKULLS, (?) WITHOUT PaARIETAL
HorN RUDIMENTS. CANINES COMPRESSED LATERALLY AND
GROOVED ANTERIORLY. LOWER MOLARS ELONGATE, CRESTS
NEARLY OR QUITE TRANSVERSE; M3 BILOBATE, NO ENTO-
CONID 2.
In 1872 Cope defined certain teeth as MZetalophodon armatus,
mistaking the posterior superior molars, m*, of two individuals
for m® and m2 of one individual, as can be proved by a compari-
son with his type of C. molestus. ‘The latter type moreover gives
us the cranial characters and constitutes a valuable cotype.
6. C. (Metalophodon) armatus.
Type, No. 4315, Am. Mus., Cope Coll. Superior m®, m2, premolars, supe-
rior canine and incisors ; two individuals, probably mingled, fully adult.
Cotype, No. 4316, Am. Mus., Cope Coll. Superior m? and m%, mg, pre-
molars, etc., juvenile.
Definition.—Upper and lower canines greatly compressed, with a deep
antero-internal groove upon the upper canine. M® with powerful anterior crest,
my without entoconid 2. Lower molars elongate, crests lunate, nearly trans-
verse. (Angle with long axis of jaw, 81°.)
Synonyms.
g. C. simus Cope. Type, U.S. Nat. Mus. Coll. Inferior m and pm=154.
Fragmentary skull, probably female. Superior canine. Mandibular rami and
teeth. Loc., New Mexico.
1898. | Osborn, Evolution of the Amblypoda. Part TJ. 209
10. C. molestus Cope. Type, U.S. Nat. Mus. Coll. Skull, dentition and
parts of skeleton. Loc., New Mexico.
11. C. domas Cope. Type, U.S. Nat. Mus. Coll. Posterior inferior molar.
Loc., New Mexico.
The slender crests and the elongate form of the posterior lower
molars in this species at once distinguish it as a type from mem-
bers of the foregoing series, since they form an angle of 81°, or
nearly a right angle, with the long axis of the jaw, and m3 is
entirely devoid of the entoconid (Fig. 16).
Specimen No. 4315, Fig. 17, gives
the most distinctive character,
shown again in Cope’s type of C.
molestus (Cope, 1877, Pl. LVI, fig.
4), which agrees with C. armatus,
namely, the flattened form of the
canines. Cope himself referred C.
lomas to C. molestus (1877, p. 237). * F
: . Fig. 25. Coryphodon armatus supe-
The type of C. semus has lower _ rior moiars, left Side (type of C. molestus
teeth of the same character, rather i
long and narrow. The upper
m./ m.2 m3
canines are, however, described by Cope as triangular and
grooved ; this raises a doubt as to the reference of this type to C.
armatus.
The juvenile type specimen of C. molestus demonstrates the
flat-topped character of the skull (Coll. U.S. Nat. Mus. No. 1119,
Cope, op. ci¢., Pl. LVI); the skull is far less expanded laterally,
when seen from above, than any of the skulls in Series IT;
but this may be in part due to its juvenile and undeveloped
condition.
“The inferior canine,” observes Cope, “has a flat interior and
convex exterior face, which are separated by anteriorly and pos-
teriorly directed cutting edges.’”’ The most distinctive feature
of the canines therefore is that the antero-posterior diameter
greatly exceeds the transverse, as in Uvntatherium. An aberrant
feature is the antero-external groove. ‘he median incisors are as
large or larger than the others.
[Way, 1898] 14
IsERTA SEDIS.
23. C. marginatus Cope.
Type, No. 4374, Am. Mus., Cope Coll. Superior molar 3, canine and pm.
Loc., Big Horn, Wyoming.
This indeterminate type resembles C. avmatus in the form of
m#, but differs from it in the form of the canine, which 1s less
compressed and may possibly represent a milk tooth. The canine
corresponds with Cope’s description of that of C. semus.
WIND RIVER TYPES.
Cope’s Wind River material of Coryphodon, all of which is now
in the American Museum (Nos. 4811, fragments of skull and
teeth ; 4812, lower molar, incisors and fragments; 4813, lower
jaw and fragments ; 4814-4817, fragmentary teeth ; 4818), merely
sufficed to determine the existence of this genus in these beds.
Our Wind River collection and the determination of manus
No. 4351 (Am. Mus., Cope Coll.) as belonging to the Wind River
Beds, is therefore of very great importance. It demonstrates that
Coryphodonts of considerable diversity and size persisted into the
Wind River period.
Owing to the general scarcity of fossil remains in these beds,
the relative abundance of these animals cannot be estimated. Of
intermediate size is the jaw of No. 2976, described below as C.
ventanus ; of smaller size there is a well-preserved skull (No. 2977),
type of the new species C. wortmani. ‘They represent respec-
tively the persistence of at least two series, namely of Series I, and
of Series IIl now discovered for the first time.
SUCCESSORS OF SERIES II.
26. Coryphodon ventanus, sp. nov.
Type, No. 2976, Am. Mus. Coll. Jaws and lower teeth. Superior incisors
and canine. L. metacarpal IV.
Definition.—Size of C. testis. Inferior mand pm series=e172. Supe-
rior canines posteriorly compressed, with antero-internal depression and long
1898. | Osborn, Evolution of the Amblypoda. Part J. 211
sharp external ridge. Lower canines with short external ridge near apex.
Second incisors enlarged; lateral incisors much reduced. Posterior inferior
molars with crests more transverse than in C. festis (angle = 74°) a persistent
entoconid 2. ( ? Cuneiform articulating with Mtc. V.)
This species is clearly distinguished from C. ¢estis by the
form of the canines, which in this animal are comparatively
straight and lance-shaped (Fig. 17), the long axis transverse
(unlike U¢ntatherium), with an antero-internal groove which is
Fig. 26. Foot structure of Coryphodon. D, external view of manus of C. ventanus (No.
4351, Coll. Am. Mus.) ; A, superior view of astragalus and calcaneum found near C. simus,
no tibial facet (Bathmodon type); 4, lower surface of astragalus, showing caleaneal and
cuboidal facets ; C, external view of calcaneum and astragalus, showing reduction of tibio-
calcaneal facet. (Coll U.S. Nat. Mus.)
worn away by the lower canine. ‘They resemble those of C. /aé-
dens Cope (except in the groove), but are much less compressed
than those of C. avmatus Cope, besides having the long axis in a
different plane. ‘The posterior crest of mz (Fig. 16) differs from
those of C. armatus and C. s¢mus in form and in the retention of
an entoconid 2, and from that of C. fests in being slightly less
oblique. Another character is the very rapid increase in size of
the molar series as we pass backward : mi=28, m3=42.
To this species belong Nos. 2982, 4813, 4812, 2774, and 2978
of our collection. The latter contains the complete lower teeth
which exhibit the marked disproportion between the second and
the first and third incisors embodied in the definition of this spe-
cies. The incisor proportions are indicated by the length of
MOOS, 1140, 12—=59, 1 3— 28.
212 Bulletin American Museum of Natural History. |Vol. X,
The metacarpal IV agrees in length (54 mm.), and lends some
probability to an association with that of the complete carpus
No. 4351 from the Wind River.’ This associated complete carpus
(Fig. 26) agrees with some specimens of U7ntathertum in the very
exceptional character that the cuneiform articulates with Meta-
carpal V.
Series I.—PRIMITIVE, NARROW-CRESTED SKULLS. CANINES
ROUNDED. INCISORS SUBEQUAL IN SIZE.
The lower teeth are unknown, and the ancestral members of
this series have not thus far been determined in the underlying
Wasatch formation.
27. Coryphodon wortmani, sp. nov.
Type, No. 2977, Am. Mus. Coll. Loc., Wind River, Wyo.
Definition.—Superior m and pm=154. Superior canines rounded. Occiput
very high and narrow. Supratemporal ridges converging posteriorly to form a
comparatively narrow sagittal crest.
Fig. 27. Coryphodon wortmanit, type. Lateral view of skull and section of superior
canine.?
The discovery of this type (Figs. 18 and 27) in the high level of
the Wind River Beds is most surprising. It is far more primitive
both in its narrow cranium and rounded canines than any of the
1 This carpus was mistakenly described by Cope as coming from the Wasatch. Dr. Wort-
man identifies it as found by himself in the Wind River.
2 Dedicated to my colleague Dr. J. L. Wortman.
= te ee
1898. | Osborn, Evolution of the Amblypoda. Part J. 213
C. testis series found in the Middle Wasatch. It appears like a
direct successor of Pantolambda cavirictus.
‘The median incisors are equal to the others, the lateral incisors
being slightly the smallest. The canines have a rounded crown
somewhat flattened in front by wear. The superior grinding
series present a rudimentary posterior crescent on m2 and an oval
m* with short oblique ectoloph. The most unique features are
the form of the occiput and the cranium as defined above, which
is intermediate between that of Pantolambda and Coryphodon
armatus.
The metatarsal V is short and robust (= 42 mm.), with the
characteristic peroneus tuberosity of the true Coryphodon. Both
femora are finely preserved (length, = 340 mm.), being of the
smallest size known.
INSERT#@ SEDIS.
The position of the following types with reference to the Series
I-III, which we have been considering, is uncertain.
4. Coryphodon radians Cope.
Type, No. 4300, Am. Mus., Cope Coll. Superior molars 1, 2 and 3. Prob-
ably associated lower jaw, No. ? 4300. Portions of skeleton. Loc., Evanston,
Wyoming.
Definition.—Third superior molar with a spur (metacrescent) upon the pos-
terior crescent of the ectoloph. Third inferior molar without entoconid 2,
hypolophid nearly transverse. Lower canines somewhat incisiform.
This classic species, which rests upon somewhat uncertainly
associated upper and lower teeth, jaws and skeleton, was the first
described in America. The structure of the last upper molar
is shown in Fig. 15. The last lower molar has crests nearly as
transverse and simple as in C. Zatidens. ‘The most distinctive
structure is the lower canine which, although badly broken,
exhibits a distinct flare at the base of the inner face, as in the
incisors, and is apparently becoming incisiform, an interesting
approach to Uintatherium.
214 Bulletin American Museum of Natural History. {|Vol. X,
15. Coryphodon hamatus Warsh.
Type, Yale Museum No. 1330. Skull and dentition much worn.
Cotype, Yale Museum No. 1334. Female skull with perfect superior and in-
ferior dentition. Loc., Evanston, Wyoming.
Synonym.
18. Coryphodon(Manteodon) subquadratus Cope. Type, No. 4340, Am. Mus.,
Cope Coll. Superior molar 2, incisors and fragmentary premolar. Loc., Big
Horn, Wyoming.
Definition.—Size large. Superior molars with quadrate crowns and well
developed hypocones upon mi and m2. Inferior molars with nearly trans-
verse crests; mg without entoconid 2.
This species was mistakenly associated with C. elephantopus
by Earle. In size it equals C. ¢es¢’s, but it 1s well distinguished by
the quadrate form of the superior molar teeth in which, according
to the figures of Marsh (Dinocerata, Fig. 55, p. 52), a representative
of the hypocone is present. This is developed from the ridge
extending backwards from the protocone. In the inferior molar
teeth the crests are nearly transverse, and there is no trace of the
entoconid 2.
The unique quadrate tooth with a prominent hypocone, type of
Manteodon subquadratus (Fig. 15), was without reason considered
by Cope as a third superior molar. It proves, upon comparison
with Marsh’s cotype made by Dr. Matthew, to resemble a second
superior molar of C. hamatus. It differs, however, from C. hama-
tus in the more distinct development of the posterior spur of the
metacone crescent, a character which may subsequently prove
to give it distinct specific rank.
The type skull of C. Aamatus is somewhat fractured. The top
of the skull of the cotype, a female, is considerably narrower than
that of C. fests, female, presenting a condition intermediate
between that of C. ¢estis and C. armatus. ‘The canines in this
animal, as in other females, are small.
28. Coryphodon singularis, sp. nov.
Zype, A hind limb, tibia, fibula and pes No. 2980. Loc., Wind River, Wyo.
A small and unique hind foot and limb from the Wind
River Beds, found upon the level of C. wortmant, is of excep-
1898. | Osborn, Evolution of the Amblypoda. Part J. 215
tional interest (Fig. 28). Associated lower tooth fragments, put
together by Dr. Matthew, resemble those of a small Coryphodon,
and clearly separate this animal from Sathyopsts. The differ-
ences from the pes of Coryphodon are very significant, as follows :
1. Navicular laterally reduced, excluded from cuboid by ectocuneiform, a
unique condition.
2. Ectocuneiform enlarged, articulating with astragalus (unique).
3. Second or middle phalanges greatly abbreviated upon all digits, I-V,
as in Uintatherium manus.
4. Front surface of astragalus widened, separating tibial and navicular facets
asin Uintatherium.
5. Tibia long and slender, unlike Coryphodon.
The measurements of the metatarsals are as follows:
Nits el —> Oe VtS ss elf oe Mts ease Mts ViR—42s VES Vie==34 0
Other measurements in Table on page Igg.
Fig. 28. Coryphodon singularis. Superior and lateral views of pes. Am. Mus. Coll.
No. 2080.
This animal thus shows one progressive character (4), two
entirely unique and distinctive characters (1, 2); the latter,
together with (5), sharply separate it from Coryphodon ; two char-
acters, 3, 4, parallel or approach Usntatherium. The other Wind
River species, C. wortmant and C. ventanus, are distinguished
from this by their typical metapodials, one of which is known
in each type.
The associated femur (No. 2970) is proportioned like the tibia,
long and slender.
216 Bulletin American Museum of Natural History. |Vol. X,
Prophetic of this type, perhaps, is the pes of Pantolambda
cavirictus (Fig. 12), in which the navicular is reduced upon the
outer side and the ectocuneiform is elongated so as to nearly
come in contact with the astragalus.
Foot STRUCTURE.
Cope (1884, 1, p. 1120) proposed the theoretical groups, P/aty-
arthra (with flat astragalus) and Amblypoda hyodonta (astragalus
without a neck) from which to derive the Amblypoda. Both groups
are superfluous now that it is clear that the ancestral Amblypoda
can be derived directly from the Creodonta, all of which possess
an astragalar neck.
Planes EER et
oie es ‘ Re Sarees Caan eeey eee
Planes of
_ Astragalo-
Navicular
Us (BL Facets.
Fig. 29. Angles formed by tibio-astragalar astragalo-navicular facets, to exhibit widening
of front face of astragalus. Ur, Ursus; P. Pantolambda,; C, Coryphodon testis; Ut, Uinta-
therium ; E, Elephas.
The transition is simple. By shortening of the neck of the astra-
galus (Fig. 29 P. and C. and W.) the tibio-astragalar facet is
gradually brought almost into confluence anteriorly with the astra-
galo-navicular facet, as in C. radians. In C. lobatus and C. sin-
gularis this space widens as in Uzntatherium.
1. The variables in these feet are the astragalar foramen and
the “biale facet. From our present knowledge both these struc-
tures (inherited in Coryphodon from Pantolambda) are useless or
vestigial, inconstantly developed and therefore not constant specific
characters.
In Fig. 26 (identical with Coryphodon III, Cope, 1877, Pl. 60),
a small astragalus and calcaneum is shown which lacks both astra-
galar foramen and tibiale facet. In C. lobatus (No. 4335, type of
C. pachypus) there is a large tibiale facet, while the astragalar
1808. | Osborn, Evolution of the Amblypoda. Part J. 217
foramen is not even grooved. In No. 2870 the tibiale facet is
irregular, and a groove represents the astragalar foramen. In
C. testis, No. 258, the ibiale facet is irregular, the astragalar
foramen is wanting; in No. 2869 it is completely bridged over; in
No. 4300 (Cope’s cotype) it is partly bridged over.
2. In therelative constancy of the tibiale facet and of the astra-
galar foramen or groove, the pes of Uintatherium mirabile is there-
fore more primitive than that of Coryphodon.
CONCLUSION OF PART I.
The phylogenetic conclusions drawn from this analysis of the
Taligrada and Pantodonta will be more fully discussed at the close
of Part II of this paper, which will treat of the Dinocerata.
The two main results thus far brought out are these: First, the
demonstration of a number of separate phyletic lines of Coryph-
odons; these lines probably represent the local differentiations
of the Coryphodon type in adaptation to different feeding ranges,
that is, swamp, plain, and upland. ‘The second result is, that
certain Coryphodons approach the Dinocerata in some structures
as Closely as they depart widely from them in others; for example,
C.armatus resembles Uintatherium in canine type, but differs from
it in skull type; C. ¢estis approaches Uintatherium in the upper
posterior portion of the skull, but differs from it widely in the
anterior portion of the skull, and in the structure of the canine
teeth ; C. radians shows the assumption of the incisiform shape
by the lower canines, so distinctive of Usntathertum. But no
Coryphodon is fully known which fills all the conditions of an
ancestor of Uzntathertum. Until the skull of Bathyopszs is known
the transition between the above types will remain obscure.
BIBLIOGRAPHY—PRINCIPAL REFERENCES.
Corr, E. D. 1877.—Report upon the Extinct Vertebrata obtained in New
Mexico by parties of the Expedition of 1874. U.S. Geol.
Sury. west of tooth Mer. Palzontology, Vol. IV, Part ii.
y 1884.—The Vertebrata of the Tertiary Formations of the West.
U.S. Geol. Surv., Vol. III, Part i, 1884.
oe 1884 (1).—The Amblypoda. American Naturalist, Nov. & Dec.,
1884, pp. I110-1121 and pp. 1192-1202. Jan., 1885, pp. 40-55.
Adie ts PI Dip ee a OTN A ASOT ath gh SLA Ba”, on te Mee
rt . Acne ae ee
218 Bulletin American Museum of Natural History. |Vol. X.]
EARLE, CHAS. 1892.—Revision of the Species of Coryphodon. Bull. Am.
Mus. Nat. Hist., Vol. 1V, 1892, pp. 149-166.
MarsH, O. C. 1876.—On some of the Characters of the genus Coryphodon,
Owen. Am. Jour. Sci. (3), Vol. XI, pp. 425-428, 1 pl.
1877 (1).—Brain of Coryphodon. American Naturalist, Vol.
a5 (De S775
(2).—Principal Characters of the Coryphodontidz. Amer.
Jour. Sci. (3), Vol. XIV, pp. 81-85, pl. iv.
(6).—Introduction and Succession of Vertebrate Life in Amer-
ica. (Vice-President’s Address before the American Associa-
tion for the Advancement of Science, Nashville (Tenn.)
meeting, Ate. 30.1577.) roc. Assoc., Vol. XOXVileapp:
211-258, 1 pl. 1878.
1884 (g).—Dinocerata: A Monograph of an Extinct Order of
Gigantic Mammals. With 56 plates and 200 woodcuts.
Washington, 4to, XVIII, pp. 237. Author’s edition.
1893 (7).—Restoration of Coryphodon. Amer. Jour. Sci. (3),
Vol. XLVI, pp. 321-326, plates v and vi. New Haven,
Oct 1803:
OsporNn, H. F. 1893 (4).—Rise of the Mammalia in North America. (Vice-
Pres. Add. Amer. Assoc. Ady. Sci., Sec. Zool., Aug. 16,
1893.) Amer. Jour. Sci,, Nov. and Dec., 1893.
(5).—Fossil Mammals of the Upper Cretaceous. Bull. Am.
Mus. Nat. Hist., Vol. V, p, 15, Dec. 20, 1893.
1895 (1).—Fossil Mammals of the Puerco Beds, Collection of
1892. Bull. Am. Mus. Nat. Hist., Vol: VII, Art. 1; pp:
1-70, Mch. 8, 1895. (With Charles Earle.)
1898.—A complete Skeleton of Coryphodon radians. Notes
upon the locomotion of the animal. Bull. Am. Mus. Nat.
Hist., Vol. X, April 4, 1898, pp. 81-91.
afte) | Ur -
tional (
By ‘Hew1 y Pamerenp Ospony..
=
y
Article XII.—ADDITIONAL CHARACTERS OF THE
GREAT HERBIVOROUS DINOSAUR CAMARASAU-
RUS.
By Henry FAIRFIELD OSBORN.
WITH THIRTEEN FIGURES IN TEXT.
This gigantic reptile was found in the famous Como Bluffs of
Wyoming by Dr. J. L. Wortman, of the Museum party of 1897,
and Prof. Wilbur C. Knight, of the University of Wyoming. The
Museum number is R. 222.) The bones include the left ilium, the
ischia and pubes of both sides, the right and left femora, the left
tibia and astragalus, the right scapula and coracoid, two shattered
cervicals, two complete dorsal vertebrae, two incomplete dorsals,
three or four incomplete ribs, coalesced spines of three sacral
vertebree and one sacral centrum, twenty caudals and twelve
chevron bones. With the exceptions stated, the bones are in a~
remarkable state of preservation, having been worked out with
exceptional skill by Mr. Granger and others, under the direction
of Mr. Hermann.
This is a large individual. The identification is provisional.
The measurements, in comparison with those taken from the
type of B. excelsus Marsh, are as follows:
Brontosaurus American Museum
excelsus. No. 222.
WenothrotitemUts. arc cies ielcy sere ers 11eF= Sy iiés (6) ial, Slit 1) abot
VB PREL DIAM Ve isie, «eke Mena nine ai te vans tes G3 titeg | (Oia Binley © teiyat
pat take PRULDIS the avo afatticas. s oheltvers: sy oye) Seer Brit. Lovin: Bt. LOM,
pomiear ISCIIUMce bs ceil cys ics sa 3} ie fey tal 3 ft. gl in
uenamS CAP UIA areastfarsrert siscercis sells ais). tities 2) tole Sey Gabe
peru iaee COLACOIC naan niaiteremmie teats) «(0h -Fe 2 ft. 216 in. 2 Ties! Feb ol
The new points of greatest importance are :
First—The discovery of the hitherto unknown characters of the
anterior caudal vertebre.
Second.—TYhe apparent resemblances of Camarasaurus Cope
to Amphicaelias' Cope, to Brontosaurus Marsh, Atlantosaurus
Marsh, and Afpatosaurus Marsh.
1This has been anticipated by Marsh (Am. Jour. Sci., Aug., 1881) in his first classification
of the Dinosauria, in which Amphicwlias is bracketed after Camarasaurus, and placed in
the Atlantosauride. In the final classification of the Dinosauria, however (* The Dinosaurs
of North America,’ p. 241), Caszarasaurus and A mphicelias are removed to the Morosauridz.
[219]
220 Bulletin American Museum of Natural History. |Vol. X,
Third.—The observation of structural analogy to certain stru-
thious birds in the anterior dorsals and posterior cervicals.
1. HABITS AND SIZE OF THE ANIMAL IN RELATION TO ITS
STRUCTURE.
The estimate given by Marsh of the total length of this animal
is nearly or quite 60 feet; the tail is figured at about 24 feet.
Since the vertebra believed by Marsh to be the third caudal is
probably the roth or rrth, the tail should be increased to over 30
feet in length, by the addition of at least seven large anterior
caudals. The total number of caudals is estimated at 4o as
against 37 in Diplodocus. Marsh has attributed to Brontosaurus 27
precaudal vertebrze, or 13 cervicals and 14 dorso-lumbars. From
reasons given below it is probable that there was a larger number
of dorso-lumbars, which would still further increase the length of
the animal to considerably over 60 feet.
We can only conceive of the Camarasaur as a great wading and
swimming quadruped, enjoying a habitat similar to that of the
Upper St. John River, Florida, at the present time, namely, a
relatively firm bottom gently graded to all depths, supporting a
richly luxuriant aquatic vegetation, the river banks bordered by
sloping shallows of sand (Colorado, Cafion City Beds) or clays
(Wyoming, Como Beds). As imagined by Cope in his picture of
Amphicelias (‘Century Magazine,’ November, 1887), the animal
could walk along the bottom, raising the anterior portion of its
body. We believe also that it could swim rapidly, propelled by
its light but long and powerful tail, which would be useless
upon land. The abundance of cartilage around all the limb
joints and the non-osseous nature of many of the carpals and
tarsals afford positive evidence that the limbs were not con-
tinuously subjected to the hard impact of the enormous weight
of the body by motion on land. Feeding was done in the
water and along the shores. Excursions upon shore were there-
fore like those of the Alligator, mainly for breeding and egg-
laying purposes, and they exposed the animal to attack by the
Megalosaurs. By means of powerful mid- and posterior-dorsal
spines and opisthoccelous vertebre, the entire anterior part of the
1898.| Osborn, Additional Characters of Camarasaurus. 221
body, whzle in the water, could be raised or lowered with the
great acetabulum acting as a fulcrum, thus presenting an analogy
to the Hadrosaurs, which exerted a similar movement fon land.
The long neck, similar in structure and almost as flexible as that
of an Emeu (Dromeus), could thus pass through a prodigious are
in the search for food either under or above water. The neck
motion apparently involved the anterior non-spine-bearing dorsals
Fig 1. Droma@us. Cervicals 13 and 14; dorsals1 and 2, showing absence of median spines.
Dorsal 3, showing large blunt median spine, resembling that of the Camarasaurus dorsal,
Fig. rr of this Bulletin, Am. Mus. Coll., No. 607.
as in Dromeus, behind which the comparatively inflexible large
spine-bearing dorsals rose to maximum height in the sacrum for
the insertion of the ligamentum nuche and elevator muscles.
The importance of such an hypothesis of function will appear
in the following description and discussion, and it applies to all
the Cetiosauria, namely, to the A7orosaurus and Diplodocus types as
well, which so far as known are wniform with the Camarasaur type
in the peculiar bird-like structure of the posterior cervicals and
anterior dorsals and in the possession of a very powerful swim-
ming tail.
Ten of the caudals in our specimen afford an interesting illus-
tration of the cause of the distribution of these large skeletons
222 Bulletin American Museum of Natural History. |Vol. X,
over a considerable surface. The dorsal spines, and in some
cases the sides of the centra are found to be deeply gashed
with the sharp teeth of a carnivorous Dinosaur. The upper
portion of several spines, in fact, is entirely bitten away, the upper
surface containing jagged transverse grooves, which prove that the
carnivore was of great power, and applied its pointed teeth with
strong effect to the gristle and muscles upon the sides of this
prodigious tail.
TasLE J.—MEASUREMENTS OF VERTEBRE.
] |
= ; a a bus Lad 7 o
ates 2 “e “2 | ae | oe a 8 oe
S) (S) S) QO |o |e H ea}
| | |
Anterior dorsal. i || BICC) 310 | 1050 470 | e864
Posterior dorsal.; ? | 368 BAOm |e SOmaneecr | e715
First caudal....) 162 383 BS 5 ll ALO Sih tear ste etersncu ees 957
Geel CanCkiIljsseao0\) WOE Noodoor CEH lls saogo|lbau céallqaodoulldoce or <3&4
e 5th caudal.... .| 168 | 360 | - 32z BOD alee. [eae e695 | <6
ercthucaudalie sc.) 9677 || B40) mgoen| meso ulesc: .: eee 615 ai
emthecandalyeee-- TOW esss 300 7@ Sullierrsveve|| eer 622 3
emeth caudal eee.) i710 305 274 FiO he lcte veh iieeietene 587 agers
Sigil Cell, 565.) uO 283 260 OBB est: Hiatere east 537
edAthcaudall sees 073 282 258 BO Sial ences <i eae BORN
eunotheeaticalener 180 272 248 BASE Mcgovern, 2. 2/lsteue wees e444 1
ex7th caudal... .| 180 256 228 B20 Me are, ces ES ereverne €395
e 20th caudal? .... 185 | 220 205 bk aa (aoe See <18&19
eP2stecatidalaene|i 1iQ2ll2rO 194 igetd |eateeae 8 172
e2od caudal?-.-..| 200 202 186 iy Meaeioe eeeisccret Te 7
em sducaudaleanncr| saloon Lod 176 8 ecencicetc | PE heareene
é 24th caudal?....| 205 184 175 Rem get 2 Ml scott ast meee eee
eriiacaudal? 4 5\ i203 hi) 7 il erOOm ee ene ll ne a ist,|| foe. eee <25&26
e 28th caudal’ 24 199s) DzOy|\ Ese ule een eae | ecece Siecle
e 30th caudal?....) 200 | 152 TA G2 la sees ct Ee A Renee <29
erg lStmcaticdalenee 203 less |, UX) IN, ogg Be (er P| Sua a a
egoth (caudal? 2), 85 |) noi || eerie rare Jocreee | pleroen leer S32&35
eido total ass oe leet elon [eee apeme lee te | Eiht tone well ote siege <37
Nove.—Many interesting facts are brought out by the exact measurements of the caudals
given in this Table. The centra decrease steadily in their transverse and vertical diameters,
but they increase steadily in the longitudinal diameter as far back as the 26th caudal, then
begins a decrease. Thus the estimated 31st caudal measures 203 mm.; the 36th caudal, meas-
uring 185 mm., probably marks the natural decrease towards the tip of the tail. A constant
ratio of decrease, however, characterized the dorsal spines.
1 Measurements agree approximately with supposed 4th caudal, Marsh.
2 Summits of spines bitten off and grooved by large cutting teeth.
1898.| Osborn, Additional Characters of Camarasaurus. 223
2. DESCRIPTION OF SKELETON.
The mid-dorsal (Fig. 2) corresponds closely with the supposed
‘posterior-dorsal,’ described by Marsh as belonging to rontosaurus
excelsus (Fig. 12). With the exception of the anterior portion of
the centrum and the extremities of the tubercular processes, this
vertebra is in a remarkable state of preservation, the neural spine
M DORSA»
PB A
Fig 2. Camarasaurus. Mid-dorsal. A, anterior view. /, posterior view. Am. Mus.
Coll. R. 222. Approximately jg; natural size.
being complete upon the anterior face, and lacking only the upper
and lateral portions. It displays the characteristic hyposphen,
the complex laminated and excavated condition of the spine, and
a pair of small cavities upon the sides of the centra.
The true posterior dorsal (Fig. 3) is a superb vertebra, in
nearly perfect preservation, lacking only the anterior portion of
the tip of the spine, parts of the extremities of the tubercular
processes, and the anterior face of the centrum. It corresponds
closely with Cope’s type of Amphicalias altus (Fig. 13) both in
form and measurement, and in the lateral cavities of the centra.
224 Bulletin American Museum of Natural History. |Vol. X,
It is possible, however, that Cope’s type represents a Diplodocus.
The mid- and posterior-dorsal vertebree are so different that
Cope supposed that they represented different genera, viz., Cam-
arasaurus and Amphicelias.
P. DORSAL,
Fig. 3. Camarasaurus. Estimated 5th caudal. Posterior dorsal vertebra. Am. Mus.
Coll. R. 222. Approximately #5 natural size.
The sacrum is represented in this specimen by a single large
Sree sacral vertebra, and by the coalesced spines of ¢éree sacrals.
The significance of these facts will be pointed out later.
The tail of Camarasaurus includes a far more powerful and
complex series of vertebree than has been indicated in the figures
and descriptions of Marsh.
Immediately behind the sacrum, the first caudal of Camara-
saurus (Fig. 4) is an entirely unique vertebra, extending laterally
into two great transverse plates, which may have come in contact
at their extremities with the posterior portions of the ilia. These
plates are concave anteriorly. The pre- and post-zygapophyses
1898.] Osborn, Additional Characters of Camarasaurus. 225
are small and obliquely inclined inwards ; the anterior face of the
spine has a prominent rugose crest, similar to the crest upon the
posterior face of the spine, but the latter (posterior crest) sinks
between two prominent laminz, which extend downward to
support the post-zygapophyses, /z.
Tz
Fig..4. Camarasaurus. First caudal. A, anterior view. /P, posterior view. Am. Mus.
Coll. R. 222. Approximately ;5 natural size.
Behind this highly specialized vertebra, three vertebrae were
probably intercalated, only one of which is preserved in our
collection, giving us the transition to the vertebra estimated as the
5th caudal (Fig. 3).
This supposed 5th caudal (Figs. 3,5) exhibits a marked reduction
of the spine as well as of the transverse lamina; the posterior
surface of this lamina being excavated by two deep depressions.
This vertebra was followed probably by two, which are missing,
so that the fourth preserved in our collection is estimated as the
8th caudal (Fig. 5); in this the transverse lamina is still more
reduced, and the lateral pockets upon the posterior face are
wanting. Behind this is the supposed gth caudal, which
[ June, 1898. | US
226 Bulletin American Museum of Natural History. [Vol. X,
‘
approaches in its structure that which Marsh has figured as the
4th, but is somewhat more complex. Here follows another
interval (Fig. 5), in which would lie the vertebrz described by
Marsh as the 4th. (Fig. 6.)
The estimated rath, 13th and 14th succeed each other with a
regular diminution in size, and a steady reduction of the trans-
TR-
atl CAUDAL 5t4 CAUDAL sth, cAuDAl, 9% canDAL. 12h caupat
Fig. 5. Camarasaurus. Anterior caudals, First caudal and sth,
8th, oth, r2th caudals as estimated. Am. Mus. Coll. R. 222. Approxi-
mately ;'; natural size.
verse processes, which gradually transforms into
a horizontal plate. The next vertebra preserved
is the estimated 16th, which exhibits a very
marked reduction in the transverse process, and
this disappears entirely in the supposed z2oth and
21st, in front of which at least two vertebre are
missing. (See also Note 1, p. 233.)
: The missing 16th and 17th vertebrae were
Fig. 6. Brontosau- probably detached in the manner described
rus excelsus. Sup-
posed 4th caudal ver- above, and the 16 or 20 vertebre at the end of
febyanl Miter Marche : P
1897, Platexxiv. Ap. the tail were also drawn away. In front of the
proximately 4; nat-
aaa 15th no tooth marks are observed.
_
1898.| Osborn, Additional Characters of Camarasaurus. 227
3. HisroricaL NOTES.
Large Cetiosauria have been named from these and similar
beds! in Colorado, as follows :
No. 1. Tttanosaurus montanus Marsh.—Am. Jour. Sc., July, 1877, p. 87.
— Golden, Colorado.
No. 2. Camarasaurus supremus Cope.—Pal. Bull. 25, Aug. 23, 1877, Proc.
Am. Phil. Soc.—Cafion City, Col.
No. 3. Caulodon diversidens? Cope.—Pal. Bull. 26, p. 193.— Cafion City,
Col.
No. 4. Tichosteus lucasanus? Cope.—Pal. Bull. 23, p. 194, Nov. 21, 1877.
Proc. Am. Phil. Soc.—Cafion City, Col.
No. 5. Atlantosaurus (Titanosaurus) montanus Marsh.—Am. Jour. Sc.,
Dec., 1877, p. 514.—Colorado:
No. 6. Apatosaurus ajax Marsh.—Am. Jour. Sc., Dec., 1877.—Colorado.
No. 7. Apatosaurus grandis Marsh.—Am. Jour. Sc., Dec., 1877, p. 515.—
Colorado.
No. 8. Amphicelas altus Cope.—Pal. Bull. 26, Dec. 10, 1877. Proc. Am.
Phil. Soc.—Cajion City, Col.
No. 9. Amphicelias latus Cope.—Pal. Bull. 26, Dec. 10, 1877. Proc. Am.
Phil. Soc.-—Caiion City, Col.
No. 10. Symphyrophus musculosus? Cope.——Pal. Bull. 26, p. 246, Jan. 12,
1898.—-Caiion City, Col.
No. 11. Adlantosaurus tmmanis Marsh.—Am. Jour. Sc., March, 1878, p.
241.-—Colorado.
No. 12. Brontosaurus excelsus Marsh.—Am. Jour. Sc., Dec., 1879.——
Como, Wyoming.
The rapid and profound transformations in the characters of
the vertebree in different parts of the back bone give these animals
an entirely unique position among reptiles, and largely explain the
diversities of view as well as the numerous generic references, held
by different authors, as indicated in the following brief résumé :
(1) The first notice of these animals in American literature was by Marsh,?*
under the name of 7v¢anosaurus montanus. This generic name had been
shortly before employed by Lydekker.+ (2) The second notice was by Cope,’
in which the name Camarasaurus was proposed. The species Camarasaurus
Supremus was based upon a number of dorsal vertebrae from the supposed
Dakota Beds of Colorado, near Cafion City. In 1878° these remains were
1 These beds are equivalent to the Wealden of England, from which many large Dino-
saurs have been described, of the same order (Cetiosauria) and possibly representing the
same genera.
* Never fully characterized or figured.
3 Notice of a new and Gigantic Dinosaur, Am. Journ. Sc., July, 1877, p. 88.
4 Rec. Geol. Surv. Ind., Vol. IV. p. 38, 1877.
5 Ona Gigantic Saurian from the Dakota Fpoch of Colorado, Pal. Bull., XXVI, Pub-
lished Aug. 23, 1877.
6 On the Saurians recently discovered in the Dakota Beds, etc.—Am. Nat., Feb., 1878,
De 7ie
228 Bulletin American Museum of Natural History. |Vol. X,
figured. It now appears that the anterior dorsal vertebra (Fig. 2, p. 73, of
Cope’s article), characterized like the cervicals, by the absence of the median
spine, was correctly determined by Cope. In this vertebra the hyposphen is
less accented than in the vertebra figured upon the following page (Fig. 5, p.
75), in which the hyposphen is sharply defined, and there isalso a median dorsal
spine. (5) The fifth notice was by Marsh,! in which the name 4//antosaurus
Fig. 7. SACRA OF CETIOSAURIA.
A, Apatosaurus ajax. ‘Three coalesced sacrals. After Marsh, 1897, Plate xvii.
B, Atlantosaurus montanus. Three coalesced sacrals and portion of a fourth. After
Marsh, 1897, Plate xvii.
C, Brontosaurus excelsus. Five coalesced sacrals. After Marsh, 1897, Plate xxiii. All
figures approximately 35 natural size.
was proposed to replace 77/anosaurus. (6) In the same paper the new genus
Apatosaurus ajax, was distinguished by the opisthoccelous cervicals with hollow
centra ; anterior dorsalis with similar characters; posterior lumbars with flat
articular faces ; sacral vertebree more solid ; anterior caudals biconcave ; chev-
rons not united above. (7) In the same paper the species 4. grandis is
described with the following measurements :
ILyerakeqiln CFE MEM G5 556005 2 aeng0a04 AAO IoOSaUd Tasesag WOO ire,
W@ransverse diameter of proximaliends= 6-25 = slits 340 mm.
(8) The eighth notice by Cope was based upon characters of the posterior dorsal
vertebree. The vertebra (Fig. 13, p. 80) with a very prominent dorsal spine, re-
' Notice of new Dinosaurian Reptiles from the Jurassic Formation. Am. Jour. Sc.,
Dec., 1877, p. 514.
1898.] Osborn, Additional Characters of Camarasaurus. 229
ferred to Amphicelias altus, agrees closely with that here shown to belong to
Camarasaurus. (12) In the twelfth notice (Marsh, 1879) a new type, Bronto-
saurus excelsus, was defined. Characters: Sacrum composed of five thoroughly
coéssified vertebree ; in other respects resembling A/orosaurus from the Como
Beds of Wyoming. The succeeding notices of these animals are fully embraced
in Marsh’s Memoir, ‘‘ The Dinosaurs of North America,” published in 1897.
Here (pp. 166-241) the order Sauropoda and family Atlantosauridze are described,
the latter, however, not including the genus Camarasaurus, which is placed in
the Morosauridz (page 241).
COMPARATIVE MEASUREMENTS OF Limps, ARCHES, VERTEBRA,
é < 3 & bea nye
Res [seg RS) Soe al SY ao
Seb ieess] S ale Vie g
SSN 8 8oe| S88] SoS] Fes
SSE [8 °4q) Sse; SPE] S23
dotallenathiot¢scapulay es. isc. ts 1517 | 159 a. ee 1600
Greatest diameter of coracoid......... .| 690 736 672
/NMISRCO OOS Ke OKs, wbiblinl 55 ESqcucondqce us 1373 fi - Ae
Wotalengthvoftischitim.. <1 2 sa.45..: Ee 1150 whe ; 1118
7 is PWNS 3565 howesodee cs cs 1168 | 1060 ab 1168
He oh FEMA Siero sicherstnisyses sous TS2ON eh 70M kOZO™| LOSOul e750
Height of ant-dorsal spine............. 830 | 1050 oe 948
oe poOst-dorsalispines a. e reer ae 1280 | II00 an 5g
Length, centrum ant-dorsal ....... See 275 ws Ps ae 276
Mrediankcatidalltrvcny ciaccsvacsacaiscre cue
It is @ priort im-
probable that so
many different gen-
era of gigantic Sau-
rians of similar size
co-existed, It is
against the princi-
ples of evolution
that closely similar
types of equal size
should occupy the
: iy Fig. 8. Brontosaurus excelsus. Posterior cervical vertebra.
same territory at the After Marsh, 1807, Plate xxi. Approximately ,’g natural size.
1 These measurements are given in Cope’s second paper upon Camarasaurus. (Am.
Phil. Soc., Dec., 1877, pp. 242-245.)
2 These measurements are approximate, as no exact measures have been published.
3 Cope gave the length of this femur as 6 feet 4 inches (=1930), and described it as longer
than that of Camarasaurus, but in his metric table it is by misprint assigned 1524 mm,
same time. It appears moreover to the writer that the evidence
which has been brought forward to demonstrate such an excep-
tional condition is inadequate and is capable of different inter-
pretation, as follows :
Amphicelias having been shown to be in all probability rela-
ted to Camarasaurus, the only absolute basis of separation of
the remaining types is the number of sacral vertebrz, as in the
definitions of Marsh and Cope :
Afpatosaurus. Camarasaurus. Atlantosaurus. Brontosaurus.
3 sacrals. 4 sacrals. 4 sacrals. 5 sacrals.
This definition does not separate At/antosaurus from Camarasau-
rus ; the type sacrum of Afantosaurus (Fig. 7B) is, however,
fractured anteriorly, and this animal may therefore have pos-
sessed 5 sacrals, like
Brontosaurus (Fig. 7C).
The three coalesced
sacral spines and the
single free sacral cen-
trum in our specimen
suggest the following
hypothesis, namely,
that Camarasaurus had
five sacral vertebra ;
three of these constantly
coalesced both by centra
and neural spines, two
Fig. 9. Afatosaurus ajax. Dorsal vertebra. After others coalesced less con-
Marsh, 1897, Plate xviii. Approximately 7 natural size.
stantly and possessed free
spines.
Such difference in the growth and degree of coalescence is shown
to be probable (1) by Diplodocus, in which at least three vertebre
coalesce by centra but only two coalesce by spines; (2) by our
specimen, No. 222, in which (closely resembling Pronfosaurus in
every other respect) one centrum is free and three spines are
coalesced ; (3) by the type of #. exce/sus, in which three spines
are coalesced, as figured. (See also Note 2, p. 233.)
The removal to a distance of one or both of the freer sacral
vertebre is quite explained by the fact that these skeletons are
; ‘
ee
1898.| Osborn, Additional Characters of Camarasaurus. 231
in the great majority of cases very much disturbed and confused
before embedding. An instance in point is the loss of all the
anterior caudals in Marsh’s otherwise excellent &. excelsus
specimen.
Let us suppose
that the three early
united vertebrae (of
the more primitive
Cetiosaur sacrum)
were found alone,
the others having
been detached and
lost, we would then
have the genus 4pa-
tosaurus ; if one of
the freer vertebrze
had united we Fig. 10. Camarasaurus supremus type. Anterior dorsal
vertebra. After Cope. Approximately 7 natural size. Com-
would have the ge- pare Morosaurus, Marsh, 1807, fig. 31, p. 181.
nus Adantosaurus
or Camarasaurus ; if both of the freer vertebra had united we
would have Brontosaurus.
4. ANTERIOR DORSAL VERTEBR# OF THE CETIOSAURS ARE OF
AVIAN TYPE.
Anterior dorsals without median spines have been described
by Cope in Camarasaurus (Fig. 10) and by Marsh in Apatosaurus
(Fig. 9), Worosaurus and Diplodocus. This condition seems so
general as to constitute almost an ordinal character of the Cetio-
saurs.
These vertebra are, moreover, remarkable in resembling those
of certain Struthious birds such as Dyromeus (Fig. 1), in the
absence of median spines, in the elevation of lateral spinous
processes above the zygapophyses, and in the abrupt development
of a median spine upon the third or fourth-dorsal.
It is also not improbable that the abrupt transition from verte-
bree without median spines to a vertebra with a strong median
spine (Dromeus, D.3) is paralleled in the blunt spine of the
Camarasaurus dorsal (Fig. 11), which may well represent D 3 or
232 Bulletin American Museum of Natural History. |Vol. X,
Fig. 11. Camarasaurus supremus.
Anterior dorsal. Supposed to be the
first possessing a median spine. After
Cope. Approximately j; natural size.
Fig. 12.
D4. Behind this would appear
the transversely expanded mid-
dorsal spines of the Brontosaurus’
(Fig. 12) type, and of our speci-
men, No. 222 (Fig. 2). Behind
this again would appear the pos-
terior dorsal of the Amphiccelias
type (Fig. 13) or the still more
posterior dorsal of our specimen,
No. 222 (Fig. 3).
The difference between these
three types of vertebre (Figs. 9,
12 and 3) is so profound that it
seems hardly possible that they
could be compressed within the
limits of 14 dorso-lumbars—the
number which Marsh has figured
Fig. 12. Brontosaurus excelsus. Dorsal vertebra. After Marsh, 1897, Plate xxi. Approxi-
mately js natural size.
Fig. 13. Posterior dorsal of Amfhicewlias altus, type. After Cope. Approximately 5
natural size.
1 This is mistakenly determined by Marsh (‘Dinosaurs of North America,’ p. 169) as a
‘posterior dorsal.’
1898.] Osborn, Additional Characters of Camarasaurus. 233
in Brontosaurus. ‘There were, therefore, in all probability, more
dorso-lumbars than 14. Cope has assigned 20 dorso-lumbars to
Camarasaurus.
So far as habit and function are involved, the transition from
the lateral spined to the median spined dorsals, as in Dromeus,
probably indicates that Camarasaurus had a comparatively free
anterior dorsal region, consisting of two or more vertebra, which
bore short ribs and moved in all directions with the neck.
The differences in proportion, in length of limb and in other
parts subject to age, sex and individual variation noted in these
different specimens are undoubtedly superseded by genuine spe-
cific differences which mark the Camarasaurs of different geo-
logical levels, also by generic differences, which, however, are still
to be positively determined by more careful and thorough explora-
tion, and by comparison of the type specimens. ‘The generic
name Camarasaurus is therefore provisionally employed here.
NOTE 1.—CENTRA OF CAUDALS.
There is also great diversity in the caudal centra.
The first caudal centrum is dzconvex.
The second to the estimated seventeenth caudal centra are procelous.
The estimated twentieth caudal centrum and all behind it are amphicalous.
NOTE 2.—SIZE SACRUM.
The four sacrals of Camarasaurus exhibit exactly the same measurement as
the four anterior sacrals of Brontosaurus.
ne of the oie resh-
e West.
By W.. De Marruew.
ty Re Be ; ia, , or : : ; AEX i
evict Driseunr oe Batural History,
yh. ‘XI Aenete ne! PP. 19-75.
et nie. \ wees y
“New York, Marth 3h, 1899.
NIE Bat
eee
cea
¥
¥
arene
Article III.—A PROVISIONAL CLASSIFICATION OF
THE FRESH-WATER TERTIARY OF THE WEST.
By W. D. MATTHEW.
The table of formations immediately following was drawn up
by the writer under the guidance of Prof. Osborn and Dr. Wort-
man, for the arrangement of the collections in the American
Museum of Natural History, and submitted to the late Prof. Cope
and to Prof. W. B. Scott for approval oramendment. The faunal
lists appended are made up chiefly from the study of these collec-
tions, covering all the formations of the Western Lakes except
the Pliocene horizons. A partial essay is made at a critical
revision of the species ; characteristic and well marked species
are in special type, and those considered invalid are in roman.
It is quite impossible to make this revision with any approach to
completeness, from the fragmentary character of many types and
the meagreness of descriptions. The object of the lists is to
assist in the correlation and classification of the various basins by
showing the characteristic fauna. The exact horizon and locality
are therefore given as fully as possible ; and to facilitate future
comparison of species, the location of each type specimen Is given,
as far as known to the writer. No bibliography or complete
synonymy of the species is included ; it would be undesirable in
this connection, as well as rendered superfluous by the excellent
bibliography of mammals recently published by Prof. Trouessart.
In this preliminary attempt, errors will doubtless be numerous.
The writer will be glad to have his attention called to any, that
they may be corrected in a subsequent edition. The synonymy
of genera and species is necessarily to a considerable extent pro-
visional, and will require revision, especially in the Creodonta
and Primates. Some of the results of Dr. Wortman’s recent
studies on the former group are embodied here.
New genera defined in this list: Pale@ictops, p. 35; Phlaocyon,
Pp. 54; Miolati Hay, p. 74.
New species defined: Paleolagus intermedius, p. 53; Phlao-
cyon leucosteus, p. 54; Ictops bu’latus, p. 55; Aceratherium pro-
fectum, p. 71.
The following names not heretofore used will be defined in
[19]
20 Bulletin American Museum of Natural History, [Vol. X11,
forthcoming articles by Dr. Wortman and the writer : Pal@osinopa
veterrima, nN. g. and sp., p. 313; Procynodictis vulpiceps, n. g. and
sp., p. 49; Oxyenodon dysodus, n. g. and sp., p. 49; Prodaphenus,
p. 49; Paradaphenus and Nothocyon, p. 62; Pachyena
intermedia, N. Sp., p. 31.
DIVISIONS OF THE TERTIARY LAKE-BASINS.
PERIODS. FoRMATIONS. Be FauNAL Divisions, | LAKE BAsINs.
ea
Pleistocene. |Sheridan. 100 |Equus. + Silver Lake and other
| localities in Oregon,
he eat ' Nevada, and. else-
i Pliauchenia. ES:
Pliocene. Blanco. 150 ae
Ho |piteha wien | Great Plains (Niobrara
| Basin).
,,|Loup Fork. 400 |Procamelus. |
vo
& aah
Deep River. | 150 |Cyclopidius. | | | Deep River, Montana.
| i
Miocene. |-~ | |
a
cs)
zg
=
iG
= John Day. 19000 /|Diceratherium. Toke Dy, Orson:
= |
; f Protoceras. { Great Plains (Sioux
Oligocene. |White River. | 800 |Qreodon. | Lake).
Titanotherium. |
ne : )
: Diplacodon.
2. Uinta. 800 P as | Uinta, Utah.
5S Telmatotherium.|) |
& 1 | )
S Bridger. 2000 |Uintatherium. | Upper Green River,
S Wyoming.
\ | | Huerfano, Colorado.
f : }
,;|Wind River. | 800 |Bathyopsis. | | Wind River, Wyoming.
Eocene. | 2 ADB |
° )
a Wasatch. 2000 |Coryphodon. | | | Big Horn, Wyoming.
yy
_|Torrejon. 300 |Pantolambda, | |
S | San Fuan, New Mex-
Ss | | ico.
|; |Puerco. 500 |Polymastodon. |
|
1899. | Matthew, Fresh-Water Tertiary of the West. 2.
NOTES ON THE LAKE BASINS, |
San Fuan Basin.—In northwestern New Mexico, on the divide
between the Rio Grande and San Juan Rivers. The beds con-
formably overlie the marine and brackish water Cretaceous
(Laramie), and contain fossils confined in the Basal Eocene to
three strata, two at the bottom of the Puerco beds, one in the
Torrejon. The Wasatch is sparingly fossiliferous throughout.’
Big Horn Basin—In northern Wyoming. No weighty faunal
distinctions have been shown to exist in this great mass of sedi-
ment, which is sparingly fossiliferous throughout. Buffalo Basin
is a subordinate basin, apparently somewhat later than the main
basin,”
Wind River Basin.—In central Wyoming. Fossils are rather
scarce in this basin, All the sediments are later than the Wasatch
and earlier than the Bridger.’
Hluerfano Basin —In southern Colorado. Osborn has recently
shown that the Wind River is represented here as well as the
Bridger. There are also overlying sediments of ? Pleistocene age.*
Upper Green River Basin—In southern Wyoming. This in-
cludes the Bridger and Washakie, with Middle Eocene fossil
mammals, and much larger areas where Green River (= Wind
River) and Vermillion Creek (= Wasatch) sediments are the only
beds not washed away by the extensive erosion which has taken
place. The Vermillion Creek sediments contain fossil mammals
in a few localities (Evanston, Black Buttes, etc.), while the Green
River shales contain fossil fish in abundance.°
Uinta Basin.—I\n northeastern Utah. The Middle and Upper
Eocene beds are underlain by a considerable thickness of sedi-
ments, probably of Lower Eocene age, in which no fossils have
been found. The three fossiliferous horizons are: C, the Upper
or True Uinta, explored for fossils by Marsh in 1870 and the
Princeton party of 1886 ; 4, the Lower Uinta or Telmatotherium
Beds; and A, probably equivalent to the upper part of the
Bridger. The last two horizons were discovered by the American
Museum Expedition of 1894."
1 Authority, Wortman. See Bull. Amer. Mus. Nat. Hist., 1892, 135; 1897, 259-
2 Authority, Wortman, Bull. Amer. Mus. Nat. Hist., 1892, 135.
3 Authority, Wortman, Z. c.
4 Bull. Amer. Mus. Nat. Hist., 1897, 247.
5 Authority, Hayden Survey Reports, etc. Also Wortman (communicated).
® Authority, Peterson, Bull. Amer. Mus. Nat. Hist., 1895, 72; Marsh, Amer. Jour. Sci.,
March, 1871. :
22 Bulletin American Museum of Natural History. [Vol. XII,
Basin of the Great Plains. —The precise relations of the fossil-
iferous horizons in different parts of this wide area stand in need
of further study. The deposits are coéxtensive with the Neocene
and Quaternary, but in no place has a continuous section been
observed, and the series of fossiliferous horizons is still more
incomplete.
The following table is an attempt at correlation of a number
of typical sections with the section published by Hayden and
Leidy thirty years ago. ‘The proper identification of these di-
visions has been prevented by a confusion as to the fauna properly
belonging to Horizon D, The descriptions given by Hayden of
the character of the rocks in the several divisions of his section
agree accurately with the corresponding divisions observed by
the writer in Colorado. But the fauna which Hayden and Leidy
ascribe to Horizon Y was found by the writer in Horizon C,
while D belonged stratigraphically and faunally to the Upper
Miocene. The fauna which Leidy ascribes to C is a mixture of
Oreodon and Protoceras Beds species.
The Middle Miocene does not appear to be represented in any
of the sections. The faunal break is least serious in Colorado
and Oregon, where a number of genera pass through without
more than specific change. Such are, in Colorado, Anchippus,
Aceratherium (sensu stricto), Merycocherus, ? Cants. In Oregon,
Mesohippus and probably others.
Another break occurs in the Pliocene, partly if not completely
bridged in Texas, but tolerably well marked elsewhere.
It is probable that the Protoceras Beds overlap to some extent
on the Lower Miocene John Day. Further study of both and
determination of the exact horizons of different species will be
necessary to find the extent of the overlap. At present it seems
that some lines of descent are more advanced in one basin, some
in the other. The Rhinoceroses seem to have run into Dicera-
therium in the John Day, while continuing the line towards
Aphelops in the Plains. The Camels are more advanced in the
western basin, but the Horses are persistently primitive, while
Anchippus had already appeared in the Leptauchenia Beds.
Side branches of Canidz appear in both basins, while the
more direct line shows little difference in age between the two.
Merycocherus and Eporeodon appear in both, but the relations of
23
of the West.
tary o
Matthew, Fresh-Water Terti
1899. |
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‘AV 'S P'SIN
24 Bulletin American Museum of Natural History. {Vol. XII,
the species to each other are not yet known. Leptauchenia is un-
known in the John Day basin, and is very close to the Upper
Miocene Cyclopidius, but if these genera were aquatic they may
have had a peculiar and limited habitat, and their absence would
not have much weight.
The so-called Loup Fork Beds are not all strictly of the same
age. The more easterly and southerly deposits in Nebraska,
Kansas, and Texas are uppermost Miocene or Pliocene. The
species described by Leidy and Marsh came chiefly from these
beds. The Colorado Loup Fork is a distinctly older horizon,
Upper or perhaps in part Middle Miocene. Prof. Cope explored
this region in 1873 and 1879 and described its fauna. He seems
to have been less familiar with the eastern Loup Fork; and this
may partly explain the discrepant views held as to the age of the
Loup Fork Tertiary. Williston and some other recent writers
entirely refuse to assign a definite age to the Kansas Loup Fork ;
and the present writer desires to avoid including with the beds to
which a definite age is assigned, any but those in which a suf-
ficient fauna has been found. The Colorado Loup Fork is here
placed as equivalent to the Deep River of Montana, the Ticho-
leptus Beds of Cope—an unfortunate name, as Zicholeptus is a
synonym of Merychyus, and Merychyus is equally characteristic of
the typical Loup Fork or Procamelus Beds. Cope placed with
the Deep River the Upper Miocene Beds of the John Day valley,
and of Laramie Peak, Wyoming. Other beds on the western and
northern margin of the Great Plains basin may be equivalent.
Scott considers the Deep River as older than any of the beds
above mentioned, laying especial stress on the occurrence of
Cyclopidius in that basin only. Cyclopidius, however, occurs in the
extreme west of Nebraska (Cheyenne Co.), and its absence else-
where may be explained, as above noted, by its aquatic habitat.
Miolabis* characterizes the Colorado Loup Fork and the Oregon
beds, and stands well in opposition to Procamelus of the upper
horizon. The lower beds have been called Cyclopidius Beds.’
They show the following characteristic faunal differences :
2 ( = Protolabis Wortman, 1898, not Cope.)
2 On account of its evident inconstancy a name derived from the fauna to designate a par-
ticular horizon is less suitable than a geographic term. Pawnee Creek Beds would be a better
name for the lower horizon in Colorado, Deep River being used correlatively with Loup Fork
for the general designation of the lower beds.
1899. | Matthew, Fresh-Water Tertiary of the West. 25.
Procamelus Beds. Cyelopidius Beds,
Pliohippus. ( Pliohippus.
Protohippus. | Protohippus.
Hipparion. | Hipparion,
? Mesohippus. ! 4 Desmatippus.
| Anchippus.
| Anchitherium.
( Pliauchenia. (peels
( Procamelus. Procamelus.
Protolabis.
Miolabis.
Cyclopidius.
Teleoceras. Aceratherium.
Cosoryx. Blastomeryx.
That is to say, Horses with short-crowned teeth and Camels
with split metapodials persisted into the lower horizon, as
did also the aquatic Oreodonts and more primitive Aceratheres.
Blastomeryx characterized the lower, as Cosoryx did the upper
horizon. The large Zeleoceras (Aphelops) fossiger, unknown in
the older horizon, is the most abundant fossil of the later one.
King’ applied the names ‘Sioux Lake’ and ‘Cheyenne Lake”
to the supposed lakes of the Oligocene and Upper Miocene of the
Plains (Miocene and Pliocene of older writers) ; and Marsh * has
called the latter the Niobrara basin. The deposits cover to a
great extent the same area, and a single geographical name may
conveniently be used, as is done in the other Tertiary basins.
The White River sediments are usually considered as lacustrine,
A similar origin has been assigned to all the later sediments, but
recent studies of Gilbert in Arkansas, Williston and others in
Kansas, and Darton in South Dakota, tend toward the view that
the Loup Fork is chiefly flood-plain sediment while the Pleisto-
cene is flood-plain and zolian deposit.* With this view the obser-
vations made by the writer in Colorado in 1898 entirely coincide.
Fohn Day Basin—In central Oregon. Two or three more or
less separate basins in the valley of the John Day River. The,
Cottonwood basin, containing a higher fauna, equivalent to the
1 Two lower molars referred to Mesohippus are described by Scott and Osborn as coming
from the true Loup Fork of Nebraska. ‘This is the only occurrence in the Procamelus Beds,
as far as Iam aware.
? goth Parallel Survey Rep.
® Am. Ass. Adv. Sci., Ann. Address, 1872. Sac
* Gilbert, 17th Ann. Rep. U. S. Geol. Survey, 1895-6, part II, 575. Williston, Kansas
Univ. Geol. Survey, Rep. 1896.
26 Bulletin American Museum of Natural History. [Vol. XII,
Deep River (Cope and Wortman) or Loup Fork (Scott), is
separated from the lower beds by a basaltic flow. Dr. Wortman
has recently found sufficient differences in the true John Day
to warrant its division into two horizons ; the upper, containing
a great abundance of JVerycocherus, being exposed at Bridge
Creek and elsewhere ; the lower, typical at the locality known as
“he Cove: *
Deep River Lasin.—In central Montana. It would seem to be
an outlier of the Great Plains basin. The upper beds, containing
the typical Deep River fauna, rest unconformably on beds con-
taining a scanty fauna which Scott considers equivalent to the
Upper John Day. The facies of the fauna does not seem, how-
ever, to forbid placing it as equivalent to the Leptauchenia fauna
of the Great Plains.’
New Mexico Loup Fork Basins.—A number of small scattered
areas, of which the Santa #é Basin in the northern part of the
state, and the Sax Francisco Basin in the southwest corner, are
the most important. Some of these deposits may be Pliocene.
The PLEIsToceneE fossil beds are scattered over all parts of the
West, and cannot be grouped into any definite areas. The Equus
Beds, largely river sediments, contain abundant remains of Zguus.
The prairie loess, so far as the writer is acquainted with it, repre-
sents a later deposit, eolian, and still in progress. The wind cuts
out all exposed places at a very considerable rate, as may be
seen by the rapid hollowing out of roads and ploughed fields.
The dust is caught by the sodded prairie and spread uniformly
over it. The effects of this mode of deposition are characteristic
and curious features of the plains. The deposits are over two
hundred feet thick in places, and contain remains of Zguwus in the
lower layers, but in the upper chiefly Azson bones in various
stages of fossilization. In the Cordilleras are many dried-up
lake basins of small size, in the deposits of which the Equus
fauna has been found. Silver Lake in Eastern Oregon is the
most noted of these.
1 Authority, Wortman (communicated). 2 Scott, ‘‘ Mammalia of the Deep River Beds.”’
1899. | Matthew, Fresh-Water Tertiary of the West. 27
CHARACTERISTIC Fosstu MAMMALS.
The more important or abundant species are in heavy type ; in-
valid species in roman. Present location of the type specimen is
stated (under the formation in which it was found) as follows :
No agin oo dic American Museum of Natural History, New York.
Whe Poosnse .. National Museum, Washington.
leeooo0cadar Philadelphia Academy of Sciences.
Virvosthemierererers Yale University, New Haven, Conn.
iSc co oooones Kansas University, Lawrence, Kan.
Iplneobebooss Harvard University, Cambridge, Mass.
eartveisiicrers .»Princeton University, N. J.
Wesasnonanse Texas Geological Survey.
Gs Gy Soacos Canadian Geological Survey.
28 Bulletin American Museum of Natural History. [Vol. XII,
iueCL RECO:
All the types and described specimens are preserved in the
American Museum of Natural History.
The localities of all are
in northwestern New Mexico, west of the divide between the Rio
Grande and San Juan Rivers.
MULTITUBERCULATA]
PLAGIAULACID,
Catopsalis foliatus Cope.
Polymastodon taGdensis Cope.
Polymastodon latimolis Cofe.
Polymastodon attenuatus Cope.
oS selenodus Osborn & Earle.
Neoplagiaulax americanus Cope.
Neoplagiaulax sp.
CREODONTA.
OXYCLAENID/.
Oxyclenus cuspidatus (Cope.)
ie simplex (Cope.)
Protochriacus priscus (Cope.)
Protochriacus hyattianus Cope.
Loxolophus adapinus Cofe.
Protochriacus attenuatus O. & E.
TRITISODONTID-®,
Triisodon quivirensis Cope.
sf heilprinianus Cope.
Triisodon rusticus Cofe.
Triisodon biculminatus Cope.
Conoryctes crassicuspis Cope.
Sarcothraustes coryphzeus Cope.
Mioclenus bathygnathus Cope,
Tritsodon gaudrianus (Cope).
INCERT, SED.
Oxyacodon apiculatus O. & E.
Oxyacodon agapetillus (Cope).
Carcinodon filholianus (Cope).
CONDYLARTHRA.
PERIPTYCHIDA.
Periptychus ! coarctatus Cope.
Periptychus brabensis O. & Z.
Ectoconus ditrigonus (Cope).
Hemithleus kowalevskianus Cope.
Conacodon entoconus (Cope).
Anisonchus coniferus Cofe.
Conacodon cophater (Cope).
Anisonchus gillianus Cope.
Hemithlzeus apiculatus Cope.
? Zetodon gracilis Cofe. (Indeter-
minate.)
PHENACODONTID,
Protogonodon pentacus (Cope).
Protogonodon stenognathus Matthew.
MIOCLAENIDE.
Mioclenus turgidunculus Cope.
EDENTATA.
STYLINODONTIDA.
Hemiganus otariidens Cope.
CONORYCTID.
Onychodectes tissonensis Cope.
Onychodectes rarus O. & E,
INDETERMINATE SPECIES.
Miocleenus interruptus Cofe.
a riitimeyeranus Cofe.
Il. TORRE/JON.
All the types and described specimens are preserved in the
American Museum of Natural History.
The localities are in
northwestern New Mexico, on either side of the divide between
the Rio Grande and San Juan Rivers.
1 The name Periptychus is perhaps preoccupied by Periptyches.
If this be the case, the
name Catathleus, proposed for the permanent dentition of the same species, takes its place,
C. rhabdodon being the type.
1899.
Matthew, Fresh-Water Tertiary of the West. 29
MULTITUBERCULATA.
PLAGIAULACIDA.
Neoplagiaulax molestus Cope.
Ptilodus medievus Cope.
Ptilodus trovessartianus Cope.
BOLODONTID&.,
Chirox plicatus Cope.
PRIMATES.
TARSIIDA.
Indrodon malaris Cope.
RODENTIA.
MIXODECTID&.
Mixodectes pungens Cope.
Mixodectes crasstusculus Cope.
CREODONTA.
OXYCLANIDA.
Tricentes subtrigonus (Cope).
Mioclznus bucculentus Cofe.
Phenacodus zuniensis Cope,
Tricentes crassicollidens Cope.
Chriacus pelvidens (Cope).
Chriacus stenops Cope.
Chriacus baldwint (Cope).
Chriacus truncatus Cope.
Chriacus schlosserianus Cope.
Deltatherium fundaminis Cope.
TRIISODONTID,
Goniacodon levisanus (Cope).
Sarcothraustes antiquus Cope.
Triisodon conidens Cope.
MESONYCHID.
Dissacus navajovius Cope.
Dissacus carnifex Cofe.
Dissacus saurognathus Wortman.
VIVERRIDE,
Viverravus haydenianus (Cope.)
Didymictis primus Cope.
ARCTOCYONID.
Clznodon ferox (Cope).
Clenodon corrugatus Cope.
Clenodon protogonioides Cope.
CONDYLARTHRA.
PERIPTYCHID-®.,
Periptychus ! rhabdodon Cope.
Periptychus carinidens Cope.
Anisonchus sectorius Cope.
Anisonchus mandibularis (Cofe).
Haploconus lineatus Cope.
Haploconus xiphodon Cofe.
ZS angustus Cope.
Laploconus corniculatus Cope.
PHENACODONTID<E.
Euprotogonia’® puercensis (Cope).
Phenacodus calceolatus Cofe.
Protogonia subquadrata Cope.
ee plicifera Cope.
Mioclzenus floverianus Cofe.
Euprotogonia minor Matthew.
MIOCLAEENIDA.
Mioclenus turgidus Cope.
Miocleenus zittelianus Cope.
Mioclenus lydekkerianus Cope.
Mioclenus lemuroides Matthew.
ss acolytus (Cope).
Mioclenus ineguidens (Cope).
Mioclenus minimus Cope.
Protoselene opisthacus (Cope).
Hemithleus baldwini Cofe.
AMBLYPODA.
PANTOLAMBDID&.,
Pantolambda bathmodon Cope.
se cavirictus Cope.
EDENTATA.
STYLINODONTID&.
Psittacotherium multifragum
Cope.
Psittacotherium aspasiz Cofe.
a megalodus Cofe.
Hemiganus vultuosus Cope.
CONORYCTID&.
Conoryctes comma Cope.
1 See note on p. 28.
2 Dr. T. S. Palmer has kindly called my attention to the fact that the generic name Exfro-
togonta is antedated by Tetraclenodon, a genus of Creodonts founded by Prof. Scott on
Mioclenus fioverianus of Cope, a wrongly interpreted specimen of Euprotogonia puercensis.
‘The generic description of Vetyaclenodon is doubtfully valid, very misleading, and founded on
error; the name is inappropriate in the extreme ; and I have therefore retained Exfrotogonia
as a well known and appropriate name substituted for the original preoccupied name
Protogonia,.
30 =©Bulletin American Museum of Natural History. [Vol. XII,
Hil WASATCH.
Hele
a | Bg
S| 2| S| #2
q S Oke
Boal Gest onl lietre
mo} a] 4
PRIMATES.
Omomys (Anaptomorphus) homunculus (Cope).....-.- eel Sc
Hyopsodus paulus Leidy........... Seecr aie ret xT xX
ts VICATINS COPE ica. crs cw oiaiseeon stesotele aiclete x || XX
ef powellianus Cope............ SeGnGon || 5 neule
LHyopsodus lemotnianus COpe.......--+2see.ssceeees dC Ihe *
a PIG TUL Copoav anc gopseaccousooaGede ate Ka eUers:
% MRE ECUAIPAAS® (AE OPC) a aie ay ai5is ete rants oie <a atonaeer ze A.M.
Pelycodus frugivorus Cope.......... bb AGIOS seat x Be
BRelycadus mun enusiCapenyassiealelelerye citer serie =ii x
Pelycodusi tutus Cope. wee on elenctsa teins cle « < x<
IEASARHDS yf GOES CON n000 a0 8h00 900005000 SoosC sail >< x
“ MORO EONS 6 00.0 bod 0 Go00e 00 CdODOOODE x A. M.
WVICCKOSYOP SENET CLUES EGLO Yrnrslefor eta aheroleter=fole(eitol valet el (ae x
m POUAILS (Coens abocobagsboogsogunnoor x Jake IN
Cynodontomys latidens Cope...... AGH OO Oren creer x os
-RODENTIA.
ISCHYROMYID-®,
Paramys buccatus Cope......... eONoL selapscoicerer es x x
eS delicationlWertyzr ce seek seme eo x
sf delicatissininsmceidy sacreimierine vcietersisiore Sui
CREODONTA.
VIVERRID#.
Viverrayus* protenus (Cope)..). sce vk -c ie see tue or <a elt <a iss
se leptonaylis (Cope) stasis <a
Viverravus curtidens (Cope)....- scaled Achebe erohal casusoxe tees x A\s Wile
i MLONSELEMUGIES: (SOME) eteraleyernetterteratelleheraitens ale 2s
eS ARIES WETS Ns opooogcoogs eo ssouonuSdcE x
Uintacyon (Miacis +) canavus (Cope) .............. x A. M.
Uintacyon (Miacis) brevirostris (Cope)..........50+-- <ul He
a es CES DECOR OB OOD G00.0 8 CU PO OOO OOO x ae
1 The premolars associated with the type and only specimen of Diacodexis laticuneus
Cope, are those of Hyracotherium index ; the upper and lower molars belong to Hyopsodus,
closely allied to H. Jowellianus. See Wortman, Bull. Am. Mus. Nat. Hist., 1896.
2 Viverravus as defined by Marsh in 1872 covers Dzdymictis Cope, 1874. See note
on p. 35.
3 The type of Didymictis leptomylus of Cope may be the same as Marsh’s Limnocyon ri-
parius of the Bridger. This Wasatch species was referred by Cope to Zeptomylus but is
probably distinct from the type.
4 These species are considerably different in type from the Bridger species to which the
name was originally given.
1899. | Matthew, Fresh-Water Tertiary of the West. a
Ill. Wasatcu.—Continued.
E/ 4) e| £8
mile ei) oe
wo / 3] o || 4
ee) |) es) bz
PROVIVERRID.
Palzosinopa veterrima Wortman!................ x A.M.
Sinopa (Stypolophus) hiams (Cope)................. x <i) WS
CG ss strenuay(Cope)haacceecn eee x < ae
ut “s Wirittze: (Cope)... co. «6.5 seer x Nadk
ae cs Viviertinay(Cope)iaqaccece vee x Sei WES
Stnopa (Stypolophus) mzdlticuspis (Cope)...........-- sé ae
se a SecuUmaarea (Cope) maces sees ers < s
up a wean (Copy oocooccé: conue Pe
OXY NIDA,
seyce ee Mia Cope sii j ais te se sees oe eas 400 b SZ SK ese Ii) tor, S
Me POUGE DREAM CONES es. ale ore. 5 oS aciererals » desate ¥0s >< x ae
ORE NGEE AGREES EO Y2508 6 G0 eG COU CDE DEO CORO OOCE OOS x BC
PALZONICTID, |
Palzonictis occidentalis Osborn and Wortman....| A.M.
AU TAUSTD TOS. LOST Goi NS6hs dO b Po OO AEM Don UC Oe x Wis:
MESONYCHID.®.
Pachyena’ ossifraga (Cope)...........0000% aatiawile x HWE Ss:
CCR CEI OLE MLCALG NVIOLEMMAM «erie = 00 iis 2 oo cies 2s x< A. M.
Pachyena gigantea Osborn and Wortman.......... x | “
LDUSTECOS UGLOTIDT IOS, (Oy ENN Be oceHeCO CORIO PCO Oe Dio. <x ob
ARCTOCYONID,
PAULAEOMOMMUTSTACH SA COME sealers) aieiele)s aie = eis) siete ieieys 614 THLE x A. M.
INCERTZ SEDIS.
DGAEL ROMS AUS OGORE CODE Weis qerya a: -/eisielsicye tee Saas) SX A.M.
PIP RYS COMESLOCRE (COPE) oo 2 o2.00.c 1s ss: s5 o iceleiais 80 de x ‘
LOWE GRUTOCHLUOS COs. och ecHeeeb Ho ba BOO MODOC ODOoS SCe lh es
2 GIMICLIG LES CO So 35 0805 Ob COO UEUON COCO DO OOE x BG
INSECTIVORA.
LEPTICTIDA,
LUNG DSO? (PETS ES (COW) Good sooo necoU aC cee nOr x<
TILLODONTIA.
ESTHONYCHID. | |
mathouyed burmeisteri Cope. ....2..2. 60. eaeves: XS OX SS:
LESUADEY CR CGE CD N66 Bob 6 Go OO UO CED OEOOGC COUGE Pd | aie
ee WESCUAE UTE EQ) a6. 6 us CUOC COU BOONE ONCOKia x a
es spits. Coyslenectoa 6 ced CORO CDOS nOUOC ile |
1 Unpublished.
2 The original type from the Bridger was indeterminate. The species properly dates from
Cope’s description in 1884 (ert. Vert.) of a specimen in the Princeton Museum referred to
S. aculeata. ‘
3 Unworn molars of Pachyena show that it possessed a vestigial metaconid, though less dis-
tinct than that of Dzssacus ; the original ground for the separation of the two therefore fails.
I retain them separate however, believing that a careful comparison will show distinctions of
generic importance between them.
4 Generic position very doubtful. a
® See note on p. 35. ® The species of Esthonyx need revision,
32 Bulletin American Museum of Natural History. | Vol. Ul.
III. Wasatcu.—Continued.
cae
n S) S
¢| a | 3) 8
See pecs oH
en os | Jan
on eS | z 4 °
oi, A | AZ |
Neate
EDENTATA. |
STYLINODONTIDA. |
Calamodon simplex Cope........ Bees eae a Me <a << {ll Un SS
CalamodonyancamoentsiCg7 cmminrlatecisictelctelaieleiale fans | S€ || oo"
Calamodon novomehicanus Cofe.............---| x io
Dryptodon crassus Marsh ........+++2+-2+0--->| x ae
CONDYLARTHRA. |
|
PHENACODONTID. | |
Phenacodus primeyvus Cope *. 00... a. ie see ns x * || AoM.
IDNR OslnS GueUhVOHNS (CGN? nooosdganoooceoo00G | | MURS:
Phenacodus nunienus Cope®......- Ao BonorDeDdaoapoa) y ; A. M.
Phenacodus wortmani Cope®........... sa0006000l| OX x
Phenacodus apternus Cope. ..........- s5H00050|| 3X A.M.
Phenacodus hemiconus Cope. ......++- podaoe einc.doll pan ee
Phenacodus brachypternus Cope ..............--| X | X | ss
Phenacodus macropternus Cope ......+eee eee a6. eral. oS a
uy SMALOS COWS 55 900000005 Seeman s59005 Y< |i) Wotse
a (Ectocion) osbornianus Cope........- soul 3X A.M.
I oloyAUe CEES WCAWE 600000 0apapdsoc2ando0bDSGaD < We
as RAMUS WH AVG ado0ccndgup9asodaoomccd0DGc x BG
MENISCOTHERIID&,
Meniscotherium chamense Cope. ............... : x |) WS.
ss terree=Tubree) COpesree).cleerieieleclsel- ee AeaNls
Meniscotherium tapiacitis Cope ....- IAA CE Ee < of
Hyracops socialis Marsh *...... sdabod0GG0G000 DOO don x We
1 Type from Evanston, Wyoming,
2 Perhaps a small variety of P. przmaevus.
3 See note on this species on p. 36.
4 Doubtfully distinguishable from P. xunxzenus.
5 Eohyus Marsh (nom. nud., 1877) is perhaps a synonym of Phenacodus; E. distans
(figured, 1894) might be taken for the very uncharacteristic m$ of that genus, and the description
of Z£. robustus (1894) corresponds as far as it goes to the lower jaw of P. prim@evus. Wortman
(Bull, Am. Mus. Nat. Hist., 1898, p. 101, foot-note) believes that Z. dzs¢tans is founded on the
last upper molar of 7rzgonolestes etsagicus ; but the tooth as figured by Marsh is too large for
that species, even on the supposition that m® is unreduced, which, judging from the reduction
of the heel of mz, is not the case.
® Hyracops, which has been identified by Osborn with Mezzscotherium, differs considerably
in its foot structure if Marsh’s figures are accurate. The large magnum, the entirely serial
carpus, and the epicuneiform seen in Marsh’s figures of Hyvacops are not present in Wenzsco-
therium, which has a carpal and tarsal structure very like that of Aufrotogonia, with small
magnum, lunar supported partly on unciform, and other normal primitive features. The
metapodials and phalanges are like those of Hyracops.
io>)
1899. | Matthew, Fresh-Weater Tertiary of the West. 3
Ill. Wasatcu.—Continued.
<
g PAY ede eel | eel
g fo) & —Q 4 § &
eee ea) Ee] ae
a | ele i si se
alma| 4
AMBLYPODA.
CORYPHODONTID&.
Coryphodon (Bathmodon) radians Cope.........- henge AS Ns
Coryphodon (Metalophodon) testis (Cope)...... x :
Conyphodonirepanditss Caper eisai iia | x ae
Corypuodoni lobatis Copesc.c sonnet 33: <ul (<A fice | ELS
Coryphodonianaxe Cope were itera ie XK Jakes Jil
Bathmodon pachypus Cofe............+..-- << t
Coryphodon elephantopus Cope............. | eal act ie eal) Oleg
Coryphodon obliquus Cofe...............-- | | x a
Coryphodon cuspidatus Cofe............... | [se x ae
Coryphodon hamatus Marsh................- eee VG
Manteodon subquadratus Cofe...........+- | |X A. M.
Coryphodon latidens Cope.................. lex Sue Ss
Coryphodony simuss Capes recess oso ate | x a
os MOLES HUSH COPE. yeneryone tole tosh 2 = | x he
Cayyphodon CURUICTISTES COPE 6 5s. veins veins seuss x A.M
e (Metalophodon) armatus (Cope). |
Black Buttes, Wyo. ares
S (Ectacodon) cézctus (Cope)........-- eee
Coryphodon EatpesnCofia.s Sa actactatens ives 4 2186 cect x ss
mareimatuss Cope tie Meneses x st
PERISSODACTYLA.
EQUIDA.
Hyracotherium (Eohippus®*) index Cope...... PONS | OS ee
Hyracotherium angustidens Cofe........... esse
a CuSpidatumeCopEen es.) -16 «1 erie’ me
HOhippusspernixeAVa7S oa. eb iecle-leye os 5 ss x fines
Hyracotherium (Eohippus) vasacciense Cope.| K | X | A. M.
HOhippUSmVvalidus e725 (vate ers seis cole eielle cc: x 1 Ye
Hyracotherium (Eohippus) tapirinum Cope... ms SR <
Hyracotherium (Eohippus) cristatum Wortman... x 1 eds, Wile
ts (Phiolophus) cristonense Cope..... | pol eet Ss
He es montanune Vortman. x | A. M.
LOPHIODONTID. |
JUG REDHEE ROSHIEDS (EOS, So 000 06 SUE DOUCHE CODE OC | Xx | A. M.
i PROMS ACO Keo e os CoC Oa aC OU OnOOAD | x | U.S:
1 Part of the same individual is in the Yale University Museum.
2 Indeterminate types, synonymy doubtful.
3In Hyracotherium proper, p? has but one cusp; in all the American species, a trittocone
is also present. ‘‘It may yet be found that there are other important differences between these
groups which will necessitate recognizing a separate genus for the American forms, in which
event the name Zohifpus, proposed by Marsh, would have to be adopted.’’ Wortman, Bull.
Am. Mus. Nat. Hist., 1896, ror.
3
34 Bulletin American Museum of Natural History. (Vol. XII,
Ill. Wasatcu.—Continued.
a
oF n .
§ ee caer lier:
#5) Boel eee
i ee ee g
a} al] 4
TAPIRID#A,
Systemodon protapirinus Wortman.......... x Ss A.M
Systemodon primevus Wortman........--...--. SK | OL
ve semeniam's\ (CODEsacereelei sie ieee a) a < ee
ARTIODACTYLA.
HOMACODONTID,
Trigonolestes chacensis Cope.*... 3... .-5....- x x Mw, S-
Trigonolestes brachystomus COpe.....+ ese eeee eee x | Jak Wil
a OOS CON aoossove gooa00n80D0 x | Pe
a MESES EQ Eso nnacobauosqoseoc x “
ue ASTUCHS Coico nogsoccooss GoGde x “
ACHENODONTID,
SASS BOOS MEIN, Gop d560050009900000506 x :
a QUEEF ATSANUATS Wepretetartetsenienst keer sieiee x ue
LV WAND RIVE.
All the types and described specimens are in the American
Museum of Natural History.
2 o¢ ®
4.8 aS Be
E sc Soh
ge oo in
i oS ia "6 °
psoas || nssS) 4
PRIMATES.
Hy opsodusipaulusiveicdynrrriee titre ir x
se Wicarius (Copes were cclsi e es: sceei sieve | xX
ELM OD SOM LSI AMICILOURLO TEU COMET eaeRrtet rte eget tere oii <<
a ROM DOS (COPSsa606b00050 00050000060 | ee
LEVIRGQI LS UTES CONS55.6 010 0506 coG0 D0 obo GDonoD Ob ADE ee | Sc
Pelycodns frugivorus| Coperjri senses ene ane an or x
Relycodus| munienus Gap ememrermiselseicie ie eierse eX JAG ML:
Microsyopsi gracilis Weidyanmaerimer sitet teil IP xX %
LCEOSH OP SISCOLELOMLUS © OD Ck: soucterat teticn taut letertee it hee | AGEN It
1 This is the common species of 7r7gonolestes. The others are founded on somewhat smaller
or otherwise different individuals, but all very closely allied except 7. etsagicus, which differs
widely and may have to be separated generically, as Wortman has suggested (Extinct Camelida
of North America, Bull. Am. Mus. Nat. Hist., 1898, p. ror, foot-note). I have seen no
specimen of 7yzgonolestes proper (our collection contains 26) in which the paraconid is not
present and twined with the metaconid in a peculiar way.
? Doubtfully distinguishable from JZ. gracilis.
1899. | Matthew, Fresh-Water Tertiary of the West. 35
IV. WiInvd RIvVER.—Continued.
|
Fs lees
A) S¢ 2
%.5 au 58
eo eeeiliige
A 35 2
Sie |r) 4
RODENTIA.
ISCHYROMYID.
Paramys delicatior Leidy...... Mahe lke alae lentes yetefers x x
se delicatissinuspWeidyaese eee eon x
CREODONTA.
VIVERRID,
Viverravus | (Didymictis) altidens (Cope).......... x A. M.
“tverravus (Didymictis) protenus (Cope) ............ SK x
ut (Limnocyon) v7parius (Marsh)....-.....
? Didymictis leptomylus Cofe, type only.......... x A. M.
Wiverraviusionacilisy Marshicre crm taie clits octets © x x
Didymuictisidawkimstanus) Cope). 4 <1 se 1hole*)s x A. M.
Uintacyon (Miacis) canavus (Cope)............... x 6
ef we brevirostris (Cope). ...... el eee “
Gintacyor ((Miacis)\efz vorax Leidy.: 22.6.0 6s. x
OTE Y OMNES [eM arayeratatatatefelanelsyeleyetereyeleletel elarete)elciel x
PROVIVERRIDA,
Sinopa (Stypolophus) whitiz (Cope). .............. A. M
Stnopa, cf. viverrina (Cope)*....... BOP ohey heroy'< oheive lait aie
OXYANID,
OnVena AUCH ANeEnStS, OSHOTM.)....)- ese +. ese cass Soe x || A. M.
LALO AUIS 1EGGUETS (COs). 58 520 as sadn dobOdenoneGs Ss 2
INSECTIVORA. |
LEPTICTID/.
Palaictops® bicuspis (Cope)... 2.00.0. eeseecees Peon i >.< | A.
“ diaelppordesa (Cope) deve cise sysics| «| oe eis oe <) x lio se
CHIROPTERA, |
Vesperugo anemophilus Cope.......+. Yee Ler Sicesenerey= x eee
NOPE RTPA EE Not etras con wise 15M Ge seed Bids cue ess ee Wie ni
1 Viverravus Marsh antedates Didymictis Cope, and refers tothe same genus. The generic
description appears to the writer to be a sufficient one, although Marsh in this as in other de-
‘scriptions uses the term ‘‘ tubercular ’’ to describe a tuberculo-sectorial tooth.
2 Apparently a different species from the Big Horn specimens referred to S. viverrina, I
have not compared either with the type, from New Mexico.
3 Referred by Cope to /cfofs, an Oligocene genusclosely allied to Leftict/s Leidy. Though
not distinguishable by the dentition from /cfofs, there are important and very constant differ-
ences in the skull characters, among which I select for generic definition the backward ex-
tension and broadening of the posterior ends of the nasal bones, and the presence of a single
median crest on the cranium instead of the two parallel crests of /ctops and Leftictis.
4 Pertinence of this species to the genus very doubtful. = Type lost.
36 Bulletin American Museum of Natural History. [Vol. Xi,
IV. WInpD RIVER.—Continued.
Era ies
2, ee || Zh
6.5 ey || 68
28 | Bao) 3a
det ES Era, | on
Seo eons
TILLODONTIA. 1
ESTHONYCHID/. |
Esthonyx acutidensp@opens).:e.ce cvs 65 see ee x ~ || A.M,
Lsthonyx spatularius Cope........ Biever Ms aieivle ie x lease
EDENTATA. |
STYLINODONTID#. |
|
Stylinodon (Calamodon) cylindrifer (Cope).......+.+5+ x |
CONDYLARTHRA.
PHENACODONTIDA.
Bhenacodus 7 primesvusiGope tan. sass deiscs «seis ae x
Phenacodus wortaianl Cope*. hen -ceessescs en. x |} A. M.
Phenacodus (Ectocion) osbornianus Cope......... 5 XS
AMBLYPODA. I}
CORYPHODONTID&. |
Coryphodon ventanus Osborm 22.24. .)cc.02 06-0. x Soll AL Me
Comphodon worimonm Osborne nerine see ears one ee x lhe sone
4 PESTNEULAHES. OSDOLM Era aeiicoie eiete oieiee eo x [Keaaiais'c
UINTATHERIID.
Bathyopsisfissidens: Copey-¢ a. oo ee oe om 5 her < A. M.
PERISSODACTYLA. |
EQUIDA. |
Hyracotherium craspedotum Cope............... nex || A. M.
Protorohippus (Hyracotherium) venticolus (Cope)., x x | “
Lambdotherium popdagicum Cope®.............. x <a os
LOPHIODONTID&. |
Heptodon calciculus Copesssticce onscreen «oie x “
LG REMI TL DUE (CE OBSo. ons bo Oo pa OHA HO AeOoHsadsocaC x Weems
TITANOTHERHD&, |
Telmatotherium (Palzosyops) boreale (Cope) .. x i|
Lambdotherium brownianum Cofe ............. x
ARTIODACTYLA.
HOMACODONTID®. |
Tpigonolestes Seca res MEODE taacinet avec ete oestoeisal aa x x | A Min
1'The single jaw fragment referred to this species differs in several particulars from the
Wasatch specimens. . $y Sod
°*'Two or three species are apparently included under this name, but the material is too
fragmentary for their separation. The Wind River specimens, including the type, are, I be-
lieve, distinct from those of Wasatch age, including the complete skeleton described by: Cope
in * Tertiary Vertebrata.’ Two points of difference are the internal instead of postero-internal
position of de on p*, and the more compressed p*—both points approximating the Wind River
species to Hctoczon, the most advanced of the Phenacodonts.
8 See Osborn, Am. Nat., 1897. le
* Upper molars being unknown this species can be placed in 7yZgonolestes only provisionally.
Matthew, Fresh-Water Tertiary of the West.
1899. |
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Bulletin American Museum of Natural History. [Vol. XII,
38
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39
Matthew, Fresh-Water Tertiary of the West.
1899. ]
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48
1899. | Matthew, Fresh-Water Tertiary of the West.
Wile CePA:
PRIMATES.
Elvopsodus otacilis Marsh.c0cs.aso0e6c2saiee ees
LL MESIEDS SOnadidas coaccopssgodaduaocosocopasHol |
VG TSGES CPG Peps OA 2665 0odn eo oD ob ob aneD OOK
RODENTIA.
RArAihySeSCIUTOLdeS S66 Ol acic amass sstelatl ate idee bs
RURAL SA ICLI LEM SS ©) SDOLIM aleve cteieisi- ao) esis) selene hs 6) 9 oer
RO LOPOY CLUS) HOLGHENL COLL yan assole sie) iva eres ee) |
CREODONTA.
CANIDA,
MUGCIS WMS SEOUE was oonsco boob su daopDGHduoeGDS
LORNA COLES CLD SOROS AOE OCOD ee
Prodaphznus (Miacis) uintensis (Osborn)..........
Brocyuodictis vulpiceps® 3): dances ce edles oes
OXY AZNIDE,
OxyeenodonidySOdusi. asco. dees an ecw case:
MESONYCHID,
ESOMV ICN CEA OULUSTCLETIS | CODES wis\sjehes Nora ahetale) ona) 6 =\~)) ole ie
Mesony x lintenSiS) ScOttiass<. s0o5 4s soensc eons:
AMBLYPODA.
UINTATHERIID,
Uintatherium sp
OC er rr ry
PERISSODACTYLA.
EQUIDA,
Espilinpus wintensis) Marsh... ....s.e+ <> os c= oes
einen puesnomactles, (NVALESI) saris cies dow sel ss oe esis, s/o 6 15
LOPHIODONTID&.
Flelaletes guyotit Scott
S/ejlelia (eis ie) «ely \uje) ms me)e/'e e)\6, 0) 6) 6.6 (6, © 80 0
TAPIRID.
Isectolophus annectens S. & O
Ce
| Horizon A.
| Horizon B,
xX
xX
x X
| Horizon C,
Location of
Type
+ Type mislaid or lost.
? Unpublished.
* Unpublished. See Osborn, Bull. Am. Mus. Nat. Hist., 1896, 78, fig. 3 (Hy@nodon).
4
50 Bulletin American Museum of Natural History. [Vol. XII
VI. Uintra.—Continued.
eee dese
< a ovo So
S ge | Se
g s | 8 ao
ofa cela sf bod
HY RACODONTID. |
Triplopus opliquidenss 160 Once eal ape Ga ne 2
Prothyracodon intermedium S, & O............ |
Triplopus sp. (? = Lephiodon bairdianus Marsh, part). x
AMYNODONTID.
Amynodon advenus Marsh.,.........+..+-0+-+-eseee- Nec Y.
Amynodon intermedius S.&O................... S< Ie
TITANOTHERIID. |
Telmatotherium cornutum Osborn................ | x | A.M.
Telmatotherium diploconum Osborn..........+...05- | ><a oe
Telmatotherium ? diploconum, var. minus!....... | < as
Telmatotherium hyognathum S. & O............+-- | Men EX< 1p.
Palzosyops manteoceras Osborn”................ ON OK | Ca ACNE:
Je ksh Oo oan =p ooobgaundogedsouodac ex te
Diplacodon elatus Marsh (e026. weer sss | x ANS
Diplacodon emarginatus Hatcher... .....:....62+-4. heed Ee
Diplacodon sp. indese-. 22.2... ieee fae eees ee eee | eh ONG ihe
ARTIODACTYLA. |
HOMACODONTID#.
Bunomeryx montanus Wortman.................. | NS late Wl
Bunomerjx elegans \WWortman....-.....---..+-2+--0- | x oe
CAMELID&. |
Protylopus petersoni Wortman.......+--++-eeee sees! SO fats Mle
Leptotragulus proavus S. & Oia Ps GOO ORE ee ORICON SNS SC] eR:
PATO INEe UES ETE oo ong o00 ahoozCasao que nSDE| SCA wees
ut ROUBIS Met os dan oadeoeaombomOOGan oS | Sule
(Cinta ncpes UM PGCAPeS SCO scoa noo op oopobduoaocucue | x P.
OREODONTID&. |
Leptoreodon marshi Wortman.................4-. | all Sai Aes
Mierycodesmanss oracilismocorl er teetetnr-i-i toler (11-1 | x :
Eomeryx (Protoreodon) parvus (S. & O.)........... | x *s
JE TUG PEA Fos HUAO)E TINGKENE, cco Soc o0csDdNCbODao ADDS x
Cite e idvemar lees NV aRSh ig eeitenete ete ryetadererett veel x Y.
Agriotherium paradoxicum Scott .........- Venton eas x Ie
VELOCE Ti RMON CULEED SEN LAGS Uetater tee ettel tel telson der el =e nee i « WG
ORORLET CUP ICHLIL Se MATS Deets eaten Peet aeheetel feted) tele t olka x sf
ACHENODONTID&,
Protelotherium uintense (Osborn).............--- x | A.M
Achenodon sp. (A. insolens, fide Osborn)............ | x
INCERT® SEDIS.
SP REHOCEIUSMUUMLETSESA @SDOTM Gea helt either x A.M
1 Unpublished. Heretofore confounded with 7. megarhinum. ? Unpublished.
1899. | Matthew, Fresh-Water Tertiary of the West. 51
VE ALTE RIVE
The localities tabulated below are :
1. Colorado. — Headwaters of Cedar, Lewis, Horsetail, and
Pawnee Creeks in northeast Colorado. Fauna described by
Cope in 1873 and later. The Lower, Middle, and Upper White
River are represented. Cope’s collections were from the lower
and middle beds, Horizons 4 and #&; the fauna of Horizon C is
determined from collections by the American Museum Expedition
of 1898.
2. Nebraska. — Hat Creek Basin, adjoining the S. Dakota
White River. Lower, Middle, and Upper White River are
present.
3. South Dakota.—This is the largest and best known area. It
lies between and about the White and Cheyenne Rivers, and
furnishes the typical section. The Lower or Titanotherium Beds
are divisible into three subzones, the Middle or Oreodon Beds
into two subzones, and the Upper or Protoceras Beds probably
into two subzones of which the uppermost one bears a generally
scanty fauna.
4. North Dakota.—A small area near White Buttes contains a
fauna apparently near to the Protoceras fauna in age.
5. MWontana.—\ have here included the lower beds of the Deep
River valley, which Prof. Scott considers as Upper John Day.
That they are later than the uppermost part of the White River
(Horizon C, Leptauchenia Beds) is, I think, not proven, and they
belong geographically to the White River.
6. Canada.—Swift Current Creek in the Cypress Hills, N. W. T.
The area is limited and the‘rocks conglomeritic, so that the speci-
mens were largely fragmentary. They are preserved in the
Museum of the Geological Survey of Canada.
Bulletin American Museum of Natural History. [Vol. X11,
52
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‘PIPNJIUOJ— AAAI ALIA AA
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62 Bulletin American Museum of Natural History. [Vol. XII,
VA ONE AY
The level at which each species occurs is known only in a few
cases, as noted.
RODENTIA.
SCIURIDA,
Type.
SCUMTS yOAsmMEyTr COPSs coons¢connocasosooseecnaeSngcoanodououes & A.M.
SOLOS: PONTLIOLOIS (OWS or 0680s Cone OO AD dO SHagoODeOOHADOONADONS ss
Ailomiys nitens) Marsha. cyte oecacnsrsurse sieges os rs lena caret ngersi sc heyket nies) steers Ve
ce (Meniscomiys) phippodusiCope sae cece rece oe reme cei A.M.
ZANT ES (MIS MIREoNS)) LOL OGN BES (CONS ooponsnoconssaeasoce0codKKC
ei a PAG HOS GROG nic Caacemajacdaceedsedoycacel as ny
ze 9 MRMINCONES oc ansoubedccopeossonanessonbucte sa
CASTORID,
SRR OT AUTHORS CONS a os obo ogo Unda sO UbasdooSbEEboOOnODONDDS A.M.
oe PE PDSMATS EQ VSG no poco eb bcocoons son ooonCooaUAOUpOOe *$
GEOMYID
LEU OVLGIES SLLLEUI TOJUS GO Cattatsteve eee ered ten nedel Tene tare okateke eae TeveneV ey ovale tarekor A.M.
eA UE HGH TMS (COVES cas aacdcaseadosnoposoeo apocrine 2d0d04 5 =
on ED MON AEDS (CONS oohoaan \ebavecet onsen onsovobonoGONt oy
Entoptychus plaiitronskCoper cesses eee ee Cee tee eer 3
Cavitrons=Copeses sac sascr wears seul: iver a Morar uneeers oy
us HUE b Oey pal Coole ANA ala omer OG CI Noo clo mabe 4 pone cd a ale a
Entoptyc OSTEO TLAES (CONS So Ho Vdn nS Coo DOOD OHO Saad SS oC UboR ROOM ee
ROXAS COWeos dacchbacngass006s000s cocHosDOdoUSS .
MURID©
Lesperomys nematodon Cope.......++.- 0 dcing 9000 pe SEONG OMONKOOe eS Pad WN
Pacicnlus locking tonianus (Cope yates ete ie iors oir
JERERCMDS PSN IES EO Neon obs Boke Dood be te sonbaGanoMUadeIOe d6Odoo0 OC oe
LEPORID
Wepusieumislans Copernic in ieee eee era aie a oe ee A. M.
CARNIVORA.
CANID.
Paradaphenus’ (Amphicyon) cuspigerus (Cope)..............+--- A. M.
SEMAN VMAS SOs WNIT: oda acaenousobcocodeaasbdadss sropeoaoncnc ‘a
Amphicyon hartshormianns s Cafes (Mant) eres yarner-seeee yar ier alate 5
Nothocyon’” (Galecynus) Seismarianus(Cope)ts cae rere seis %
aS Lemrs(C ope) ir se younae essere aoe tes ene oe ae pas
G ss latidems; (Cope) sacs tvs so Senn os -
sluemmnocyonraltigenis| € ope mere re mr eer eer ey tal ieee eee eran ie
LED ENTO CLMIT LOM ILS (ONS soo as ono ho pne Sooo deE nae euddoonedGoUSS be
i“ jer Ox Evermann Be RB eG A Ooo ies tee CRS RRR IL OO Veh
1 Includes all specimens from John ian referred by Cope to Pale@olagus.
2 Unpublished.
1899. | Matthew, Fresh-Water Tertiary of the West. 63
VIII Joun Day.—Continued.
Type
Hypotemnodon (Temnocyon) coryphzus (Cope).........-.-..---. A.M.
Hypotemnodon ( Temnocyon) josepht (COpe)..... eee eee eee eee eens “
Hyenocyon basilatus Cope.......+-.+----.-:- Falta nah ciato.g Aa oRIae u
ae VAGUS COMSs. coo 8 deoec 00 Sons Sn yan od oo aon os OG uo +
Oligopumis ceasstvaltus Copel. <ci.c sala. eo te cs wep ve stone se = ee
Enhydrocyou stenocepialus Cope. 2. fin- eee es cs abet seme cere: es
MUSTELID#.
PUTS je DGS NEO 95 o oeero co Uo obo OUDSU DOD OGObUoSeO0ndeooScS 1
FELIDZ.
Dinictis cyclops Cope........-.. sss eect eee eee e eee ee eee ee d 5 Wil
/A\neingallians CIDE (Cees casbeosnossobancueooucDono obo un OSD oe. *
Nimitavils)o-OmpHrodiusiG open. sie-j1lser lacie l= rere el-et= = hee ere craae a
NOLES GE AUDS COSo0s Sno ne oo con eH OROOeG DOOD SOK SHConCoUSNS as
BOMOUMUOON pla VvEOpis: COME: a.ya0 cle sales alu «sine ne sees ne cin cseeles 5s
LOH AIEDE HAMM ODS CODE, oa. 056050065 000c0TdnOo DO DUnOOOnb ena ooaG oe a
PieeSONONeUS CeLeDFalis Cope vitae wig couse eanele venta oa feat woreeels SY
Fioplophoneus strigidens Cope (Sp: gmdeterm:)). 2. 2... eee incense Ze
PERISSODACTYLA.
EQUID.
Mesohippus (Anchitherium) prestans (Cope)’....-.....-...+-..-- A.M.
equicepsa(Cope) nace sceecere eerie =o
Mesohippus (Anc hither tum) brachylophus (Cope)... +++ e.eeeceeeeeeees <
(WREE TOSS (COMM oc coceoucdsocacosudnoan e
se Be ROG. NGO acigioe og 1000 Ocodo Cooronods U.S.
os (CUDDLE FLO) CH QRHETOS (WUBI) Soc Gabbbbuooneocno600 4n00C We
SS a PAGS (MATA Yio 6oocug abode noes sasao0ca0 a0 es
LOPHIODONTID&.
me@uinaen (Lophiedon)ucctdcntalts Leidy?. ... 02. +.. cee een ss esse ns U.S.
RHINOCEROTIDA,
AGE TIER LTOTEOND. WANG 3 5 96.6 COORD OU BUGC dO OD aU DO oUbU IO Dob aoe Weis:
7 Paka POE NNW oo wos oo oobcoosdoKOueCo dose DondbOC a
MCCrALMeMImMs tulbitetsGope) (IMGet. SPs) .1s 1c alele eis selves fe ieiel i= ent= lool A.M.
GET OMLED UMM TUG ULEMUIL (COME selale = :\- 1s) 2s «ciel + 0 «ee he = oes) * fede re
o ChipeHeos MIME Nees com ope DIDO woo oTOG So NeCaedascc w.
PRCELALUCTIUIM ATINAtUIM MAGSH: 6125 66 vice nie see eels os erm 8 aioe nee misials is
e mara MENA, 6605 da daspaacb anual” Soesuidnhoocdoot *
INDETERMINATE.
DIE MGFIRG LOS LOMEZESE SA GOD Cieareycdayeieleheislel ele els, 2ha\* -hoie) ele) + nfelene (aleve) otels/efeh selene AGM
? CHALICOTHERIID (aut EDENTATA).
Luis Epes IMIENES No See. op oho de DOUDDCOUSBE EDO AOdOrOnooLOoGoURCC Yi,
SCIECICANUATS Mopars ro aie oicvayeie «+06 Fi heen ioe det as eee a Reo eNO a ae ae
1 Bridge Creek (Merycocheerus Beds). 2 Bridge Greeks
64 Bulletin American Museum of Natural History. (Vol. XII,
VIII. Joun Day.—Concluded.
ARTIODACTYLA.
ELOTHERIID.,
Type
Boocherus humerosus Cope! is..52 ies <01c = aluicios over tanerandaneran epee ALE
SIMU OOT OOD UP NGDON) 6 6.5 608 80S ONO HOODOO FG naS DOsadan OU G0 6K Ui :Ss
SUID.
Bothrolabis (Dicotyles)'pristinisLeidy.™... 2% 10: eu sei tree ee Wrasse
BOCHYOLQUUS UZLCR EVES COPE rete Wate erstele eels isle) ie ects eee AY Me
oF VOSULALUS AS OWE persphne aaye tts «iets ocd eNO) Mee TOE oi
os SOLU UAMSA COP eieysei tan \-ls Sieaaye me orate erohenetereveeeepe teers
CHENG USIAECEAEISA CODE MI tite eretie eho eer ieee Tee
Lihidaeo leyaes erates, NVATS Inn efencnc orerevaie's clots 6.c ti eini sl oosie wees lates nem Vi
us GEUALPSANTATS Nay. yorershNarsove sai oaversitsussabs cle a's op eee oro tee i
OREODONTID#.
AG TOC CETUS PEIICONS CODE is... sele Shalala wees) es oie ctl eel ares A. M.
Up guyotianus Cope........ Spee hove oho rameter eae i
LER ADE HCL ALS. TUTTE SCO) \Sevn wid hag 6.9 DODeO HOOD OUAGS Endo OOOnOSS x
Agriochcerus (Coloreodon) ferox (Copel ssi ssiro tne Secelauave ere oe
macrocephaluss (Cope) sire tera 7
PE pOreod ou. OCCicentalis a Marsh w.% setae esas s/o. «1-0. seicteeiatar son ete Ws
ISERIES EKO AKO IATA, ENE op cob odiao Good On oD GD OMOodSc Sc
7" OE Oe. JECHELTE Elding an aig ROO dOOoGe ddedo.0 da60a00 00
Eporeodon occidentalis var. leptacanthus (Cope)*........-2.,.:<- 20 ming
ne var. pacificus (Cope)?....... Lomein no lesa “
EON PEM ALAA HOINS (COS). .000 ncogn con aenb bb dodn eon. onon Onde s
Ee SOGUQULS gE ATS Dears hoc ienay pine hsielee Pelsactsre. eraicce Olan ea cient Ne
as EG ies LO UL (CO QS) sig Selon 6 Sloan How san and Dodie da. Neuve
Merycochcerus|(Oreodon) superbus Weidy*.: 5. -..:5-.-.sc2-ses he UES:
sf chelydra Cope AD er Cera IG eee CNA Oat A. M.
Merycocher ERUGTES Waal SKE NY 85'S char SoeHON ONE FO AeD HOO DOOD OOsan oc
TILAGHOSLE RU CODE cer aye rape Sieve sislteas oe ion ude eKee Fev Wile
CAMELID&.
Protomeryx (Gomphotherium) Sleruberga, Cones. coco <i annie ate AC iS
cameloides Wortman*... 0.25... 4. a
Hypertraculus calcaratis (openness acer er aero Meine ce
1 Perhaps the same as Elotherium imperator Leidy.
2 Bridge Creek.
®* These may probably be distinct species.
4 Bridge Creek Beds.
1899. | Matthew, Fresh- Water Tertiary of the West. 65
Exe LOOP FORK.
The localities tabulated below are as follows :
1. Vortheastern Colorado. Headwaters of Cedar, Horsetail, and
Pawnee Creeks, in Logan and Weld counties, north of the South
Platte River. Sands and gravels overlying the White River clays,
usually unconformably with heavy conglomerates at or near the
base. Another unconformity of erosion occurs sometimes within
the formation dividing it into horizons D and E (p. 23); but no
important faunal distinctions appear. These unconformities of
erosion in fluviatile beds are not of much importance. Collec-
tions made by Prof. Cope in 1873 and 1879, and Amer. Mus.
Exped. of 1898.
2. Laramte Peak, Wyoming. A small collection chiefly of
Merychyus, made for Prof. Cope in 1880.
3. Deep River, Montana. A limited area in the valley of Deep
Creek or Smith River, near Great Falls. Collections made for
Prof. Cope in 1877, and Princeton Expedition of 1892.
4. Cottonwood, Oregon. Cottonwood Creek in the valley of the
John Day River. Collection made for Prof. Cope in 1879 by Dr.
J. L. Wortman.
5. Oregon Desert. See toot-note, p. 70.
6. Nebraska. In the basins of the Loup River (Loup Fork)
and Niobrara River, and elsewhere in the .central and northern
part of western Nebraska. The fauna, described mainly by
Leidy and Marsh, is the typical and largest Loup Fork fauna.
7. New Mexico. This fauna was described by Cope in 1874
from the collections made by the Wheeler Survey, and now pre-
served in the National Museum. Most of the species came from
the Santa Fé basin.
8. Kansas—Nebraska. Northwestern Kansas and southwestern
Nebraska in the drainage basin of the Republican River. The
latest of the Loup Fork faunas, some of it Pliocene according
to Prof. Scott. The short-legged Rhinoceroses are the most
abundant fossil.
9g. Zexas. In northern Texas, near the head of the Red River,
along the northeastern border of the Llano Estacado.
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1899. | Matthew, Fresh-Water Tertiary of the West. 75
Mee LE LIOCENE.
A PALO URO:
(GOODNIGHT BEDS.)
This fauna, except for the presence of Zguus, corresponds with
the later Loup Fork fauna, which may also be Pliocene according
to Prof. Scott. It is too scanty for certain correlation. The
formation overlies the Texas Loup Fork unconformably.*
PERISSODACTYLA.
RHINOCEROTIDA.
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EQUID
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* Texas Geol. Sur. Rep., 1892
2 One specimen in Accican AMigeuth of Natural History.
A er DB \ y ns
\
Ancestry of Certain Members of the
1 ide, the Viverride, and Procyonide.
AUTHOR'S EDITION, extracted from BULLETIN
OF THE
ee ie
vican Museum ot Aatural History,
Bry Vow. XI, ARTICLE VI, pp. 109-138.
. Ween York, Sune 21, 1899.
7
Article VI.—THE ANCESTRY OF CERTAIN MEMBERS
OF THE CANID4, THE VIVERRIDA, AND PRO-
CYONIDA®.
By J. L. WorTMAN AND W. D. MaTTHEW.
PLATE VI AND TEN TEXT FIGURES.
The Eocene deposits of North America have yielded from time
to time a large number of remains of primitive flesh-eating mam-
mals whose nearest affinities appear to be with the modern
Carnivora, although not a few of them, so far as their dentition
is concerned, exhibit resemblances to certain of the Insectivora.
This group has been held by some to represent a distinct order,
and is considered by many authors under the name Creodonta,
originally proposed for it by Cope. By others it has been placed
as a suborder of the Carnivora, while still others have referred
to it under the name Carnivora Primitiva.
Without entering into a discussion of the merits of this group
to rank as a separate order or even a suborder, it 1s sufficient to
observe in the present connection that it is not capable of very
exact definition, and if it is desirable to retain a separate grouping
of these forms it must be done with the understanding that it is
purely as a matter of convenience.
That they stand in direct ancestral relationship to the later ap-
pearing families of the modern Carnivora, there can be no ques-
tion whatever, but whether these families arose from one or
several points among the Creodonta has never been demonstrated.
In fact no actual contact between the two groups has ever been
satisfactorily traced, and while many suggestions have been made
in regard to these connections, they have, up to the present, been
unsupported by any direct proof.
It is the object of the present paper, therefore, to trace the an-
cestry of two important modern families of the Carnivora directly
backward into the Creodonta. We will also take occasion to
point out the descent of several somewhat aberrant species of the
existing Canide.
Another important discovery included in the present paper
deals with the origin of the American family Procyonide, which
is now shown to have existed as far back as the upper Oligocene.
109
110) Bulletin American Museum of Natural History. (Vol. XH,
I.—TuHe SHORT-JAWED Docs OF THE EOCENE.
Uintacyon’ Lezdy.
This group of species has been found thus far in the Wasatch,
Wind River, and Bridger beds, and is represented so far only by
more or less complete specimens of the lower jaws. In one
species, Usntacyon vorax from the Bridger, a few fragments of the
skeleton are known, but beyond this the materials are very defi-
cient. Most of the species which we place in this genus have
been referred to AZzacis, a genus originally established by Cope
upon a jaw fragment from the Bridger.’ It now appears, how-
ever, that this first described species, and therefore the genus
Miacis, is synonymous with the genus Vudpavus, previously estab-
lished by Marsh upon a first superior molar.*. The name AZzaci's
will therefore have to be abandoned and the name Uintacyon,
given by Leidy, substituted. Leidy’s type of Usntacyon edax is
anomalous, in that it possesses five premolars instead of the nor-
mal number four, but this is not open to very serious objection,
since the shape and position of the extra tooth will convince
almost any one at a glance of its abnormal nature.
As the genus stands at present no very striking or trenchant
characters can be assigned to it which will separate it sharply
from its Bridger contemporary, Vulpavus. Although the upper
dentition is completely unknown, there can be very little doubt
that the most important distinctions between the two will be found
to consist in the number of superior molars. In Vudpavus they
are reduced to two, as in the modern genus Camis, while there can
be very little doubt that the number was three in Uzntacyon. The
only positively known character by which the two genera can be
separated at present, is seen in the thick, heavy lower jaw,
together with the abruptly rounded chin. In Vu/pavus, on the
other hand, the jaw is relatively slender and without the abrupt
chin, quite as in the modern Canide.
There are apparently several distinct lines of species in this
genus reaching from the Wasatch into the Bridger. In one pre-
cocious series the jaw is remarkably deep and short, the lower
1 Proc. Acad. Nat. Sci. Philada. 1872, 277.
2 Proceed. Amer. Philos. Soc. Aug. 1872, 740.
3 Amer. Jour. Science, Aug. 1871.
1899.| Wortman and Matthew, Ancestry of the Cantde, ete. 1T
canines laterally flattened, and the anterior premolars much
reduced. In another line the jaw is somewhat more slender but
still very short, the lower canines laterally compressed, but the
premolars are not especially reduced in size. A third section in-
cludes a single very small species from the Bridger in which the
jaw is very short and deep, the canines not flattened, and the pre-
molars unreduced. In a fourth section the jaw is much more
elongated and shallower, the canines not flattened, and the pre-
molars are not reduced.
The dentition of the lower jaw is I.3,C.z, Pm.z, M. 3. The
second pair of incisors is much displaced, being pushed back out
of the transverse line. ‘The sectorial is made up of an elongated
trigon and a basin-shaped heel ; the second molar exhibits a
crown of similar composition, but the trigon is much lower and
the anterior cusp considerably reduced. In some species the last
molar is very small and single-rooted, while in others it is larger
and is implanted by two distinct roots. One peculiar feature of
all the short-jawed species of the genus is the great length of the
molar as compared with the premolar series. In many the
molars occupy quite one half of the tooth-line of the jaw.
Uintacyon promicrodon, sp. nov.
This species is represented in the collection by a single specimen
of an almost complete lower jaw (No. 83), in which the sectorial
and fourth premolar are in good state of preservation ; portions
of the third premolar and second molar, as well as the roots and
alveoli of all the remaining teeth, are present. The species be-
longs in the first section of the genus mentioned above, and is its
oldest known representative, coming as it does from the Wasatch,
The chief distinctions between it and U. canavus from the Wind
River is seen in the size of the fourth lower premolar in com-
parison with that of the first molar. In the species under con-
sideration it is of normal size and holds about the same relations
to the succeeding tooth in this respect as is commonly observed
in the modern Canide, whereas in U. canavus this tooth is much
more reduced.
The remaining premolars were remarkably small and weak, if
we can judge by the size of the roots; the first was implanted by
a single root, while the second and third each had two, The
112 Bulletin American Museum of Natural History. (Vol. XU,
crown of the fourth premolar has a prominent submedian cutting
heel. The length of the lower dental series from the alveolus of
the first premolar is 41 mm.
Uintacyon canavus' (Co/e).
This species is represented by the jaws of four individuals in
the collection, all from the Wind River beds [Nos. 4783 (type),
4784, 4786, 4798]. The type consists of a lower jaw bearing the
first premolar, but displaying the roots of all the remaining teeth.
In the other specimens the crowns of the other teeth are moder-
ately well preserved, so that the lower dentition can be determined.
As compared with the preceding species, the jaw is not quite
so deep and heavy, especially in the region of the symphysis, the
anterior premolars are smaller and more spaced, and the fourth
much more reduced than in the preceding species. Two of the
specimens show the alveolus for the last molar, and indicate that
this tooth was implanted by two distinct roots, whereas in the
preceding species the alveolus is not divided.
In this connection it is proper to mention a specimen from the
Wind River, consisting of the last two molars (No. 84), which does
not agree with either of the Wind River species, U. canavus and
U. brevirostris, in that the last molar is single-rooted, as it is in
the Wasatch U. promicrodon. The premolars belonging to this
type are entirely unknown, and it would not be at all surprising
if another species is indicated by this specimen, If the anterior
premolars are reduced, it undoubtedly represents the direct suc-
cessor of U. promicrodon in the Wind River. Until better
specimens are known, we refrain from proposing another specific
name.
The entire length of the lower molar series of the present
species as indicated by the type is 42 mm.
Uintacyon brevirostris’ (Cofe).
The type of this species consists of a lower jaw (No. 4785)
from the Wind River beds; it is the only specimen known from
this horizon. The jaw is short and deep, as in the preceding
species, and the canine is laterally compressed. The premolars,
1 Bull. U. S. Geolog. Surv. Terr. VI, Feb. 1881, 186.
2 Bull. U. S. Geolog. Surv. Terr. VI, Feb. 1881, 190.
1899.] Wortman and Matthew, Ancestry of the Canidae, etc. 13
of which the crown of the fourth is preserved, show little or no
reduction, differing in this respect from the preceding species in
a marked degree. The second molar, whose crown is preserved
in perfect condition, has a squarish outline with low tubercular
cusps ; the trigon is very slightly raised above the heel, and the
anterior cusp is much reduced. The last molar was distinctly
two-rooted. The length of the molars and premolars is 38 mm.
There are five specimens of lower jaw fragments in the collec-
tion from the Big Horn Wasatch which agree very closely with
the above type and which we provisionally refer to this species.
The premolars Were apparently unreduced, the jaw of the same
depth and shortness, and the last molar strongly two-rooted. In
one specimen (No. 4224), however, the anterior portion of the
second molar is less elevated, and the anterior cusp of the trigon
more reduced than in the type specimen from the Wind River.
Uintacyon vorax Zevay.'
This species 1s not represented in the Museum collection and
the description here given is from the type of U. dathygnathus of
Scott,” which name is undoubtedly synonymous with that of the
above species.
The specimen, like that of the type, is from the Bridger, and
consists of the greater part of both lower jaws, one of which con-
tains the heel of the sectorial and the second molar in perfect
condition. There are also present the proximal end of the femur,
distal end of the tibia and fibula, part of the proximal end of the
ulna, distal end of radius, a complete fifth metacarpal, a part of a
metatarsal, and two phalanges. ‘The original type of the species
consists of a fragment of the lower jaw bearing the second molar,
preserved in the collection of the Philadelphia Academy.
As in the preceding species the premolars are not especially
reduced, and the jaw is relatively deep and short. The last molar
is single-rooted, and the size is considerably greater than in U.
brevirostris ; the length of the lower molar and premolar series
is 51 mm.
The fragments of limb bones are not certainly known to belong
with the jaws, but they seem to agree so well in every respect
1 Proc. Acad. Nat. Sci, Phila. 1872, 277.
?Some Little Known Creodonts. Jour. Acad. Nat. Sci. Phila. Vol. IX, 172.
Fune, 1899. | Po
114 Bulletin American Museum of Natural History. [Vol. XII,
that there is comparatively little doubt that they do. They do
not display any special characters which will serve to distinguish
them from either the early Cats or Dogs. The more salient
features may be stated to be a moderately well-developed third
trochanter on the femur, a very slight grooving of the astragalus,
as well as a well-flattened distal end of the radius, all of which
belong to certain members of the early Canidee and Felide.
Uintacyon pugnax, sp. nov.
This smallest known species of the genus is represented in the
collection by a single nearly complete lower jaw (No. 1744), bear-
ing the second and third molars in good preservation, together
with the roots of the remaining teeth. ‘The jaw is remarkably
short and heavy, especially in the region of the symphysis, the
canine is not laterally flattened, and the premolars apparently
unreduced. ‘The crowns of the second and third molars display
the usual cusps, but the trigons are more elevated than in any of
the preceding species. The length of the molar and premolar
series 1S 27 mm.
Uintacyon edax Zevdy.'
The type of this species and that of the genus is represented by
a moderately complete lower jaw preserved in the collection of
the Philadelphia Academy. ‘The chief characters of this species
are seen in the comparatively shallow, elongated jaw, as well as
its smaller size. As already stated, the type displays an extra
premolar, which is undoubtedly abnormal, since it is placed with
its long axis transverse to the long axis of the jaw. Length of
molar and premolar series, including the extra premolar, 33 mm.;
canine not laterally flattened and premolars unreduced.
Several fragments of jaws from the Big Horn probably represent
this species, but the specimens are so imperfect that the reference
is uncertain.
Prodaphenus scotti,* gen. et sp. nov.
We propose this genus upon a series of upper molars (No.
11,238) of the Princeton collection, together with a lower jaw (No.
1 Proc. Acad. Nat. Sci. Phila. 1872, 277. he
2'This species is dedicated to Professor W. B. Scott, of Princeton, whose contributions to
Palzontology are so well known.
1899.]| Wortman and Matthew, Ancestry of the Canidae, etc. 115
2510) of the American Museum collection, both from the Uinta.
The distinctions between it and Daphenus of the White River
Oligocene are especially seen in the characters of the superior
molars ; in Prodaphenus the external cusps are flanked by a broad
cingular ledge which anteriorly is developed into two distinct
cusps. The unusual extension of this ledge serves to increase the
transverse diameter of the tooth, especially upon its anterior bor-
der, and on this account it resembles
the corresponding tooth of Viverravus
more than that of the Dogs in general.
Another important distinction is the
small development of the postero-
internal cusp, which in Daphenus is
as large asit is in the modern Canide. Me Gass. pee ee RA
In the specimen under consideration #7¢.,,, 1 7Pe specimen. No. 11238,
the tooth is broken in such a way as
not to show this postero-internal ledge very distinctly, but there
can be little doubt that it was present though small.
Its technical distinction from Uzntacyon cannot at present be
given on account of our lack of knowledge of the upper teeth of
this later genus. That it is a direct descendant of Usntacyon, and
the immediate forerunner of Daphenus, there can be little doubt.
The most striking similarity to Daphenus is seen in the low
rounded cusps of the molars, the absence or very small develop-
ment of intermediates, the reduced size of the third superior
premolar, as well as the molar formula above, which is 3.
Until it can be more clearly distinguished from Uzmtacyon the
genus must be regarded as provisional only, but at the same time
there is little probability that the genus Usntacyon continued into
the Uinta without change.
teeta sANCESTRY OF THE DIHOLES.
Cyon, TEMNOCYON, DAPHANUS, PRODAPHANUS, UINTACYON.
1. Cyon and Temnocyon.
The Dhole or Red Dog of India (Cyon) can be confidently
considered as the living representative of the John Day genus
Temnocyon, and through this genus is probably descended from a
116 Bulletin American Museum of Natural History. [Vol. X11
line which we can trace back with tolerable accuracy into the
lower Eocene.
This important connection has been entirely overlooked by
previous writers, who have considered Zemmnocyon to be an abor-
tive side-branch of the Canidz.
The evidence is briefly as follows:
t. Heels of lower molars trenchant. This unusual character is
shared by the recent /cticyon and the John Day genera Oligobunts,
Enhydrocyon and Hyenocyon. All of these have dental formule
excluding them from the ancestry of Cyon.
Fig. 2. Cyon alpinus. Upper and lower teeth, three fourths natural size.
From a specimen in the American Museum collection.
2. Postero-internal cusp of superior molars reduced in Zem-
nocyon, obsolete in Cyon, antero-internal cone reduced, separated
from external cusps by an unusually deep valley, corresponding
to the high trenchant heel of the lower molars.
3. Last lower molar reduced in Zemnocyon, absent in Cyon.
Last upper molar somewhat reduced in Zemnocyon, considerably
1899.] Wortman and Matthew, Ancestry of the Canidae, etc. 17
so in Cyon. Premolars unusually large; m® unusually small,
with internal cusp of the trigon reduced in Zemmnocyon, absent in
Cyon, Jaw unusually deep under premolars.
The two are separated by the usual progressive characters seen
in all modern Canide ; the hallux is much reduced, mt. 11 fall-
ing from $ to + the length of mt. m1. The pollex suffers a
similar reduction from $ to+of mc. 1. The foot is elongated
and narrowed, the brain cavity increased, the skull shortened.
Premolar 2 has acquired a posterior cusp in Cyon, lacking in
Temnocyon ; the internal cusp of the superior sectorial is more re-
duced in the modern species. In our skull of Cyon alpinus the
incisors show a small basal lateral cusp. All these are acquired
characters, most of them being developed in almost all species of
Fig. 3. Zemnocyon ferox Eyerman. Upper and lower teeth, three fourths natural size.
After Eyerman.
dogs. We find no acquired characters in Zemmnocyon that Cyon
has not, nor any primitive characters in the latter that the former
has not.
Cyon, with its congener /cficyon, retains more than any of the
other dogs the primitive short legs and long body. ‘The tail is
much reduced in both genera.
2. Related Genera.
Related to the Dhole and Zemmnocyon, and distinguished like it
by the trenchant heels to the molar cusps, are the modern /efzcyon,
118 Bulletin American Museum of Natural History. (Vol. XVI,
and the John Day genera Lnxhydrocyon, Hyenocyon and Oligobunts,
all with a more reduced dental formula. They form the most
carnivorous division of the family, approximating the Viverride
or Felide, while Vothocyon and AMega/otis stand at the other ex-
treme, approximating the Procyonide.
3. Daphenus and Temnocyon.
The relationship between these two genera has been pointed
out by Prof. Scott’ and Dr. Eyerman.’ All our evidence tends to
confirm the view that Zemnocyon is a descendant of Daphenus,
although the gap between them is in some respects considerable.
Both have the same strong, round-edged, massive cusps, but in
Daphenus the molars are large and low, while in Zemmnocyon they
are small and high-cusped. Daphenus is just beginning to de-
velop a trenchant heel on m 7-3; it has the same deep jaw, and
does not show any intermediates on the upper molars.
Hypotemnodon has a somewhat problematic position intermediate
between Cynodict’s and Daphenus in most of the skull characters
but with trenchant heel on m-+ only. Scott derives it from
Daphenus ; there is, however, a species of Cynodictis in the upper
White River, which presents a closer approximation in character
of teeth, and if our association of specimens is correct, has de-
veloped a low trenchant heel on mz only. For this reason it is
placed provisionally in the Cynodictis line. :
Prof. Scott derives Cynodesmus also from Daphenus, basing the
derivation apparently on the presence of frontal sinuses in both.
Whether the presence of these in the larger species of Canidz is
of much phylogenetic value seems uncertain ; on other grounds
Cynodesmus might well be connected with Cynodictis.
4. Prodaphenus and Daphenus.
5. Uintacyon and Prodaphenus.
These have already been considered.
I1]1.—Tue Eocene ANCESTORS OF CYNODICTIS.
Vulpavus palustris * J/arsh.
With this species we come to consider the phylum which im all
probability terminated in the modern genus Camis. Its oldest
1 Notes on the Canidz of the White River Oligocene. Trans. Am. Phil. Soc. 1898.
2 Amer. Geol. 1896.
3 Amer. Jour. Sci. Aug. 1871, (p. 16 of separate),
1899.| Wortman and Matthew, Ancestry of the Canida@, etc. 119
known representative comes from the Wind River beds, and is
known from only an imperfect fragment of a lower jaw in the
Museum collection. In the Bridger the genus is represented by
at least two species, one of which, V. palustris, was described by
Marsh from a single superior molar and forms the basis of the
genus ; and the other, V. parvivorus, was described from a frag-
ment of a lower jaw by Cope and made the type of his genus
Miacis. In the Museum collection is a specimen which upon
careful comparison with Marsh’s type of V. palustris we identify
with this species ; 1t consists of two superior molars of the right
side, two nearly complete mandibular rami, together with some
fragments of the skeleton of the limbs (No. 2305). It was found
by Mr. O. A. Peterson of the Museum party of 1895, and was
obtained from the middle horizon near the extreme southern
limits of the Washakie Basin.
Fig. 5. Vulpavus pa-
Zustrts Marsh. Upper
Fig. 4. Vulpavus palustris Marsh. Lower jaw, natural teeth, twice natural
size. No. 2305. size, No. 2305.
The superior molars are two in number and display about the
same proportions as those of many of the existing Canide. They
present a number of features, however, very different from the
modern Dogs. The first molar is remarkable for the great elonga-
tion of its anterior side in comparison with the posterior, and the
drawing out of the antero-external angle, as well as the unusually
broad ledge intervening between the base of the external cusps
and the outer border of the crown. Another marked feature of
this tooth is the great disparity in size between the anterior exter-
nal and the posterior external cusps. ‘The antero-internal cusp is
large and lunate and there is a distinct anterior and posterior in-
termediate. The postero-internal cusp is represented by a strong
cingulum in the usual position but it does not rise up into a dis-
tinct cusp as in the later Canide. The second molar is similar
120 Bulletin American Museum of Natural History. [Vol. XII,
except that there is no representative of the postero-internal
cusp. The general appearance of these teeth is more like that of
many Creodonts than that of the modern Carnivora.
The lower jaw is very dog-like in its general proportions ; the
symphysial region is slender and the ramus is elongated. The
teeth resemble those of the early Dogs, especially Cynodictis ; the
canine is long and pointed, being separated from the anterior
premolar by a short diastema. The premolars have the usual
form in the Canidz, with compressed pointed crowns and basal
cingula ; the fourth has a distinct posterior accessory cusp. The
sectorial closely resembles that of Cynodictis. The anterior and
external cusps of the trigon form an effective shearing blade ;
the internal cusp is large and the heel basin-shaped. The second
molar is similar in structure but its cusps are lower and there is
no distinctive blade-like shear produced by a union of any of the
cusps of the trigon. The last molar is much reduced, as in the
later Canide.
Some fragments of the limb bones are preserved and they in-
dicate, as do the teeth, a clear affinity with the early Dogs. The
distal end of the radius is present, but the facet is cup-shaped
Fig 6. Vulpavus palustris Marsh. Humerus, natural size. No. 2305.
and exhibits no distinct ridge separating scaphoid and lunar
facets. This is not, however, conclusive proof that the scaphoid
and lunar were not separate, since in Viverravus, Oxyena, and
other Creodonts in which these bones are distinct, the distal end
of the radius gives little or no indication of the facet.
Vulpavus parvivorus’ (Cofc).
This second species is represented in the collection by a frag-
ment of a lower jaw from the Bridger of Wyoming, and has been
figured by Cope as the type of his genus A/zacis. The fragment
1 Proc. Amer. Philos. Soc. Aug. 1872, 470.
1899.| Wortman and Matthew, Ancestry of the Canide, etc. 121
carries the second molar in an excellent state of preservation, and
exhibits the alveolus for the last molar and a portion of that of
the first. The species can be readily distinguished from V, palustris
by its smaller size and the less laterally compressed character of
the second molar.
Procynodictis vulpiceps, gen. et. sp. nov.
This genus is proposed upon two specimens in the Museum
collection, one of which (No. 2514) includes one upper and both
lower jaws, together with the greater part of a hind foot, and the
other (No. 2506) includes a part of the skull and the greater
portion of the right fore foot. Besides these there are two frag-
ments of jaws (Nos. 1895 and 1995) which we refer to the same
species.
The dental formula is the same as in Vudpavus and Cynodictis,
there being but two true molars above, and the systematic posi-
tion of the genus is entirely intermediate between these two. In
the structure of the superior molars it agrees with Vu/pavus in
that there is a great extension of the antero-external part of the
Fig. 7, Procynodictis vulpiceps W.& M. Upper and lower teeth, three halves
natural size. Type specimen No. 2514.
tooth and a comparatively small development of the postero-in-
ternal cusp. In Cynodicti’s this cusp is as well developed relatively
as in the modern Dogs, but the great antero-external extension is
122 Bulletin American Museum of Natural History. [Vol. XI,
lacking. The only important character in which it differs from
Vulpavus is seen in the nearly equal size of the two external cusps
of the first superior molar, whereas in Vu/pavus the anterior
greatly exceeds the posterior in size. This at first sight would
seem to be of little significance, but the primitive condition was
undoubtedly the one displayed by Vudpavus, and the more ad-
vanced one that of Procynodictis and Cynodictis. It therefore
represents a distinct advance. The other teeth are very similar
in their structure to those of Vu/pavus and need no special
mention.
The hind foot contains five toes; the metapodials are shorter
and somewhat heavier than in Cyzodictis, in which the hind foot
had already begun to assume the elongated form so characteristic
of the modern Canide. The tarsal bones are very like those of
Cynodictts, as are also the remainder of the podial elements. The
claws are much compressed laterally, with heavy subungual pro-
cesses, and the middle phalanges are unsymmetrical, indicating
some degree of retractility of the claw.
The fore foot also contains five toes, but, as in Cynodictis, it is
much shorter than in any of the modern
Canide. The scaphoid, lunar, and cen-
trale are completely co-ossified as in the
modern Dogs, without any trace of suture,
into a scapholunar, which differs from that
of Cynodictis, according to Scott’s figure,
in the greater vertical depth anteriorly,
which is again a more primitive condition,
The other carpals have practically the
same proportions as in Cynodictts. The
fore foot of Vudpavus is unknown, but it is
more than probable that the scaphoid and
Der Rr a iin ees Ade lunar were separate in that genus, which if
ae halvesnaturalsize. No. true will constitute an important difference
between it and the present genus.
IV.—NOoTES ON A SKELETON OF CyYNODICTIS
GREGARIUS.
In the American Museum Collection, Expedition of 1898, is a
nearly complete and very well-preserved skeleton of Cynodictts
1899.] Wortman and Matthew, Ancestry of the Canide, etc, 123
gregarius (No. 8774). Prof. Scott has already fully described
nearly all parts of this species, but we are able to fill out the
description in one or two points for which his material was in-
complete, and to give a series of measurements, which, being
taken all from one individual, give the relative proportions with
greater accuracy. Some of the individuals which Prof. Scott
includes under C. gregarius may be referable to C. “éppincottianus.
Hind Foot.—Vhe tarsus has already been described in full by
Prof. Scott. There are five mefatarsals, the first slender, its
length two thirds that of mt. 1, diameter of the shaft half as
great. Head rather small, shaft nearly straight, slender, rather
compressed. Second, third, fourth, and fifth metatarsals sym-
metrical, the central pair a little longer and about one fourth
greater in diameter of shaft. The distal articular ends are
spheroidal, not at all of the square-cut style seen in all modern
Dogs. ‘The upper part of the shaft is somewhat compressed, the
distal part not at all, and the distal ends are much more enlarged
than in Canis, the diameter being one third more than that of the
shaft, which in Cams increases in diameter at the distal end and
is not over one tenth of the shaft diameter. The foot was therefore
more spreading, as well as 15 per cent. shorter. The bones are
as slender, however, asina modern Dog. The phalanges are long
and slender, quite as long as in modern Dogs. The second pha-
lanx of each toe is asymmetrical and excavated, as in Daphenus,
indicating, according to Scott, some retractility of the claws. No
strong basal sheath appears to have been on the unguals, which
are much as in Canis.
As the measurements indicate, the hind limb bones were much
nearer to their present proportions, while the fore limbs were still
quite short. The difference in the feet is yet more marked, for
while the fore foot was scarcely more elongated than the Creo-
dont ancestors, the hind foot was already long and slender,
though not as narrow nor quite as long as in modern species. In
both feet, however, the tetradactyl symmetry is already distinct,
and both pollex and hallux are reduced.
The measurements of the skeleton, No. 8774, as far as at
present obtainable, are given in comparison with those of the
Fennec, the nearest living Canid in point of size.
124 Bulletin American Museum of Natural History. [Vol. XII,
Measurements of CYNODICTIS GREGARIUS (Cofe), in Comparison
with CANIS ZERDA.
Cy. gregarius. Ca. zerda.*
Skull, length, premaxillz to condyles... est. 76 mm. 8r mm.
“= » lengthy of palatel(dentition)/ se -) ) 9) 42 45
“Sey breadth yyy se") | « stauscmrssissieraere tee aoe D7 24,
ay ae brainicasevesiaccas cies ers 29 35
Mertebrecy allipre-caudalsiera-e ere nie 291
oS Cervicalls uaa wipes tune cecetert 72
“s GOLSAIS 5 sreyateneyerh. oraetetes stele acche 113
InN ENS o God ado aod sde00de oOe 105 go
os SACHA SWereye fen eersvetave aitersice tate) ens) bee 23 16
e caudal 35 \s's.tyeee the cise i etecsits Oe 350
Hore limbritotalislencthyeerrrisreirncer 180 230
ba INU ETUS Sy jare et sccyeiversncheseueye.s oe 71 80
% TAGIUS sjeyerss eat ate setae tue misters 56 79
ne POOLE ai. sys aiattete-tyaletste, sevetsiers oun elle 53 7
© CED ss ono odode oCnDadaC 6 esi h2
a MNELACATPSaehetete steerer Nelsieialeye 22 28
es PhalanCeswerrtmrey ttle Git & est. 25
Find limb, itotal men oths s.r). ei cise 241 272
oe PEMMUT hie metas cocteaeweecee sare 80 81
is tibialis mrecpoe ste seuerele cao eres 81 98
oe foot (astragalus to ungues). .. 80 93
a3 LATSUS cotashs is eyeielsees eeu, « 1g est. 22
ea MMEEATATSUS rete piu cask apeieereree 34 43
3 PIENNESo ts oonsanaanaeoaac 27 OF
Taking the above measurements in order, we see that the skull
of Cynodictis is shorter and with smaller brain-cavity. The neck
and back vertebre are nearly of the same size as in the Fennec,
but cannot be measured at present. ‘The lumbars and sacrum
are considerably larger, and the tail was probably as long (no
other living Dog has a tail of this length). The fore limb is one
fourth shorter, while the hind limb is only one ninth shorter.
The greater part of the elongation has been in the lower limb and
foot bones. The phalanges have not lengthened at all, there be-
ing no tendency to become unguligrade.
V.—Tue ANcESTRY OF CERTAIN SOUTH AMERICAN
FoxEs.
Nothocyon, gen. nov.
A genus or subgenus of Dogs distinguished by :
(1) Short muzzle; (2) upper carnassial very small, trigon of
lower carnassial reduced, shear partly transverse; (3) molars
1 Measurements taken from Mivart’s Monograph of the Canidz.
1899.] Wortman and Matthew, Ancestry of the Canid@, etc. 125
large, not extended transversely, subquadrate ; (4) lower carnas-
sial broad-heeled, and with an accessory cusp at the postero-ex-
ternal corner of the trigon; (5) canines slender; (6) otic bullee
large.
Cants urostictus Mivart, and C. parvidens Mivart, both from
South America, belong to this genus, to which we also refer pro-
visionally the three John Day species, /attdens Cope, lemur Cope,
and getsmarianus Cope. ‘These latter seem to be directly ances-
tral forms, and, like all the earlier Dogs, have shorter feet, longer
lumbar region, and smaller brain-case than the modern species.
The distinctions above noted may seem hardly to be of generic
value; they are, however, tolerably constant, Urocyon cinereo-
argenteus being the only intermediate type. And if the group
was first separated in the Oligocene, as seems probable, it gives
it an ancestry that deserves full generic recognition. The alter-
native to uniting the John Day and modern species is to place
the one as a subgenus of Cynodictis, the other of Canzs ; the pres-
ent method seems, however, to recognize more clearly the actual
genetic affinities ; for if the living Cans parvidens and urostictus
are lineally descended from the John Day Cynodictis latidens
lemur, and geismartanus, the two modern species are more nearly
related to the three Miocene species than to any living Canide.
The best way to express this fact is to remove the Miocene species
from Cynodictis, to the typical forms of which they are not more
nearly related than the half-dozen or more distinct genera of the
Phosphorites of France, and to unite them with two modern
species, disregarding the considerable modernization of the latter,
which retain, however, an unusual amount of the Tertiary facies.
Nothocyon urostictus (M/vvar?).
Canis urostictus MIVART, Proc, Zoél. Soc. Lond, 1890, 112; Monograph
of the Canidz, 81.
The type and only specimen hitherto known is in the British
Museum. No. 391, Dept. Osteol., Amer. Mus. Nat. Hist., we
refer to this species and figure here. In size it is not very dif-
ferent from Canis azare, but the size and characters of the
molars and carnassial easily distinguish it. The lyrate area on
top of the skull is shared by many species of Dogs. Limbs and
feet about as in modern species of Canis ; the lumbars appear to
126 Bulletin American Museum of Natural History. [Vol. XII,
be larger in proportion. Locality, Chapada, Matto Grosso,
Brazil.
v8 on Hie
+)
nea v7
Wy i i.
Rae y. y
“UG
YW),
©, \y
Ze oS
Fig. 9. Wothocyon urostictus (Mivart). Skull and jaw, five sevenths natural size.
No. 391.
Nothocyon parvidens (J/7var‘?).
Canis parvidens MIVART, loc. cit.
The only specimens hitherto known are three in the British
Museum. We are enabled to add another skeleton, No. 349,
1899.] Wortman and Matthew, Ancestry of the Canidae, etc. 127
Dept. Ost., Am. Mus. Nat. Hist., on the somewhat uncertain den-
tal distinction separating it from JV. wrostictus. P-* is shorter,
canines not,so slender, mg more reduced, m+ and m2 seem to be
variable inshape. No. 2091 also belongs to this or the preceding
species. Locality, Chapada, Matto Grosso, Brazil.
As another division (Primeevi) of the same genus, distinguished
by relatively small brain, short feet, and long lumbar region, we
place the three John Day species.
Nothocyon latidens (Cofe).
Galecynus latidens COPE, Bull. U. S. Geol. Surv. VI, 1881, 181. '
Cynodictis latidens ScoTT, Trans. Amer. Phil. Soc. 1898, 400.
Distinguished by exceptionally broad upper molars. Bulle
very large.
Nothocyon lemur (Coc)
Galecynus lemur Cork, Bull. U. S. Geol. Surv. VI, 1881, 18r.
Cynodictis lemur Scott, Trans. Amer. Phil. Soc. 1898, 400.
Molars narrower, cusps marginal, double entoconid on mj, pos-
tero-external accessory cusp on mg doubtful. Bulle very large.
Nothocyon geismarianus (Cofe).
Canis geismarianus COPE, Pal. Bull. No. 30, 1879, 9.
Cynodictis geismartanus SCOTT, loc. cit.
Larger species, deuterocone of p* more reduced than in the
others. Bulle of more moderate size.
The skeleton of this species is fortunately known and _ has
been described by Cope. It stands intermediate between that of
Cynodictis gregarius and that of Wothocyon urostictus. The follow-
ing comparisons show the advance in the especially progressive
characters of (1) length of feet, (2) length of lumbar region,
(3) size of brain.
Cynodictis gregarius, width of carpus, 15 mm.; length of mc. III, 22 mm. ;
proportion, 1: 1.5,
Nothocyon geismarianus, width of carpus, 16.5 mm.; length of met. II,
33 mm. ; proportion, I: 2.
Nothocyon urostictus, width of carpus, 18 mm.; length of mec. 11, 48 mm. 5
proportion, I: 2.7.
In the same species we have :
128 Bulletin American Museum of Natural History. [Vol. XII,
Cynodictis gregarius, length of skull, 76 mm. ; width of brain-case, 29, height,
22, length, 43 mm.; proportion of length of skull to width of brain-case,
1 :0.37; length of seven lumbars, 104; proportion of length of skull to that of
lumbar vertebrez, I :1.37.
Nothocyon geismarianus, length of skull, 111 mm. ; width of brain-case, 40,
height, 33, length, 52 mm. ; proportion of length of skull to width of brain-
case, 1: 0.37; length of seven lumbars, 155; proportion of length of skull to
that of lumbar vertebrz, I : 1.39.
Nothocyon wrostictus, length of skull, 112 mm. ; width of brain-case, 45,
height, 35, length, 52 mm. ; proportion of length of skull to width of brain-
case, 1: 0.41; length of seven lumbars, 129; proportion of length of skull to
that of lumbar vertebre, I : 1.15.
From the above proportions it appears that the John Day
species exhibits an important advance on the White River Cyno-
dictis in the increase in length of feet and decrease in their width.
The increase in the size and robustness of the species marks the
advance in brain capacity, and the lumbar region has not suffered
any proportionate reduction. In the modern species there is a
further increase in length of feet, the brain increases in capacity,
and the lumbar region is greatly reduced. The length of the
femur and humerus does not increase in proportion; that of the
tibia increases slightly and of the radius considerably.
Proportions, length
C. gregarius, N. getsmartanus. N. urostictus, of skull as unity.
Length humerus.. 71 98 106 -93: .gO: .94
“OS radius 55 ? 105 572 eeLOd!
Se femureer 79 110 116 1.03 : 1.00; 1.04
ce Stiblae uses 80 116 120 1.04) 1, 04021 O77,
The above measurements are from the complete skeletons of
C. gregarius (No. 8774), WV. geismartanus (No. 6886), VV. urostictus
(No. 391, Dept. Ost.).
The longer-legged specimens referred by Prof. Scott to C. gre-
garius may be referable to C. “ippincottianus. These have the
limbs, especially the tibia and fibula, much longer than in C,
gregarius proper, but the feet apparently of the same size.
VI.—ANALYSIS OF THE MIOCENE GENERA OF CANIDA.
It is doubtful whether some of these genera should rank as
such, the distinctions being probably of only subgeneric value.
They certainly represent well-defined groups, and it would be
difficult to invalidate one without invalidating nearly all. The
mere zumber of the teeth cannot be considered as a more im-
1899.] Wortman and Matthew, Ancestry of the Canida, etc. 129
portant character than their form, and unless certain characters
are arbitrarily selected as generic, the representation of natural
groups is the only valid foundation for generic distinctness. As
a mere matter of convenience, the groups are here used as genera,
with the proviso that they are not all of generic value. Phlaocyon
stands widely apart from the rest, as does also Oligobunis ; but
whether two, three, or several genera are made out of the remain-
ing species is a matter of arbitrary selection rather than natural
grouping.
I. PROCYONIDA,
Skull short and wide, orbits well forward ; carnassial teeth imperfectly or
not sectorial; a hypocone on p*. Pentadactyl, plantigrade. Dentition typi-
ae
3-I-4-2
A. Third lower molar absent, second elongated. Modern genera Bassaris-
cus, Procyon, Bassaricyon, Nasua, Cercoleptes.
8B. Third lower molar present, second not elongated.
3-1-4-2
cally
Phlaocyon. WDentition
II. CANIDz.
Becoming tetradactyl, digitigrade. Skull long, orbits not advanced, teeth
sectorial and tubercular.
A. Short-footed, pentadactyl, with shear of sectorial partly transverse, small
brain and other primitive characters.
Sntsds3
3-1-4-3,
a. Daphenus Leidy. Upper molars transversely unsymmetrical (paracone
more external than metacone). M® oval, aligned with inner cusps of anterior
molars. Heels of lower molars low-ridged, with low entoconid crest, mg a
1. Dental formula
convex nub.
D. vetus Leidy. Amer. Mus., Nos. 1388, 1390.
D. hartshornianus (Cope). Amer. Mus., Nos. 6811, 1387.
D, felina Scott.
D, dodtget Scott.
- The first three species are from the Oreodon beds, the last from the Titan-
otherium beds.
6. Paradaphenus, gen. nov. Upper molars much extended and symmetrical
transversely ; m® aligned with outer cusps of anterior molars. Heels of lower
molars wide and deep, basin-shaped ; mg and mg with two anterior cusps and
basin-heel.
P. cuspigerus (Cope). Nos. 6852 (type), and probably) 6853 (type of
Amphicyon entoptychi Cope).
P. transversus, 1. sp.
[ Fune, 1899] 9
130 Bulletin American Museum of Natural History, (Vol. XII,
Size one third larger (lineal). Upper premolars compressed.
Type No. 6851 (referred to Amphicyon hartshornianus by Cope).
Both species are from the John Day,
3-1-4-2
3-1-4-9)
a. Temnocyon Cope. Heels of lower molars trenchant, without internal ridge.
Trigon of ms trenchant, without internal cusp.
T. altigents Cope. Nos. 6855 (type), 6856.
T. wallovianus Cope. No. 6858 (type).
T. ferox Eyerman. No. 6857.
6. Hypotemnodon Eyerman. Heels of lower molars trenchant, internal ridge
vestigial or wanting. Trigon of mgs with prd and med of equal size.
1. corypheus (Cope). Nos. 6859 (type), 6860, 6862, 6922, and doubtfully
Nos. 6861, 6920, 6909, etc.
#1. josephi (Cope). Nos. 6878 (type), 6863, 6908, 6921.
All from the John Day,
c. Cynodictis Bray, and Pom. Heels of lower molars basin-shaped. Upper
molars extended transversely. The American species are :
C. gregartus (Cope). Nos. 5297 (type), 5298, 5299 (fig. sp.), 1004, 1383,
1472, 8774, 5300. No. 6879 is probably a distinct species but does not pre-
sent any well-marked specific characters.
C. lippincottianus (Cope). Nos. 5327 (type), 8757-63, 1384, 1389, etc.
Teeth one fifth greater in lineal dimensions, somewhat more robust. Sectorial
proportionately larger, shear less transverse.
? C. temnodon,n.sp. Size of Hypotemnodon josephi ; lower premolars smaller
than in that species and trigon wider ; p+ (in associated upper jaws) with internal
cusp almost obsolete, shear rather more longitudinal than in Cyzodictis ppin-
cottianus, m+ well extended transversely.
C. gregarius and C. lippincottianus come from the Oreodon beds (except
the one skull No. 6879, of doubtful reference, which comes from the John Day).
C. temnodon is from the Protoceras beds.
ad. Nothocyon (section Primzvi). Upper molars short transversely ; p*
small; lower molars with wide basin heels, trigon of mz reduced, and a small
accessory cusp at its postero-external corner.
LV. latidens (Cope). Nos. 6896 (type), 6897-99.
WV. lemur (Cope). Nos. 6888 (type), 6889-94.
All from the John Day.
3. Premolars reduced.
a. Enhydrocyon Cope. Dentition 3-1-3-2.
2. Dental formula
E. stenocephalus Cope. Nos, 6901 (type), 6902.
John Day.
3-I-3-1
ee
fT, basilatus (Cope). No. 6904 (type).
fH, sectorius (Cope). No. 6905 (type).
John Day.
b, Hyenocyon Cope. Dentition -
1899.]| Wortman and Matthew, Ancestry of the Canide, etc. 131
4. Molars reduced.
Oligobunis Cope. Dentition setae
3—-I-4—2
O. crassivultus (Cope). No. 6903 (type).
John Day.
B. Modernized species. Long-footed, tetradactyl (functionally), with shear
of sectorials nearly longitudinal, large brains and other advanced characters.
Salsa.
3-I-4-4
3-1-4-2
35154-3
3. Dental formula 34”. Cyon.
3-1-4-2
t. Dental formula Otocyon,
2. Dental formula Canis, Nothocyon, Lycaon, Urocyon.
4. Dental formula seer
3-I-4-2
Icticyon.
VII.—Awn ANCESTRAL Raccoon.
Phlaocyon leucosteus J/atthew’
Founded on an exceptionally perfect skull and jaws, with a
nearly complete skeleton, discovered last summer by Mr. Handel
T. Martin of the American Museum Expedition of 1898 in north-
eastern Colorado. The level is the uppermost beds of the White
River formation, associated with Merycocharus, Anchippus, Lep-
tauchenia, Hyracodon, etc. ‘The specimen was found in the rock
mixed with skulls and skeletons of two adult and three young
Merycocheri, all within a space of six feet square.
It represents a new and aberrant genus of Dogs, the characters
pointing clearly in the direction of the Raccoons, so that if we
adopt the genealogical conception of a family it must be placed
in the Procyonide, although it is nearer to such primitive Dogs
as Cynodictis than to the modern Raccoons.
Dentition—(1) There is a small but clearly marked postero-
internal cusp on the upper sectorial, which, however, (2) retains
the triangular shape characteristic of the early Canide. (3) The
dentition is that of Cynodict’s and Canis, (4) but the cusps are low
and rounded. (5) Premolars small, stout, and crowded ; (6)
upper molars short transversely and subquadrate, as in Cynodon,
(7) the lower molars broad and low. (8) Canines short, the upper
ones not dagger-shaped, but curved as in the Dogs. (9) Incisors
in an even, transverse row. Yaz (10) short and thick, deep in
i Bull. Am. Mus. Nat. Hist. 1899, 54.
132 Bulletin American Museum of Natural History. (Vol. XII,
front, (11) condyles very wide, (12) coronoid process short and wide
with deep fossa. Sku// (13) short and wide, (14) orbits placed as
far forward as in Procyon, (15) jugal process of squamosal not
reaching as far forward as postorbital process of malar. (16) Shape
and proportions of skull and jaws resemble those of Procyon lotor,
except that (17) the brain-case is much smaller proportionately.
(18) An alisphenoid canal as in Cynodictis. (19) A median and
two lateral foramina in palate between canines. (20) Palate not
extended posteriorly. (21) Base of skull much less broadened
than in Procyon, (22) paroccipital process not developed.
Of the above Nos: st; 4,5;°0; 10, bi, 112,13, 14,505, 16, andere
are progressive characters in the direction of the Raccoons. Nos.
2, 3, 6, 7, 8, 17, 18, 20, 21, 22, are primitive characters, shared by
all early Canide.
Skeleton.—The hyoid bones were found perfectly preserved,
but offer few characters for distinction.
The atlas is like that of Cynodicti’s ; the posterior opening of
the vertebral canal presents a little upward instead of directly
upward as in recent Canide, or directly backward as in Procyon,
where the opening is bounded by a strong ridge passing outward
from the axial cotylus.
Concerning the other vertebrz no exact account can be given
at present, as they are not yet removed from the matrix; they
resemble in most respects those of Cynodictis. ‘The ribs are
somewhat larger ; the dorso-lumbar formula is not yet known.
Fore Limb.—Vhe humerus is of the same length as in Procyon
Zotor, but more slender throughout. The greater tuberosity is
higher and the deltoid crest is more marked and extends farther
down. The radius and ulna are 15 per cent. less in length and
of about the same diameter in the shaft. Shafts of both trihedral
or irregular in cross-section, instead of regularly oval as in modern
Raccoons. Carpus with co-ossified scapholunar, thin cuneiform,
large unciform, rather small magnum, rather large trapezoid, and
small trapezium. Five metacarpals, of which mc. 1 is only half
the length or diameter of 111; me. Vv is three fourths as long, but
of considerably greater diameter than mc. 111; mc. 11 and Iv
equal in diameter to, but a little shorter than, mc. III.
Hind Limb.—TVhe femur is of nearly the same length as that of
P. lotor, and one fifth slenderer. The shape resembles Cynodic-
1899.] Wortman and Matthew, Ancestry of the Canide, etc. 133
tis ; the greater trochanter is much less prominent than in Pro-
cyon, the lesser trochanter more prominent, the neck longer and
more slender. The distal end of the femur is only three fourths
as wide as in Procyon ; condyles projecting more than in that
animal, less than in Canis and Cynodictts.
The fdéa is one sixth shorter than that of P. dotor, somewhat
more slender in proportion, stouter than that of C. gregartus and
of the same length. Its distal trochlea is narrower antero-poste-
riorly, deeper and more oblique than in Procyon ; the fibular
facet also is twisted around so as to face postero-externally instead
of externally as in Procyon. The fibula is somewhat less reduced,
its shaft being as stout as in the Raccoon, the ends somewhat
smaller.
The hind foot, though uniformly smaller, is proportioned like
that of the Raccoon, with the following differences worth noting.
The astragalar trochlea is narrower, sharper, less entended back-
ward. The internal hook of the zavicular is smaller. The cuboid
is wider and has a small facet for the astragalus. The evtocuner-
form is not nearly so high. The first sefatarsal is only two
thirds as long, though somewhat stouter in proportion ; and the
fifth is as long as the second, and nearly as large in the shaft ; in
the Raccoon it is intermediate between mt. 1 and mt. 11 in length.
The hind feet of PA/aocyon have thus a tetradactyl symmetry,
while in Procyon they are intermediate between tetra- and penta-
dactyl symmetry. The second row of phalanges shows a distinc-
tion of some importance, the superior external surface of the
shaft being excavated, especially toward the distal end, a
character which Scott has observed in Daphenus and considers
a probable indication of slight retractility of the claws. It is also
present in Cynodictis and in some Creodonts, and is probably a
primitive character. In the Raccoon this excavation of the
second phalanx has entirely disappeared.
The ungual phalanges are a little larger and less compressed
than those of Procyon lotor.
AFFINITIES OF PHLAOCYON.
PLATE VI.
Phlaocyon, in both skull and skeleton characters, seems to
point towards Procyon, standing intermediate between that genus
34 Lulletin American Museum of Natural History. |Vol. XII,
and Cynodictis in almost every character. The entirely intermedi-
ate character of the skull and teeth is well shown in our drawings:
especial attention should be drawn to the postero-internal cusp
on p*; short jaws with deep muzzle and round blunt premolars ;
broad low sectorials, with additional postero-external cusp on
trigon of mz; reduced mz; lower incisors in a straight row.
Eyes set far forward, skull short, arches wide. On the other
hand the resemblance to Cymnodictis is closer in the form and cusp-
arrangement of the teeth ; palate terminating opposite m4, ali-
sphenoid canal; occiput not so much expanded ; retention of
mz; My not lengthened out.
In the ‘skeleton the same story appears. The hmb bones are
like those of Cynodictis with varying amounts of change to Procyon:
Only in the hind foot is there any difficulty. Here the first toe
is more reduced in Ph/aocyon than in Procyon, which makes it
seem probable that P. /eucosteus is slightly off the line from
Procyon lotor.
The resemblance to Bassariscus is even closer. But Bassariscus
cannot be considered as the living representative of Phlaocyon.
It shows less departure from Cymodictis in many characters, es-
pecially in the shape of the upper molars and lower sectorial and
slender jaw ; the postero-internal cusp of the upper sectorial is
not developed in one species, and although it is present in the
other, the tooth retains much more of its trenchant function.
The loss of mg is an advance on Ph/aocyon ; on the other hand,
the eyes are not set so far forward, nor is the muzzle so much
shortened or the arches so wide. The brain is very much larger
in proportion, as might be expected, but the feet are precise
copies of those of Cynodictts.
With WVasua the comparison is not so close as with Procyon.
The long skull, large dagger-shaped canines, comparatively nar-
row, square occiput, reduced and peculiarly shaped bulle, carry
it out of the probable line of evolution pursued by PAlaocyon.
The limbs and feet are somewhat nearer to Ph/aocyon in some
characters ; the feet have the internal toe unreduced.
Cercoleptes is much more aberrant in the skull and teeth and
tHlurus has a different pattern of teeth. Altogether Procyon
seems to come nearest to fulfilling the conditions required of a
descendant of Phlaocyon.
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1899.] Wortman and Matthew, Ancestry of the Canidae, ete. 135
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136 Bulletin American Museum of Natural History. [Vol. XII,
VIII.—VIvVERRAVIDA, fam. nov.
This family is proposed to include the species which have been
formerly arranged under the genus Didymictis of Cope. That this
genus is synonymous with the one previously established by
Marsh, there cannot be the slightest question whatever. A care-
ful comparison of the type of Viverravus gracilis Marsh,’ with
the type of Didymictis dawkinsianus* Cope, reveals the fact that
they are identical in every detail; the name Didymictis must
therefore be abandoned and the name Viverravus substituted for
the genus.
It has been customary to associate this group of species, together
with that which has been referred to Uzutacyon (Mziacts), in the
family MWiacide. We have already shown that this latter is clearly
related to, and no doubt ancestral to the Daphenus, the Cyon
section of the Canidz. It is also evident from much additional
material now in the Museum collections that Viverravus belongs
to an entirely distinct line and has no relationship with the Canidz
whatever, but in so far as the skeleton can be depended upon for
evidence of affinity it finds its nearest relationship with the living
Viverridz. This opinion is not a new one, having been first ex-
pressed as long ago as 1886,° and later in 1891.*
The new evidence consists of the greater part of a skeleton of
Viverravus protenus from the Wasatch Eocene of Wyoming, to-
gether with a considerable portion of a skeleton of V. leptomylus
from the same horizon. ‘The skull, of which there is a fairly well-
preserved example, shows many striking resemblances to the more
typical Viverrines, especially Viverricuda ; it is long and narrow,
with prominent overhanging occiput and high sagittal crest. The
muzzle is long, the basilar portion of the skull is narrow, and the
mastoid not very prominent. ‘The tympanic bulla is not preserved,
but the general conformation of this part of the skull is very
Civet-like. ‘The lower jaw is long and relatively slender, with
high coronoid and prominent angle. The dental formula, as is
well known, is that of the Viverride, and the pattern of the teeth
resembles most astonishingly that of the more typical members of
1 Amer. Jour, Sci. 1872 (p. 7 of separate).
2 Bull. U.S. Geol. Surv. VI, 1881, rgr.
3 Wortman, Teeth of the Vertebrata, 457.
4 Flower and Lydekker, Mammals Living and Extinct, 539.
1899.]_ Wortman and Matthew, Ancestry of the Canida, etc. 137
this family. This is especially seen in the superior sectorial, in
which there is a deep vertical notch separating the two halves of
the blade, and a prominent anterior basal cusp, which Is invariably
present in the carnivorous Civets. The superior molars, more-
over, have the same characteristic pattern, and the lower teeth,
with the exception of some unimportant details, are very viver-
rine. The atlas has the same arrangements of the perforations
for the vertebral artery as is found in the Civets, differing in this
respect from the other known families of the Carnivora. The
remaining vertebre agree with those of the Viverride, and the
lower end of the radius has the same characteristic triangular
form in cross-section as in Viverricula. ‘The feet and limbs are
also very like those of the Viverrines, with the exception that the
scaphoid and lunar bones of the carpus are free and not united
as in the modern family.
Altogether we think it may be stated with considerable certainty
that the group represents the forerunners of the Viverrine phylum
whose members towards the close of the Eocene migrated to Asia.
This view receives strong additional support from the fact that
the typical genus Vzverra runs backward without change into the
Upper Eocene of Europe, showing a remarkable degree of per-
sistence of structure, which also characterizes the genus Vzver-
ravus, passing as it does with only slight specific modifications
from the Torrejon beds of New Mexico through to the Bridger.
The technical definition of the group rests upon the exclusive
development of the fourth superior premolar and the first inferior
molar into enlarged typical sectorials, thereby distinguishing it
from all the other known Creodonts except the early members of
the Canide. From this latter family it is distinguished by having
only two molars in the lower jaw and an anterior basal cusp upon
the superior sectorial. From the Viverride it is separated by
the free condition of the scaphoid, lunar, and probably the
centrale as well.
The species are numerous and range in time from the Torrejon
to the Bridger.
138 Bulletin American Museum of Natural History. [Vol. X11.
SUMMARY.
The principal points brought out in the foregoing paper may
be briefly summarized as follows:
(1) The tracing of the Daphenus ancestry back into the
Wasatch Eocene by way of Prodaphenus and Uintacyon, which
latter has always been considered a Creodont.
(2) The establishment of the relationship of. Daphenus to
Temnocyon, and that of Zemnocyon to the living genus Cyon.
(3) Tracing the ancestry of the Cants-Cynodictis line back into
the Eocene through Procynodictis and Vulpavus.
(4) Some additional points in the structure of Cynodictis.
(5) The descent of certain South American Foxes from North
American Miocene species, with the establishment of a new genus,
Nothocyon, for their reception.
(6) Classification of the Miocene genera of the Canide,
(7) The discovery of the origin of the Procyonide from the
Canidz through the new genus PA/aocyon.
(8) The discovery of the genus Viverravus and the establish-
ment of a new family.
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P sro! XII, ARTICLE VII, pp. 139-148.
New York, Fune oat 1899.
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Article VII.—RESTORATION OF OXYANA LUPINA
COPE, WITH DESCRIPTIONS OF CERTAIN NEW
SPECIES OF EOCENE CREODONTS.
By J. L. WortTMan.
PLATE VII AND THREE TEXT FIGURES.
FamMity OXYAINIDA.
This family of the Creodonta was the most specialized in cer-
tain ways of any of the primitive flesh-eaters of Eocene times,
and apparently occupied the same position with reference to the
remainder of the fauna that the modern Felidz do to the existing
fauna, In the matter of dental equipment it is surprising to find
that even as early as the Wasatch this family had developed a
sectorial dentition almost if not quite as effective as that of the
modern Cats. In other respects, however, they were far in-
ferior ; this is particularly seen in the small brain capacity, as
well as the comparatively smaller, decidedly weaker limbs with
shorter, more spreading feet, provided with flatter, fissured, non-
retractile claws.
The origin of the family is completely unknown at the present.
They begin with several species abruptly in the Wasatch deposits
of both the San Juan and Big Horn beds without any known
predecessors in the underlying Torrejon. For this reason there
can be very little doubt that they represent migrants from another
region, probably northern Asia, which came with the Corypho-
donts, Artiodactyles, Perissodactyles, Primitive Dogs, as well as
many other types whose existence begins so abruptly in the
Wasatch beds of North America.
According to our present knowledge the family represents a
perfectly natural grouping of the known species, and is easily
distinguished from the other groups of typical Creodonts that
had developed a more or less perfect sectorial dentition. A con-
venient distinction of these families may be made upon the
enlargement of a special molar in the lower jaw; it is as follows:
1. Third lower molar enlarged,—Hy NODONTID&. Includes the genera
Pala@osinopa, Sinopa, Proviverra, Cynohyenodon, Pterodon, Hyenodon,
139
140 Bulletin American Museum of Natural History. [Vol. XII,
2. Second lower molar enlarged,—Oxy 2NID&. Includes the genera Oxyena,
Patriofelis, Oxyenodon.
_ 3. First lower molar, enlarged,—PALONICTID&. Includes the genera
Peres. Amblyctonus, Alurotherium.
Oxyzna lupina Cope.
By a fortunate circumstance I am now enabled to give a rather
full account of the skeleton of this species, which has hitherto
been only imperfectly known. At the time of my first trip into
the Big Horn Basin in 1880 the country was a wild, uninhabited
region, save for the occasional visits of roving bands of hostile
Indians, and any explorations there by a small party were at-
tended by no small amount of risk to one’s personal safety. In
fact, I was advised by the commander of Fort Washakie, at that
time the base of our operations, that the trip was a hazardous
one, and that he would not undertake to answer for our safe con-
duct. We went through, however, without serious inconvenience,
but at the same time the collecting was not as thoroughly done
as it probably would have been under less trying circumstances.
On this expedition, among other things, I secured a part of the
skeleton of this species, which was erroneously referred to Cope’s
somewhat larger but closely allied species O. forcipita, in his
volume ‘ Tertiary Vertebrata.’ In 1891, I conducted another
expedition into this same region for the American Museum,
at which time, through a general settlement of the country, the
former more hazardous conditions had been entirely removed and
undertakings of this sort were not accompanied by the same
risks of violence at the hands of savage Indians as formerly pre-
vailed. Our knowledge of methods of collecting had, moreover,
materially increased, and while the actual number of specimens
secured was perhaps less, yet a somewhat greater success attended:
our efforts, especially in securing those parts which had already
been washed out of the matrix in which they had been originally
imbedded. In this category comes the present specimen, a por-
tion of which had been collected by the expedition of 1880, and
the remainder by the expedition of 1891.
During this latter expedition, a new method was employed for
securing the missing parts which had been washed out and cov-
ered up again by the accumulating debris. Wherever possible
1899. | Wortman, Restoration of Oxyena lupina. TAI
the loose dirt containing the fragments was gathered up and trans-
ferred to the nearest stream where it was washed out after the
manner of the placer miner. In this way, wherever conditions
were favorable, all the fragments were recovered, but in the case
of the present skeleton, after every possible exertion, a large
number of pieces necessary to complete the skeleton remained
missing. When the Cope collection was purchased by the
Museum and the two collections were brought together it was
accidentally discovered by Dr. Matthew that the specimen of the
Cope collection furnished the missing parts of one and the same
individual of the Museum specimen collected in 1891 ; in this
way an unusually complete skeleton for an Eocene fossil results
and furnishes all the more important characters.
Skull—All the details of the form of the skull cannot be
accurately made out owing to the great amount of crushing and
fracture which the specimen had sustained during the process of
Fig.1. Oxyena lupina Cope, Skull and jaws, 34 natural size. No. 107.
fossilization ; but enough is preserved in an uninjured condition
to indicate that the facial portion was very short and broad in
proportion to the total length of the cranium. That part of the
skull posterior to the orbits is proportionately much elongated
and, owing to the small size of the brain-case, it appears to be
unusually so. The sagittal crest is prominent and extended well
forward ; the zygomatic arches are wide and heavy, and the
glenoid cavity has a distinct pre- and post-glenoid process, The
142 Bulletin American Museum of Natural History. | Vol. XH,
mastoid is prominent, and the paroccipital process has a more or
less backward direction. The foramina of the base of the skull
cannot be determined, but it appears probable that there was
a distinct post-glenoid present.
The dentition, the formula of which is I. 3, C. 4, Pm. 4, M. 8,
has been quite fully described and requires but a brief mention,
It appears from the very perfect lower jaws that there were ¢hree
incisors present upon each side, although this is not entirely
demonstrable. In the upper jaw, the outer incisors are much
larger than the two inner pairs, which are subequal. ‘The canines
are long and pointed, being slightly compressed at the base. The
first premolar is small, single-rooted, and has a simple crown,
The second is two-rooted and has a prominent posterior heel.
The third is three-rooted with two external and one internal
cusp. The fourth premolar, while possessing all the elements of
Fig. 2. Oxyena lupina Cope. Upper and lower teeth, 34 natural size. No. 107.
the superior sectorial of the typical Carnivora does not form
a very perfect shear between the rudimental blade and that of the
inferior first molar. The great shearing function was transferred
to the first molar above and the second molar below; the first
superior molar is curiously modified, in that the two original
external cusps are placed very close together, and a large post-
ero-external cusp added, which, in conjunction with the true
postero-external cusp, form a very effective blade, the internal
cusp being small. This is, at least, the interpretation which has
1899. | Wortman, Restoration of Oxyena lupina. 143
been placed upon the structure of this tooth, and the evidence in
favor of such a view is found in the gradual assumption of a
similar if not identical structure seen in /7y@nodon from the more
generalized pattern of the tooth exhibited by Szwopa. The last
molar is transverse, and its crown consists of a single external
cusp, with a great extension of the antero-external angle, together
with a low, smaller internal cusp.
In the lower jaw the first premolar is small, the succeeding
teeth gradually increasing in size to the last molar. Premolars
two, three, and four have prominent heels, and the fourth has a
small anterior basal cusp in addition. The molars have the
typical sectorial pattern, with elevated trigon and a low, relatively
small, basin-shaped heel. The internal cusp and heel are reduced
in the last molar, foreshadowing its almost complete loss in
Patriofelis.
Vertebrea.—The usual seven cervicals are present which, so far
as the imperfect state of their preservation will permit one to
judge, resemble closely those of Patriofelis. The atlas has rather
wide, roomy cotyles for articulation with the occipital condyles
and the transverse processes are perforated at the base quite as
in Patriofelis and the modern Cats. The axis has a peg-like
odontoid, a prominent inferior tubercle on the inferior posterior
surface of the centrum, and distinct transverse processes, which
are perforated at the base for the passage of the vertebral artery,
The spine is not preserved. The remaining cervicals have
rather flattened centra and resemble those of Patricfelis.
The dorso-lumbar formula is 20, of which 13 are dorsals and 7
are lumbars. They increase in size gradually from before back-
wards, the posterior lumbars being the largest. The lumbar
articulations are not so complex as those of Pasriofelts, the convex
postzygapophyses fitting into concave prezygapophyses with no
evidence of the double, concavo-convex arrangement of certain
of the other Creodonts. There are distinct anapophyses, but no
metapophyses appear to have been developed. The sacrum is
not preserved. Of the caudals enough are preserved to indicate
that there was a long and powerful tail. The vertebrae resemble
those of Patriofelts in all their details of structure.
Fore Limb.—Very little of the scapula is preserved, only the
proximal portion of both bones being present. The glenoid cavity
144 Bulletin American Museum of Natural History. [Vol. XII,
has an oval form; the coracoid is prominent, and the relatively
strong spine arises a short distance behind the glenoid border.
There is evidence of a well developed acromion and metacromion,
about as in Patriofelis. The humerus, like that of many of the
Creodonts, has a powerful deltoid crest which occupies somewhat
more than one half the entire length of the shaft. The head is
pyriform, the tuberosities well developed, and the bicipital groove
is deep. Distally the bone is broad, as in many of the modern
Carnivores, especially the felines. There is a very prominent
internal condyle; an entepicondylar foramen, and a somewhat
reduced supinator ridge. The articular surface is very much as
in the Cats. The ulna has a very prominent incurved olecranon,
the shaft is straight, and the two distal articular facets are well
separated as in the Cats. The head of the radius presents an
oval outline with an unusually prominent tubercle. The lower
end of the shaft is triangular in cross-section, and the scapho-
lunar facet is concave. The fore foot has already been fully
described,’ and there is little of importance to add. The foot is
relatively broad and spreading as in Patriofelis; the scaphoid,
lunar, and centrale are free; the first phalanx of the thumb is
relatively large, and all the claws are deeply fissured with the
possible exception of the fifth. There is reason to believe that
the habitual position of the foot was digitigrade, but there is no
evidence of any retractility of the claws.
Hind Limb.—The pelvis is not very well preserved but enough
is present to indicate that it was very similar to that of Patrio-
felis. The femur has a well-rounded head, a nearly straight
shaft, and a small though distinct third trochanter. ‘The tibia
equals the humerus in length and has a slightly curved shaft ;
the cnemial crest is prominent, the internal malleolus is unusually
thick and heavy, and the distal trochlea is directed obliquely
inwards and is little excavated. The fibula is complete and
comparatively little reduced. ‘The hind foot is more slender
than the fore foot. The tibial facet of the astragalus is but
slightly grooved ; the astragalar foramen is distinct, and the head
of the bone is flattened from before backward, rounded and
oblique ; it articulates with both cuboid and navicular. The
1* Fossil Mammals of the Wasatch and Wind River Beds.’ Bull. Amer, Mus. Nat. Hist.,
Vol. IV, 1892, p. 108.
1899. | Wortman, Restoration of Oxyena lupina. 145
three cuneiforms have about the same relationship and arrange-
ment as in the modern Cats. Of the metapodials there is a dis-
tinct interlocking, although the degree is much less than in the
Felidz. The third is the longest but the second is slightly the
heaviest of the series. As in the fore foot, the claws are fissured
and non-retractile.
Oxyznodon dysodus, gen. et sp. nov.
This genus is founded upon an unusually perfect half of a skull
from the Uinta Eocene, collected by the writer in 1896. It indi-
cates an animal of the size of a Gray Fox and is, therefore, if
properly referred to the Oxyzenidz, the smallest member known.
The dental formula is the same as that of Oxyena, viz.: I. 3, C.
1, Pm. 4, M. 3, and the last superior molar is, moreover, trans-
verse. The chief distinctions between the two are seen in the
form of the skull and the character of the premolars, as well as the
great enlargement of the symphysis in the Uinta genus. In
Oxyenodon all the lower premolars are simple and have only rudi-
mental posterior heels, whereas in Oxyena the heels of the
Fig. 3. Oxyenodon dysodus. Skull and jaw th natural size. Type specimen, No. 2515.
second, third, and fourth are large, and form important elements
of the crown. In Oxyenodon the third superior premolar has no
internal cusp, while in Oxyava a well developed inner cusp is
present. The heel of the last lower, as well as the internal cusp
of the trigon, are more reduced in the Uinta genus than in
Oxyena. In Oxyenodon the mandibular symphysis is enlarged
and extends back under the third premolar, the lower jaw being
relatively shallow and thick, while in Oxyena the mandibular
[Fune, 1899] 10
146 Bulletin American Museum of Natural History. [Vol. XII,
symphysis is of much less extent and the jaw is deeper. The
facial portion of the skull in Oxyenodon is longer.
I have suggested * that this family has some affinities with the
Seals. The skull Oxyenodon strengthens this view in certain
particulars. ‘The most important resemblances to the Sea Lion |
are seen in (1) the great postorbital construction of the-skull ;
(2) the pressure of a distinct lachrymal tubercle ; (3) the ex-
clusion of the frontal from any share in the anterior boundary
of the orbit ; (4) early union of lachrymal with maxillary; (5)
forward extension of sagittal crest; (6) absence of anterior
process of frontal between nasal and maxillary, and (7) an
elongated mandibular symphysis. While the skull is not strik-
ingly seal-like in its general appearance yet when we know the
immediate ancestors of such a form as the Sea Lion it will not be
surprising if it is found to connect with such a form as
Oxyenodon.
Famity MESONYCHID.
The Mesonychidz undoubtedly represent a distinct line of
the primitive Carnivora whose later representatives came to be
characterized by elongated and specialized limbs for a running
habit, as well as by a peculiar reduction and simplification of cer-
tain cusps of the molar teeth. Somewhat contrary to the generally
accepted arrangement of this group, I unite with them the so-
called Triisodontide as a well marked subfamily. This latter
series is the older and displays the same tendency towards the
peculiar reduction and rounded, conical form of the cusps of the
molars as is seen in the Déssacus-Pachyena-Mesonyx line. The
differences between the subfamilies are not great and, as observed
by Matthew,’ consist in the deep, heavy jaws with powerful sym-
physis, and the much wider, more distinctively tubercular charac-
ter of the molars in the Triisodontide.
The single genus Z7z7sodon, with three species, comes from the
Puerco and is succeeded in the Torrejon by the two genera
Sarcothraustes and Gontacodon. Matthew has pointed out that
Sarcothraustes antiguus is without doubt the direct descendant of
Triisodon and in this opinion I entirely agree. As noted by this
1‘ Osteology of Patrzofelis,’ Bull. Amer. Mus. Nat. Hist., Vol. IV, 1894, p. 157-
Bull. Am. Mus. Nat. Hist., Vol. 1X, 1897, p. 278.
1899. | Wortman, Restoration of Oxyena lupina. 147
author, the same cusp reduction and simplification is observed
in this succession as occurs in the D¢ssacus- Pachyena-Mesonyx
series.
The second subfamily, Mesonychinz, so far as at present
known, finds its oldest representatives in the Torrejon beds in the
single genus Dzssacus, with two well marked species. It is, in-
deed, very doubtful, if not impossible, that any of the known
species of Zvzisodon can be placed ancestral to the present genus,
although it is not at all improbable that some slender jawed type
in the Puerco having the cusp pattern of Zyizsodon will be found
to have commenced an early modification of the teeth terminating
in Dissacus. The evidence for the view that the inferior molar
pattern in this genus is a degenerative one, is found in the fact
that in the succeeding Wasatch Pachyena the postero-internal
cusp of the trigon is still more reduced than it is in Déssacus,
while in the Bridger AZesonyx it exists as the merest vestige,
the cusps of all the molars at the same time assuming a very
rounded and characteristic conical form.
Pachyzna intermedia, sp. nov.
It has been shown by Osborne and Earle’ that neither of the
known species of Pachyena (gigantea and ossifraga) can stand
directly in the line of descent leading to Mesonyx on account
of the greater reduction of the last upper molar in the two species
of Dissacus (navajovius and saurognathus) than in Pachyena.
Scott has shown’ that JZesonyx has only two superior true molars,
so that any species of Pachyena which exhibits a less reduction of
the last upper teeth than YDyzssacus cannot be placed ancestral
to AZesonyx. Fortunately the collections of the Museum contain
a specimen of a Pachyena from the Big Horn, obtained by the
expedition of 1896, which shows, proportionately, as great or a
little greater reduction of the last upper molar than Déssacus
navajovius, ‘The specimen consists of the last two upper molars
and bears the Museum number 2854. ‘The teeth in question are
somewhat smaller than the corresponding ones in P. ossifraga
and exhibit a considerably greater reduction of the last molar in
comparison with the tooth in advance than in this species. It
* ‘Fossil Mammals of the Puerco,’ Bull. Amer. Mus. Nat. Hist., Vol. VII, March, 1895, p. 39.
Z ‘Some New and Little Known Creodonts,’ Jour. Philad. Acad. Nat. Sci., 1886, Vol. I,
148 Bulletin American Museum of Natural History. [Vol. X11,
may be that the specimen represents only a smaller variety of P.
ossifraga and is not entitled to a specific rank, but in view of the
fact that it furnishes just the character which, upon general
grounds, one would be led to anticipate, I have thought fit to
give it the above name.
This virtually completes the series of the Dyssacus-Pachyena
Mesonyx phylum and establishes a closely connected specific
descent reaching in time from the Torrejon to the White River
without any important break. The Wind River representative of
this series, however, yet remains to be discovered, but in accord-
ance with what we already know its characters can be most
accurately predicted.
EXPLANATION OF PLATE VIL.
Oxyena lupina, Cope. Skeleton, %th natural size. No. 107, Am. Mus. Coll.
The specimen described and figured as *‘ No, 2” by Prof. Cope in * Ter-
tiary Vertebrata of the West,’ (p. 319, pl. xxive, figs. 12-15, and pl. xxivd,
figs. I-18) is a part of this individual.
Wien SAN, yin, WHY,
Nid
M.
BULLETIN A.
Sia
a
i
(
e
IS
pp Eisele ee
v \ iy)
Ki Weber del
OXYENA LUPINA Cope.
One-sixth natural size,
- assic of Wy voming. Dinosaur Contrt-
a butions, No. 2.
_ AUTHOR'S EDITION, extracted from BULLETIN >
OF FHE ‘ ; . ee
American Musenn of Ratur al History, ig
é Vou: XI, hohe. XI, pp. 161-172. a. :
New York, “ October age 1899. | } my a
Article XI.—FORE AND HIND LIMBS OF CARNIVOR-
OUS AND HERBIVOROUS DINOSAURS FROM THE
JURASSIC OF WYOMING. DINOSAUR CONTRIBU-
TIONS, NO. 3.
By Henry FAIRFIELD OSBORN.
In the Bone Cabin Quarry, opened by the American Museum
Expedition of 1898, under the direction of Dr. J. L. Wortman,
assisted by Mr. W. W. Granger, were found mingled together
bones of all the most characteristic Upper Jurassic Dinosaurs, in-
cluding six nearly or quite complet limbs and three fore feet,
which form the subject of this paper. Four of these are shown,
as mounted, upon a subsequent page. ‘They are not only grand
objects of their kind, but they bring out a number of new and
important facts relating to the limbs of Dinosaurs of the Mega-
losaur-and Cetiosaur divisions or Ornithopoda and Sauropoda.
1. Hinp LIMBs OF CARNIVOROUS DINOSAURS.
FIGURES I-5,
The late Professor O. C. Marsh described’ five genera of flesh-
eating Dinosaurs from our Jurassic, namely: A//osaurus, the
largest of the flesh-eaters; Creosaurus, a smaller allied form ;
Labrosaurus, of another type; Calurus, a very small animal dis-
tinguished by hollow bones (related to the somewhat older Ha//o-
pus); and finally Ceratosaurus, a large animal with horned nasals.
Of these, the most nearly allied to Weya/osaurus of the English
Purbeck is Ad/osaurus, distinguished by possessing less than five
vertebrze in the sacrum, also by other characters of doubtful
value (of. c7¢., p. 239), and figured (of. cz, Pl. XI.) as possessing
three digits in the pes, while Mega/osaurus is assigned four digits.
in the pes (of. cit, p. 239). Whether or not AVosaurus is distinct
from Megalosaurus cannot be positively determined at present.
The distinction based upon the number of digits is here shown
not to hold good.
Two hind limbs from the Bone Cabin Quarry, which corre-
spond in other respects with those of A//osaurus, appear to show
that the hallux had been detached in Marsh’s specimens. ‘They
; j 1 The Dinosaurs of North America, 1896.
[Oct., 7899.] [161] II
162 Bulletin American Museum of Natural History. [Vol. XII,
(Nos. 290 and 324) both possess four digits in the pes; the first
digit or hallux exhibits an interrupted metatarsal, and is turned
inwards as a subsidiary grasping toe.
The proportions between these limbs and those of the two
Sauropoda described below are well shown in the accompanying
photographs (Fig. 1). The heights in the Table include the
curvatures ; they give us therefore the usual height of the ace-
tabulum, or head of the femur, not the total length of the limb.
Fig. 1. Right Hind Limbs of Dinosaurs, A, Herbivorous Dinosaur, probably Dzplodocus, No. 25t.
B, Herbivorous Dinosaur, Brontosaurus, Nos. 309, 353. C, Carnivorous Dinosaur, A ddosaurus, OF
Megalosaurus, No. 290. D, Carnivorous Dinosaur, AZlosaurus or Megalosaurus, Nos. 324, 275-
Approximately +: nat. size.
1899. | Osborn, Carnivorous and Herbivorous Dinosaurs. 16 8
The larger of the limbs (No. 290) is the more perfect, except
that it lacks the proximal portion of Mt. I. The smaller (No.
324) has the calcaneum and astragalus restored, and a femur
(No. 275), which was found at some little distance, is arbitrarily
placed with it. Except for differences in size, due to age or sex,
the limbs are practically identical in character.
Measurements.
No. 290 No. 324. No. 275
WenotyOltemute rie siie ee cist eels OB imaataiak: Wie eas gio
Bote poe MULDIA Ss iaiatersi et es aersr scale SMO SHEE 698
OO SOUS rion Tae aadkei a meres eae ane TOMER eh cs. s 665
seers CLALATS Alig Iil wiesye, am sieve BS Va IRC 315
: ‘ ees SAS oc ADU ES shar sders 352
# oh IN Rac crerccoxer ats ZOOW vaca cee 330
Wadthvotvanklesjomet fa... =).
Calcaneum and astragalus....... 241
orluhersht of limb pei. + 1.0.1. 2B. corsets y Banaue 1980
Of greatest interest is the first digiz,
or hallux, not before described. Prox-
imally (Fig. 4a) its metatarsal fits in a
shallow groove of the upper portion of
the large metatarsal II. The shaft is
entirely interrupted or composed of
cartilage in the middle portion. Dis-
tally it is fitted to the rounded posterior
shaft of Mt. II, demonstrating that this
digit was directed inwards like the
small hallux of Afpferyx (Fig. 5). It
possesses, however, a complete and
functional phalanx and claw, which un-
doubtedly were of service in grasping. a
The digits II, III, 1V, with phalanges | eens -
numbering 3, 4,-and 5, réspectively, are Fig. 2. Carnivorous Dinosaur,
well shown in the photographs (Figs. 4, showize Same nine 35 Mie
4a). ‘The lower row of tarsals is repre- “#°%™°(D8 J. ©. WJ» ah nat size:
sented by two bony elements only, probably tarsalia 3 and 4.
The upper tarsals, astragalus and calcaneum are closely con-
joined if not actually co-ossified.
164 Bulletin American Museum of Natural History. [Vol. pil) ix
The tibia forms the en-
tire back portion of the
ankle joint, the fibula
lying in front of it; this
bone is further distin-
guished by its curved
shaft, and powerful mus-
cular crest projecting to-
wards the fibula at its
upper third. There is a
very prominent cnemial
crest, which has a deep
groove upon its outer side
towards the fibula; this
groove is entirely con-
cealed from the front.
This bone is very different
from that assigned to
Megalosaurus by Owen.’
The fibula has a slender,
subrounded shaft, but ex-
pands proximally and dis-
tally upon the tibial side.
The femur is distin-
‘ guished by its marked cur-
vature and well-rounded
head, by the laterally com-
pressed and inferiorly
placed great trochanter,
by an internal trochanter
much more elevated than
in the Iguanodontia, and
by a large rugose area on
the inferior front face of
the shaft above the in-
ternal condyle. The inner
trochanter is broken off in
Fig. 3. Right Hind Limb of Carnivorous Dinosaur, the smaller femur, No. 275-
No. 290. Oblique internal view. yg nat. size.
™ Fossil Reptilia of the Wealden and Purbeck Formations, Pt. III, 1857, p. 18, Tab. Ix.
1899.| Osborn, Carnivorous and Herbivorous Dinosaurs. 165
2, Hinp LIMBS OF CETIOSAURS.
FIGURE T.
The smadler of these limbs (Fig. 1, A, No. 251) was fortunately
found nearly complete and in position, all parts being preserved
excepting the head of the femur. It was removed and worked out
with the greatest care by Mr. Granger. ‘There is thus absolutely
no doubt about the position of the phalanges.
This limb is of a type distinct from the larger one, which is
undoubtedly a Brontosaurus. We should unhesitatingly refer it
to Diplodocus, were it not that the fourth trochanter has a more
elevated position than in the DzAlodocus femur recently described
by the writer,’ and that the pes differs from the pes of Dzplodocus
figured by Marsh (’96, Plate 28). None the less it is a Zong-limbed
type, and this we believe to be a characteristic distinction of
Diplodocus. it has the following characters :
1. ‘Tibia and fibula very long and slender.
2. Femur long with relatively slender shaft.
3. Metatarsals I and II as in Arontosaurus.
4. Metatarsal III much more slender than in Bronfosaurus.
5. Metatarsal IV still more slender, two osseous phalanges.
6. Metatarsal V more slender, no osseous phalanges.
This limb may prove, therefore, to represent merely a form of
Brontosaur distinct specifically from #&. exce/sus. But, in spite
of the exceptions noted above, the probabilities are that it belongs
to Diplodocus, for this animal is abundantly represented in the
Bone Cabin Quarry by parts of several series of caudal vertebra,
besides pelvic and other bones.
Measurements.
No, 258. “No. 309; =No. 353.
piGralpherioitofalimalyy si.) - « 2a crsstaee eye bs e€ 2920 3040
SCRA ES lige DEO Maes D COC Oe TCO ks | e 1430 1640
Femur, circumference below tr. 4....... 560 €= 730
Mstintarwlen Mth yer. -\sieaye.o sree one 1060 1080
Whibiar ‘circumference: .j...~.6s.000.e8e 390 e 480
i ulaeelenpthtrcin ce grat. esis yeeaets.} « 1120 1130
ANSEL AUS WIGth (Of...c.5 -|¢ s/s sie sitet « 22 280
Pes, width of proximal metatarsal surfaces, 410 445
The circumferences of the various shafts are very important
* Osborn, ‘tA Skeleton of Diplodocus.” Mem. Am. Mus. Nat. Hist., 1, Pt. IV, 1899.
166 Bulletin American Museum of Natural History. {Vol. XII,
when compared with the total lengths. In each case the least
circumference is taken. The total height of the limbs is measured
as mounted—that is, including the curves.
The larger limb (Brontosaurus) is to feet high. The smaller
limb (? Diplodocus) is 9 feet 7 inches high.
The /arger limb (Fig. 1, B, Nos. 309, 353) is of the true Bronto-
Sas oes Fd FA ES ESSE
Fig. 4. Right Hind Foot of Carnivorous Dinosaur, probably A Zllosaurvus (No.
324, Same specimen as Fig. 1, D). Front view. 4 nat. size.
1899.] Osborn, Carnivorous and Herbivorous Dinosaurs. 167
saurus type. All of the bones of the lower leg and pes belong to
one individual, excepting the terminal claws I and II. ‘The
femur was found at some distance and 1s arbitrarily associated.
The phalanges are complete and osseous upon D. I, II, III.
This and the foregoing specimen prove conclusively that Marsh
was in error in restoring a complete series of bony phalanges and
Fig. 4a. Right Hind Foot of Carnivorous Dinosaur, probably 4 2osaurus ( No. 324, same
specimen as Fig. 1, D.) Oblique side view. } nat. size.
168 Bulletin American Museum of Natural History. [Vol. X11
claws upon digits IV and V. The phalanges upon digits IV and
V were cartilaginous or incomplete ; these outer digits functioned
only in supporting the foot pad. Every known Sauropod pes
shows the same deficiency upon the outer side.
3. Fore FEET OF CETIOSAURS.
FIGURES 6, 7.
‘wo restorations of the fore feet of the Sauropoda have been
published by the
late P nehes stom
Marsh, namely those
of ATorosaurius (96,
Plate 38) and Bron-
tosaurus (op. ctt.,
Pl ate.4q2)s + Tt fais
pears probable that
both are incorrect ;
the error apparently
has arisen, first, from
supplying all the
digits with a com-
plete series of pha-
langes ; second, from
placing the largest
phalanges upon the
inner or tst digit
(as in the pes) and
grading off the small-
er phalanges to the
Fig. 5. Afteryx. Internal yiew of right pes. 5th or outer digit.
It is now practically
certain that in the manus as in the pes a number of phalanges
were either cartilaginous or missing entirely. It also appears prob-
able that the three central digits LI, III, 1V bore a full series of
phalanges and claws, while the outer digits were deficient in
phalanges.
ForRE FEET OF UNDETERMINED CETIOSAUR.
The strongest evidence comes from the two fore feet of one
individual (No. 332), one of which was found with all its parts in
‘azIs [eInjeu ? ‘a}eUTUIOJapUT JUaSoId yW “ZEE ‘ON ‘INeSOUICT SNOIOATGIA}Y JO Joaq V1Og (q) eT pue Cy) Iysry 9 BI 691
170 Bulletin American Museum of Natural History. [Vol. XI,
position (Fig. 6). The right manus is more complete in certain
parts, the left manus in others. The animal is possibly a
Morosaurus agilis.
The Metacarpal of the first digit is much longer and more
slender than in either the typical Worosaurus or Brontosaurus.
A single phalanx was found near it, rounding off distally and
quite distinct from the first phalanges of digits II, III, IV.
The second Metacarpal is still longer and is supplied with the
characteristic broad phalanx 1, the narrow phalanx 2, and the
terminal claw or phalanx 3. This digit is slightly heavier than
the 3d.
The third or middle digit is the longest; the 1st phalanx is
broad, the 2d and 3d are restored, the 4th is complete, and was
found associated in the left manus.
The fourth digit presumably had four phalanges, but only
phalanx 1 is preserved in each case; and it has the same broad
form asin D. II and III. The fifth digit is of about the same
length as the first, and is likewise supplied with a single phalanx
unformed. or rounded distally, without any distinct facet for a
second phalanx.
It thus appears certain that in this animal (No. 332) the middle
three digits were fully functional and provided with claws, while
the lateral digits were incomplete distally and served only to
support the weight of the body.
ForE FEET OF BRONTOSAURUS.
FIGURE 7.
The question now remains whether the fore feet of Brontosaurus
were also similarly constructed upon a mesaxonic plan.
Marsh restored (Brontosaurus, Morosaurus) the manus like the
pes with a complete series of claws, the largest being on the
inside.
The evidence is very positive in the two feet here mounted
that there was a deficiency of phalanges. The evidence, however,
that the actual construction of the foot is as we have mounted and
photographed it (Fig. 7), namely, mesaxonic, is by no means posi-
tive. The question, in fact, cannot be definitely settled until a
manus is found with all the digits in position, as in the pes above
described.
1899-| Osborn, Carnivorous and Herbivorous Dinosaurs. 171
The large fore foot (No. 268) was found with the metacarpals
in position, and the phalanges scattered. The two terminal claws
of digits III, IV, were not found with this foot, but some dis-
tance from it. Phalanges 2, 3, 4, of digit IV are restored in
plaster. The foot is therefore nearly but not quite complete.
The broad 1st metacarpal has no distinct distal articular facet,
such as are observed upon digits II, III, IV, and it seems probable
.
Fig. 7. Left Fore Foot of Large Brontosaurus, No. 268. The position of the phalanges is some-
what conjectural, 3 nat, size.
172 Bulletin American Museum of Natural History. [Vol. XII,
that we are correct in associating with it the short phalanx with
an imperfectly rounded distal extremity.
The 2d metacarpal is very powerful, with a heavy rounded
shaft and distinct distal articulation, from which three phalanges
extend, the terminal a heavy claw, if our placing is correct.
The 3d digit affords the strongest support for the mesaxonic
hypothesis, for it is by far the largest metacarpal, and is obviously
the centre of the foot. With it must have been associated (as in
the present mounting) the heaviest phalanges and claw.
The 4th metacarpal is considerably lighter and longer, and un-
fortunately its complement of phalanges is represented only by
phalanx 1, the others being added or restored.
The 5th metacarpal is stout but slightly shorter than the 4th.
With it was apparently associated the remaining imperfectly
formed phalanx 1, which was found with this foot.
If these two imperfect phalanges are not placed upon digits I
and V, it is very difficult to place them at all. It thus appears.
probable that digits I and V were deficient in all the phalanges
except the first or first and second, and served merely to support
the weight of the animal.
The admirable methods in the field, which have been largely
developed by Dr. J. L. Wortman, whereby every piece is kept
and transported in the position in which it was found, sup-
plemented by admirable museum methods, will soon render our
knowledge of the Dinosaurs not only far more complete, but far
more accurate than ever before. The writer is indebted to Dr.
Wortman for many hints in the discussion of the carnivorous.
limbs above described. The writer desires also to express his.
indebtedness to Mr. Adam Hermann, not only for the skill
displayed in mounting these specimens, but for the excellent
judgment and knowledge he has shown in locating and bringing
together the scattered parts. Owing] to the imperfect ossification
of the joints, the bones of Dinosaurs are much more difficult to
place than those of mammals.
ike Brees Plains ‘of le CXQS.
as evs Z ; ) i ; !
‘ f i :
By i. W. GiwLey.
AUTHOR'S EDITION, extracted from BULLETIN
Fee ah it a Meee i
Article XIII.—A NEW SPECIES OF PLEISTOCENE
HORSE FROM THE STAKED PLAINS
OF TEXAS.
By J: W»GIDLEY.
Equus scotti, sp. nov.
The type of this species is a nearly complete skeleton (No.
10606), consisting of the skull and lower jaws, the cervical ver-
tebrz, the three anterior dorsal vertebra, both fore limbs and
feet complete, and one hind limb and foot, besides several other
vertebreze and some ribs probably belonging with it.
This skeleton was found by the writer associated with four
other skulls and parts of skeletons of the same species, in a bed
of compact Pleistocene sand at the head of Rock Creek, Briscoe
Co., Texas. The bed in which the bones were found is about
the middle of the Equus, or Sheridan, beds, which are about 100
feet in thickness at this place.
Bones from the associated individuals have been substituted
for the missing bones of the skeleton (No. 10606) which has
been admirably mounted by Mr. Adam Hermann, and placed on
exhibition in the Tertiary Mammal Hall of the Museum.
The writer has made a very careful study of all the types of
the species of Aguus in this country and has found that an ex-
tensive revision is necessary ; this will be published in a subse-
quent paper. It appears that the horse from the true Equus beds
of the Plains has not been taken as a type but has been mistak-
enly identified with other species. A new term is therefore nec-
essary and this is selected in honor of Prof. W. B. Scott, of
Princeton University.
The species &. scott? differs from Z. caballus in proportions and
size as follows: (1) the skull is relatively larger, (2) the neck
is shorter, (3) the body is longer, (4) the lesser curvature of the
belly ribs near their heads indicates that the back was not nearly
so wide, (5) the limbs are shorter and more slender in proportion
than the larger varieties of the recent horse.
Comparing the skeleton of &. scofti with the skeleton of a
larger draught horse (No. 528) in the osteological collection of
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1900. | Gidley, A New Spectes of Pleistocene Horse. Et?
the Museum we observe some striking differences. While the
skulls are about equal in length and the series of dorso-lumbar
vertebrz is only about 1 inch longer in the recent horse, the cer-
vical series of &. scotté is about 4 inches shorter and the fore
limb, in the standing position, is about 6 inches less in length.
Thus, this skeleton represents an animal with a head about the
size of that of a large draught horse, but with the height of body
and length of limbs of an ordinary western pony, and with a
length of body very similar to that of the Zebra or Quagga.
A comparison of the separate bones of the skeleton reveals
very few and unimportant differences, hence it is to the skull and
teeth that we have to look for specific differences.
DENTITION.
Unfortunately all the bones of these skeletons, while adult,
are of young horses, none of them having shed their last milk
molars or external milk incisors, As
far as can be made out the teeth differ
from those of £. caballus only in their
much larger size. M7’ of the type skull
(Fig. 2), indicates perhaps a little
greater degree of complexity of the
enamel folding on the triturating sur-
face than is usual in &. caballus, but it
should be taken into account that this
is only a slightly worn tooth and shows
a greater degree of complexity than it would at a more advanced
stage of wear.
The measurements of the teeth that are enough worn to give
their true diameters are as follows :
Fig. 2, M? of £. scottz.
. Diameters of crown of p? eel aaa 33 mm,
transverse 31 mm.
: ro- i
Diameters of crown of m! ; BOS pete ee Oe
transverse 30 mm.
Long diameter of i ! 23 mm.
Total length of molar-premolar series I9g0 mm.
Diameters of the corresponding teeth of the large draught
horse (No. 528):
Diameters of crown of p? antero-posterior 29 mm,
transverse 27 mm,
August, 1900. | 8
I14 Bulletin American Museum of Natural History. [Vol. X11,
Diameters of crown of m! { antero-posterior 25 mm.
) transverse 25.5 mm.
Long diameter of i! 18 mm.
Total length of molar-premolar series 172 mm.
Thus it is seen that while the skulls of these individuals rep-
resenting two species are of about the same size, the teeth of
Fig. 3. Skull of 2. scott7.
£. scottd (Fig. 3) are much larger and the whole masticating ap-
paratus is proportionately shorter and better adapted to grazing
than in &. caballus (Fig. 4); although the proportion of teeth
to skull is not much greater than in the pony. The teeth seem
to differ from £. eous Hay,’ a species from Louisiana described
by Cope under the name of &. zxtermedius,’ only in their some-
what larger size.
1 Science, 1899, p. 593-
2 Proc. Am. Phil. Soc., Vol. XXXIV, p. 463.
1900. | Gidley, A New Species of Pletstocene Horse. IS
COMPARISON WITH E,. CABALLUS.
The nose of £&. scotti, from the anterior premolar forward, is
as much elongated as in Z. cada//us and in this character it differs
entirely from Z, eous which has a very much shortened nose. To
make up apparently for the longer molar-premolar series, the
Fig. 4. Skull of Z. scoftz.
skull is very much shortened in the portion between the last mo-
lar and the occipital condyles and most of this shortening seems
to be in the basioccipital region, the orbits being placed much
farther back. The maxillary ridge commences about the middle
of p*, while in Z. cadallus this ridge does not usually extend far-
ther forward than the anterior portion of m’, The vertical
.
thickness of the skull measured immediately behind the last mo-
lar is about the same as in the large skull of 4. caballus, but the
thickness of the forward portion of the skull from the face of
116 Bulletin American Museum of Natural History. [Vol. XIII.
the anterior premolar to the top of the nasals is much greater in
£. scottt, The processes of the frontals inclosing the posterior
portion of the orbits, slope more backward and are much nar-
rower than in &. caballus. The occiput seems to be more over-
hanging ; this is due probably to the great shortening*of the
basioccipital bone. The basioccipital ridge is not so compressed
and the fossz inclosed between the paroccipital processes and the
condyles are much deeper. In these last two characters Z. scotti
(Fig. 5, A) is like &. occidentalis Cope (not of Leidy).’?. The pos-
terior region of the skull, the posterior nares and the palate are
narrower than in &. cadallus.
Fig. 5. A, occipital view of £. scottz; B, occipital view of £. caballus.
s
The lower jaw is more massive especially in the dental region.
The jaw is much deeper, to accommodate the very long crowns
of the molar-premolar series. This, together with the increased
vertical thickness of the anterior portion of the skull, gives the
whole head a more massive and less graceful form than that of
E. caballus, Owing to the greater antero-posterior length of the
molar-premolar series, the jaw seems to curve upward much more
abruptly from the posterior molar. The symphysis mandibuli is
heavier and longer than in &. cabadlus, extending back of the
mental foramina. The jaw seems compressed laterally at the
posterior part of the symphysis, owing to the wide expansion,
anteriorly, to accommodate the wide incisors.
The other bones of the skeleton, taken separately, seem to be
indistinguishable from those of £&. caballus.
1 Proc. Am. Phil. Soc., Vol. XXII, p. 11.
a a
er J
uf
| Rhinoceros Contributions, No. 5.
By Hery ‘FAIRFIELD Osporn.
New York, Dec. 11, 1900.
:
Article XIX.—PHYLOGENY OF THE RHINOCEROSES
OF EUROPE.
RHINOCEROS CONTRIBUTIONS, No. 5.!
By HENRY FAIRFIELD OSBORN.
By far the most striking generalization of recent mammalian
paleontology is the early separation, absolute distinctness, and
great age of numerous phyla leading up to modern types. If con-
firmed by more detailed research, the phylogeny here proposed
will bring the Rhinoceroses also under this daw of early diver-
gence ; the supposed original or stem forms having been pushed
steadily back into the older Cenozoic. It sets aside several
homoplastic characters heretofore employed in Rhinoceros evo-
lution and attempts to establish a firmer basis zz the fundamental
proportions of the skull, whether dolichocephalic or brachycephalic, in
the correlated proportions of the body, and tn the location of the horn-
cores. These characters are found to be more distinctive of
phyla than the pattern of the molar teeth.’
Our present hypothesis is that, as distinguished from the
Amynodonts and Hyracodonts, the true tertiary and modern
Rhinocerotide belong to at least six* genetic series or phyla which
have no known relation to each other. By Flower and Lyddeker the
Rhinoceroses have been placed in one genus, RAcnoceros, and
divided into five groups, which correspond approximately to our
phyla. A characteristic subfamily name is herein given to each
phylum, for the sake of clearness, brevity, and convenience, since
several of these series have a prodigious range in time, as shown
in the following table.
Eocene. |Oligocene.} Miocene. | Pliocene. |Pleistocene.| Recent.
Il. Diceratheriine eras ae ||
Il. Aceratheriine | Lake AU oe ?
(? Elasmotheriinz) | |
Ill. Brachypodine
IV. Ceratorhine | |
V. Atelodine | Et ACh eA cay
VI. Rhinocerotine
1 See Contributions 1-4, in Bibliography.
? The grouping proposed by Depéret (’85, p. 268) and by Lydekker (86) is partly upon
homoplastic characters of the teeth.
* See Osborn, ’98, pp. 77, 121; a division of the Rhinocerotide into four subfamilies.
CW
Sr ad ok a
230 Bulletin American Museum of Natural History. |Vol. X1II,
If this or some similar phylogenetic hypothesis can be estab-
lished, it will not elucidate the origin, which remains an enigma,
but it will at once simplify the whole problem of the succession,
development, migration, and taxonomy of this’ hitherto baffling
group.
: PHYLOGENY AND TAXONOMY.
A clear conception of phylogeny is an essential preliminary to
taxonomy ; the nomenclature is still, as my friend Schlosser ex-
presses it, “ein wahres Elend”; in no European or American
museum are the Rhinoceroses properly identified or catalogued.
This paper therefore, besides setting forth an hypothesis of
descent, is a preliminary statement of very interesting systematic
and comparative results obtained by visits in 1898 and 1900 to
the collections of London, Paris, Lyons, Munich, Darmstadt,
Stuttgart, Augsburg, Vienna, St. Petersburg, and Moscow.
Many kind friends aided in this work, especially the following
palzontologists: Messrs. Lydekker, Woodward, Andrews,
Gaudry, Boule, Thévenin, Depéret, Filhol, Zittel, Schlosser,
Roger, Lepsius, Fraas, and Fritsch. The recent writings of
Lydekker, Pavlow, and Roger have been of great service.
This extended comparison was undertaken before writing
Part II of ‘The Extinct Rhinoceroses’ memoir, because in
studying the American Rhinoceroses I soon learned that their
close relations with those of Europe rendered it necessary for me
thoroughly to understand the types of both countries.
The stratigraphical or geological basis is of the utmost import-
ance and is set forth in recent correlation papers (Osborn, ’oo).
As regards nomenclature: first, the discovery that the type
Acerathere, the classic Aceratherium incisivum Kaup, has @ rudt-
mentary median frontal horn, does away with the application of
the generic term Aceratherium to many of the ancestral hornless
types; second, valid reasons are found for reviving the discarded
generic terms Afelodus, Ceratorhinus, etc., and, third, the final
nomenclature will be an expression of phylogeny. The first
steps towards clearly attacking the taxonomic problem are :
(1) To conceive of the early adaptive radiation of the Rhinoc-
eroses from an, unknown stem,
(2) To conceive of the possibly independent origin of certain
1900.| Osborn, Phylogeny of the Rhinoceroses of Europe. 231
phyla in North America, Europe, Asia, or Africa, and the sub-
sequent intermingling of these phyla by migration.
(3) To recognize the succession of species in separate phyla
or lines of descent, designating them as subfamilies by the ter-
minal zv@.
(4) To sharply mark off each subfamily or phyletic series of
species from its contemporaries as soon as its earliest members
appear.
(5) To anticipate within each phylum the probable develop-
ment of collateral as well as of direct lines of species, by the laws
of local adaptive radiation.
Among the main divergent characters for the discrimination
between subfamilies or series of species are:
1. Proportions:
a. Long-skulled (dolichocephalic), and long-footed (doli-
chopodal), or long-limbed types, ¢. g., Atelodus simus.
6. Short-skulled (brachycephalic), short-footed (brachy-
podal), or short-limbed types, e. g., Zeleoceras fossiger.
2. Reduction of digtts :
a. Precociously tridactyl types, e. g., Canopus tridactylus.
6. Persistently tetradactyl types, Acerathertum tetradac-
tylum.
3. Development of horns:
a. In lateral pairs on nasals, ¢. ¢., Diceratherium pleuroceros.
6. Single on nasals, a, on tips, ¢. g., Zeleoceras, 6, on centre,
e. g., Rhinoceros.
c. In longitudinal pairs on nasals and frontals, e¢. g.,
Ceratorhinus.
d. Single on frontals, ¢. g., Aceratherium incisivum, Elas-
mothertum,
4. Cutting teeth :
a. ‘Megalodine types,’ in which the cutting teeth persist,
é. g., Rhinoceros indicus.
b. “Atelodine types,’ in which they degenerate, ¢. ¢.,
Atelodus simus.
Some of these divergent characters also become convergent or
homoplastic and are employed to distinguish the generic and
232 Bulletin American Museum of Natural History. [Vol. XIII,
specific stages of several distinct subfamilies or phyla. Thus
several ‘megalodine’ types gradually pass into ‘ atelodine.’
Correlated with the above
fundamental divergent
characters are numerous
minor characters which
are of great service ; for
ll prefossette €Xample, dolichocephalic
and brachycephalic Rhi-
noceroses frequently ex-
hibit also distinctive types
of auditory meatus, of
occiput, of premolar and
molar teeth, and of limbs.
crochet An early division is ob-
Fig. 1. Typical Rhinoceros molar, showing ter- seryed into /Aeavier and
minology.
medifossette
'
lighter types, correlated
with speed ; while collateral brachyodont (shrub-eating) and hypso-
dont (browsing) species may arise within the same phylum ; example,
A, simus and A. bicornts.
Pamily RAINOCEROTIDAS,
Oligocene phyla-—Two similar lines appear simultaneously in
the Oligocene of Europe ; the most precocious of these is the
subfamily Diceratheriinz, represented in Europe and America ;
the less precocious is the Aceratheriinz, probably represented in
both countries also. The characters of both are sharply defined.
It is probable but not yet demonstrated that the smaller Rhinoc-
eroses throughout the Oligocene chiefly represent the Dicerather1-
inz ; nevertheless it is best to leave certain species znucerte sedis
(R. velaunum, R. gaudryi), one or both of which may belong to
the Amynodontide.
Subfamily DICERATHERIIN4:. PHYLUM I.
Smaller Oligocene Rhinoceroses ; dolichocephalic, with paired nasal horns,
full-sized cutting teeth ; cursorial, long-limbed, with relatively slender bodies
well raised from the ground.
General characters.—1, Manus precociously tridactyl (as observed in Ameri-
can species), correlated with swift motion. 2. Horns developed in lateral pairs
1g00.] Osborn, Phylogeny of the Rhinoceroses of Europe. 233
on the nasals, beginning in the Middle and Upper Oligocene stages. 3. Lower
canines sub-triangular in section, flattened on outer and upper sides, slightly
convex on lower side (as observed in Middle and Upper Oligocene American and
European types). 4. First lower premolar early reduced or wanting, as ob-
served in European and American types (also in XR. gaudryi and R. velaunum).
5. Molars quadrate, frequently exhibiting a conical cingule or cusp at the open-
ing of the median valley, 6, Narrow skull, with narrow elevated occiput, ex-
panding and notched above. Zygomatic arches suddenly expanding posteriorly.
These are some of the characteristic features which are ob-
served in both European and American types and reach their
full development in the Upper Oligocene. The nomenclature is
Fig. 2. Ronzotherium gaudry?, Type: Paris. A, Internal view of left ramus. 4},
External view. A?, Superior view. 3,
still uncertain ; to the Lower Oligocene forms, which probably
possessed upper canine teeth, the generic name Ronzotherium
Aymard possibly applies. It is possible that the type species,
R. velaunum, belongs to the Amynodontide, in which case it may
anticipate the genus Amynodon Marsh. If, however, it belongs to
the Diceratheriinz it may anticipate the genus Cenopus Cope
(primitive hornless Rhinoceroses with precociously tridactyl feet),
or the genus Leptacerathertum Osborn, or Triggnias Lucas (primi-
tive hornless Rhinoceroses with persistent upper canine teeth).
To the Upper Oligocene form, Diceratherium Marsh is applicable.
234 Bulletin American Museum of Natural History. |Vol. XIII,
1. LOWER OLIGOCENE. INCERT SEDIS.
Ronzon, Argiles du Cantal, Phosphorites, Cadibona.
A most interesting primitive Lower Oligocene type is:
Ronzotherium gaudryi Rames. Type: alower jaw, Paris Museum. Local-
ity, Brons, Cantal. Definition: dentition -.- =~ -3 -3 ; second pair of lower in-
cisors greatly reduced ; median or first pair typical ; lower canines erect, laterally
compressed ; first lower premolars wanting ; premolars 3-4 with internal and ex-
ternal cingulum ; molars 2-3 without internal cingulum ; premolars 2-4 much
worn but apparently simple in pattern, z.e., without complete posterior crests.
SysTEMATIC PostTion.—The erect lower canines indicate the ex-
istence of upper canines, as in the Amynodontide or in Lefpfacer-
atherium trigonodum Osborn ; the laterally compressed shape of
the canines resembles that in Leptaceratherium and is distinct from
the more triangular form seen in Amynodon, but, if a member
of the Rhinocerotide, this animal was very primitive. Since it is
certainly not a member of the genus Aceratherium it may be pro-
visionally referred to the genus Ronzothertum Aymard, the type
of which is a lower jaw from Ronzon, similar in some respects.
The absence of the first lower premolar in &. gaudryi and RX. ve-
Jaunum is also distinctive of the Diceratheriine.
According to M. Boule the Argiles du Cantal, containing &.
gauadryt, are, if anything, a shade older than the Marnes de Ron-
zon, containing &. velaunum.
The jaw is slightly smaller than that of #. velaunum,' there is
a wider space behind the third molar ; the dentition is similar in
the simplicity of the premolar teeth ; in fact it may subsequently
prove that R. velaunum and R. gaudryi are allied.
Space occupied by lower |
rinding series, premolar -
s g Sue r (estimated)
2 to molar 3 inclusive
Ronzotherium gaudryt..... 170 mm.
Ronzotherium velaunum...194 mm.
The lower grinding series of A. gaudryZ is closely equal in size
to that of the Upper Oligocene D. minutum series (p2-m3 =
173) in the Paris Museum.
Ronzotherium velaunum 4ymard—Type: A lower jaw,
Collection Aymard, Puy Museum. The writer has not personally
examined the type and must rely upon the descriptions and figures
1M. Filhol gives no measurements but figures the jaw of 2. velawnum as = natural size
(Plate xii, figure 69, pp. 75, 266, Mammiféres Fossiles de Ronzon).
1g900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 235
(Fig. 3) given by M. Henri Filhol; as above stated the incom-
plete condition of the jaw leaves it uncertain whether this animal
3,
Wieerarsesess
>
Loey
a) ‘A ‘
Sar ee
3
=
3
is
|
to
S| Se
Fig, 3. Ronzotherium velaunum. Type: Puy. External view of left ramus. X 3, after
Filhol.
from the Lower Oligocene of Ronzon is a primitive member of
the Diceratheriine, Aceratheriine, or Amynodontide.
Characters.— Premolars 2, 3,
4, with incomplete crests ; pre-
molar I missing in the type
specimen ; coronoid and condyle
greatly elevated (as in Amyno-
don); tetradactyl, fifth digit of
manus believed to be present (as es
in Amynodon and Aceratherium). & 7) I.
WIMAGY) IN
2 S|
It should be noted that
the American Diceratheres
Fig. 4. Superior grinding teeth. A, Third and
of the Lower Oligocene fourth premolars, Cadzsoxa, Municu. 8, First
; s and fourth premolars, first molar, Phosphorites,
are tridactyl ; the Ameri- Municu. C, First and second molars, Phosphorites,
can Amynodonts (Cadur- eam a ee |
cotheriide or Amynodontide) are tetradactyl, but with a much
larger fifth digit than that associated by Filhol with R. velaunum.
236 Bulletin American Museum of Natural History. [Vol. XIII,
The most primitive species of American Rhinoceros, Z77rigonias
osborni, recently described by Lucas, presents an entirely differ-
ent type of cutting teeth from that seen in &. gaudryt,
SUPERIOR MOLARS OF DICERATHERIIN.
In London, in Paris, and in Munich are numbers of small ex-
tremely primitive molar and premolar teeth from the Middle and
Lower Oligocene Phosphorites of Mouillac, Quercy, and Bach,
also from the Lignites of Cadibona, which are for the most part
erroneously catalogued as D. minutum and its synonym, D.
croizett, specific names which were applied originally to much more
highly evolved Upper Oligocene types. In point of evolution all
these upper grinding teeth resemble the Lower Oligocene Dicer-
athere types of America, especially such species as Cenopus (Ac-
eratherium) copet ; but, as in the case of the lower jaws (of 2.
gaudryi and R. velaunum) above described, it is not possible to
determine their phyletic relations or exact systematic position at
by
)
HH]
|
Fig. 5. Diceratherium minutum. Type: Paris. Fourth premolar, first and second
molars. <4. After Cuvier.
present. It is probable that all these teeth belong either to 2.
velaunum, R. gaudryt, or some allied species. (See Fig. 4.)
Characters.— Dentition: premolars unlike molars; premolars 2-4 with
proto- and metalophs confluent upon wear ; a crista in premolars 3-4 ; m2dk pre-
molars dp?—dp* with complete crests resembling the molars ; #zo/ars with rudi-
mentary antecrochet (protoconule fold) ; more or less marked external cingulum,
rudimentary metaconule fold; variable cingulum suggesting a ‘cingule’ at
entrance of median valley (Fig. 4). ,
Lonpon, British Museum Collection: No. M. 1732, superior molars 1 and 2,
Loc., Phosphorites, Bach, Lalbenque (Lot), France (see Fig. 17, Lydekker,
1900. | Osborn, Phylogeny of the Rhinoceroses of Europe. 237
Cat. Foss. Mamm. Pt. III, p. 142); also No. M. 4507, superior deciduous pre-
molars 1-4, Loc. Phosphorites, Mouillac, France. In the MUNICH collection
from Cadibona are two small upper premolar teeth, pm 3-4, which answer this
description ; others from the Phosphorites, Quercy, are found in the Munich
collection (m1, m?, pm!, pm®). In Paris! from Quercy (Coll. Massénat) are
also small isolated premolar and molar teeth (p'—m*) of the same character,
found both in the Jardin des Plantes and the Ecole des Mines collections.
Conclustons.—The small European Lower Oligocene species of
Rhinoceroses, although zucerte sedis, are partly Diceratheriine,
partly Amynodontidz ; they should be referred to &. gaudryi or
RR. velaunum or to new species. The /arge Lower Oligocene spe-
cies of the Phosphorites should be referred to the Aceratheriine.
2. UPPER OLIGOCENE.
St. Gerand-le-Puy, Moissac, Gannat, Eselsberg (Ulm), in part.
Diceratherium minutum Cuvier.—Type: Upper premolar 4 and molars
1-2. Parts, No. 2346, Loc., Moissac, Upper Oligocene (Fig. 5).
Definition : Dentition + ¢ ;4, 3; upper incisors small ; ower canines sub-tri-
angular ; with flat outer and sharp inner edge, procumbent ; first lower premo-
lars variable ; upper and lower premolars 2-4, with two crests resembling the
molars ; upper premolars with small antecrochet, with crista and crochet ; up-
per molars with crista (soon disappearing), pointed crochet (disappearing in old
age), antecrochet and postfossette ; cusp or cingule at entrance of median val-
ley. Measurements: type p+-m* = 100; pp—m, = 173; p*-m® = 173-180.
This represents the Upper Oligocene species of the French
and German Museums, which are readily distinguished from the
Lower Oligocene species by the complication of the teeth; but
exhibit little or no increase in size. In Paris are Cuvier’s types
described in the ‘Ossemens Fossiles’; also a cotype lower jaw
No. 2343; also Duvernoy’s R. pleuroceros (synonym) type skull
from Gannat. The finest series of upper and lower teeth are
those in the Muntcu collection from Eselsberg and Eckingen
near Ulm, which are catalogued as A. croizeti Pomel. In the
STUTTGART collection from the same localities we find espe-
cially Nos. 4757 and 9861, rightly identified as D. minutum.
* Unless otherwise stated Paris refers to the Galerie de Paléontologie, Jardin des Plantes,
under the direction of Professor Gaudry.
238 Bulletin American Museum of Natural History. {Vol. XII,
Additional Characters.—PAris: Cuvier’s type: Fourth superior premolar
with protoloph and metaloph confluent in old age, small antecrochet; molars
with antecrochet, crochet, metaconule fold, postfossette, and median internal
cingule. Duvernoy’s type (R. pleuroceros): molars agreeing precisely in size
with above ; skull and jaws of dolichocephalic type, paired horn-cores on nasals,
occiput narrow elevated ; zygomatic arch convexity as in Cenopus tridactylus
Osborn. In the Paris jaw (Gannat) the first lower premolar is wanting, in
Munich and Lyons specimens it is vestigial, indicating that, as in the American
Diceratheriine, this tooth was variable ; m1-m3= 100, Tibia (Gannat) = 260.
MUNICH and STUTTGART (Eselsberg, Ulm, specimens): first lower premolar very
small ; pm 2-4 molariform with
elevated posterior crests ; lower
canines sharp, subtriangular
with flattened outer surface,
flat upper face partly destitute
of enamel, slightly convex lower
face and sharp inner edge ; ca-
nines large in males, small in
females (this tooth is very sim-
ilar to the canines of the
American species C. ¢ridactylus
Osborn) ; unworn premolars
Fig. 5a. Diceratherium minutum. A, Type (catalogued A. croizeti, Mu-
premolar and molars. Xj Paus 5, Par of left nich) as im Cuvier's type with-
out antecrochet, but with crista
and pectinate crochet (see Nos. 4757, 9861, STUTTGART, Eggingen) ; wnworn
molars exhibit crista, crochet, and antecrochet, while woz molars lose crista
and show greater prominence of antecrochet and crochet, and postfossette
(especially in p 4-m 2), also an internal cingule or cusp in the median valley as
in American Diceratheres, This animal is exactly the size of Cenopus mite of
the American Lower Oligocene. Lyons: A small jaw (catalogued A. croize¢?)
with vestigial pm 1.
A finities.—By this comparison there is little question that all
these teeth belong to the Upper Oligocene Diceratherium and, so
far as we know, to the single species D. minutum Cuvier, which
presents many features of close resemblance to the American
Diceratheres. In Paris the skull of the Upper Oligocene D.
(Pleuroceros) minutum is now placed in the case side by side with
that of the Middle Oligocene Canopus occidentalis from South
Dakota; it exhibits a remarkable similarity in the form of the
occiput, the zygoma, and the paroccipital region.
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 239
3. LOWER MIOCENE.
Sables de l Orléanats, Eselsberg (Ulin, in part), Bugti Beds.
The Eckingen, Ulm, formation also contains a Lower Miocene
fauna, indicating that the Diceratheres may have persisted into
this period.
Other indications as to Miocene persistence are those afforded
by a juvenile lower jaw and a maxillary series in the Ecole des
Mines collection, Paris, from the Sables de I’ Orléanais, typical
Lower Miocene; these were kindly shown the writer by the Cu-
rator, M. Douvillé ; they are of about the size of A. platyodon Mer-
mier, but they almost certainly constitute a new species which
probably belongs in the Diceratheriinze. The animal is consider-
ably larger than D. minutum.
Incerte Sedts.
Diceratherium douvillei,’ sp. nov.
Type: A maxillary series Coll. Ecole d. Mines, Paris. Definition : Type:
Upper premolars with crenulated anterior border of metaloph, and reduced
antecrochet ; upper molars with large crochet and antecrochet, crista not ap~
parent in worn teeth.
Fig. 6. Diceratherium douville?. Type: Paris. After a photograph by M. Douvillé.
This species is placed zncerte sedis, phyletically. The indica-
tions that it belongs to the Diceratheriinz are, first, the crenulated
or pectinate anterior border of the metaloph in the upper pre-
molars as in C. ¢ridactylus ; second, the tubercle in the valley of
* Dedicated to M, Henri Douvillé, to whose kindness the author is indebted, both for the
permission to describe the type and for the accompanying photograph (Fig. 6).
240 Bulletin American Museum of Natural History. (Vol. XIII,
m%, It is also possible that it represents an ancestor of 2. san-
santensis, which is placed in the Ceratorhine below. It certainly is
not Zeleoceras aurelianensis, and it apparently cannot be referred
to Aceratherium platyodon ; these are the only strictly Lower Mio-
cene (Burdigalien) Rhinoceroses hitherto described in France.
Subfamily, ACERATHERIINA:, PHYLUM.
Large Oligocene and Miocene Rhinoceroses of Europe ; dolichocephalic with
long, narrow nasals ; smooth or with rudimentary horns at sides of the tips ;
Sfrontals finally developing horns ; large cutting teeth ; relatively persistent tet-
radactyl manus ; long-limbed,
Contemporary with the small Diceratheres is this phylum of
large Rhinoceroses which appears to rise in a large but primitive
species in the Lower Oligocene, A.
fiholi, and pass through A. lem-
anense and A. tetradactylum into A.
incistvum of the Lower Pliocene,
which in turn is possibly the ances-
tor of Alasmotherium and the Elas-
motherine. The European Lower
Oligocene Acerathere is exactly
similar in size to A. platycephalum
Osborn, which is_ possibly the
poe Pine Ce el ee merican. "representative! of hic
premolars andsirsimigier, type; but it differs widely in the
mode of transformation of the upper premolar teeth ; for this
reason it is referred to a new species.
I. LOWER OLIGOCENE.
Phosphorites, Quercy, Cazark, Escamps.
Aceratherium filholi,’ sp. nov.
Type: Left maxilla containing second premolar to third molar inclusive ;
Paris, Coll. Rossignol, Loc. Phosphorites. Cotype: Paris, lower jaw, contain-
ing pm’, pm4, and m!? (Figs. 7, 84).
Definition: Large upper premolars, simple, unlike molars, with incompletely
formed crests ; upper molars with internal cingulum and strong protoconule fold,
small antecrochet, no crochet ; depression in posterior face of metaloph of third
1 Dedicated to my friend M. Henri Filhol, who has contributed so extensively to our knowl-
edge of the fauna of the Phosphorites.
1g00.| Osborn, Phylogeny of the Rhinoceroses of Europe. 241
molar; third and fourth lower premolars with depressed and incomplete
posterior crests. Measurements: pm?—m*=224,
This new species is well represented by teeth in the Paris,
Munich, and British Museum collections from the Phosphorites of
Quercy and Cazark, mistakenly catalogued as the Upper Oligo-
cene A. lemanense, from which it differs widely. No true A.
Zemanense remains are found in the Phosphorites, which is believed
by the writer not to extend into the Upper Oligocene. None
the less A. filhold is probably ancestral to A. /emanense and repre-
sents the first known member of the tetradactyl, dolichocephalic
phylum which directly or through collaterals leads up to Ace-
ratherium incisivum of the Lower Pliocene.
Thé distinctness of this species was independently recognized
by M. Boule, who in recent lectures has compared it with 4.
platycephalum. M. Filhol and M. Depéret also both concur with
the writer that it is distinct from A. Jemanense, and M. Filhol as-
sures me that it is equally distinct from 2. velaunum, which is a
much smaller animal. The name Aceratherium may be retained
for all members of this phylum, although technically the names
Badactherium Croizet or Aphelops Cope might be applied to the
ancestral truly Aornless Aceratheres.
The entire absence of a crochet and the non-molariform pre-
molars distinguish this species sharply from A. demanense ; the
internal cingulum is partly a sexual character ; it varies in differ-
ent specimens, although strongly marked in the type.
Besides the admirably preserved and highly characteristic
Paris types, in MUNICH we find two large molar teeth, m®? and m3
from the Phosphorites (Escamps, Lalbenque, Dép. Lot) ; also a
single well worn molar, m?, and two isolated upper molars, m!
(Phosphorites, Cazark, Dép. Lot), of exactly the same size as the
A. platycephalum from our Lower Oligocene ; also from Cazark
two upper premolars, p*, p*, which exhibit imperfectly formed
crests and a crista. In LonpoN (British Museum) are lower pre-
molars and molars (Phosphorites, Caylux, Nos. M. 1457, 1458,
1459, also upper molars M. 1455, m!—m*) all catalogued &. Zeman-
ensis. There can be no question that all these teeth belong to
the same species, 4. f/ho/i, which is far more primitive than the
Upper Oligocene A. /emanense to which they have been referred ;
not only the premolars but the molars are simpler. The premolar
November, 1g00.] 16
Fig. 8. Evolution of the grinding teeth in the Aceratheriine. A, Aceratherium filholi.
Type: Paris. Lower Oligocene. 8, Aceratherium lemanense (reversed). PARIs. Upper
Oligocene. C, Aceratherium lemanense. Ulm, Municu. Upper Oligocene. D, Ace-
ratherium tetradactylum. Paris, No. 2379. Middle Miocene. £, Acerathertum incist-
vum. Type: DarmMsTApT. Lower Pliocene. All contours, X 3
[242]
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 243
evolution differs from that of the American 4. flatycephalum as
shown in the diagram above (Fig. 8) ; other resemblances, how-
ever, are so strong that one is tempted to consider the fosszd:lity
that these animals belong to the same Aceratherine race which is
distinct from, but contemporary with the Diceratherine race. A
fossa representing the vestige of the postvallum in m® is seen in
these specimens, also in A. platycephalum.
2. UPPER OLIGOCENE STAGE.
St. Gerand-le- Puy, Gaillac, Gannat, Randan, Eckingen, (Ulm).
A. lemanense Pome/.—This well known species is represented
by superb materials in Paris, Lyons, Munich, Stuttgart. These
specimens represent different (early or primitive and later) stages
of development ; some are less progressive and probably of Mid-
dle Oligocene age, others are more modernized and probably of
Lower Miocene age, but in all the premolars and molars are far
more advanced than in A. filholt.
Sexual differences are pronounced as in the Dicerather-
iinee and Rhinoceroses generally ; the females have small canine
tusks and long very narrow or slender nasals; the males have
large tusks and rudimentary rugosities or horn supports on the
sides of the extremities of the nasals. Adaptive radiation is also
marked and probably certain collateral species are given off from
the direct line.
All these animals are readily distinguished from the Dicer-
atheriinze by much larger size.
General Distinctions of Teeth.—Lower canines lance-shaped, lenticular in
section ; first lower premolar small, usually present ; sapertor premolars with
crista, producing a medifossette upon extreme wear ; large antecrochet ; premo-
lars 2-4 with complete crests, which in the more primitive stages unite in
extreme wear; an internal cingulum. J/o/ars with strong antecrochet, becom-
ing stronger in wear ; with crista becoming weaker in extreme wear ; crochet
also becoming weaker with extreme wear ; metaconule fold becoming stronger
with wear; protocone small; postfossettes indicated in both m‘ and m*,
with internal cingulum reduced or confined to median valley. Measurements :
pm!—m*=26s5,
Liffect of age upon molar pattern.—It is extremely important to
observe that, exactly as recorded above in the Diceratheriine,
244 Bulletin American Museum of Natural History. |Vol. XIII,
the newer characters, namely the crista and crochet, are formed
near the summit of the crests and are thus worn away in old
teeth ; while the older characters, such as the antecrochet, are at
the base of the crests and thus become bolder in extreme wear,
The same law applies to the newer and older characters in the
molar teeth of the horse.
General Distinctions of Skeleton.—Skull and jaw of dolichocephalic type
(measurements, symphysis to condyles=630) ; nasals long and narrow, more or
less separate, notched at sides, slender in females ; tetradactyl, a well developed
5th metapodial, lunar wedge-shaped distally ; symphysis of lower jaw varying
with sex, short in females, longer in males.
These characters may be verified in the following specimens:
Paris, No. 2372 (Badactherium* borbonicum Croizet, type, loc.
Auvergne), an old individual with well worn molars. Duvernoy’s
fine type skeleton of A. gannatense (Gannat, Allier), probably a
female, with small lower canines and short symphysis of lower
jaw, large and powerful skeleton; skull measuring 630 from
symphysis to condyles ; superior teeth partly worn and finely
preserved ; pm!—-m*=265; femur measuring 460. Also J.
randanense, No. 2302 (Randan, Auvergne), lower jaw containing
pm 2-m 1, with a very long symphysis (unlike the 4. gannatense
type) ; this is possibly a sexual or made character. Also a com-
plete jaw (Gaillac, Tarn) with small lower canines, probably
female, small pm 1 on left side. Portion of left anterior foot,
No. 2373 (Gannat, Allier), showing characteristic tetradactyl-
ism. Lyons: (1) A. lemanense (Gannat), skull, nasals long and
thickened at the ends, but separate in median line, notched at
the sides ; this type represents an early stage, because the premolar
crests are bridged internally and would unite upon extreme
wear. (2) Large lance-shaped lower canines of lenticular sec-
tion, unworn (Allier). (3) Two maxillz from Gannat exhibit
molar and premolar characters entirely agreeing with those
above described. (4) A complete skull and skeleton, probably
female, lower jaw with small canines, medium-sized upper
canines, molars agree with Pomel’s type in character, size below
that of Pomel’s type ; nasals extraordinarily long, slender, extend-
1 This is possibly a MS. name. It is not recorded in Trouessart’s *Catalogus Mam-
malium.’
2 See Duvernoy’s Memoir, Plate viii.
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 245
ing over premaxillaries, with smooth surfaces (this length, slender-
ness, and smoothness is also a female character in C. occzdentalis
and C. ¢ridactylus) ; a sagittal crest, occiput high and narrow.
(5) Another skull (loc. Pyremont between Lyons and Geneva) has
the same general characters but the nasals exhibit distinct and
quite well marked rugosities at the sides of the tips; this is
evidently a male ; the digits are somewhat shorter than in 4,
Wemanensée, namely, Mtc. Ill=140, Mts. Il]=125; Depéret
regards the animal as a distinct species and will describe it as
such. MunicH: (1) The maxilla from Eselsberg, Eckingen,
near Ulm, is beautifully preserved ; it belongs to an early stage
because the premolar crests are bridged and unite when worn ;
the premolars exhibit medifossettes ; the molars show the strong
crochet, antecrochet, and metaconule folds. (2) An unworn
molar from Eckingen shows a crista, antecrochet, and crochet,
and the characteristic small protocone of this species. STUTT-
GART: (1) Molars of a /ater stage (Eggingen, Ulm) show a more
prominent crista and crochet which unite to form a medifos-
sette. (2) A fine pair of lower jaws (Ulm) with large lower
canines and no traces of pm 1 probably also represent a later
stage ( 7. e., Lower Miocene).
We thus find that 4. Zemanense is the characteristic Upper Oligo-
cene species, presenting various stages of premolar transforma-
tion and probably giving rise to some collateral species.
3. LOWER MIOCENE STAGE.
Sables de 2 Orléanais, Royans.
A. platyodon AZermier, represents this stage-—Lyons: The
type skull, probably belonging to a small female, pm!—m*=207,
exhibits unique, extremely elongate, slender, and slightly separate
_nasals ; the lower canines, as the specific name indicates, are ex-
cessively flattened toward the extremities but exhibit a triangular
mid-crown section ; the premolars (Mermier, ’96, Pl. IL) have a
prominent crista and medifossette ; the crests unite early upon
wear. The teeth may be readily distinguished from those of the
contemporary Brachypodinz by the small size of the protocone.
A. blanfordi Zydekker.—A jaw is ascribed to this species
246 Bulletin American Museum of Natural History. (Vol. XIII,
(Lydekker, ’86) from the Lower or Middle Miocene Bugti Beds
of Sind. Itis zwcerte sedis here.
4. MIDDLE MIOCENE STAGE,
Sansan, Simorre.
A. tetradactylum Zar‘e/.—This is the noble species of Sansan
(Nos. 3378 male, 2379 female, 2389 female, etc.) and Simorre,
represented finely in the Paris Museum. It shows striking re-
semblances to A. /emanense, together with all the progressive char-
acters which we should expect to find in a descendant, and
unquestionably belongs to the same line. The scapula is high and
narrow as in dolichopodal types generally. The hind limb
Fig. 9. Aceratherium (incisivum) tetradactylum. Georgensgemiind. MunIcn.
(femur and tibia) is of approximately the same length as in 4.
Zemanense, but the metapodials are longer, and more stilted (Mtc.
I1l=160-180, Mts. IJJ=135-165), indicating that the phylum
was developing a progressive running power. ‘The progressive
and retrogressive changes in the skull and jaws are most interest-
ing and significant as seen in a magnificent male specimen (No.
3378, Loc. Sansan). As compared with A. /emanense note the
following
SKELETAL DISTINCTIONS.
Skull.—A slight loss of size, symphysis to condyles of skull = 559, A. Zetra-
dactylum ; symphysis to condyles of skull = 630, 4. /emanense. Nasals, males
(No. 3378) slightly less elongated, similarly notched at sides, roughened or ru- -
gose distally, but not thickened (a sexual character) ; females (Nos. 2379, 2389,
Coll. Lartet) very narrow and elongate, separate in median line, not expanding
distally. Premaxillz slender. Occiput elevated, spreading superiorly. Sagit-
tal crest lower but still well marked. Premaxillz slender. Jaw with elon-
gate symphysis, wide diastema, angle deep, projecting backwards. Manus
tetradactyl (Coll. Lartet, Nos. 2518, 2537), with reduced metacarpal V ; lunar
of tridactyl type (foreshadowing the loss of metacarpal V, which is now reduced
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe, 247
in length to 75), PArts, Coll. Lartet, Nos. 2518, 2537; long, stilted digits ;
metacarpal III enlarged ; metacarpals II, IV relatively smaller. Scapula ver-
tically elongate, with long neck, as in all long-limbed, speedy types.
Dental distinctions.—Inferior canines less lance-shaped, with internal flare
and flattened external section. First lower premolar sometimes present (in the
slightly older Sansan specimens). Sup. premolars and molars (No. 2379) with
somewhat reduced antecrochet and very strong crochet placed near ectoloph ;
this unites with the ectoloph when well worn and forms a conspicuous medi-
fossette (No. 2388). Crista conspicuous in unworn premolars and molars. An-
tecrochet somewhat reduced and becoming conspicuous only in old or worn
teeth. Molars with cingulum entering median valley between crests, a crest in
the bottom of median valley (No. 2388), or embracing protoloph only ; with
postfossette in m? ; and traces of external cingulum ; posterior cingulum of m*
(also observed in A. platycephalum and A. lemanense) persisting. Measure-
ments, female, pn? — m = 230.
The scapulz and limb bones of large size in the Paris Museum
(which are catalogued R. sansaniensis) undoubtedly belong to
A. tetradactylum. Some of these indicate an Aceratherine race
as large as or larger than the A. demanense type.
Paris: This species is also represented in the slightly higher
levels of Simorre, especially by a very large jaw of a female, with
small lower canines, first lower premolar absent, formula: p3;—
m3 ; of marked dolichocephalic type. Lonpon: The jaw from
Sansan (Hastings Collection, No. 27454, catalogued 2. goldfusst )
also represents this species; it is large and dolichocephalic in
type; the lower canines are flattened with a marked internal flare.
5. Upper MIOCENE STAGE.
Georgensgemiind.
MunicH: Upper teeth with closely similar characters (cata-
logued A. iucisivum, Georg-
ensgemiind, Bav.) are seen
here in a shade earlier stage
of evolution, because we
observe more of a bridge
between the premolar crests
and somewhat greater
prominence of the antecro- Fig. 9a. Aceratherium tetradactylum. No.
: 3378. X §%. Paris.
chet as well as of the internal
cingulum of the molars. fAnother specimen is a characteristic
248 Bulletin American Museum of Natural History. [Vol. XIII,
long and straight lower jaw of this race. (A maxilla, mistak-
enly catalogued A. cncisivum, Georgensgemiind, belongs to the
Ceratorhine or C. sansaniensis race.) VIENNA: A large maxilla,
containing pm 1—m 3 (without label) has all the distinctions of
the Aceratherine race.
6. LOWER PLIOCENE STAGE.
Lippelsheim, Maragha.
Relations of A. tncistvum to Elasmotherium.
Aceratherium incisivum Aauwp.—Darmstapt: In cranial
characters this classic species is less dolichocephalic. In dental
characters it follows closely upon its predecessors (Fig. 8 #) ; in
fact, most writers, beginning with Kaup, have not hesitated to unite
the A. tetradactylum with this animal. The cranial characters, how-
ever, are much more progressive, the nasals are shorter and more
upturned, the frontals are thickened and bore a rudimentary horn
in the males at least. The latter character (Osborn, gg, p. 162) is
very significant. One can imagine that this phylum, having failed
in the development of horns upon the mechanically weak nasals (as
indicated in the Lyons specimen), began to evolve frontal horns.
There is no question that
the frontals are not only
thickened to support a horn
(they are very thin in the
contemporary 7. drachypus),
but that they show a well
marked rugosity with the
characteristic converging
depressions of nutrient ar-
teries (Fig. 10). It is this
character which led the
writer to advance the idea that this animal is an ancestor of
Elasmotherium, an hypothesis which depends upon the future
discovery of intermediate forms. It may be observed here, more-
over, that Elasmothertum has long, narrow, smooth nasals of a type
found only in the Aceratheriine and that there is theoretically no
‘f
<, Frontal, j
Fig. 10. Aceratherium incistvum. Type:
DarmstapT. Rudimentary frontal rugosity, with
lines of convergent nutrient arteries.
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 249
difficulty in deriving the enormous frontal horn of the Pleistocene
species from the vigorous rudiment in 4. zncisivum.; or the pty-
chodont Pleistocene molars from the simple Lower Pliocene stage.
There is a fine skull of A. zuc¢stvwm in HALLE as well as the
two in Darmstadt.
7. EASTERN TYPES,
Incerte Sedis.
The Siwalik Aceratheriinz have not yet been carefully com-
pared by the writer. A. perimense is a very large animal from
Perim Island with a skull which, as restored by Lydekker (’81,
Pl. X), suggests this phylum, although higher and shorter.
Subfamily BRACHYPODINA. PHYLUM III.
Brachycephalic Rhinoceroses, short broad skulls. Teleocerine, horns when
developed appear on tips of nasals, Megalodine, large cutting teeth. Brachypodal,
short spreading feet, short limbs, body and trunk near the ground. Tridactyl,
probable early reduction of lateral digits. Known Geological Distribution,
Lower Miocene to Lower Pliocene, inclusive, Europe and America.
These Rhinoceroses, short and broad in all their proportions,
including their spreading grinding teeth, represent, so far as we
know, the sudden occurrence of a new type in the Lower Mio-
cene of Europe ; for they have no known prototypes in the Oli-
gocene of either Europe or America. Either the original home
of this type is Africa, and if so, they came into Europe with the
Mastodons, or they represent an offshoot of the Aceratheriine.
Typical species are 7. aurelianensis Nouel; 7. brachypus Lartet ;
T. goldfusst Kaup ; T. fossiger Cope. Doubtful species are A.
persia and A. blanfordi. The phylum Brachypodine takes its
name from one of the oldest known forms, 7°. drachypus Lartet,
although it first appears geologically in the 7. aurelianensis Nouel
of the Lower Miocene (Sables del’ Orléanais of France), and in-
cludes a great variety of European and American types, extend-
ing to the Lower Pliocene, 7. goldfussi Kaup. The feet in 7.
brachypus and 7. fossiger become extremely short. Associated
with the shortening of the skull is a shortening and broadening
of the grinding teeth—the very broad fourth upper premolar dis-
tinguishes the higher members of this series, notably as developed
250 Bulletin American Museum of Natural History. [Vol. XIII,
in the Lower Pliocene 7°. goldfussi Kaup. In the superior molars
the protocone is very prominent and rounded, giving a circular
form in extreme wear. The lower and upper cutting teeth at-
tain an enormous size, hence the adjective megalodine is appro-
priate. The shortening of the skull lowers the middle portion
of the cranium and in the typical species causes the nasals to
project upwards at the tips; thus the irresistible tendency of
every Rhinoceros to develop a horn finds expression in the later-
ally compressed rugosities of the tip of the nasals (7. aurelianen-
sis, T. fossiger), while an abortive horn may appear on the frontals
(7. aurelianensis). The strong resemblance of 7. fossiger to this
series was noted by Mme. Pavlow.
The generic name 7Ze/eoceras Hatcher is the first applied to a
member of this series and will be of service to distinguish its
members throughout. Valid specific differences are found be-
tween the Lower, Middle, and Upper Miocene and Lower Plio-
cene stages ; there are certainly three and possibly four species
in Europe.
1. LOWER MIOCENE STAGE.
Sables de f Orléanats.
Teleoceras (R.) aurelianensis MVoue/—Type: A skull.
Loc. Neuville-aux-Bois, Loiret.
Paris: Characters of type. (1) Three lower premolars in jaw
associated with skull ; also observed in an isolated better preserved
jaw, thus: pg2z, m#; flattened outer face of inferior molars is
Fig. 11. Teleoceras aurelianensis. Type: Superior molars. X §. Paris.
another characteristic ; superior premolars 3—4 shorter than the
molars and provided with antecrochet, as seen also in maxillary
series (Collection Vibraye). The strong simultaneous develop-
ment of antecrochet and crochet distinguishes the molars, as well
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 251
as the stout cylindrical protocone. Occiput broad, in a vertical
plane ; jaw with a decided angle. Front view of nasals is very
characteristic (Fig. 12, D), a rugosity appears upon each tip with a
cleft between (as in 7. fossiger); there is also a very slight swell-
ing and faint rugosity upon the fronta/s immediately above the
eyes which may indicate the rudiment of a median horn. As
compared with the Aceratheriine the fore and hind limbs and
feet in the Paris Museum are relatively short, but they are longer
than those of its Upper Miocene relative 7. fossiger mounted be-
side them ; this species is also distinguished by narrower pre-
molars. (2) A fine lower jaw (Loiret) exhibits premolars with
flattened outer wall, also a very small, single-fanged pm,. (3) The
maxillary series (Coll. Vibraye) above alluded to shows a cement
layer on the molars as in some Upper Miocene American types.
Lartet in a letter to Nouel expressed the opinion that this spe-
cies was identical with the Middle Miocene 7. brachypus. A
close comparison of the teeth of these two species in the Paris
Museum reveals the following resemblances and differences :
T. aurelianensis. T. brachypus.
Premolars broad Premolars broad
Antecrochet strong in p3, p4 Antecrochet reduced or wanting
Metaloph of p* long Metaloph of p* long
Molars, internal cingulum wanting Cingulum strong
2. MIDDLE MIOCENE STAGE.
Simorre, Sansan (?),
T. brachypus Zarte¢—Paris: A fine maxillary series from Simorre, Gers
(Coll. Lartet No. 2386); fourth superior premolar (p+) compressed antero-
posteriorly and extending transversely more than in 7. aurelianensis ; superior
molars with strong internal cingula ; inferior molars with extremely flattened
outer faces; enormous upper incisors. This species is generally said not to
occur at Sansan, but a single lower canine tooth (catalogued 2. tetradactylum,
Paris Museum) probably represents it on this level.
3. UppER MIOCENE STAGE.
Grive-St.-Alban, Steinheim.
T. brachypus.—The finest examples of this species are found
in the Muséum d’Histoire Naturelle of Lyons and have been
described and figured by Depéret. Relying upon his determin-
ation (’87, p. 178) we observe the following dental characters :
252 Bulletin American Museum of Natural History. [Vol. X11,
Superior premolar I simple ; premolars 3-4 without antecrochet (thus differing
from 7. aurelianensis); premolars 2-3 with small crista and crochet ; premolar 4
with crista and forked crochet ; molar I with small crista, strong crochet, ante-
crochet reduced (as compared with R. awrelianensis); molars 1-3 with internal
cingulum extending around inner face. /ferior premolars with flattened outer
faces. Measurements, Mts. III = 110.
Depéret observes that the true 7. drachypus always has an in-
ternal cingulum upon the upper molars. I do not, however, feel
convinced that this specific determination 1s correct.
Hate: A distinct variety of this type occurs at Steinheim,
and was shown to me in this fine collection through the kindness
of Professor Fritsch ; it is distinguished by very thick enamel,
square posterior fold of ectoloph (due to the antero-posterior
compression of the dentition correlated with the brachycephalic
skull), crista, crochet, and antecrochet all showing in well worn
superior molars; cingulum only around protoloph of molars
(in typical 7. drachypus it embraces metaloph also). This vari-
ety may become known as a distinct species, 7. eurydactylus, for
there certainly are some minor differences between this and the
typical Z. drachypus.
MunicH: The foot bones of manus and pes, Mtc. III = 145,
Mts. III = rio, astragalus = 50 (types of 2. eurydactylus Haushal-
ter), are almost identical in size and proportions with those of our
Upper Miocene species, 7. fossiger Cope, of America. A lower
jaw (Steinheim) exhibits the following characters : symphysis,
short ; diastema very short, first lower premolar vestigial, single-
fanged, close to canine; second lower premolar comparatively
simple, reduced, single-lobed. There is also a fragmentary skull
from the Dinotherium Sands near Giinzburg with occiput low
and broad as in 7. fossiger. Also from Steinheim a large col-
lection of isolated upper molars, with the following characters :
superior fourth premolar broader than first molar (as in 7. gold-
fusst); superior first molar with very thick enamel, a crista, large
antecrochet, and broad internal cingulum extending around pro-
toloph only. The Steinheim teeth of Munich therefore agree
closely with those in the Halle collection and indicate that the
northern (? 7. eurydactylus) variety differed in a definite particular
from the southern typical Z. drachypus race, namely : cingulum
extends around protoloph only ; this character (cingulum around
1900.|] Osborn, Phylogeny of the Rhinoceroses of Europe, 253
protoloph only) is also observed in a cast of four molar teeth
(Mantscha bei Graz) in the Munich collection, but it is not seen
in the Augsburg skull.
H ca Ye
Fig. r2. Characteristic types of nasals and nasal rugosities. A, Aceratherium tetradac-
tylum, Paris. B, 4. demanense, Lyons (Pyremont) C, A. zecistzum, HALLE. D, Teleo-
ceras aurelianensis, Paris. E, /ucerte Sedis, AUGSBURG. F, Rhinoceros sansaniensis
(restored). G, R. Alatyrhinus. HH, Atelodus bicornis. 1, A. simus. Mostly after rough
sketches by the author.
AuGsBURG: Other characters of this variety are given by the
fine specimens in this museum shown to the writer through the
kindness of Dr. Otto Roger, also communicated by letter (May
3°, 1899), but especially in his very full and valuable paper
254 Bulletin American Museum of Natural History. [Vol. X11,
(Roger, ’0o), received just as this paper was going to press.
Locality : base of lower Dinotherium Sands, near Augsburg.
Very aged skull (Roger, ’oo), found at base of sands, short and
massive ; premolars and molars with strong cingulum surround-
ing three sides of the crown ; p'—-m? = 260. Measurements:
p-1 2 Zi 4 m. I 2 3
Length 27 29 > 35" 37 40 4oy 58h
Breadth 18 35 48 57 56 56 53
Breadth greatly exceeding length throughout upper grinders ;
superior diastema i-p = 85; zygomatic arches strong; occiput
as broad above as below; supra-temporal crests separated by a
median groove ; in mid-frontal region bones as thin as paper (an
important distinetion from A. zzcistvum); nasals a single, compact,
laterally compressed bone without trace of median cleft or suture
(thus differing from 7. aurelianensis), thickened at the extremity
but not laterally compressed as in 7. aurelianensis and 7. fossiger ;
meatus auditorius open below (resembling A. swmatrensis). Iam
inclined to regard this as a female skull and to believe that a
male would show rugose nasal tips; in fact, nasals with rugose
tips were described and figured by Roger (’85) (Fig. 12, &).
This animal shows decided specific differences, but an unmistak- |
able racial resemblance to both 7. aurelianensis and T. fossiger.
Other characters of this species observed in Augsburg specimens |
are, lower jaw : short symphysis ; very short diastema (10); small
coronoid process bending sharply forward ; small incisors ; first
lower premolars always wanting; outer face of lower grinders
flattened ; total premolar series = 111.5, molar series = 165 ; den-
tal formula, + 9 4 2; grinding teeth large in proportion to skull.
STUTTGART: Teeth measurements: ~~ Pm*, breadth, "6e
length, 40. Limb measurements: Steinheim collection (Roger,
‘oo, pp. 16-17), humerus, length, 350 to 420; radius, 290 to 370;
femur 390 to 540; tibia, 290 to 340; metatarsals, I = 108,
’
lil = 112, IV = 963 "these measuremetits: indicate thats tm.
limbs are somewhat longer than those of 7. fosstger (cf. Os-
born, ’98, p. 57). The pointed vestigial first lower premolar is
preserved in one jaw; lower incisors small and sharply pointed.
A maxillary series (Steinheim, No. 6314) is referred by Roger
(‘oo, p. 14) to this species, mistakenly, I believe, because the
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 255
long narrow measurements of the grinding teeth indicate that
they belong to a dolichocephalic type, probably A. ¢etradactylum.,
A jaw recently excavated by Professor Fraas himself (Steinheim)
exhibits small pointed incisors and a vestigial pointed pm,.
LOWER PLIOCENE STAGE.
Lippelsheim.
T. goldfussi Kawp.—The foregoing studies enable us to de-
termine that the tooth which Kaup selected from the sands of
Eppelsheim for the zype of this species is not a molar, as he sup-
posed, but a fourth superior premolar ; this tooth has a broad in-
ternal cingulum (‘ Ossements Fossiles,’ Darmstadt ; in ‘Akten d.
Urwelt,’ 1841, he adds as cofyfes, a lower molar, and upper incisor ;
in ‘ Beitr. z. Naher. Kennt.’ he figures an upper molar, Taf. II, fig.
20; a lower molar, fig. 15). In the same Eppelsheim sands are
found other teeth with characteristic peculiarities of this brachy-
cephalic-megalodine phylum, viz.: greatly enlarged upper in-
cisors, upper molars with crochet and antecrochet projecting into
median valley, lower molars with flattened outer wall.
T. goldfussi is very imperfectly known ; it cannot now be dis-
tinguished specifically from 7. drachypus, except by its larger size.
So far as we know it was the last member of the subfamily
Brachypodine.
Types. Incerte Sedis.
The Siwalik Rhinoceroses have not yet been carefully examined
by the writer. The Aceratherium blanfordi Lydekker, type, re-
sembles the Brachypodine in the structure of its superior molars.
From the Lower Pliocene or Maragha is another remarkable in-
teresting form, A. ferste Pohlig, which appears to be distinct
from A. dlanfordt.
Aceratherium persiz Poflig.—This species is richly repre-
sented in VIENNA (Collection Polak) by ten more or less complete
skulls ; there is also a fine skull in the HALLE Museum. Charac-
ters: Last superior molar quadrate with an exceptional extension
of ectoloph, and a vestige of posterior valley ; ectoloph of molars
in a nearly straight line; antecrochets and crochets of molars
very prominent, giving a complex pattern upon extreme wear ;
256 Bulletin American Museum of Natural History. (Vol. XIII,
enamel thin ; protocone large, strongly constricted off. These
dental characters approach those of Ze/eoceras fossiger.
In some of its cranial characters
it approaches A. zucisivum, except in
the extraordinarily broad chin which
is hollowed out in the median line
: (see also lower jaws referred to A.
Pres —entivcoonee Blan fordi in British Museum); thus
le a ey ee the lower canines diverge and are set
perond and (itd euperncr wnbiate: widely apart, with the persistent alve-
oli of lower incisors between ; nasals
short, straight and smooth; a sagittal crest; occiput higher
than broad ; zygomatic archslender. A strong rugosity upon the
molars beneath the eyes for the origin of the masseter muscles,
which are inserted in a strong ridge on the outer border of the
angle of the jaw. Tibia and fibula closely united (as.in Brachy-
podine). Limbs of medium length.
erochet erista
7
Subfamily CERATORHINAt. PHYLUM IV.
Middle Miocene to recent Rhinoceroses ; dolichocephalic, with frontal horns,
and nasal horns upon a distinct mid-nasal convexity, not terminal ; nasals more
or less pointed and recurved anteriorly ; cutting teeth large in early members,
gradually reduced in certain branch phyla ; cursorial limbs.
The first known of this series, 2. sansaniensis of the Middle
Miocene, appears to represent a new arrival and a new phylum in
Lurope ; it certainly has no ancestors among the previously
known Diceratheriine, Brachypodine, or Aceratheriine, for the
structure of the entire upper portion of the skull is different ; it
is barely possible that some of the teeth referred to 4. minutum
from the Upper Oligocene may represent its ancestors; but
this is not probable. Its successors or collateral descend-
ants, however, are probably determined as the &. szmorrensis of
Simorre, the 2. steznheimensis of the Upper Miocene of Steinheim
and Grive-St.-Alban ; these animals apparently gave off: (4) a
smaller race, the last of which appears in the Lower Pliocene, Ep-
pelsheim ; in Eppelsheim and Pikermi, however, there also ap-
pears (B) the larger race of R. schleiermacheri possessing many of
the same characters as &. sansaniensis, but with certain notable
distinctions.
1900.|] Osborn, Phylogeny of the Rhinoceroses of Europe. 257
The generic name RAinoceros may be retained at present for
members of this series, but in case a relationship to the Sumatran
1o| Ce
Fig. 13. Superior molar series. A, R. sansaniensis. Type: Paris. B, R. simorrensis,
No. 2380. Paris. C, R. schleiermacheri. StTutTTGart, after Kaup. All x 3.
Rhinoceros should subsequently be demonstrated, it would be
well to apply Gray’s term Ceratorhinus throughout.
A. Smaller Race. Middle Miocene to Lower Pliocene.
I. MiIppLE MIOCENE STAGE.
Dental Characters.—Large lower canines (males) ir Miocene ; first lower pre-
molars relatively persistent (unlike Diceratheriinz),1 tained to Middle Miocene,
then reduced ; upper premolar transformation retarded, crests confluent upon
wear in X. sansaniensis, free in RK. stmorrensis ; upper molars and premolars
with internal cingula reduced or absent ; molar-premolar series of moderate
length (pm? — m® = 190 in 2. sansaniensis), much shorter than in the contem-
porary Aceratheriinz, proportionately longer and narrower than in the Brachy-
podinz ; molars retaining a feeble antecrochet.
[November, rg00.] LT
258 Bulletin American Museum of Natural History. (Vol. XIII,
Cranial characters.—Nasals short and broad, triangular when seen from
above ; median horn precociously (Middle Miocene) developed upon both nasals
and frontals; occiput both broad and high (very distinct from Aceratherine,
Diceratherine, or Brachypodine types).
a. Lower Level, Sansan.
R. sansaniensis.—Paris: The type skull (No. 2395, Coll.
Lartet, Sansan) is that of a made, a small animal; it is very much
crushed antero-posteriorly, disguising its real dolichocephalic char-
acter, which is strongly marked in the uncrushed lower jaw; the
first lower premolar has a broad double or grooved fang, while in
R. simorrensts this tooth is small and single-fanged ; the premolars
are greatly worn so that the median valley has almost disappeared
Fig. 14. Lower grinding series. A, 2. sansaniensis. Paris. B, R. simorrensis. Paris.
and the crests are quite confluent : although a male (because of
its well developed horns) the inferior canines are smaller than in
the 2. simorrensis jaw ; the simple character of the molar crests
in this specimen is deceptive, and is due to extreme wear, the
crochet (a superficial fold) having been worn off, the protocone
and antecrochet are indistinctly marked (quite unlike the Ace-
ratheriinz and Brachypodine of this geological period) ; the in-
ner face of the molars is without cingulum (unlike Aceratheriinz
and Brachypodine) ; there are indications that young teeth would
show both crista and crochet ; the nasals and occiput have a very
characteristic shape, somewhat similar to that of Gaudry’s &.
schleiermacheri of Pikermi; the nasals are especially distinctive,
being broad and rugose behind, where they carry the horn,
but converge to a smooth point anteriorly; (See Paris, Nos.
2395, 551); metatarsals (erroneously catalogued A. setradactylum),
of moderate length, probably belong to this species.
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 259
b. Higher Level, Simorre.
R. simorrensis Zarte¢—Paris: (1) In this animal from a
higher level (100 metres) we observe distinct specific progression ;
the size is the same; the lower canine is larger; the first lower
premolar is single-fanged, reduced or wanting ; the crests of the
upper molars (Lartet Collection: No, 2380, catalogued as A.
tetradactylum) are more distinct; the younger molars have a
feebly indicated antecrochet and a very strong crochet ; upon
extreme wear the antecrochet comes out more strongly. (2) A
handsome lower jaw shows the vestigial pm, persisting on the
left side, wanting on the right (Fig. 14); the lower grinders are
small, fine, and delicately built ; the diastema is rather short.
(3) A maxillary series (Coll. Lartet, No. 2380) is beautifully
preserved ; the fourth superior premolar is fully molariform,
with a prominent antecrochet ; the superior molars show a re-
duced antecrochet and a very prominent crochet. Measure-
ments: pm! — m3 = 193 (Coll. Lartet, No. 2380); pm, —.m;
— 195.
Lyons: Two fine maxille of &. s¢morrensts are found in the
Muséum d’Histoire Naturelle and present characters exactly
similar to the above ; strong postfossettes are observed in pm? —
m! and strong and prominent crochets on pm” — m§; the molars
have the internal cingula feeble or wanting. Lonpon: Upper
jaws and teeth (No. 33525, Villefranche, d’Astarac, Gers, France)
a beautifully preserved apparently fema/e skull with no trace of
median horns on frontals ; molar teeth with the same characters,
p! —m? =193. A lower jaw (No. 33526, same locality) ex-
hibits a single-fanged and evidently much reduced first premolar.
The 2. austriacum Peters is represented in Munich by the third
superior molar tooth. The type of this species from Eibiswald-
Leiding is doubtfully distinct from the foregoing.
2. Upper MIOCENE STAGE.
Steinheim, Grive-St.- Alban.
The Upper Miocene stage of this small race is the so-called
R. steinheimensis Jager, from Steinheim. (1) The finest example
of this stage is a maxilla in SrurTGarrT (Steinheim, No. 6032);
260 Bulletin American Museum of Natural History. [Vol. XIII,
pm! is quite simple ; pm? shows a crista and small antecrochet ;
pm ® shows a prominent crochet ; pms °* differ from molars in
the absence of antecrochet fold; the molars show a crista, re-
duced antecrochet, and very prominent crochet. (2) Another
maxilla (No. 4230) shows a larger size, pm! — m® = 200,
Lyons: similar teeth are found from Grive-St.-Alban. It is
probable that this stage represents a distinct species, in which case
it should be termed &. stecnheitmensis Jager ; at present, however,
we know no means of distinguishing it from 2. s¢morrensis. Mu-
NicH: A fine example of maxillary series from Georgensgemiind
(catalogued A. znciscvum) exhibits premolars and molars without in-
ternal cingulum ; the premolars have complete internal crests.
Conclusions.—T here is a gradual advance in size (molars from
190 to 200) and in the evolution of the premolars, as we pass
from the Middle to the Upper Miocene Ceratorhine.
3. LowER PLIOCENE STAGE. LasT OF SMALLER RACE.
Lippelsheim.
R. steinheimensis.—Kaup referred the smaller teeth of
Eppelsheim to A. minutum Cuvier ; this was an error. One of
these Eppelsheim teeth, a third superior molar, is
in Lonpon (British Museum, No. 1257); it agrees
closely in every particular with those of &. sémor-,
rensts both in size and character; it is a much
worn tooth and shows a large antecrochet. Casts
Tie eae of the Eppelsheim molars (M. 2739, 2740, 2742) are
steinheimensis. also identical with those of /. stmorrensis. DaRM-
ast superior s, a A a os é 2
molar. Stein- sTrapT: An examination of Kaup’s originals in this
heim, Municu. F :
Museum confirms the above determination (see
Kaup, ’62, Taf. II, figs. 6, 10, 11, 13). There is little doubt, there-
fore, that this smaller race of Ceratorhine persisted in the Lower
Pliocene ; the specific characters of this stage are undetermined.
BL. Larger Race.
4. LOWER PLIOCENE STAGE.
Lppelsheim, Pikermi.
We can imagine that the smadler race arrived in Europe (either
from Asia or Africa), was arrested in size development and
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 261
terminated in the smaller Eppelsheim species; then from the
same original stock, by subsequent migration, a collateral larger
race arrived, which in general had developed along the same
lines, but had retained certain primitive characters.
Such a collateral species is Kaup’s 2. schletermacheri of Eppels-
heim (Fig. 13, C). It exhibits molar-premolar teeth measuring
260; it is thus nearly one-third larger than A. steinhetmensis ; it
resembles the #. sansaniensis series in the following points:
superior molars: antecrochets reduced ; a crista (progressively
bifid); a prominent crochet; skeleton: metapodials of medium
length ; tridactyl manus. It differs as follows: premolars with
crests internally confluent upon wear (primitive); first lower pre-
molar persistent (primitive); a sagittal crest (primitive); small
cutting teeth (progressive); very large nasal and frontal horns
(progressive); no postfossettes in the molars; wide distance be-
tween orbit and naso-maxillary notch (this space is somewhat
shorter in 2. sansaniensts, indicating a progressive lengthening of
the skull in 2. schletermachert ).
Therefore, as placed together in the Paris Museum the Middle
Miocene &. sansaniensts and the Lower Pliocene (Pikermi) 2.
schleiermacheri exhibit first a striking racial similarity in form;
second, a difference in size exactly such as one would expect in
the progression from a Middle Miocene to a Lower Pliocene type;
third, certain primitive and progressive differences which render
the theory of direct descent of one from the other impossible. If
one compares the skulls closely one sees the striking racial like-
ness in the form, and especially in the proportions and positions
of the horns upon the nasals ; the occiput of &. schleiermacheri is
relatively lower and is somewhat broader below. In both speci-
mens the infraorbital foramen is very close to the naso-maxillary
notch ; thus it is evident that these species, although not geneti-
cally related, represent collateral branches of a similar race. The
growth of the skull between the orbit and anterior nares points
to progressive dolichocephaly and to correlated elongation of the
limbs and feet.
The successors and relatives of this Ceratorhine phylum are,
apparently, 7. deptorhinus Cuvier, Middle Pliocene, represented
by a fine skull (Paris Museum, Montpellier, Hérault); the long-
limbed &. etruscus from the Upper Pliocene of Italy, France, and
262 Bulletin American Museum of Natural History, [Vol. XIII,
England, with a nasal septum; 2. platyrhinus of the Pliocene
Siwaliks of India (which Lydekker has mistakenly associated with
the Atelodinz); finally, the smaller and somewhat primitive
living species, A. suwmatrensis.
Sublamily ALELODINAL PHYLUM. W.
Lower Pliocene to recent Rhinoceroses. Dolichocephalic, long low skulls,
moderately broad, depressed, backwardly inclined occiput; two large horns
developed upon nasals and frontals; nasals square or blunted anteriorly,
horns extending to the extremities; Atelodine, cutting teeth vestigial or
wanting ; mesopodal, moderately long limbs and digits, similar to those of R.
indicus.
1. LOWER PLIOCENE STAGE.
Pikermi, Maragha.
In the Lower Pliocene of Pikermi there suddenly appears in
Europe a fifth type which cannot be derived from any of the
preceding ; the cutting teeth are precociously vestigial or wanting
(hence the term Atelodinz) ; the skull is easily distinguished by
the form of the temporal fossa and occiput, by the form of the
nasals and by the absence of front teeth in the dolichocephalic
megalodine &. schleiermacheri, which appears in the same beds.
The species is not found in the more northern Eppelsheim beds,
and in view of the many resemblances which the Pikermi type,
R. pachygnathus, bears to the existing African species (2. simus,
R. bicornis), we may not consider as unreasonable the hypothesis
that this is an African phylum which entered southern Europe
with the numerous Antelopes and Giraffes of Pikermi; the later
members of this phylum are the Pleistocene &. hemitechus and
R. antiquitatis (= tichorhinus),and the recent &. simus and R.
bicornis. The main characters of this phylum are given above.
R. pachygnathus Wagner.—Paris: A fine skull and skeleton
of this type have been described and figured by Gaudry. Even in
the young skull there is a decided thickening for a frontal horn ;
the nasals are very broad and thick at the extremities ; the lower
jaw is without distinct angle, and a single convex sweep from
condyle to angle is very characteristic ; correlated with this we
observe a weak zygomatic arch and early reduced front teeth ;
the most distinctive feature is the backward sweep of the tem-
poral fossa, the low, backwardly inclined occiput. The molars are
brachyodont. In the older jaw the formula is: igcegpsmg.
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 263
Duvernoy actually attributed 2. pachygnathus to the Pleistocene
species A. antiguitatis, from resemblances in the limb bones, and
Gaudry remarks (’62, p. 177) that this was very natural because
the bones are extremely similar, Again, as originally remarked
by Gaudry, 2. pachygnathus resembles 2. bicornis (the smaller
brachyodont shrub-eating species of Africa), and (’62, p. 178)
closely also &. stmus (Burchell’s Rhinoceros, the larger hypso-
dont, grass-eating species of Africa); I have verified these
remarks by very careful studies of specimens in Paris and Lon-
don. &. stmus has a square upper lip, with broadly truncate
upper nasals, the horn rugosities being carried to the very ex-
tremity, and its cranial resemblance to &. pachygnathus is
remarkable. &. dicornis has, on the contrary, a pointed prehen-
sile upper lip, and its somewhat more pointed nasals may be
correlated with this narrower snout, but the horns are carried to
the very extremity (at which there is sometimes a slight cleft,
British Museum specimen).
Atelodus neumayri, sp. nov.’
Type, a large male skull, Vienna Museum, from Pikermi or Maragha (Persia),
(erroneously catalogued as R. schleiermacheri). This skull resembles 2.
pachygnathus as follows : large frontal and nasal horn cores ; auditory meatus
closed ; zygomatic arch slender (correlated with reduction
of angle and masseteric muscles); lower border of jaw
convex ; dentition : aS It differs from R. pachy- \ ch
gnathus as follows: molars elongate, tending to hypso-
dontism ; cement covering sides of molar crowns; the
pattern of the premolar and molar teeth unique and with-
out precedent ; there is no true antecrochet on the pro-
toloph, but a fold, which might be considered as an aberrant
crista, projects into the median valley from its outer portion,
that is, external to the crochet (whereas the antecrochet oe
always appears internal to the crochet); the prominent Bigeeh vets
crochet is placed internally to this; strong hypostyle fold eumayri. Type:
4 S d i =
and postfossette on p® to m?, een eG po)
‘
a
An apparently similar fold is observed in R. antiguitatis, and
connects the protoloph diagonally with the metaloph; J.
neumayrt therefore resembles 2. antiguitatis more closely than R.
pachygnathus, both in the presence of this fold and in the greater
hypsodontism of its molar teeth.
* Dedicated to the late distinguished Austrian geologist, Melchior Neumayr.
264 Bulletin American Museum of Natural History. [Vol. XIII,
2. PLEISTOCENE AND RECENT STAGES.
An adaptive parallel to these types is presented in the Middle
and Upper Pleistocene species: &. antiguitatis resembles R. simus
(and less closely &. newmayri) with broadly truncate nasals,
slender zygoma, and hypsodont, small, very narrow molar teeth ;
while &. hemitechus* resembles R. pachygnathus and R. bicornis,
with brachyodont molar teeth. In both &. hemitechus and R.
bicormis the nasals are somewhat narrower and the upper lips
more prehensile and pointed. These large Pleistocene animals
(which co-existed for a while) thus differed in details of dentition
in adaptation to local differences of feeding ranges and habits, but
resembled each other in (1) extreme dolichocephaly, (2) back-
ward inclination of the occiput, (3) powerful nasal septum, (4)
horns on extremities of nasals.
The existing African species, &. stmus and R. bicornis, like R.
sumatrensis, in the Ceratorhine series, are, however, both less
specialized than the Pleistocene types.
Subfamily RHINOCEROTINA. PHYLUM VI.
Brachycephalic or intermediate between extreme dolichocephalic and brachyce-
phalic types ; occiput inclined forwards. Single horns upon mid-nasals ; nasals
pointed and generally smooth at the extremities. Megalodine, large upper and
lower cutting teeth.
No representatives of this phylum have been found in Europe.
In Asia, however, the Pliocene Siwaliks yield species which are
probably ancestral to the typical Ahinoceros unicornis of India.
Lydekker (’81, Pl. X) shows that 2. paleindicus leads into the
hypsodont or grass-eating A. unicornis type, while R. stvalensts
leads into the brachyodont or shrub-eating &. sondaicus type. All
these four species exhibit a skull with forwardly inclined occiput,
concave and hornless in the frontal region, nasals with a large
horn in the middle portion which does not extend to the smooth
and pointed extremities ; well developed cutting teeth.
The origin and relationships of this phylum are unknown; it
will be noted that it is exclusively south Asiatic in distribution
and this (Oriental Region) may ultimately prove to be its home
and exclusive centre of adaptive radiation.
1See Geol. Mag. (2), Vol. I, Pl. XV, as figured by Davis.
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266 Bulletin American Museum of Natural History. [Vol. XIII,
CONCLUSIONS.
This phylogeny leaves many species untouched and unsettled.
It certainly contains both errors and omissions, and I set it for-
ward mainly as a method of solution of the Rhinoceros problem.
1. It disregards homoplastic or convergent characters, which
are often entirely misleading.
2. Great stress is laid upon exactness as to stratigraphical or
geological succession, the neglect of which has been a fertile
source of error.
3. According to our present knowledge, none of the six phyla
can be connected by European stem forms, as in the phy-
logenies previously attempted.
4. The newer hypothesis of foreign (African or Asiatic) invasion
into Europe of certain phyla has at present more in its favor
than the older hypothesis of the derivation of all Upper
Tertiary from Lower Tertiary types of Europe.
5. It is a fact that the earliest known members of each phylum
show substantially all its fundamental characters ; subse-
quent modifications are adaptive and may be more or less
convergent to other phyla.
6. Generic, specific, and subfamily terms are simply our symbols
1885.
1887.
1877.
1881.
for clear thinking and description. If the hypothesis of six
or more distinct phyla is correct, and these breeds or races
have been distinct since the Middle, and in some cases since
the Early Tertiary Period, then the actual remote relation-
ships of the individual members of said phyla will be most
truthfully and clearly expressed both by the revival of cer-
tain disused generic names, and by the use of subfamily
names.
BIBLIOGRAPHY.
DEPERET, CH. Descriptions Géologiques du Bassin Tertiare du Rous-
sillon. Annales des Sciences Géologigues (Hébert et Milne Edwards),
XVII, pp. 1-272.
DeEPERET, CH. Recherches sur la Succession des Faunes Vertébrés
Miocénes de la Vallée du Rhéne. Lyon, 1887.
FILHOL, HENRI. Recherches sur les Phosphorites du Quercy. Ann.
d. l. Soc. Géol., pp. 126 et seq.
FILHOL, Henri. Etude des Mammiféres Fossiles de Ronzon (Haute
Loire). Annales des Sciences Géologigues, 1881, XII, 5, Art. 3.
1900.] Osborn, Phylogeny of the Rhinoceroses of Europe. 267
1862. GAUDRY, ALBERT. Animaux Fossiles et Géologie de l’Attique. Paris,
1862-67.
1894. HATCHER, J. B. Ona Small Collection of Vertebrate Fossils from the
Loup Fork Beds of Northwestern Nebraska ; with Note on the Ge-
ology of the Region. American Naturalist, March 1, 1894.
1832. KaAup, JEAN-JACQUES. Description d’Ossements fossiles de Mammi-
féres inconnus jusqu’a présent. . . . Premier Cahier. Darm-
stadt, 1832.
1862. KAup, JEAN-JACQUES. Beitrage zur Naheren Kenntniss der Urwelt-
lichen Sdugethiere. Erstes Heft. Darmstadt and Leipzig, 1862.
1881. LYDEKKER, RICHARD. Siwalik Rhinocerotide. Mem. Geol. Surv.
India, Paleontologia Indica, Vol. II, Pt. II, Dec., 1881.
1886. LYDEKKER, RICHARD. Catalogue of the Fossil Mammalia in the
British Museum of Natural History. London, Apr. 5, 1886, pt. iii.
1896. MERMIER, Evie. Etude Compléementaire sur l’Aceratherium platyo-
don de la Molasse burdigalienne supérieure des environs de Saint
Nazaire en Royans (Drome). Ann. d. 1. Soc. Linnéenne d. Lyon,
Tome XLIII, 1896.
1893. OsBorRN, HENRY F. Aceratherium tridactylum from the Lower Mio-
cene of Dakota. Aull. Am. Mus. Nat. Hist. Vol. V, Apr. 29, 1892.
1898, 1. OsBORN, HENRY F. A Complete Skeleton of Teleoceras fossiger.
Notes upon the Growth and Sexual Characters of this Species.
Bull. Am. Mus. Nat. Hist. Vol. X, Art. IV, March 18, 1898, Pl.
IV and IVA, pp. 51-59.
1898, 2. OSBORN, HENRY F. The Extinct Rhinoceroses. Memoirs of the
Amer. Mus. Nat. Hist. Vol. I, Pt. III, Apr. 22, 1898, pp. 75-
165, Pl. XITA—XX.
1899, OsBoRN, HENRY F.. Frontal Horn on Aceratherium incisivum. Re-
lation of this Type to Elasmotherium. Sczence, Feb. 3, 1899, pp.
16n..25
1900, OsBoRN, HENRY F. Correlation between Tertiary Mammal Horizons
of Europe and America. . . . Ann. N. Y. Acad. Sct. Vol.
XIII, No. 1, pp. 1-72, July 21, 1900.
1892. PAvVLow, MARIE. Etudes sur I’Histoire Paléontologique des Ongulés.
VI. Les Rhinoceridz de la Russie et le dévelopement des Rhino-
ceride en général. Szll. d. 1. Soc. d. Nat. d. Moscou, 1892.
1866. RAmeEs, B. Note sur l’age des argiles du Cantal et sur les débris fos-
siles qu’elles ont fournis. Bud. d. 2. Soc. Géol. d. France, Tome
XIV, 1866, p. 357.
1900. RoGER, Orro. Ueber Rhinoceros Goldfussi, Kaup und die anderen
gleichzeitigen Rhinocerosarten. Separatabdruck aus dem 34. Bericht
des Naturwissenschaftlichen Vereins fiir Schwaben und Neuburg in
Augsburg.
By Henry Farrvietp Osgor
EDITION, extracted fou hb age ie
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by
New York, Dec. 21, 1900.
Article XX.—OXYAANA AND PATRIOFELIS RESTUD-
IED AS TERRESTRIAL CREODONTS.
By Henry FAIRFIELD OSBORN,
PLATES XVIII AND XIX.
Comparatively little was known
of the skeletal structure of these
animals until the American Mus-
eum Expeditions of 1891 and 1893
secured complete skeletons of each,
which Dr. J. L. Wortman carefully
described and figured. After a
searching comparison with modern
land and water Carnivora he con-
cluded that Patriofelis was prob-
ably aguatic in habit and posstbly
ancestral to the modern Pinnipedta
and that the much older type Fig.1. Didelphys virginiana, left
Oxyena and the more recent type rea
Oxyenodon, bore similar testimony to affinities with the Seals. In
describing Patriofelis
he remarked: “ The
broad, flat, planti-
grade feet with their
spreading toes sug-
gest at the first glance
their use for swim-
Sming,” «(/94,.p. 10 ne
Recently, under
the direction of the
present writer, the
Oxyena lupina skele-
ton has been mounted
and the Patriofelts
ferox skeleton taken
apart and remounted
by Mr. Hermann,
head preparator. At
Fig. 2. Patriofelis ferox, left fore foot, from mounted skeleton. the same time several
[269]
270 Bulletin American Museum of Natural History. (Vol. XIII,
alterations were made in the restored parts of the skull of Patrio-
felis, the teeth were restored, one dorsal vertebra added, and, for
reasons stated below, the feet reset in an angulate subdigitigrade
instead of a plane plantigrade fashion. A more thorough study
of the dentition of this animal was also made from all the mate-
rials in the Museum.
In this connection a careful restudy of all the evidence led the
writer to the opposite conclusion, that these were powerful ferres-
trial, or partly arboreal, animals, analogous to the Cats in habits of
feeding, with analogous (not homologous) sectorials, clumsy in
limb structure, without prehensile claws, and presenting no evi-
dence of successors among the modern Carnivora. The reason-
ing upon which this conclusion is based is, in brief, that the
alleged points of resemblance to the Pinnipedia are in part per-
sistent primitive characters due to the descent of the Oxyznidz
and Pinnipedia from a common Insectivore-Creodont ancestor,
in part homoplastic adaptive characters due to similar habits or
uses of certain parts of the body, while the main trend of adapta-
tion is divergent from the Pinnipedia as seen both in the teeth
and feet, and from all other modern Carnivora, especially as seen
in the teeth,
In this paper the principal osteological and dental characters .
will be briefly restated with a number of corrections and addi-
tions, referring the reader back to Wortman’s fuller papers for
details.
I. EvIDENCE FOR TERRESTRIAL HABITS.
1. TDhe Feet.
It was claimed (Wortman, ’94, p. 161) that the plantigrade feet
with spreading toes indicated a webbed (’94, p. 146) or swimming
foot. It will be observed by comparison of the photographs
(Figs. 1, 2) that the feet of Patriofel’s are no more widely
spread than those of the terrestrial and arboreal Didelphys.
Moreover, the planes of the articular facets of the metapodials
and phalanges in Patriofelis entirely forbid the supposition that
this animal was plantigrade.
The writer has pointed out (’oo, p. gr) that the angulation of
the limbs in Ungulates is expressed in the angles which the prox-
1900. | Osborn, Oxyena and Patriofelis Restudied. 271
imal and distal facets make with the long axes of the shafts ;
considering the shafts as perpendicular, facets tn horizontal planes
indicate straight limbs ; facets tn oblique planes indicate angulate
limbs. Exactly similar principles apply to the hand and foot
of Unguiculates, as shown in Fig. 3. In the passage from Ofarta
U
lV Il HY
Fig. 3. Angulation of facets in feet of (1) Ursus, (11) Procyon, (111) Patriofelis, (IV)
Felis, showing increased obliquity in relation to increased angulation.
A, distal facets of metacarpals ; B, distal facets of 1st phalanx; C, distal facets of 2d phalanx.
(secondarily plantigrade), Ursus (primarily plantigrade), Procyon
(subdigitigrade), and Fe//s (digitigrade), we see that the planes
of the distal facets give certain indication of the modes of
progression.
1. As regards angulation, Patriofelis is shown to occupy a posi-
tion intermediate between Procyon and Felis, with a decidedly
angulate foot, the angles between phalanges 1 and 2 being espe-
cially acute. This proves that the metapodials, as well as phalan-
ges 1 and 2, were raised off the ground by palmar and plantar
pads asin Fe/ts. Taking a conservative view, the feet of Pa-
triofelis may be described as subdigitigrade in position. The
straight terminal claws indicate that they entirely lacked the
grasping and tearing power developed in Fedis.
272 Bulletin American Museum of Natural History. (Vol. XIU,
2, As observed by the writer and Wortman in Oxyena (’99, p.
144) ; “ There is reason to believe that the habitual position of the
foot was digitigrade, but there
is no evidence of any retrac-
tility of the claws.” Begin-
ning therefore with a subdigi-
tigrade foot, the progression
from Oxyena to Patrtofelts
does not indicate an advance
toward secondary plantigrad-
ism, as would be the case if
these animals were becoming
more and more aquatic in
habit. On the contrary, the
analogy of the feet of the
known Oxyzenidz with those
of Procyon and Didelphys
would indicate that they were
pe meee ae typical lumbar ver- ysed mainly in slow terrestrial
or arboreal locomotion, and
exceptionally if at all in swimming.
2. TDhe Dentition.
The lack of prehensile power in the feet of Patriofelis is com-
pensated for in an extraordinary manner by the increased pre-
hensile power in the progressive evolution of the teeth ; this
again is analogous to that of the Felide in its extreme heterodon-
tism or specialization, whereas the key-note of dental evolution
among the Pinnipedia is a secondary homodontism or reduction of
the premolars and molars to a common triconodont pattern.
The dental parallelism of the Oxyznide with the Felide is well
stated by Wortman (’99, p. 140).
The progression of both skull and teeth in the Oxyenide is
towards a raptorial type with increasing temporal and masseter
muscles, deep zygomatic arch and large temporal fossa, heavy
jaw, deep and broad symphysis, with the biting power concen-
trated at three points, namely, the canines, the fourth lower
premolar, and the enormous carnassial teeth; the carnassials
1900. | Osborn, Oxyena and Patriofelis Restudied.
273
[December, 1go0. |
Left lower jaw, external view (No. 1507 coll. A. M.N. H.).
Specimen included in mounted restoration,
Patriofelis ferox.
Fig. 5.
18
274 Bulletin American Museum of Ne atural History. [Vol. XIII,
are adapted to flesh and bone cutting by the loss of the talonid
exactly as in Felis. We should emphasize the contrast however :
Carnasstals
Oxyenide First upper molar Second lower molar
felide Fourth upper premolar First lower molar
Il. Common CREODONT CHARACTERS OF THE OXYANIDA,
As stated above, many primitive Insectivore-Creodont charac-
ters are found in this family which are also found in other types.
Among these are:
An alisphenoid canal ; exposure of mastoid (Pa¢riofelis) ; small
brain ; large temporal fossa ; cranium constricted behind the or-
bits (as in Mesonychide and Arctocyonide) ; powerful caudals ;
elbows everted ; prominent deltoid crest and entepicondylar for-
amen of humerus; separate scapho-lunar ; free centrale; large
trapezium ; small trapezoid ; femur with 3d trochanter and shaft
expanded distally; astragalus with flat oblique tibial trochlea and
astragalar foramen (as in Creodonta and Pinnipedia); small
mesocuneiform (functionally analogous to the small trapezoid) ;
distal phalanges cleft distally (as in many Creodonts and Con-
dylarths) ; metapodials I—V relatively well developed.
III. SPECIALIZED CHARACTERS OF THE OXYANIDZ&.
Progressive shortening of the face and elongation of cranium
with reduction of teeth and development of jaw muscles; high
sagittal crest ; occiput narrow; a preglenoid process; a large
postmastoid foramen; no postglenoid foramina; mandibular
condyles scroll-like (as in Felidae) ; atlas with form and verte-
brarterial canal as in Felide (Wortman, ’94, p. 137) ; axis with
elongate spine ; certain dorsals and lumbars with progressively
revolute zygapophyses (as in Mesonychide and certain Pinnipe-
dia, Phoca) ; lumbars with progressively developed anapophyses ; -
scapula, humerus, and ulna of about equal length ; scapula very
large, spreading superiorly (imperfectly known in Oxyena),
supra- and infraspinous fossz subequal; powerful acromion and
metacromion processes; humerus with exceptionally elongate and
prominent deltoid crest, powerful supinator ridge, large entepicon-
1900. | Osborn, Oxyena and Patriofelis Restudied. 275
dyle and entepicondylar foramen ; olecranon process of ulna
elongate, ulna grooved anteriorly ; limited rotation of forearm
owing to proximal expansion of radius; feet spreading ; trape-
zium extended transversely (as in Pinnipedia, Wortman) ; dorsal
portion of distal metapodial facets hemispherical, ventral portion
keeled (as in Fissipedia, Wortman) ; digits angulate, the second
Talonid
\
Fig. 6. Patriofelis ferox. Lower jaw, left side; A, crown view; A’, external view;
A? internal view of 4th premolar (No. 1508 coll. A. M. N. H.).
phalanges strongly flexed upon first phalanges; subungual
(retractile) processes of distal phalanges well developed (as in
Pinnipedia ; in Fissipedia subungual processes small, foramen
vestigial, Wortman); ilium expanded on superior (post-iliac)
border into a broad lamina; pubic symphysis not anchylosed ;
patella large ; fibula unreduced, articulating with side of astraga-
lus but not articulating with calcaneum (progressive) ; tibia with
twisted shaft and -cnemial spine; tibio-astragalar facet flat,
obliquely placed ; calcaneo-cuboidal facet very oblique; large
astragalo-cuboidal facets ; external calcaneal tubercle large (as
in many Creodonts and Amblypods).
276 Bulletin American Museum of Natural History. (Vol. XIII,
IV. PROGRESSIVE AND SPECIFIC CHARACTERS,
The above and the following characters show probably the
main trend of evolution in the Oxyznide ; they are derived from
a comparison of Oxyena lupina (Wasatch, Sparnacien) and Pa-
triofelis ferox (Bridger, Bartonien) ; it is important to note that
O. lupina is an exceptionally slender species, some of its Wasatch
contemporaries were more robust, whereas P. ferox is an excep-
tionally robust species. The differences between these species
are therefore partly such as we should expect to find in the com-
parison of any slender and robust types, and partly truly pro-
gressive for the family.
Oxyena lupina.
Incisors #
Canines sub-oval.
Premolars 4.
Molars §.
First lower molar tuberculo-sectorial ;
second ditto sub-sectorial with talo-
nid preserving three reduced cusps.
Second upper molar transverse.
First upper molar sub-sectorial, with
protocone relatively prominent.
Skull relatively slender.
Dorsals 13, lumbars 7 = 20 D.L.
Caudals numerous.
Limbs, chest, and back slender.
Lumbars typical with simple zyga-
pophyses ; small anapophyses on
L 1-2,
Tibia with short cnemial spine.
Patriofelis ferox.
F 2
? Incisors =
Canines laterally compressed.
Premolars $3.
Molars 4.
First lower molar reduced, tuberculo-
sectorial; second ditto truly sec-
torial with vestigial talonid.
Ditto absent.
First upper molar blade-like, sectorial,
with protocone reduced and para-
and metacones greatly elevated and
connate.
Skull broad and massive.
Dorsals 14, lumbars 6 = 20 DL,
Sacrals 3; 2 sacrals uniting with
ilium.
Caudals 28, large chevrons,
Ditto robust. '
Lumbars, massive, heavy, with ex-
tremely revolute zygapophyses ;
anapophyses on L 1-4.
Tibia with elongate cnemial spine.
Six sternebre.
Ribs heavy.
V. Systematic REvIsION.
Family Oxyenide,
in habit ;
Terrestrial or arboreal Creodonts ; strictly carnivorous
enlarged canines ; second upper molars when present transverse ;
powerful sectorials formed of first upper and second lower molars ; incisors, -
277
Osborn, Oxyena and Patriofelis Restudted.
1900. |
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31
yd
278 Bulletin American Museum of Natural History. (Vol. XIII,
premolars, and molars progressively reduced and specialized ; subdigitigrade,
metapodials 5-5 spreading, not interlocking.
Lower Eocene. Middle Eocene. Upper Eocene.
Oxyena Cope. Patriofelis Leidy. Oxyenodon Wortman.
Syn: Limnofelis Marsh.
Protopalsis Cope.
Dentition #%.1.4.3.=40. 4.4. $. 5. = ? 32. 8.4. 4. 3. = 40.
Premolars with large talonids. Premolars with talonids. Premolars with small
talonids.
Fig. 8. Patriofelis ulta, Lower jaw, left side (No. 2691, coll. A. M. N. H.).
Dentition of Patriofelts.
The teeth of the animal are so distinctive that they deserve a
detailed description ; they exhibit a very decided evolution be-
yond those of Oxyena.
Incisors.—T he jaw (No. 1508) exhibits alveoli for two incisors,
one of which is placed directly behind the other (Fig. 6).
Canines.—The alveolus and fang embedded in jaw No. 1508
indicate marked lateral compression of both fang and crown.
Lower premolars.—First, entirely wanting (Figs. 5, 6); Second
(No. 1508), close to canine, two-fanged, with crown obliquely
placed (see Fig. 6, A); Zhird, a somewhat larger tooth longi-
tudinally placed, two-fanged, crown not preserved; Fourth
(No. 1508, Fig. 6, A, A’, A’), atriconodont type of tooth ;
enlarged, antero-posterior diameter of fangs = 210 mm.; a large
central protoconid, an anterior cusp (broken away in No. 1508),
1900. | Osborn, Oxyena and Patriofelis Restudied. 279
and a posterior cusp, or talonid, with a small basal postero-
internal cusp.
Lower molars.—First, a relatively small tooth ; antero-poste-
rior diameter of fangs = 170; in No. 4805 the posterior half of
the crown only is preserved ; it exhibits the protoconid, a small
elevated metaconid ; the talonid is narrow and feebly tri-cuspi-
date ; in No. 1508 the paraconid is partly preserved; Second,
a powerful shear (No. 4508, Fig. 7, B) formed of an out-
wardly placed paraconid and a sharp elevated protoconid ; the
metaconid vestigial or represented by a very low ridge; the
talonid reduced to a cingulum.
First upper molar.—(No. 2303, Fig. 7, A, and No. 1508, A,
both teeth of the left side.) This is a powerful carnassial ; the
elongated shear (No. 2303) consists of the greatly modified
trigon and metastyle ; the protocone is depressed and reduced to
a basal spur; the paracone and metacone consist of a pair of ele-
vated connate subequal cusps ; the metastyle is an elongate less
elevated shear. In the greatly worn condition seen in No. 1508,
A, as well as in No. 2691, P. u/ta, the crown of the tooth retains
its sharpness, proving that these teeth were employed as in Fe/rs.
BIBLIOGRAPHY.
Ig00. OsBorN, H. F. The Angulation of the Limbs of Proboscidia, Dino-
cerata, and other Quadrupeds in Adaptation to Weight. Amer.
Nat. Vol. XXXIV, No. 398, pp. 89-94. Boston, Feb. 1goo.
1894. WorTMAN, J. L. Osteology of Patriofelis, a Middle-Eocene Creo-
dont. Bull. Amer. Mus. Nat. Hist. Vol. VI, Art. V, pp. 129-
164. New York, May 24, 1894. ;
1899. WorTMAN, J. L. Restoration of Oxyena lupina Cope, with De-
scriptions of Certain New Species of Eocene Creodonts. Azd/,
Amer, Mus. Nat Hist, Vol. XII, Art. VII, pp. 139-148. New
York, June 21, 1899.
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By W. D. Marruew.
AUTHOR'S EDITION, extracted from BULLETIN
OF THE _
Mmevican Baseum of Ratural History,
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ae e Vou. XIV, Articie I, pp.1-38.
e New York, Fanuary 31, 1901.
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‘The Tnickerbocker Press,
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Ee RES ON et; ee
Article I—ADDITIONAL OBSERVATIONS ON THE
CREODONTA.
By W. D. MaTTHEw.
CONTENTS.
I. Classification. Arrangement of the Creodonta, chiefly on
the basis of the specialized carnassial.
II. Family Viverravide. Characters of the skull and skeleton
of Viverravus Marsh (= Didymictis Cope), as shown in two species,
V. protenus and V. leptomylus Cope.
III. Family Arctocyonide. Fore and hind feet of Clenodon,
with a discussion of its relations to the Urside.
IV. Family Hyznodontide. Revision of the classification.
Revision of the genus Szzopa (Stypolophus) with descriptions of
two new forms.
V. Family Mesonychide. The oldest known mammal skull
(Zriisodon heilprinianus). Dentition and skeleton of Pachyena
gigantea. Pseudo-marsupial characters of the Mesonychide.
I.— CLASSIFICATION.
In classifying any group which includes a large proportion of
primitive types we must use some combination of two methods.
A division may be considered either as a group of forms resem-
bling a given type within certain limits, or as including a particu-
lar line of development. Osborn has termed these two principles
of classification the “‘ horizontal” and “vertical” ; it appears to
[ Fanuary, 1907. | [z] 1
2 Bulletin American Museum of Natural History. (Vol. XIV,
the present writer that group and race would more truly ex-
press their intent. By the former method the unspecialized an-
cestral types of several families are placed in a separate family,
from which the later families are derived. ‘This is on the ground
that the distinctions between the different lines of descent
were not at first wide enough to be of family value, and that the
different primitive types resembled each other more than they
did any of the later specialized types. The other method of
classification divides up these primitive types among the various
specialized families to which they are ancestral or approximately
ancestral. The group conception— the Latin familia = house-
hold —is, of course, the original use of the term; the race con-
ception —the English famly — has been introduced to meet the
new conditions brought in by the doctrine of evolution and the
development of paleontology.
It seems to the writer that either principle, used exclusively, re-
sults in obscuring, or at least imperfectly indicating, the real
relationships the expression of which should be its chief pur-
pose. The group method ignores parallelism, and fails to properly
emphasize the lines of descent. The race method equally fails
to emphasize the near relationship of the primitive root types,
and in practice causes much confusion and apparent variance of
opinion by the attempt to divide into different families species
among which the distinctions have not yet become of generic
value.
Among the fossil Mammalia these two methods are used in
very varying proportion. In some groups, such as the Perissodac-
tyls, the divisions have been drawn ‘ vertically,’ all the ancestors
of a family being placed in that family, so that the primitive Peris-
sodactyls of the Lower Eocene, exhibiting no more difference in
the sum of their characters than the different species of mod-
ern “Rhinoceros, are divided among six different families, the
typical forms of which are extremely different from any of these
primitive forms ; and the various species are moved about from
one to another of these families with the utmost facility by every.
author who attacks the problem anew. In fact, constant specific
distinctions are not always easy to find among them. In other
groups, such as the Creodonta, all the ancestors of the modern
types, together with those ancient types which have left no
1go1.| Matthew, Additional Observations on the Creodonta. 3
descendants, are placed in a separate suborder, distinguished by
the absence of a specialized character common to all the modern
families. Similarly among the Creodont families, the primitive
types have been grouped separately from the specialized ones
by almost all writers, the family distinctions being on points of
specialization or non-specialization.
The present writer is in a position to appreciate very much the
evil effects of unnecessary changes in nomenclature and is in-
clined to deprecate most strongly the perpetual changes caused
by a too strict adherence to laws of priority which were not de-
signed for paleontology, and do not entirely fit its requirements.
With the progress of the science it becomes more and more evi-
dent that a very large proportion of the earlier species were
founded on indeterminate material ; and the vain attempt to find
reasons for identifying these fragments with one or another of the
species distinguishable by more perfect material has wasted a vast
amount of time, and, however valuable from an antiquarian stand-
point, adds not one iota to our knowledge of the fossil animals
themselves and their place in nature. Scientific names are not
different from other names—they are merely arbitrary terms
representing each a certain conception; and it appears to the
writer that custom should be the chief rule to govern their use.
With classifications the case is different. A classification is
not arbitrary, but an expression of the natural relationship as
nearly as our knowledge of the species and the limitations of the
method used will permit. Conservatism in classification would
mean cessation in the increase of our knowledge of a group ;,
uniformity in it would be impossible, unless all observers attached
the same relative importance to the same characters. A com-
_ paratively slight divergence of opinion on the last point may in-
volve an entire upset in the classification scheme. A graphic
method (such as is used on p. 21 in illustrating the relations
of three of the Creodont families) would more accurately express
the conception; but in such a method it is seldom possible to
include all members of a group — the physical difficulties of space
of two dimensions prevents the complete representing of poly-
dimensional relationships.
In venturing, therefore, on very considerable changes in classi-
fication from that generally accepted, and embodied especially in
4 Bulletin American Museum of Natural History. (Vol. XIV,
so accurate and recent a text-book as Prof. Zittel’s, I must dis-
claim any really radical changes in relations, except in a few
cases. The recent study of far more complete Creodont material
than has ever before been accessible has led American students
to perceive the fundamental importance of certain lines of ge-
netic cleavage somewhat less appreciated hitherto, and to make
these the primary basis for division rather than differences which
are now perceived to be in large part only contrasts between
primitive and specialized types.
First in importance among these characters of divergent spe-
cialization is the position of the carnassial teeth. In the primi-
tive types the shear between upper and lower molars is absent
or rudimentary and about equally developed on all three molars.
In some of the specialized types it is chiefly between p* and m,,
as in all modern Carnivora. In others it is chiefly between
m* and m,, or between m* and m,._ A large number of the primi-
tive genera of the Creodonta show, in some of their species at
least, a more or less evident tendency towards one of these three
specializations.
A primary division on these lines, worked out by Dr. Wortman
and the writer, and published by Wortman a year ago (’9q, p.
139), had been previously independently elaborated though not
published by Prof. Osborn for use in his University lectures,
in either case chiefly the result of study of the large series of
Creodonta in the American Museum collections.
Another line of cleavage as yet imperfectly known, but which
may prove to be of importance equal to or greater than the above,
lies in the character of the claw-phalanges. In one group of
Creodonts they are short, wide, and split, indicating a more or
less hoof-like claw, probably used for locomotion only. In the
other they are like those of most modern Carnivores, sharp,
laterally compressed, bearing apparently a sharp curved claw
which could be used as a weapon. This character is known in
the following :
No carnassial Arctocyonide Clenodon (both species)
4 Sharp-clawed.
ee os : of :
Carnassial m, Veverravide Viverravus (two species)
1
“4 ms Hyenodontide Hyaenodon
a m4 Oxyenide Oxyena, Patriofelis Blunt-clawed.
No carnassial Mesonychide Pachyena, Mesonyx
1go1.] Matthew, Additional Observations on the Creodonta. 5
Thus, as far as it goes, this distinction coincides with that
. . 4 I 5
between the Creodonta adaptiva (carnassial °=) and Creodonta in-
adaptiva (carnassial not e). Arctocyon must be included with
1
the adaptive types and AMesonyx with the inadaptive, although
they have no true carnassials, an arrangement which seems to be
supported by a variety of minor skeletal characters. But the
evidence is entirely inadequate at present to use this as a primary
basis of division. Its constancy in both the small and large
genera of a family has not been proved, and until this is proved
it is always possible that the blunt claws are correlated with the
size of the species, not with its true relationships.
Zittel (’94), following Schlosser and Scott in large part, di-
vides the Creodonta as follows (I omit invalid or unimportant
genera) :
I. Oxyclenide. Molars tritubercular, pointed cusp, no shear; premolars
trenchant. Oxvclenus, Chriacus, Protochriacus, Tricentes, Mioclenus,
Protogonodon.
Il. <Arctocyonide. Molars quadritubercular, low-cusped, no shear ; premolars
trenchant. Avrctocyon, Clenodon, Anacodon,
Ill. 7Z7ttisodontide. Upper molars tritubercular, lower molars with two high
cusps and talonid. Noshear. Premolars highand pointed. 7Z77zsodon,
Goniacodon, Microclenodon, Sarcothrausten.
IV. Mesonychide. Upper molars tritubercular, lower molars with high and low
cusp and talonid. No shear. Premolars high and pointed. Dzéssacws,
Pachyena, Mesonyx.
V. Proviverride. Upper molars and p# tritubercular, protocone far in and
well forward. Lower molars with high tricuspid trigonid and low
talonid. Myenodictis, Deltatherium, Sinopa, Proviverra, Quercythe-
rium, Didelphodus, Prorhyzena,
VI. Paleonictide. Jaw shortened. Third upper and lower molar absent.
Molars tritubercular, p* and mj; large. Premolars thick. Padgonictis,
Amblyctonus, Patriofelis.
VII. Myenodontide. Upper molars with external blade and internal tuber-
cle. Third upper molar transverse or absent. Lower molars sectional
with low, trenchant heel. Oxyena, Protopsalis, Hemipsalodon, Pterodon,
Dasyurodon, Thereutherium, Hyenodon.
VIII. Miactde. p*and mj; carnassial. Last upper molar small, transverse.
Miacis (= Uintacyon), Didymictis (= Viverravus), 2? Vulpavus.
The following changes, mainly dependent on recent discoveries,
seem advisable :
6 Bulletin American Museum of Natural History. [Vol. XIV,
Oxyclenide. Mioclenus is either Primate or Condylarth, JZ.
acolytus certainly the former.’ /Protegonodon is Condylarth (al-
though difficult to separate from Protochriacus in the lower
dentition).
Triisodontide should, as Wortman suggests (’99, p. 146), be
considered as a subfamily of Mesonychide.
Proviverride is a heterogeneous group of genera, chiefly un-
specialized types of different phyla. I believe it can be broken
up with advantage, the more primitive types being placed with
the Oxyclenide, to which they are very close; the more ad-
vanced genera being distributed among the specialized families.
Hyenodictis has the lower molars of the Mesonychide, but the up-
per teeth are like those of Pterodon ; I am unable to see how the
two styles of tooth could work together in the same jaw, and com-
pelled to suggest incorrect association. Deltatherium is very
close to Chriacus of the Oxyclenide. Some of the species of
Sinopa, though not all, show the specialization of m% as carnas-
sials, and all show characters, in the upper molars especially,
allying them with either the Oxyznidze or Hyznodontide.
Proviverra is rather closer to Sénopa than to the Oxyclenide ;
Didelphodus and Prorhyzena are unspecialized types with a lean-
ing towards the Szwopa characters. Quercytherium seems to be a
very aberrant Hyznodont.
Paleonictide. With this and the succeeding family an entire
upset of the classification seems necessary, for Patriofelis is
rather closely related to Oxyena, which is very far from being
closely related to Pferodon and Hyenodon ; the resemblances
between them are strictly parallelisms. We therefore divide
these families as follows :
PALAONICTID. P4 and mj; carnassial, posterior teeth disappearing. Pa/e-
onictis, Amblyctonus, ? Atlurotherium.
OxyaniD&. M+ and mg carnassial, posterior teeth disappearing. O2xyena,
Patriofelis (= Protopsalis), Thereutherium, Telmatocyon.
HY4NODONTID&, M3 carnassial, m® disappearing. Myaenodon, Pterodon,
Quercytherium, Cynohyenodon, Sinopa, ? Didelphodus, ? Paleosinopa.
ats he kT eae ; ;
Miacitde.’”’—W ortman and the writer have shown in a recent
paper (99, p. t10) that AZzacis is a synonym of Vudpavus and
* The primate skeleton from the Puerco described by Osborn and Earle in 1895, and referred
to Indrodon, is M. acolytus. M. turgidunculus, lemuroides, tneqguidens, ? opisthacus, and
? turgidus may also be placed provisionally among the Primates.
1go1.] Matthew, Additional Observations on the Creodonta, 7
Didymictis of Viverravus. The name Miacide therefore cannot
be used, although the family Viverravide there proposed by us
may perhaps be extended so as to take in Vadpavus and Uintacyon,
should these prove to have the Creodont carpus.
The analysis of the Creodonta on the primary basis of the
specialized carnassial will stand thus :
I. CREODONTA PRIMITIVA. No specialized carnassial ; tritubercular upper
and lower molars, shear rudimentary or absent. Claws unknown.
Oxyclenide, Includes some genera with Lemuroid affinities in the dentition.
II. CREODONTA ADAPTIVA. Carnassial when present on p* and mj.
Claws, where known, of modern type, and probably carried more or less free
of the ground. Scapholunar-centrale early uniting (podials tending towards
true Carnivore type).
i Post-camassial teethidisappeatingec... 2a. -o- one ae = Paleonictide,
2. Post-carnassial teeth becoming tubercular............... Viverravide.
3. No carnassials, molars becoming flat-crowned, premolars disap-
PEN? Coa eo modo COnG oh FOOD CUD OCHO SUCRE CO EMOOn Oe Arctocyonide.
III. CREopONTA INADAPTIVA. Carnassial when present not on p+ and
mz. Claws, where known, blunt, hoof-like, resting on the ground. No ten-
dency to union of the carpals (podials tending towards Ungulate type).
Ton Cannassialsnmsg, SHeaAmiMa teeth). 1). a1 tialevey! clei c terete oie) Oxyenide.
2. Carnassials m3 - ee aioe od boc OO Hyenodontide.
3. No carnassials, teeth with high, round, blunted cusps, upper molars tri-
tubercular, lower molars premolariform................ Mesonychide.
There is probably a definite correlation between the hoof-like
claws and the lack of union among the carpals in Oxyena,
Mesonyx, and Hyenodon ; but that the division in foot-characters
coincides with that based on carnassial specialization is not yet
proven.
The Creodonta Adaptiva early gave rise to the true Carnivora
(by union of the carpals), and are not known to have existed
later than the Wasatch, although the Middle Eocene of Vudpavus
and Uintacyon may, when the carpus is known, prove to be
Creodonts. The Creodonta Inadaptiva persisted side by side with
true Carnivora until the close of the Oligocene, but none of
them can be admitted as ancestral to any true Carnivore, unless
the supposition that has successively been raised about each of
the three families, connecting them with the Pinnipedia, should
be proven by satisfactory evidence.
Of the modern Carnivore families we may derive the Viver-
ride, Canide and Procyonide, and perhaps the Mustelidz, from
8 Bulletin American Museum of Natural History. (Vol. XIV,
the Viverravide ; the Felide and ? Hyznidz probably from the
Paleonictide ; the Urside either from Arctocyonide or, with
the Canide, from Viverravide—as discussed later in this paper.
As for the Pinnipedia, it appears to me that there is no evi-
dence sufficient to justify our deriving them from any known
type of Creodont or Carnivore, and that their ancestors are not
very likely to be found in the almost exclusively terrestrial fauna
of the fossil beds of the Bad Lands. (See also Osborn, ’oo.)
Il.—Family VIVERRAVIDA! Wortman & Matthew.
Viverravus JWZarsh.
Syn., Didymictis Core.
‘“The lower jaws in this genus are long, very slender and compressed ; the
last two molars are tubercular.' Both have the posterior part of the crown
quite low and the anterior half elevated and composed of three angular cusps.
The four teeth anterior to these? are much compressed. The upper flesh
tooth closely resembles that in some of the Viverridee, and the genus should
probably be referred to that group.”—AZarsh, ’72, p. 7, of separate.
‘“Inferior molars six, consisting of four premolars and two molars. True
molars, a posterior tubercular and an anterior tuberculo-sectorial, z. ¢., with three
elevated cusps and a posterior heel. Premolars with a lobe behind the principal
cusp. Canine teeth directed forward, and close together, so that it is doubtful
whether there were any incisors. An ungual phalange of the typical species is
strongly compressed. Humerus distally expanded transversely and margin
pierced by humeral artery. Astragalus exhibits two entire trochlear faces, the
wider external and directed interosuperiorly, the inner presenting superointeri-
orly. They are separated by an obtuse longitudinal angle and are little or not
at all concave transversely. The form is depressed. The head supports a
single transverse convex facet for the navicular, and with the neck is as long as
the trochlear portion.” —Cofe, '84, p. 304.
A fragmentary skeleton found in the Big Horn Valley by the
Princeton Expedition of 1884 was described by Prof. Scott
(87, p. 169) under the name of Dédymictis altidens. As this
species is not otherwise known to occur in the Wasatch, it is
possible that the specimen should be referred to the closely
allied V. (D.) protenus. Two somewhat more complete skele-
tons found in. the Wasatch of the Big Horn Valley by the
American Museum Expedition of 1896 further illustrate the
* Meaning, apparently, with tubercular heel. The context favors this interpretation. Marsh
apparently meant at this time to include in the genus forms like Telmatocyon.
* I, é., premolars 1-4,
1901.] Matthew, Additional Observations on the Creodonta. 9
genus, and by means of the information now available concerning
other Creodonts we are able to summarize the characters of
Viverravus as follows :
Fig. 1. Viverravus protenus Cope. _ Fragmentary skull and jaws, one-half
natural size. No. 2830, Wasatch Beds, Big Horn Basin, Wyoming. Amer. Mus.
Exp. 1896.
I.—GENERAL CREODONT CHARACTERS.—Skull very large in proportion to
skeleton, brain very small and of low type, tail large (presumably long), neck
longer than in most modern Carnivora, Limbs rather short with large joints
and heavy muscular attachments. Feet small, spreading, scaphoid and lunar
separate. Fibula large, separate from tibia. Radius with oval proximal facet,
and trihedral simply concave undivided distal facet. Part of a presternal bone
preserved is like that of Phenacodus wortmant, Humerus with prominent
deltoid crest reaching far down on the bone; distal extremity broad, not deep,
with entepicondylar foramen.
Boral
2.—FAMILY AND GENERIC CHARACTERS.— carnassials. Lower car-
1
nassial with high triangular trigonid shearing against posterior edge of p4 and
anterior edge of m+, which are extended externally into shearing blades,
Molars 1? tubercular, heel of mz and all of mg tubercular or becoming so,
premolars trenchant with
posterior cusps in front of
heel. Jaws long and slen-
der, incisors reduced and
canines approximated. Fe-
mur with second trochanter
interior instead of poster-
ior, third trochanter well
developed on external side, Fig. 2. Viverravus protenus, No. "2830. Cervical verte-
one fourth the length from bre, Seon above. One-half natural size.
the head of the bone. A
clearly marked facet between fibula and calcaneum. Trochlea of astragalus very
flat, distal facet strongly convex from above downward, almost flat transversely,
10 Bulletin American Museum of Natural History. |Vol. XIV,
neck rather long. Tarsals serial; a lunar facet on unciform. Entocuneiform
deep, but remarkably short for a Creodont, not as long as the ectocuneiform.
Toes 5-5, the first short and stout, not opposable. Claw phalanges compressed,
sharp, without hood, middle phalanges asymmetrically excavated on upper side
of shaft (? hence claws slightly retractile).
3.—GENERIC CHARACTERS.—Premolars ¢. Molars §. Trigonid of mz very
high, with antero-external and posterior shearing surfaces. Postero-external
shearing blade of p* and antero-external blade of m+ strongly developed.
4.— SPECIFIC CHARACTERS of V, protenus and V. leptomylus var.— These
two species are distinguished from all others of the genus by the elongate low
tubercular mg with very small trigonid and very long heel. The teeth of
protenus and leptomylus var, show no very constant differences, but may be
separated more or less completely on the following characters (averaged from a
number of specimens) :
V.. leptomylus var. V. protenus.
Genet Oe pgs Ny MM 20.3) LO ater OO 25.03: 2h Aa leon
Oo 8 wanes M5 AS 20.00 14.5) 2 LOy7k—= 0288
Blade p* to length OVI. 5) O40 6. Assi 0:72
Transv. to longit. diam. m+ Leche Onl = eA 5 E563) LOS == Olas
The above proportions show V. protenus as nearly one fourth
larger, with the premolars larger in proportion, somewhat longer
shearing blade on p4,
and m+ more extended
transversely. The same
character and extent
of distinctions are
found between differ-
ent species of Canide,
but they are much
more constant.
In the skeleton some
striking differences in GY’
Fig. 3. Viverravus protenus, Discee oc eppeet: a
No. 2830. Right humerus; 4’ Protenus has limb bones Peon yh Raha:
from in front; B, from outside. f tl l | parts of left aie a
One-half natural size. of the same ength as ulna, from in front; B,
_ right unciform, from
V. leptomylus but aver- ? fiout at CG peeeente
aging nearl ] it] [7 view, and WD, distal
ging y a third stouter (the dentition of V. Vig, ane aaa
protenus being one fourth larger). The mod-_ tadius., One-half nat-
5 ural size, except B,
ern Civet, intermediate between the two in Which is natural size.
size of skull, has limb bones one fourth longer than the
1g901.] Matthew, Additional Observations on the Creodonta. IT
larger species, and stouter in propor-
tion, while the neck is over one half
longer.
The cat, in which the back and up-
per limb-bones are of nearly the same
size and the lower limb bones consid-
erably longer, has a head and jaws of
about half the dimensions of V. prote-
nus, and much shorter neck.
The following table’ will give an
idea of the proportions of these ani-
mals :
Fig. 5. Viverravus protenus,
No. 2830. A, parts of right fe-
mur, from in front; 8, proximal
end of right tibia ; C, left calcan-
eum. One-half natural size.
Viverra civetta.
Felts catus.
Viverravus protenus.
Viverravus leptomylus var.
inhmnerns, lenetiucogcntsoenogsssaqqv00esooUr
a circumference of shaft........-..-.-
MG diameter of distal end..............
Wiareie, lege ooadasoéangmooRguoodasscAdaeoL
: circumference or sShattess.). .lnles + oo
iReGhie, Wendie saeco Soba gruRe+ uso 6 udoDoUe
ss circumference of shaft.............---
Uji. lsieticcacngacesncdososnseoo00ho0cmbdd
‘circumference of olecranon....... ......
@alcaneumlenetht years vee aeternere cleretcieya te elect
Tarsus, length (including calcaneum)...........
‘¢ width across navicular and cuboid.......
Sikmilll, Wematitis.o.. Sgob Aboud adicAncsconnoocooouE
ljeite Leal do dbookdapéngeagennenousopooamed
IDYefnthaials MSNA sooo eocda banc 6 GoD D EO OOpOl
Seven cervical vertebrz, length..............
21 19
yell Cy) 136 103
27 21 36 26
63 103 gI
i7/ 12
S55 | E3r-4)| Tos
30 23.2
27 22.5 29
29
12.6
167.5 158 gt
130 TU 2) OS
83.7 |e.66.4| 75 34
102 164
1 The measurements are given in millimetres. e = estimated.
12 Bulletin American Museum of Natural History. (Vol. XIV,
IlI.—Family ARCTOCYONID (Gervais) Cope.
The Manus and Pes of Clenodon.
In a previous article (’97, p. 289) I have given some account of
the foot structure of this interesting Creodont, now more fully
known than when first described by Cope (’83, p. 547). This
description can now be supplemented by figures of the fore and
hind feet.
It was mainly on the characters of Clenodon (Mtoclenus)
ferox that Professor Cope based his views as to the position of
the genus A/zoclenus Cope (’84, p. 324), referring it to the Arcto-
cyonidz, which included this and many other species of diver-
gent affinities, separated by Scott (’92) as distinct genera.
Judging from Cope’s descriptions of the skeletal material of J/.
ferox at his command, he was evidently strongly impressed with
its resemblance to Marsupials, an idea which led him astray in
parts of his description. I do not think that this resemblance is
wholly a case of parallelism, for all the Creodonta of the Basal
Eocene were quite nearly allied, and certain of them show
primitive marsupial characters that it is very difficult to explain
without admitting a closer connection between Marsupials and
Placentals than their modern differentiation would lead one to
believe. C/enodon has, however, no marsupial characters except
such as must be considered an inheritance from the common
stock which gave rise to both Marsupials and Placentals. Its
progressive characters are placental carnivore.
Several genera of this group have been described. Without
discussing the validity of Hyodectes and Heteroborus, defined by
Cope on the basis of descriptions of European Arctocyons, we
may say that three genera are sufficiently known for discussion of
their relationships. These are :
Arctocyon Blv., type A. primevus Blv.; referred species, A. ger-
vaist, A. deulit Lem., all Basal Eocene of Europe.
Anacodon Cope, type A. ursidens Cope, Lower Eocene of
America.
Clenodon Scott, type C. (Mioclenus) ferox Cope; referred
species, C. corrugatus (Cope), Basal Eocene of America.
To these should perhaps be added Arctocyonides Lem., type A.
sp. tnnom. Lem., Basal Eocene of Europe; referred species ?
1go1.] Aatthew, Additional Observations on the Creodonta. 13
A. (Mioclenus, Clenodon) protogonioides Cope,‘ Basal Eocene of
New Mexico.
The last genus is distinguished only by the smaller quadrate
teeth—simpler than those of C/enodon if the reference of Mocle-
nus protogontoides be correct. The type species is little known.
If the foot bones figured by Osborn (’go, p. 60) are correctly
referred to Arctocyon, there are important differences in foot-
structure between it and Cl/enodon, the former having no fibular
facet on the calcaneum, while the ungual phalanx is much less
compressed. The figure given by Lemoine (’78, pl. IV) of a cal-
caneum of A. gervais?, is much more like those of Clenodon ferox
and C. corrugatus and appears to indicate a fibular facet. I think it
probable that the bones figured by Osborn are those of some
other Creodont. The skeletal structure of Anacodon is unknown.
The dental distinctions are :
Clenodon.
Arctocyon. Anacodon,
Upper premolars mod-
erately reduced, the
first one-rooted,
second two-rooted,
third and fourth three-
rooted with high tri-
hedral protocone and
weak cingular cusps
at the bases of the
three solid angles.
Upper molars with three
low subequal cusps,
strong metaconule,
somewhat weaker hyp-
ocone and very small
paraconule. Metacon-
ule weak and hypo-
cone absent on m2.
Upper premolars consid-
erably reduced, first
one-rooted, second
and third two-rooted,
fourth large with
strong deuterocone.
Upper molars as in Cle-
nodon.
Opper premolars much
reduced, the first and
second minute or ab-
sent, third two-rooted,
fourth small with a
strong deuterocone
and rudimentary third
and fourth cusps.
Molar cusps obsolete.
These distinctions are based on the typical species of each
genus. It is probable that the other species referred to Arctocyon
are closer to C/enodon, but the present evidence seems to warrant
holding the three genera separate. Arctocyonides if correctly
represented by the American species A. (MWioclenus, Clenodon)
protogonioides (Cope) is the most primitive of the family, but
hardly deserves more than subgeneric separation from C/enodon.
14 Bulletin American Museum of Natural History. [Vol. XIV,
This genus, while close to Arctocyon, is distinctly less specialized,
and the differences seem to warrant generic separation. Ana-
codon, of the Wasatch, is much more specialized than any of the
Basal Eocene species. All the genera, as far as known, have
long, sharp, serrate canines, somewhat flattened and very little
worn, premolars in various stages of disuse and reduction, molars
becoming flat-topped and quadrate. The little-used premolars
and canines may be contrasted with,those of the Mesonychidz
and Oxyzenide, in which they were well worn, often to enamelless
stumps. The canines may have been chiefly for defense against
enemies, the animal not being to any extent carnivorous.
The fore and hind feet here figured are of much interest,
showing for so early a period an unusual degree of specialization
in certain characters.
The Fore Foot.
The carpus is distinguished especially by the large rectangular
centrale, lying half under the lunar, half under the scaphoid and
fused to the latter bone.
The lunar-centrale fa-
cet still persists but the
lunar - scaphoid — facet
has disappeared, re-
placed by a roughened
bony surface ; this pro-
bably points to an ap-
proaching union of
scaphoid and _ lunar,
The dunar is keeled,
and faceted for cen-
trale and magnum on
one side, for unciform
on the other. The mag-
7 num has a broad cen-
_Pig.6. Clenodon corrugatus (Cope). Fore foot, natural
size. Specimen No. 2456. Torrejon Beds, Wyoming. trale facet and narrow
lunar facet separated
by a sharp keel. This arrangement would, upon union of the
scapho-centrale and lunar give a proximal keeled facet on the
magnum fitting into a correspondingly grooved facet on the
1g01.] Matthew, Additional Observations on the Creodonta. 15
scapholunar, exactly as is found among Urside, instead of a flat
facet such as characterizes Canide and Felide.
The unctiform, cunetform, and pistform are of the usual primitive
type, the former having a broad lunar facet separated by an angle
from that for the cuneiform. The ¢vapezoid has not been found,
but was evidently small, ovate, with a narrow facet for the magnum
and pinching out on the radial side so as barely to touch the
trapezium. The ¢rapeztum is large with triangular upper surface
and projects in a blunt point underneath the centrale and
trapezoid.
There are five metacarpals, the first only two thirds as long as the
rest, but stout in proportion, and divergent though hardly oppos-
able. Its proximal facet allowed considerable play, and is
keeled toward its superior surface. The other metacarpals are
narrower, longer, and much closer together, with but little play on
the carpus.
The phalanges are
keeled about as in the
bears, a little curved and
somewhat angulate in sec-
tion. The wngual phalan-
ges are more specialized
than in any known Creo-
dont and the majority of
Carnivores. They are
large, high, and much
compressed, quite asmuch
so as in Urside but with
no trace of basal sheath.
The Hind Foot.
The ¢arsus shows some
interesting characters.
The tibial and fibular ar- Fig. 7. Clenodon ferox (Cope). Hind foot, three-
: : fourths natural size. Type specimen No. 3268. Tor-
ticulating surfaces are of _ rejon Beds, Wyoming.
nearly equal width, and
the fibula has a considerable facet on the calcaneum. The ectal
astragalo-calcanear facet is oblique as is usual among the earlier
16 Bulletin American Museum of Natural History. (Vol. X1V,
Creodonts, and is separated by a slight angle from the fibulo-cal-
canear facet. The trochlea of the astragalus is very shallow, the
foramen? is present, and the notch for the flexor digitorum wide
though not very deep. The head of the astragalus is consider-
ably flattened, about as much as in Urside or Felide. The
cuboid has a rather narrow astragalar facet, concave, and sepa-
rated by an angle from the rather flat calcanear facet. The
corresponding facet on the astragalus is not separable from the
navicular facet. The extocunetform is large and metatarsal J short,
stout, and divergent, though with less play than the correspond-
ing metacarpal. The other four metatarsals are of about equal
length, the second being wedged in proximally between the
ecto- and ento-cuneiforms.
Comparisons.
The union of the centrale with the scaphoid is an interesting
stage in the consolidation of the carpus. The position and size
of the centrale are peculiar. In Mesonyx it lies entirely beneath
the scaphoid (Scott, ’87, p. 161, pl. VII, fig. 1). In Déssacus
(Osborn and Earle, ’95, p. 33, fig. 9) it is less completely so, but
the shape is rhombic instead of rectangular. In Hyenodon (Scott,
’87, p. 182, pl. VII, fig. 5) the centrale lies partly under the lunar,
but is small and triangular, hardly touching the magnum, while
the trapezoid is very large, and the unciform has no lunar facet.
In Oxyena the carpus is less strongly interlocking, the centrale
smaller and rhombic, trapezoid larger, trapezium different in
shape. In Viverravus the carpus is narrower, foot probably
slenderer—little is known as to the arrangement of the carpals.
Comparing this carpus with the various primitive Creodonts, Con-
dylarths and Amblypods, shows a great deal of similarity among
them all. The progressive characters in Clenodon seem to be:
1. Union of centrale with scaphoid ; ? large size of centrale, almost excluding
magnum from lunar on upper surface ; sharp keel of magnum.
2. Grooving and inward facing of distal facet of trapezium.
3. Stout semi-opposable first metapodial, and length of fifth.
* The use of this foramen seems to be unknown. Prof, Osborn has suggested that it may
have held an extension of the interosseous ligament,which lies between the two astragalo-
calcanear facets and connects the astragalus with the calcaneum. If this ligament originally
passed up to the tibia, its disappearance would be directly connected with the keeling of the
proximal and flattening of the distal end of the astragalus, all being due to the transference of
the main ankle-joint from the distal to the proximal end of the astragalus, 7. ¢., the evolution
of the mammal from the reptilian stage of development in this character.
ae
1901.] Matthew, Additional Observations on the Creodonta. 17
4. High compressed claw; extension upward of distal facets of Ist and 2d
phalanges and to some extent of distal metapodial keels.
5. Slender, serrate, unworn canines.
6. Reduction of premolar dentition.
7. Low-cusped quadrate molars, m,; somewhat reduced.
The first of these characters leads towards the Carnivora gen-
erally, except the sharp-keeled magnum, which seems to point
especially towards the Bears. It may, however, be merely the addi-
tional brace required for large-clawed feet. ‘The semi-opposed
thumb is a character pointing towards arboreal types, hardly to
any known Urside. The proportions of the phalanges agree with
those of the Ursidz and of some other Carnivora (Procyon, etc.).
The high compressed claw-phalanx suggests the Urside very
strongly ; the greater extension of the facets is a modernization
common to all the sharp-clawed types.
In the dentition all three characters point to the Urside and
to no other Carnivora. The elongation of the second molar after
the disappearance of the first is exactly paralleled by the Raccoons
in evolving from the Canide through PA/aocyon — a very similar
adaptation not carried so far as in the Urside.
The argument for placing the Arctocyonide ancestral to the
Ursidz is, then:
1. They belong by their foot structure to the adaptive Creo-
donta, the teeth not placing them in eiics the adaptive or
inadaptive group.
2. The three chief progressive characters of the teeth are those
which distinguish the bear dentition,
3. The claws are far advanced towards the type found in ihe
Ursidz and in hardly any other Carnivora.
_4. The one marked distinction that I can see to separate the
bear-carpus from that of other Carnivora is found in process of
formation in Clenodon.
5. The proportions of the digits show a specialization found in
the Bears and in hardly any other Carnivora.
Against this may be urged:
1. Clenodon has the thumb more opposable than in the modern
Urside.
2. Anacodon has the premolar dentition as much reduced as in
the modern Urside.
[ Fanuary, rgoz. |
18 Bulletin American Museum of Natural History, [Vol. XIV,
(In either case, this may indicate that neither Clenodon nor
Anacodon is precisely in the line of Bear descent, but does not
seriously weaken the arguments given above as to the derivation
of the Bears from the family Arctocyonidz.)
3. The real objection to the theory lies in the remarkable
approximation of certain of the Dog and Bear families in Upper
Miocene time. Although, as Schlosser has recently shown,
Hyenarctos and Amphicyon cannot stand in the direct line of de-
scent, yet they exhibit a striking gradation between the two
families and might well be considered as survivals of primitive
links connecting the two. Cephalogale also, among the more
ancient Canidz, shows several approximations to the Urside, and
is placed by Dr. Schlosser as ancestral to the family (’99, 146).
There are some characters, however, that have not been
bridged, and these are perhaps more important than they appear
at first sight.
r. All Canidz have triangular upper molars. All Urside have
quadrate upper molars.
2. All Canidz have the inner cusp of p4 anterior. All Ursidze
have it medial.
The trigonal molar and the anteriorly placed trittocone are in-
dications of a formerly tuberculosectorial dentition, and appar-
ently very difficult to get rid of. Procyon, however, seems to
show us an earlier stage of their disappearance in a line descended
from the Canidz ; so that we may yet discover the intermediate
stages in the Ursid phylum.
To sum up—the Arcéocyons were progressing towards the Bear
line in all the most distinctive characters of both teeth and feet.
But the wide gap between Lower Eocene and Middle Miocene
makes any connection between the two somewhat uncertain. In
the Canid line, on the other hand, we have a number of appar-
ently intermediate stages known. But these intermediate stages
cannot have been actually in the line of descent, and even if
they are unaltered descendants of more ancient types we still
have a gap of some importance unbridged. The connection,
moreover, 1s based on teeth alone. Professor Osborn, in discuss-
ing the evolution of the Mammalia, remarks: “ The teeth and
feet, owing to the frequent parallels of adaptation, may wholly
mislead us if taken alone; while if considered together they give
tgo1.] Zatthew, Additional Observations on the Creodonta. 19
us a sure key ; for no case of exact parallelism in both teeth and
feet between two unrelated types has yet been found or is likely
to be.” (Osborn, 1893, p. 10.) If Clenodon be totally unrelated
to the Ursidz it is an exception to this statement, and, as far as
I recall, the only one. And yet the evidence is very strong for
deriving the Urside from primitive Dogs.
IV.—Family HY HNODONTID Cope.
In this family are included, as Wortman has already indicated
(99, p. 139), most of the genera grouped by Schlosser under the
name Proviverride. De/tatherium belongs with the Oxyclenide ;
the position of Didelphodus and Paleosinopa is uncertain, as they
fail to show the progressive characters of either group. Sopa
and Cynohyenodon clearly belong in it, Proviverra somewhat less
clearly, while Quercytherium appears to be an aberrant member.
The progressive characters of the two families may be contrasted
thus :
Hyenodontide,
Upper molars with connate exter- |
nal cusps, developing a shear, espe-
cially between m2 and mg. Last
upper molar becoming transverse and
disappearing. Heels of lower molars
disappearing, me? disappearing, pad
advancing to form a cutting blade.
| ally
Oxyclenide.
Upper molars becoming quadrate.
Fourth lower premolar becoming mo-
lariform. A considerable diastema
developing behind canines with disap-
pearance of pt. Dentition function-
Ga Wemiuins!,
Opossums, and some Insectivores).
insectivorous
Dentition functionally carnivorous (c/.
Felide).
All the above Hyznodont characters are exhibited to a greater
or less extent by various species of Szzopa, though always in an
early stage of progress. Most of them are exhibited by Cynohy-
@nodon and Proviverra, which besides are extremely close to
Sinopa. The same is true of Quercy terium except for the very
peculiar premolars (which are approximated in Cynohyenodon
minor if the reference to this species be correct of a fine speci-
men in the Paris Museum). Padl@osinopa and Didelphodus can
be placed here only on account of general resemblance to the Pro-
viverrines and lack of the progressive characters of any other
group. The generic definitions will be:
Hyznodon. M+ absent; m1—* without protocone, para- and metacone united ;
metastyle forming a shearing blade larger than the united fa. and me.
m,—, without metaconid or heel. Jaw long, slender.
20 Bulletin American Museum of Natural History. [Vol. X1V,
Pterodon. M® transverse reduced ; m!~® triangular with well-developed pro-
tocone, fa. and me. connate, metastyle blade smaller ; m,_, without met-
aconid but with small trenchant heel. Jaw short and deep.
Sinopa. M2 transverse; m+ * sharply triangular with widely separated proto-
cone, fa. and me. connate to varying degree, metastyle and parastyle ex-
tended into small shearing blades. My,-_, with high triangular trigon and
low basin heel, pa? and me? of equal size, pr4 much higher,
Didelphodus. M# subtransverse unreduced. M1~* sharply triangular with
widely separate crescentic protocone, fa. and me. somewhat connate, para-
and metastyles moderately developed. Lower molars with triangular tri-
gonid of three equal cusps (pa? internal), and large heel.
Paleosinopa, M+? subtransverse; m'~® somewhat quadrate, external styles
not much developed. Lower molars with low trigonid of three equal cusps
(pa? anterior) and large bicuspid basin heel.
Quercytherium. Molars much as in Stzofa. Premolars distinguished by ex-
treme robustness and large size, p being the largest.
The genera Lzmnocyon Marsh, Stypolophus Cope, and Prototomus
Cope are, as Scott holds, synonyms of Szxopa Leidy, except LZ.
riparius, subsequently (Marsh, ’99) separated under the name
Telmatocyon. Trtiacodon Marsh is perhaps founded on incomplete
molars of either Szzopa or Viverravus. The species placed in it by
Cope in 1872 was afterwards correctly referred by him to Szyfo-
lophus. Proviverra is very close to Sinopa, but, though very im-
perfectly known, seems to differ from any of the species of S¢zopa
in its short jaw and reduced premolar region, approximating
Prorhyzena. Cynohyenodon cayluxt is hardly distinguishable in
dentition from Szzopa, but seems to be somewhat more advanced
in skull characters. It is hardly worthy of generic separation.
C. minor, if correctly represented by the upper and lower jaws
referred to it in the Paris Museum, is more nearly allied to Quer-
cythertum, having the same proportions in the premolars, which,
however, are not nearly so robust. Hemipsalodon does not seem
separable from Péerodon; and Pseudopterodon, according to Scott,
is probably founded on milk teeth of Prerodon (Scott, ’92). Thy-
lacomorphus is known only by the back of a skull described by
Prof, Gervais in 1876, but not figured. It was conjecturally re-
ferred by Schlosser to Cynohyenodon. It is, however, not a Cre-
odont at all, but an Anoplothere, probably Déplobune quercyt.
The accompanying table will show the conception which this
paper attempts to explain of the relationship of the genera and
limits of the families of Inadaptive Shear-toothed Creodonts,
the Oxyzenide, Oxyclenide, and Hyzenodontide :
1901. | Matthew, Additional Observations on the Creodonta. 21
ae) ice]
SM etipaing | SoS
SM Betice |e plate es
z ta] > a ra ie
Q & 1S) > I
3° Ss 5 3
Z| * Zz
<
&
7
—_———"
—
Dow
Os
2 ap a ope
2 re) oa ae
=] vA iat oF no)
~ n x. oO =
“ 2} fal = ”
cen ae tayo ot
o S) oO
=] a si a
Hyanodontide
Quercytherium
Hyznodon
Pterodon (Hemipsalodon)
Sinopa
Cynohyzenodon
Proviverra
EN
Palzosinopa
Didelphodus
Oxyclenide
\
Deltatherium
Chriacus
Protochriacus
Tricentes
Oxyclzenus
Oxyenide
’ Thereutherium
=
Telmatocyon
Oxyzenodon
Oxyzena
Amblyctonus
Patriofelis
ae
22 Bulletin American Museum of Natural History. [Vol. XIV,
Palzosinopa veterrima, n. g. et sp.
Palaeosinopa veterrima “* Wortman,’”’ MATTHEW, Bull. Amer. Mus. Nat.
Hist., 1899, p. 31. Name only.
Stypolophus whitie OSBORN & WORTMAN, "92, p. 110, not S. whitie Cope.
Generic characters ; Dentition : em Metastyle moderately extended on
m2, rudimentary on m1. Trigonid of three subequal cusps; heel larger than
trigonid, hy? and en‘ strong, hl‘ present on all molars, strong only on mg. The
worn state of the upper molars in the type specimen precludes exact compari-
son with Dide/phodus, to which the genus is most closely allied.
Specific characters.
Upper incisors ra-
ther large, roots not
compressed, the third
placed behind and
within the second,
leaving a deep notch
for the lower canine
between i2 and ct,
Canines rather small,
first upper premolar
one - rooted, second
two-rooted, third
. type, No. 150a. Both natural size, from the Wasatch Beds Big
two-rooted, but with Horn Basin, Wyoming. Amer. Mus. Exp. 1891. ;
the posterior root the
wider, crown with robust protocone and rudimentary postero-internal heel ;
fourth three-rooted with well-separated internal cusp. First and second molars
with rudimentary hypocone or postero-internal cingulum, third molar transverse
apparently large. First lower premolar one-rooted, others two-rooted, trenchant,
with small heels and minute anterior basal cusps.
Type No. 95, upper and lower jaws, teeth badly worn. Associated type, No.
150a, a lower jaw with unworn teeth. Referred specimens Nos. 2849, 2851,
2852. All from the Lower Eocene Wasatch Beds of the Big Horn Valley,
Wyoming.
Measurements of Type Specimen.
Upper dentition (approximate) itm® 49
‘* molars (approximate) 15
Seepremolarsiae ss 21
First upper molar, antero-post. 5.3, tansy. 7.4
Second ‘ rs eo 8 oy 8.1
Lower dentition, c,-m, 41
“« true molars 16.6
‘* premolars 18
Depth of jaw below p», 6
ae ae se «ee
Me 12
1g01.] Matthew, Additional Observations on the Creodonta. 23
Palzosinopa didelphoides (Cofe).
Ictops didelphoides Cope, Bull. Hayden Survey, VI (1881), p. 192; Tert.
Vert., p. 268.
Paleictops didelphoides MATTHEW, ’99, Pp. 35-
The lower jaw fragment from the Wind River Beds, referred
to Jctops by Prof. Cope, more probably belongs in this genus.
In absence of characteristic parts its reference is provisional.
Sinopa Zeidy, 1871.
Stypolophus Cork, Pal. Bull. No. 2, Proc. Am. Phil. Soc. 1872, 466.
Prototomus COPE, Report on Fossil Vert. N. M., Ann. Rep. U.S. G. S. W.
of rooth Mer., 1874.
Limnocyon MARSH, in part, Am. Journ. Sci, 1872, Vol. IV, p. 122.
The original types of Stnopa, Stypolophus, and Limnocyon were
from the Middle Eocene of Wyoming. Cope afterwards described
a number of species from the Wasatch under the name of Profo-
tomus, which he afterwards (1877) united with Stypolophus. Scott
(1892) united both with Sopa, and an examination of the
types of Zzmnocyon shows that they also should be referred to this
genus.
Besides the characters given on a previous page the following
may be noticed :
Dentition : = 43 . First premolar two-rooted. Brain small. Lower pre-
molars composed of principal cusp and trenchant heel (no second posterior
cusp). Ae. of me reduced or absent. Incisors small, the upper ones in a row.
Sinopa rapax Leidy, 1871.
From the Bridger Beds, Wyoming. Type, a lower jaw with
more or less broken molars, figured in Extinct Vert. Western
Terrs. The species is of medium size with apparently a rather
large heel on mj, but with vestigial heel ( fide Cope, ’84, p. 289)
on the third. I am unable to find other specific characters in
Leidy’s figure. Length of pg-mg, 31mm.
Sinopa vera (J/arsh).
Limnocyon verus MARSH, Am. Jour. Sci. 1872, Vol. IV, p. 122.
From the Bridger Beds, Wyoming. A large species with proto-
cones of m+ and m? much compressed. Third premolar nar-
row and trenchant. Length of pg—mg, estimated, 4omm.
24 Bulletin American Museum of Natural History. [Vol. XIV,
Sinopa agilis (JZarsh).
Limnocyon agilis MARSH, Am. Jour. Sci. 1872, Vol. IV, p. 202 (published
Aug. 7).
Stypolophus brevicalcaratus Core, Pal. Bull. No. 3; Proc. Am. Phil. Soc.
1872, p. 469 (published Aug. 7); Tert. Vert., p. 291, pl. xxiv, fig. 9.
From the Bridger Beds, Wyoming. I have preferred Prof.
Marsh’s name for this species as based on a much more com-
plete specimen, including upper and lower teeth and many parts
of the skeleton. The type of S. drevicalcaratus is a fragment of
the lower jaw with m,_3. Prof. Cope’s determination of the
teeth as mj_, is, I think, incorrect.
This species, of medium size, is very close to S. white but
may be distinguished by the smaller heels of the lower molars,
especially of mz. It may prove to be identical with S. rapax
Leidy.
Sinopa pungens (Cofe).
Stypolophus pungens Cope, Pal. Bull. No. 2; Proc. Am. Phil. Soc. 1872
(Vol. XII), p. 460 (Aug. 3); Tert. Vert., p. 291, pl. xxiv, fig. 8.
Type of the genus Stypolophus. Bridger Beds, Wyoming. A
rather primitive species of medium size with long tricuspid heel
on mz, approaching Paleosinopa. Trigon not very high. Rep-
resented by a lower jaw fragment, No. 5015, Cope Collection,
Amer. Mus. Nat. Hist.
[? Sinopa insectivora (Cofe). |
Stypolophus insectivorus COPE, Pal. Bull. No. 3; Proc. Am. Phil. Soc., Vol.
XII, 1872, p. 469.
Bridger Beds, Wyoming. ‘The type of this species has been
lost or mislaid and its position in the genus cannot now be de-
termined. Prof. Cope’s figures in Tertiary Vertebrata are evi-
dently incorrect, the outline of the tooth as viewed from within
and from without being different.
[Sinopa aculeata (Cofc.)]
Triacodon aculeatus Cork, Pal. Bull. No. 1; Proc. Am. Phil. Soc. 1872,
Vol. XII, p. 460.
Stypolophus aculeatus Corr, Ext. Vert. N. M. Rep. Wheeler Survey, IV,
ii, 112. Not S. aculeatus, Tert. Vert., Pp. 299.
1901.] Matthew, Additional Observations on the Creodonta. 25
Bridger Beds, Wyoming. This species was founded on part of
the crown of a lower molar, subsequently lost or mislaid, and the
tip of a premolar crown. It is indeterminate specifically, gener-
ically or even in family, and the name has no standing. The
upper and lower jaws in the Princeton Museum figured and re-
ferred to this species by Prof. Cope in 1884 more probably belong
to S. strenua.
’
Sinopa hians (Coe).
Stypolophus hians Core, Rep. Wheeler Survey, Vol. IV, pt, ii, p. 118, pl.
XXXVili, figs. 12-20.
Wasatch of New Mexico and Wyoming. The type is an
extremely fragmentary and badly preserved skeleton from the
Wasatch of New Mexico. The anterior premolars are spaced
and the last two lower molars subequal. Fourth premolar wider
posteriorly. The other characters given by Prof. Cope are
common to the genus.
The most important distinctive character seems to be the sub-
equal second and third molars. On this ground I place here No.
2850 Amer. Mus. Coll., which agrees fairly well in size, has the
anterior premolars somewhat less spaced, and a narrower heel to
pz. It is a young individual, with teeth scarcely worn, hence
perhaps the less spacing of the premolars. It consists of upper
and lower jaws, fragments of the skull, a few vertebree, and parts
of nearly all the limb bones.
Third lower molar as large as second, first considerably smaller.
Angle between anterior and posterior shear of lower trigonids
averaging 45°. Incisors present but number uncertain. Meta-
style on p* small. Pa. and me. rather closely connate. M#
considerably reduced with vestigial metacone.
The brain is smaller than that of Cynohyenodon cayluxi although
the dentition is one third longer. Compared with that of Zhy/a-
cinus the brain is much smaller in all dimensions, smoother in
surface, and with olfactory and cerebellar lobes smaller in pro-
portion, but not in any degree covered by the cerebral lobes.
The limb bones are for the most part a little stouter than those
of the domestic cat, not very different in size from those of
S. whitte. They are apparently shorter and stouter in propor-
tion, resembling those of Viverravus protenus on a smaller scale.
The vertebre are too poorly preserved for exact comparison,
26 Bulletin American Museum of Natural History. (Vol. XIV,
Measurements, No. 2850.
CHP 54500, FO. Qraijooas (se iyo dno 25
Diameters of m1, anteroposterior 8, transverse 9
es m2 a 9 10.5
“eé m2 ae 4
ee Ms ae 9.5
mg ca 9
Depth of jaw under mz 19
Sinopa whitiz (Cofe).
Stypolophus strenuus Cork, Bull. Hayd. Sur. No. VI, p. 192.
Stypolophus whitie Corr, Proc. Am. Phil. Soc., Vol. XX, 1882, p. 161 ;
Tertiary Vertebrata, p. 292, pl. xxvb, figs. 8-14.
Wind River assise, Wyoming. This species is somewhat
smaller than S. Azans, and is distinguished from it by the reduc-
tion in size of the last lower molar and the more anteroposterior
direction of the trigonid shear. In the upper molars the pro-
tocone appears to be more compressed and placed further back-
ward relatively to the outer cusps, and the metacone of m2 has
entirely disappeared. The species is close to S. ages of the
Bridger, which has the heels of the lower molars more reduced.
It is the best known species of the genus and was fully described
and figured by Prof. Cope in 1884.
Nos. 4780 (type), 4781, and 4782.
Measurements.
CHiN a5 OF Optnwoan 8) Why sysac55 DAE
Diameters of m+, anteroposterior 8, transverse 7
m?” 8 10
m?® 3 9
Ms 2 8.3
Ms 8.3
Length of heel of m, 3
Depth of jaw below m, 19
Sinopa strenua (Cofe).
Prototomus strenuus Corr, Syst. Catal. Eoc. Vert. N. M., Rep. Wheeler
Survey, p. 10.
Stypolophus, Final Rep. Wheeler Survey (U. S. G. S. W. rooth Mer.), IV, ii,
Pe DL7hy pla xxix toes
Wasatch assise, New Mexico and Wyoming. Type from New
Mexico, in National Museum, Washington. Referred specimens
1go1.]: Matthew, Additional Observations on the Creodonta. 27
Nos. 97 and 98, from the Big Horn Valley, Wyoming, Am. Mus.
Coll. 1891. All fragments of lower jaws.
It is quite impossible to separate this species from S. wAzt7@ on
our present knowledge, but when more complete material is found
it will probably show some distinctions. In particular I should
expect to find the metaconid present on m4, as it is on all known
Wasatch and no known Wind River species of the genus. I
therefore hold the species separate provisionally and refer the
Wasatch specimens to S. strenua. The characters are: size
medium, mz smaller than mg, shear of trigonids 55°-60° from
transverse ; p2 with small internal and somewhat larger postero-
external cusp ; teeth compressed with angulate cusps.
A somewhat smaller variety or distinct species has mg less
reduced and shear more transverse. It may prove to be S.
multicuspis. (Am. Mus. Nos. 2815, 4220, 96, and 4218, Wasatch
assise, Wyoming.)
Sinopa multicuspis (Coe).
Prototomus multicuspis COPE, Syst. Cat. Eoc. Vert. N. M., Rep. Wheeler
Survey, 1874 (1875).
Stypolophus, Ext. Vert. N. M., Final Rep. Wheeler Survey, IV, ii, p. 116.
Tertiary Vertebrata, p. 290.
Wasatch assise, New Mexico. The type is an upper jaw,
separable by the much reduced m2 (if this tooth be complete).
The co-type is a piece of a lower jaw, with which the small
specimens referred to above most nearly agree.
[Sinopa secundaria (Cofe).|
Prototomus secundarius Cope, Syst. Cat. Eoc. Vert. N. M., Rep. Wheeler
Survey, 1874 (1875), p. 9:
(Stypolophus), Ext. Vert. New Mex., Final Rep. Wheeler Survey, IV, ii, 115.
Wasatch assise, New Mexico. All the characters given in the
descriptions are common to the genus. The specimen has
not been figured. The measurements hardly separate it from 5S.
multicuspts.
Sinopa viverrina (Cope).
Prototomus viverrinus COPE, Rep. Foss. Vert. N. Mex., Rep. Wheeler Survey,
1874, 13(125); Syst. Cat. Eoc. Vert. N. M., Rep. Wheeler Survey, 1874 (1875), 9.
(Stypolophus), Ext. Vert. New Mex., Final Rep. Wheeler Survey, IV, ii,
(1877), p. 112, pl. xxxviii, figs. I-11 ; Tertiary Vertebrata, p. 290.
¢
28 Bulletin American Museum of Natural History. (Vol. XIV,
Type of the genus Prototomus. Wasatch assise, New Mexico
and Wyoming. The small size distinguishes it from any other
Wasatch species. Judging from Professor Cope’s figures it is
also distinguished by the simplicity of p*, which is little more
molariform than p® of the larger species. The species appears
to be quite primitive in other characters as well, and I am dis-
posed to place with it a few lower jaw fragments in our collec-
tions containing molars of appropriate size in which, while the
trigonid is high, the metaconid is better developed, the shear
more transverse, and the whole tooth wider than is usual in the
genus. (Am. Mus. Nos. 94 and ? 2971.)
Sinopa opisthotoma, sp. nov.
Stypolophus sp. innom, OSBORN & WoRTMAN, Bull. Am. Mus. Nat. Hist.
1892, 110. ‘‘ No. 99 is much larger than any described species of Sipolophus.”
Wasatch assise, Wyoming.
Fig. 9. Stnopa opisthotoma Matthew. Upper and lower jaws, three-
fourths natural size, type specimen No. 99, Wasatch Beds, Big Horn Basin,
Wyoming. Amer. Mus. Exp. 1891. A, upper jaw, from below; B, lower
jaw, from outside ; C, from above. :
With the type (No. 99), upper and lower jaws, teeth well pre-
served, I associate provisionally No. ror, upper and lower jaw
tg01.] Matthew, Additional Observations on the Creodonta. 29
fragments not well preserved, which either is a very large indi-
vidual of this species or represents a distinct species undescribed.
Anterior premolars spaced; p® without distinct accessory cusps; p* with
main cusp conical-lenticular ; internal cusp conical, postero-external cusp not
extended into a shearing blade. Outline of m1 and m2 obtuse-angled triangles,
metastyle especially extended on m2 fa. and me. but little connate; m® with
small metacone, transverse diameter greater than that of m2. Mg with well-
developed heel ; mg in type with much reduced metaconid, advanced paraconid
forming a shearing blade more nearly anteroposterior than in any other Szzopa,
and narrow heel. In No. 101 mz is normal in character, with more transverse
shear and larger heel.
The depth of the jaw is moderate in the type ; in No. ror it is
excessive, although the wear of the teeth does not indicate a
much older animal. Both have the same rounded cusps, com-
paratively slight approximation of fa. and me., simple p*, wide m3,
extended blades on m+-2 and other characters, so that if distinct
the two are evidently closely allied.
Measurements.
No. 99 No. 101
Length c1-m2 83
‘« mi-3 26
“ my-3 31 32
Diameters of p +, anteroposterior 10 II
transverse Io 10
a aa anteroposterior 9 pian
transverse IO Ruta
of * anteroposterior 12 12
m=
transverse 13 I4
i a anteroposterior 6 6
transverse 14 14
Length pg-qg anteroposterior 18.5 23
oe my 8
EL IO 10
wits 13 13
~ eel ofm- 4 6
Width ay kee Ge 4 5
Depth of jaw at mg 19 31
30 Bulletin American Museum of Natural History. (Vol. XIV,
V.—Family MESONYCHID Cofe.
Triisodon heilprinianus Cope.
Two very incomplete and poorly preserved skulls of this spe-
cies are of especial interest as the first skulls described from the
lower or true Puerco Beds, and hence the oldest mammal skulls
known.
My
IG. We
mn
LD
Fig. 10. Triisodon heilprinianus Cope. Part of skull, from above, one-half natural size.
No. 764, Lower or True Puerco Beds, San Juan Basin, New Mexico. Amer. Mus. Exp. 1892.
The skull compares most nearly with those of Avctocyon (Cer-
naysien), AZesonyx (Bridger and Uinta), and Periptychus (Torre-
jon). All have many characters in common :
Brain small, of low type ; zygomatic arches broad ; occipital and sagittal crests
very high ; palatal and basicranial axes parallel (2. ¢., face not at all bent down
on basicranial axis); mastoid well exposed, tympanic bulla rudimentary or
absent ; basisphenoid broad and slightly convex downward ; glenoid fossa deep
and long, post-glenoid process moderately developed ; paroccipital process stout
not long, confluent with mastoid, projecting laterally rather than downwards ;
muzzle thick and heavy, premaxilla with wide ascending process and long con-
tact with nasals. (Nares terminal.)
All the above characters are primitive ones which will proba-
bly be found in nearly all Basal Eocene Placentals. The follow-
ing characters, more or less peculiar to 7yd/sodon, I judge to be
also primitive.
1901.] Matthew, Additional Observations on the Creodonta. 31
Zygomatic process of the squamosal moderately stout, as in
Periptychus, placed well back on the side of the skull (its anterior
-- 2,
Fig. 11. Tritsodon heilprinianus. No. 764. Side view of skull, one-half natural size. Outline
of occiput from No. 3181.
edge one fourth the skull-length from the occipital condyle) as in
Mesonyx. In Arctocyon it is placed further forward, besides being
more massive, reducing the length of the zygomata, which are
increased in width though not proportionately.
The postorbital process on the frontal appears to have been
less developed than in Mesonyx ; on the malar it appears to have
been absent. The postorbital constriction of the skull is very
marked, as much so as in Arctocyon, more than in JZesonyx or
Periptychus.
Fig. 12. Triisodon heilprinianus, No. 764. Upper dentition, crown view, natural size.
The occipital bones seem to have been firmly sutured together,
breaking away rather easily from the rest of the skull.
32 Bulletin American Museum of Natural History. |Nol. XIV,
With the skull were found a few fragments of the skeleton. A
humerus resembles that of Avctocyon, but is smaller and shorter,
deltoid crest hardly as prominent, supinator crest somewhat
more so, distal trochlea wider but quite as deep. The form of
the distal trochlea approximates that of Pantolambda and Perip-
tychus, but the bone is smaller and slenderer than in either of
these, the skulls being of nearly equal size.
Pachyena gigantea 0.& VW.
‘‘ Founded upon a series of finely preserved upper cheek teeth lacking only
the first premolar. The specific distinctions from P. osstfraga are very
marked— (a) the presence of a metacone upon the third upper molar; (b) the
more complex structure of the third and fourth premolars ; (c) the very broad
crenate external cingulum ; (d) the relatively smaller size of the metacone in the
molars.” —Osborn & Wortman, 1892, p. 113.
This is one of the largest of the Creodonta, being exceeded in
size of skull only by some undescribed specimens of A/esonyx in
the Museum collections. Hemipsalodon grandis Cope, though
much larger in skeleton,’ seems to have had a somewhat smaller
skull. Part of a skull and jaws (No. 2823) and a fragmentary
skeleton (No. 2959) are here described. Both are from the
Wasatch beds of the Big Horn Valley, Wyoming, collected by
the Expedition of 1896.
The skull preserves the second premolar and third molar, and
alveoli of all the upper teeth except the incisors, also most of one
ramus of the lower jaw. The fragmentary skeleton includes
parts of the upper jaws with the molars in place, and parts of
the lower jaw with the molars in position and the canines and
third premolar emerging from the jaw. With it are several loose
teeth, most of the hmb bones, but all more or less broken, the
astragalus and calcaneum and several vertebree. The epiphyses
are missing from the majority of the bones.
Upper Teeth.— First premolar one-rooted, the second and third
two-rooted and simple crowned, the fourth three-rooted, molari-
form, with two well separated cusps of equal size and a widely
separated internal cusp. Its peculiar shape, small size, and un-
usual wear suggest that it is a persistent milk tooth; this view is
strengthened by the fact that the corresponding premolar in the
* If the femora referred to it by Professor Cope really belong to the species, which I consider
very doubtful, as the proportion of skull and body would be most unusual for a Creodont.
1g01.] Matthew, Additional Observations on the Creodonta. 33
lower jaw has dropped out early, its alveolus being closed up.
Median (? upper) incisors with large very much compressed root
and small crown. Lateral incisor
with large root of oval section and
small pointed crown. Canine large
of round-oval section.
Lower Teeth—Canines large,
moderately stout, regularly oval in
section, set near together (inter-
space about half width of canine).
First premolar one-rooted, second
: Fig. 13. Pachyena gigantea Osborn
two-rooted, third true premolar & Wortman. Three upper molars in
5 5 : place. Crown view, one-half natural size.
emerging from the jaw, stout, with Young individual.’ No. 2959, Wasatch
4 : Beds, Big Horn Basin,Wyoming. Amer.
protoconid directed strongly back- — Mus. Exped. 1896.
ward, wide, square- based, low tren-
chant heel and no anterior basal cusp. No indication of a fourth
permanent premolar could be found in the immature individual,
but the state of preservation of the specimen does not make it
possible to disprove the existence of one. The temporary pre-
molar is indicated by some remains of an alveolus. In the adult
Q mJ m.2 m*
Fig. 14. Pachyena gigantea, No. 2959. Fragmentary lower jaw, external view, one-half natural size.
jaw, as already observed, the temporary fourth premolar had fallen
out and its alveolus closed without replacement. The first and
second molars have a vestigial metaconid ; traces of it are dis-
cernible also on the third. All three are shaped like those of /.
[ fanuary, rgor. | 8
34 Bulletin American Museum of Natural History. [Vol. XIV,
ossifraga, rectangular in outline when seen from above with high
stout protoconid ridged before and behind, large well separated
trenchant heel, and smaller less trenchant paraconid. The pro-
toconid is vertical in mg, projects slightly forward in mg, and
somewhat more backward in mz. The second molar is the
largest and the third the smallest of the three.
The angle of the lower jaw is inflected, more so than in Dis-
sacus or P. ossifraga, less than in Mesonyx uintensts. In M.
obtusidens ““the angle is prolonged into a stout hook much like
that of Stypolophus”’ (Scott, 87, p. 157), hence is quite different
from the other Mesonychide, in which there is a progressively
increased inflection of the angle, reaching a maximum in the
Upper Eocene Mesonyx uintensts where it is almost like that of a
Marsupial.
Fig. 16. Pachyena gi-
gantea, No. 2959. Right
Kj A ulna and radius, one-
ig.15. Pachyena gigantea, No. 2959. Left femur fourth natural size.
and right tibia, from in front, one-fourth natural size.
The skeleton is very massively proportioned ; the limb bones
are somewhat shorter than those of P. ossifraga, but one-
fourth thicker in the shaft, although the animal is not full
grown. The few dorsal vertebra preserved are short and thick
1901.] Matthew, Additional Observations on the Creodonta. 35
in the centra, and heavy in the arches — apparently short-spined.
The calcaneum has a small fibular facet. The astragalus is very
short and wide, with flatter trochlea, shorter neck, and smaller
sustentacular process than in P. osstfraga; it has the distinct
cuboidal facet common to the Mesonychide, and the navicular
facet is flat transversely (slightly concave in P. osstfraga, flat in
Dissacus, convex in M. obtusidens).’
The most interesting point in regard to this animal is the
question of the succession of the teeth. If, as our specimens
seem to indicate, the fourth premolar belongs to
the milk series, and is not replaced by a permanent AO
tooth, then ¢h7zs species approximates the marsupial | ]
dental formula, with four true molars and three I
premolars. The progressive inflection of the angle == Sx ‘
of the jaw in the Mesonychid phylum is another Os
character of some importance approximating them SS
to the Marsupials. From what we know of the ;
phylum we must regard both these characters as
acquired in the Mesonychide ; the question then
arises — are they not also acquired in the Marsupi-
alia? This is a matter hardly suitable for discus-
sion within the limits of this paper ; I present the
: Bae é Fig. 17. Pach-
evidence for what it is worth. Thatthere is a pro- yena gigantea,
2 : 5 No. 2959. Left
gressive inflection of the lower jaw in the Meson- calcaneum and
ychide is certain; the evidence for the retention of se a
the fourth milk molar may be summarized as fol- seein tir
lows :
1. In the upper jaw the last premolar has the ordinary charac-
ter of a milk molar (small size, short roots, thin enamel, precocious
molarization) and is more worn than the first molar or the third
premolar, hence probably appeared before either of them.
2. In the lower jaw the fourth premolar has disappeared and
its alveolus closed up in an adult specimen, while in a young in-
dividual indications exist of an early protruded tooth, but none
of a replacing tooth although the third permanent premolar is
completely formed and emerging from the jaw.
This evidence cannot be considered satisfactory, especially in
1 Professor Scott's statement regarding the last species is that the navicular is concave in
both directions, This involves apparently an astragalus convex both ways.
36 Bulletin American Museum of Natural History. [Vol. XIV,
view of the fact that in Z7rt¢sodon, the only other Mesonychid
in which the replacement of the teeth has been observed, there
is a permanent fourth lower premolar, though protruded rather
late in life, considerably after mz (Cope, ’84, 270).
COMPARATIVE MEASUREMENTS
M.
Pachyena gigantea . | Mesonyx|uintensis
P. osst- Wa
Sraga dens
No. 72 2823 2959 No. 1892
Benethiof skull? ...-... e. 450 e. 380 279 | e€. 440
Pe Soman oe e. 162 163 135 98 SI)
EN es vere revere ieusepene Ce 7/9) 78 75 54
Breadth palate at m2... 135 e€. IOL 78 121
Depth jaw below ms... 94 7B 68 50 ;
TMengthi my gs. aces ns 99 63 51 (881)
Humerus, width distal
CN eceiss a aches Teer 93 62
Hemmunrwlenetheentr 1-1 e. 290 315
‘* circumference of
Shaft) streaccare 108 86
Winawlenstheee eee os €. 293 286 207
“* circumference of
Shattaeeeereeee 87 65
Radius, diameter of head 49 34 23
‘* circumference of
Shatter nates €7 53
Mibia;slength! s.5 secs e. 265 275 205
““ diameter of distal
@nd is teactyeeee 58 43 25
“* circumference of
Shaltemepccmor 87 74
Calcaneum, length..... 98 We) 56
ue iwadlt hinvaeverers 49 41
Astragalus, length...... 61 44 31
s WGI Sg dis oe 58 41 18
Posterior dorsals, av’ge
length of 4 centra... 34 €. 30
Posterior dorsals, av’ge
width of 4 centra.... 58 e. 25
REFERENCES CITED.
Cope, ED:
’
the Eocene.
72. 1. Descriptions of some New Vertebrata from the Bridger Group of
Proc. Amer. Philos, Soc. Philadelphia, Vol. X11,
p. 460 = Paleontological Bulletin No. 1, published July 29.
2. Second Account of New Vertebrata from the Bridger Eocene.
fbid., p. 466 = Pal. Bull. No. 2, published Aug. 3.
* From type specimen, ‘e.’ = estimated from an incomplete specimen,
1g901.] Matthew, Additional Observations on the Creodonta. OT.
3. Third Account of New Vertebrata from the Bridger Eocene of
Wyoming Terr. Jéid., p. 469 = Pal. Bull. No, 3, published
Aug. 7.
74. Report on the Fossil Vertebrata of New Mexico. Annual Report
U.S. G. G. S. West of tooth Meridian.
75. Systematic Catalogue of the Eocene Vertebrata of New Mexico,
Annual Report for 1874 U. S. G. G. S. W. of tooth Mer.
77. Report upon the Extinct Vertebrata obtained in New Mexico by
Parties of the Expedition of 1874. Final Report U.S. G. G.
S. W. of 1ooth Mer., Vol. IV, part ii.
81. 1. Eocene Vertebrata of the Wind River Valley. Azdl/, U. S. G.
S. Terr., Vol. Vie¥Feb. 26.
2. The Temporary Dentition of a New Creodont. American
Naturalist, Vol. XV, p. 667. August (published July 27).
’82. Contributions to the History of the Vertebrata of the Lower Eocene
of Wyoming and New Mexico made during 1881. Proc. Amer.
Philos. Soc., Vol. XX, pp. 139-194 (printed Mar. 11, 1882).
’83. First Addition to the Fauna of the Puerco Eocene. Proc. Amer.
Philos, Soc., Vol. XX, pp. 545-563. Paleeontological Bulletin
No. 36.
84. Vertebrata of the Tertiary Formations of the West. Final Report
WsSuGeass lems Vol; LI:
EID, I.
71. Proc. Acad. Nat. Sct. Phila., 1871, p. 115.
LEMOINE, V.
78. Recherches sur les Ossemens Fossiles du Terrain Tertiaire Infer.
de Reims ; 1° Partie— Etude du Genre Arctocyon. Aznales des
Sciences Naturelles ( Zool.) 6 Sér., Vol. VIII.
Mars, O. C.
’72, Preliminary Description of New Tertiary Mammals. American
Fournal of Science, Ser. 3, Vol. IV, pp. 122-140, 202-224.
Parts I, II, III, and IV issued as separata July 22, Aug. 7, Aug.
13, and Aug. 17, respectively.
’99. Note on a Bridger Eocene Carnivore. American Fournal of
Science, Ser. 4, Vol. VII, May, 1899.
- MATTHEW, W. D.
’97._ A Revision of the Puerco Fauna. Bull. Amer. Mus, Nat. Hist.,
Vol. IX, pp. 259-323, Nov. 16, 1897.
’99. A Provisional Classification of the Fresh-Water Tertiary of the
West. Bull. Amer. Mus. Nat. Hist., Vol. XII, pp. 19-75, :
published April 8.
Ossorn, H. F.
’90. A Review of the Cernaysian Mammalia. Proc. Acad, Nat. Sct.
Phila., 1890, pp. 51-67.
’93. The Rise of the Mammalia in North America. Vice-Presidential
Address, Amer. Assoc. Ady. Sci., Sec. Zodlogy.
38 Bulletin American Museum of Natural History. [Vol. XIV,
’95. Fossil Mammals of the Uinta Basin, Collection of 1894. Bul.
Amer. Mus. Nat. Hist., Vol. VII, pp. 71-105, pub. May 18.
‘oo. Oxyzena and Patriofelis Restudied as Terrestrial Creodonts. Azil.
Amer. Mus. Nat. Hist., Vol. XIII, pp. 269-279.
OsBorn, H. F., and EARLE, CHAS.
’95. Fossil Mammals of the Puerco Beds, Collection of 1892. BzdZ.
Amer. Mus. Nat. Hist, Vol. VII, pp. 1-70, published
March 8.
Oszorn, H. F., and WorTMAN, J. L.
’92. Fossil Mammals of the Wahsatch and Wind River Beds, Collection
of 1891. Bull, Amer. Mus. Nat, Hist., Vol. IV, pp. 81-148,
published Oct. 20. ,
’94. Fossil Mammals of the Lower Miocene White River Beds. Col-
lection of 1892. Bull. Amer. Mus. Nat. Hist., Vol. V1, pp.
199-228, published July 28.
SCHLOSSER, MAX.
’87. Die Affen, Lemuren, Chiropteren, Insectivoren, Marsupialer,
Creodonten und Carnivoren des Europdischen Tertiars. Bez?,
zur Paldont, Oesterr.-Ungarns, Bd. VI, Wien, 1887-90.
99. Ueber die Baren und Barenahnlichen Formen des Europaischen
Tertiars. Paleontographica, Bd. XLVI, pp. 95-146, Taf. xiii,
Xv.
Scott, W. B.
*87. On Some New and Little Known Creodonts. ourn. Acad. Nat.
Sct. Phila., Vol. IX, pp. 155-185, pl. v—vii.
‘92. A Revision of the North American Creodonta. Proc. Acad. Nat.
Sct. Phila., 1892, pp. 291-323.
WormTMaAN, J. L.
’99. Restoration of Oxyena lupina Cope, with Descriptions of Certain
New Species of Eocene Creodonts. Bull. Amer. Mus. Nat.
ffist., Vol. XII, pp. 139-149, published June 21.
WortTMAN, J. L., and MATTHEW, W. D.
99. The Ancestry of Certain Members of the Canidez, the Viverride,
and the Procyonide. Bwll. Amer. Mus, Nat. Hist., Vol. XII,
pp. 109-139, published June 21.
ZITTEL, K. v.
’94. Lehrbuch der Paldontologie, Pt. I, Vol. IV, Mammalia.
,
°
‘
'
7%
+
*
o
= y
ce
d
‘By J. W. Grwiey.
AUTHOR’S EDITION, extracted from BULLETIN
OF THE
merican Flusenm of Hatural History,
Vou. XIV, ARTICLE IX, pp. 91-141.
New York, May 31, 1901.
Article IX.—TOOTH CHARACTERS AND REVISION
OF THE NORTH AMERICAN SPECIES OF THE
GENUS EQUUS.
By: Wo GIDLEY;
PLATES XVIII-XXI AND 27 Text Ficurks.
CONTENTS,
: PAGE
SUL STi 0) Le Shes) dee 90) 3S a Cola eae 43
sooth! Charaerersas; Altered Dy AGE.) 2305 [24 Veck fost c csc. ode ede 94
Mea yiGUal VARADINEG. carseat aPidsl beat sedee Pose te eatebat TOI
Revision and Diseussion. of the Species. 2.650... i040 saacded ac seceeesan 108
Silesia) Ben coda as Beet pen woe ah aoek ec areee See ee ark eee: & 139
In the successive seasons of 1899 and 1900 the writer was sent
by Professor Osborn to study more carefully the Paloduro
(Goodnight) and Mt. Blanco beds of Texas, both of which Pro-
fessor Cope had placed as typical American Pliocene. This study
resulted in proving that the Paloduro beds are typical Loup-
Fork or Miocene. The Blanco beds are rightly regarded as
Pliocene but do not, as Cope supposed, contain any true remains
of EAguus. In the overlying true Lower Pleistocene or Sheridan
Beds the writer found several skeletons which have already been
described as Zguus scotti, sp. nov. In connection with this work
it appeared that the Pleistocene American Horses were greatly
confused, and Professor Osborn detailed the writer to a thorough
restudy and comparison of all the types, which have been carried
on under his advice. The general results of this work are
summarized at the close of this paper.
Fossil remains of the genus Aguus (indigenous to North
America) were reported as early as 1826 by Mitchell,’ who re-
ferred to this genus some teeth and vertebre found near the Never-
sink Hills in New Jersey, without assigning them to a species.
Since that time about twenty different species have been
proposed by various authors, based on material found in different
localities of the United States and Mexico.
Although the remains of this genus have been found widely
1 Catalogue of Organic Remains, New York, 1826, pp. 7 and 8,
[or]
92 Bulletin American Museum of Natural History. (Vol. XIV,
distributed, and very abundant in some sections, it was not until
the discovery by the writer in the summer of 1899 of the excellent
material upon which he founded the species Lgwus scott that
anything like a complete skull had ever been found in the United
States; nor had the teeth and skeleton of a single individual ever
been found certainly associated. The materials upon which the
North American species were founded, and which formed the
basis of the descriptions and discussions of Owen, Leidy, and
Cope, were very fragmentary, for the most part consisting of dis-
associated teeth and bones, among which portions of maxillary
bones containing several teeth were rare specimens; hence a
number of the species proposed were founded on single disasso-
ciated teeth. Owing to this lack of good material and to an im-
perfect understanding of the characters presented in the teeth,
errors have been frequent and several of the species of this genus,
as they now stand, are practically indeterminate.
This is the conclusion reached by the writer after a care-
ful study of the abundant Aguws material from the Niobrara
River (collected by the American Museum expeditions of
1893 and 1897’), together with the skulls and associated partial
skeletons from the Staked Plains of Texas, and of numerous
specimens of the living species.
The object of the present paper is to present the results of this
study and to attempt a revision of the species that have been
proposed.
Before proceeding with the discussion, however, the writer
wishes to extend his thanks to Prof. Henry F. Osborn for his
valuable advice and aid in the systematic arrangement of this
paper; to Mr. F. A. Lucas of the National Museum and Dr. C. R.
Eastman of the Museum of Comparative Zodlogy, Cambridge,
Mass., for making possible the examination of some type speci-
mens and associated material; to Mr. Witmer Stone of the
Academy of Sciences, Philadelphia, for assistance in examining
the Hguus material in the Academy that was used or identified
by Dr. Joseph Leidy and Prof. E. D. Cope; and to Dr. W. D.
Matthew and Dr. O. P. Hay of the American Museum, for valu-
able suggestions and assistance. The drawings for this paper
his material consists of a large number of mostly disassociated bones, teeth, jaws, and
seul cere There are a few complete and several nearly complete molar-premolar series
in the lot,
tgo1.] Gidley, The North American Species of Equus. 93
were very carefully and accurately made by Mr. Bruce Horsfall.
The photographs were taken by Mr. A. E. Anderson.
The nomenclature of the elements of the molar teeth employed
in this paper is that introduced by Prof. Henry F. Osborn and
is clearly indicated especially in Figs. 1 and 2a, pp. 94, 95.
I. PRINCIPLES OF TOOTH STRUCTURE. CHARACTERS DUE TO
INDIVIDUAL VARIATION AND TO AGE OR DEGREE OF WEAR
DISTINGUISHED FROM TRULY SPECIFIC CHARACTERS.
The following are the chief characters used in definition by
Owen, Leidy, and Cope:
Tooth Characters.
1. Degree of complexity of enamel foldings (Owen, Leidy, and
Cope).
2. Degree of incurvation of superior molars (Owen, Cope).
3., Degree of curvature of superior molar-premolar series
(Owen).
4. Relation of the antero-posterior and transverse diameters of
the triturating surfaces of the superior molars (Owen, Cope).
5. Ratio of antero-posterior diameter of protocone to antero-
posterior diameter of whole triturating surface (Cope).
6. Comparative size (considered unimportant unless difference
was considerable) (Owen, Leidy, Cope).
7. Presence or absence of cup in external lower incisor (Cope).
8. Relative size of third superior molar (Owen, Cope).
g. Crimpled or ptychoid enamel of fossette (Cope).
ro. Character of the separation of the lobes of the metaconid-
metastylid columns (Cope).
Skull Characters Relative to the Teeth.
11. The position, in relation to the teeth, (a) of the anterior ex-
tension of the maxillary ridge, (b) of the anterior extension
of the palatal notch, (c) of the post-palatal foramina; also
(d) the amount of the posterior projection of the maxillary
beyond m2 (Cope). :
More importance has been attached to most of these characters,
apparently, than their merits warrant; evidently, therefore, the
94 Bulletin American Museum of Natural History. [ Vol. XIV,
only way of forming an estimate of their true value is to find out,
in‘a large series of specimens, in what way and how much these
characters may be affected in the individual and in different in-
dividuals of the same species.
The conditions which affect the tooth character of the Horse
may be classed under two heads: (A) Age and (B) Lndividual
Variability.’
A.—TootTH CHARACTERS AS ALTERED BY AGE OR DEGREE OF
WEAR.
(1). Effect of Wear on Complexity of Enamel Folding.
When a molar or premolar tooth first comes into use, the face,
as well as the sides of the crown, is completely covered with
mesostyle metastyte enamel (Fig. 2, 4),
I which folds in and
out, and (though
somewhat hidden by
cement) presents the
same general appear-
ance as that seen
in the much more
‘metaloph primitive forms, An-
chitherium and Me-
sohippus. Soon the
enamel on the tips of
the cones and along
the ectoloph wears
through, and_ small
patches and ridges of
dentine completely
surrounded by a bor-
der of enamel are ex-
povaatyud metastylid entostylid posed. (Fig. 2, B:)
Upper molar, Anchitherium. B, Loner molar, Merjeligpa, “+S the tooth is further
(After,Osborn.) worn away, these
patches and irregular ridges broaden and rapidly lengthen until
when about one-half to three-quarters of an inch of the crown
parast yle
hypostyle
protoloph--
a
_ * Sex seems to affect the tooth characters only in the canines, which are always prominent
in the adult males, but absent or at best vestigial in the females ; hence this factor is left out
of the discussion.
tgo1.] Gidley, The North American Species of Equus. 95
has been worn away they have all become united by narrow
isthmuses, and the fun-
damental tooth pattern
of the horse is pre- > Mia fe
sented. (Fig. 2, C, D.) 800 MN aad p27 2cone
The five prominent ae ad
points or cusps on the
triturating surface of a
newly erupted molar or
premolar tooth are the
paracone, metacone,
protocone, protoconule,
and a cusp apparently
formed by the union of
a ridge thrown out from
the hypostyle meeting
the metaloph at about
its middle point. (Fig.
2, 4.) Thecusps form-
ing the ectoloph are very tm Oo nh
early united, as is also
the protocone with the
protoconule. The next mene 4i PAP -sntecrochet
points of union are ~ >
usually at the anterior
and posterior walls of
the tooth, where the pro-
toloph unites with the
parastyle, and where the
_hypostyle unites with the D
metastyle ; usually then
the antecrochet and
mesostyle
parastyle
metastyle ‘
2
hypostyller~
hypocona=- a :
. vi
crochet unite, and final- 7
ly the metaloph extend-
ing across from. the neiloer: eed
33 protoloph
metaconule unites with
: : Fig. 2. Cusps, Crests, Styles, Crochets, and Fossettes
the ectoloph directly OP- in the molar teeth of the Horse (Z. complicatus). A,
posite she mesostyle unworn|crown. A-D, successive stages of wear.
this is because the metaloph is usually much depressed opposite
96 Bulletin American Museum of Natural History. [Vol. XIV,
the mesostyle, leaving the pre- and post-fossettes continuous
after all other connections have been formed. Thus a continuous
irregular wall of dentine is formed entirely around the tooth
leaving a deep valley or inlet on the internal face of the crown and
two large fossettes or lakes separated by the narrow transverse
wall of the metaloph. The outside wall of dentine is bounded
externally by a thin sheet of enamel, as are also the isolated
fossettes. Just below the point where all these ridges and cusps
are well united, appear the most elaborate enamel foldings, of
the fossette borders, that are to be seen at any stage of wear.
From this point, which is usually about one-half to three-quar-
ters of an inch below the face of the unworn crown, the triturat-
ing surface presents a gradually less complex pattern of enamel
folding as the tooth crown is worn away, until in the very much
worn tooth the simplest pattern of enamel folding is presented.
Thus it may happen that a tooth will present the most elaborately
complex enamel foldings when it is little worn, and become most
simple in this respect when in old age the crown is worn very
short, exhibiting all the intermediate stages in the course of wear.
This principle is demonstrated in Fig. 3. A represents the little
worn crown (No. 2726, Nebraska Coll.) of a fossil tooth of
moderately complicated pattern. A’, 4’, and A® are three sections
of the same tooth, and 4* indicates where each of these sections
was cut. Note the difference in general of the fossettes and
also the changes in the character of the outside enamel foldings.
(II). Zffects of Wear on the Proportions of the Teeth.
1. The Teeth Taken Individually.
Unlike the degree of complexity of the enamel foldings, the
corresponding diameters are affected differently by wear in differ-
ent teeth of the molar-premolar series. The same general rule
for the change in ratio of the antero-posterior to transverse diam-
eter may be applied to the intermediate teeth p2 to m2 in-
clusive, but the most anterior and posterior teeth (p2 and m3)
are affected differently, in this respect, from the intermediate
teeth of the series and from each other as well.
a. Laws Governing the Changes of Diameters of the Tooth Crowns.
There seems to be no exception to the following laws for the
changes of diameters of the tooth crowns as they are worn away
by use.
-_
tgot.]| Gidley, The North American Species of Equus. Q7
Fig. 3. Superior molars of Aguus complicatus, sectioned to illustrate changes in dimensions and
pattern at different stages of wear. 4, triturating surface of mt, A', A*, and A%, sections cut
parallel to triturating surface. 4+ shows where sections were cut ; &, triturating surface of m®; AB, sec-
tion ; B? shows where section was taken.
[May, 1901.] >
98 Bulletin American Museum of Natural History. [Vol. XIV,
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tgo1.] Gidley, The North American Species of Equus. 99
(1) The antero-posterior diameters of the grinding surfaces of
all the intermediate teeth are greatest at the stage when the tooth
has just fully come into use, that is, when about one-half of an
inch, or less, of the crown has been worn away; from this point
the antero-posterior diameter diminishes very rapidly for a short
distance and then continues to diminish more gradually to the
roots of the tooth.
(2) The antero-posterior diameter of the first premolar (p?)
remains about the same for the whole length of the crown, except
that sometimes it narrows slightly near the roots.
(3) The antero-posterior diameter of the last molar (m#), how-
ever, is relatively small at first, and increases continually as the
tooth 1s worn away.
(4) When the teeth first come into use the ¢ransverse diameters
of all the teeth of the series are quite narrow, owing principally
to the rapid incurving of the ectoloph; this diameter increases
very rapidly for about one-half to three-fourths of an inch, but
from this point to the roots of the teeth the transverse diameters
of p= to m2 inclusive remain about the same, diminishing slightly
near the roots; p2 gradually diminishes while m® increases in
transverse diameter as the crown wears away.
(5) The antero-posterior diameter of the protocone in all the
teeth of the series remains the same for the whole length of the
crown.
(6) The antero-posterior or long diameter of the incisors di-
minishes with age while the transverse diameter increases.
b. Effect of Wear on the Relative Measurements of Tooth Crowns.
(1.) Ratio of the antero-posterior to the transverse diameter.
It will be seen from the foregoing that owing to the very slight
variation of the transverse diameters of the crowns of p* to m®
inclusive, for almost their entire length, and to the great shorten-
ing of their antero-posterior diameters, the ratio of these diameters
in these teeth is very different in old and in young individuals of
the same species. Thus in the little worn condition of these
teeth in a young horse, especially before the teeth have worn to
that stage where the transverse diameter is greatest, the antero-
posterior diameter is always greater than the transverse. As the
crown wears away, the antero-posterior diameter diminishes and a
100 Bulletin American Museum of Natural History. [Vol. XIV,
stage is reached where the two diameters are about equal, then,
as the antero-posterior becomes still more shortened, the trans-
verse exceeds it. In every series this variation in ratio seems
always to be most advanced in m1 and m*. This is evidently
due not only to the order in which the teeth of the horse come
into use, whereby the first to appear would at a given stage be
most worn, but also, as is shown by an examination of Table I
(p. 98), because the range of reduction of the antero-posterior
diameters is greater in the molar than in the premolar teeth.
M+ is always the most advanced, as it comes into use before any
of the others of the permanent set.’
(2.) Ratio of antero-posterior diameter of the protocone to the antero-
posterior diameter of the crown.
The antero-posterior diameter of the protocone, being, like the
transverse diameter of the crown, practically unchanged through
wear, also holds to the ever-changing antero-posterior diameter
of the crown in the old and much worn tooth a very different
relation from what it did when the tooth first came into use.
Thus, it may happen that in a little worn tooth the antero-posterior
diameter of the protocone is much less than ha/f that of the entire
crown, but may become greater than half this diameter when the
tooth has become much worn in consequence of this shortening
of the antero-posterior diameter of the crown.
2. On the molar-premolar series as a whole.
The shortening of the antero-posterior diameters of all the other
teeth in the series, except p®, is not nearly compensated by the
lengthening of this diameter in m3, hence it results that the series,
as a whole, becomes much shortened and the teeth from behind
crowd forward toward p® which retains the same relative position
in the skull, so that m2 shows the greatest displacement and the
discrepancy in length is all taken from the posterior end of the
series. The gap which would otherwise be left in the maxillary
bone behind m#? becomes gradually filled in with a new growth
of bone, as the teeth shift forward, leaving a flattened ridge
which is continuous with the rugose prominence or ridge which
WOE 3 ;
7p The order of appearance or eruption of the permanent teeth of the large species from
exas (4. scott7) is the same as Owen has given for £. caéadZus, and is: first, m4, second, m?,
hind, p*, fourth, p%, fifth, p*, sixth, m’. This is probably the order in all other fossil species of
isgenus.
rgor.| Gidley, The North ‘American Species of Equus. IOI
marks the posterior extension of the maxillary bone beyond m2;
hence the length of this posterior extension of the maxillary
depends principally upon the age of the horse. In passing from
the young to the old stage, there is also a marked change in the
relative position, with respect to the molar teeth, of the anterior
projection of the maxillary ridge, the post-palatal foramina, and
the anterior projection of the post-palatal notch,—all appearing
relatively more posterior in the old individual?
B.—INDIVIDUAL VARIABILITY.
a. Degree of complexity of the enamel foldings of the teeth.—The
examination of a large number of specimens of £. caballus, many
of them of about the same age, shows a great variability in this
character aside from that caused by wear; this indicates that in-
dividual variability also has a very wide range, and no two speci-
mens can be found, even of the same age and species, in which
the enamel foldings of the corresponding teeth are exactly alike,
and even corresponding teeth of the opposite sides of the same
skull often show slight differences in the number and style of the
A
Fig. 4. Superior molars m2, Z. cabal/us. A and B of different
individuals of about the same age, showing variation in plication.
minor enamel folds. Fig. 4(4 and 4) represents the correspond-
ing teeth (m2) of two very large draught horses of the same variety,
with apparently only about a year’s difference in age, the one
represented by 4 being the younger. These teeth show consider-
able difference in the degree of complexity of the enamel folding,
which may be due in part to their difference in age, but specimens
can be found in individuals of the same age in which the differ-
ence is just as great. The corresponding tooth of the Texas
pony (Pl. XIX, Fig. D) is much simpler than either of the above,
yet the degree of wear of the teeth shows that this tooth was
apparently intermediate in age between those of 4 and B, Fig. 4.
102 Bulletin American Museum of Natural History. (Vol. XIV,
Although domestication may have made the range of individual
variability greater in Z. caballus, yet a careful examination of a
large number of fossil teeth indicates that in the extinct species
as well the range for this character was great.
b. Lffect of individual variability on dimenstons of the teeth.—It
has been shown under the topic of age variations that the trans-
verse diameters of all the superior molars and premolars, except
p2 and m8, the antero-posterior diameters of the protocones of
all the teeth and the antero-posterior diameter of p* are measure-
ments which change but slightly for much the greater length of
the crown; hence, unless specimens of the same age are taken
for comparison, it is in these measurements that one should look
for evidences of individual variability. Careful measurements of
the teeth of more than ten specimens of £. caballus have led to
the following conclusions: (1) The transverse diameters of the
corresponding teeth for p2 to m® inclusive are remarkably con-
stant, especially in skulls of nearly the same size; the greatest
difference in a certain series of four skulls of large draught horses
examined not exceeding 2.5 mm., and in another series of three
skulls belonging to animals about the size of carriage horses being
less thani1 mm. (See table of measurements, p. 98.)
The transverse diameters of m2 of the large series vary only
.8 mm., while in the small series all the transverse measurements
for this tooth are the same. ‘The greatest difference in the trans-
verse diameter of m2, including both series of skulls, is only
I.5 mm., and adding a skull of the Texas pony to the list the
extreme difference between the transverse diameter of m2 of this
whole lot of skulls, ranging in size from the large draught horse
to the small Texas pony, is only 2.5 mm. It will be seen by
reference to the table of measurements (p. 98) that the average
variation of the corresponding transverse diameter for all these
teeth is very small considering the great difference in size of the
animals represented.
It seems reasonable that much greater variations of the com-
paratively constant characters of the teeth would be found in Z.
caballus than in the extinct species, since in this species domestica-
tion and breeding have caused such a very wide range in size and
proportions of the individuals; hence, when, in two lots of fossil
horse teeth, the difference between the transverse diameters of
tgo1.] Gidley, The North American Species of Equus. 103
corresponding teeth is on the average greater than that between
the large and small varieties of £. caballus, it would seem that
the teeth of the two lots could scarcely belong to the same species,
and although the character of size, alone, could hardly be con-
sidered sufficient ground for establishing a species, yet where this
difference exists, it seems reasonable to expect that when skulls
which represent such two lots of teeth are known, other differ-
ences will be found which will clearly mark them as distinct
species.
It has been shown that the antero-posterior diameter of the
protocone is very little affected by wear; Table I (p. 98), makes it
clear, however, that the range of individual variability of this
diameter is very great, and cannot be depended upon as a distin-
guishing character even in corresponding teeth of individuals of
the same size.
The Lower Teeth.
The characters of the lower teeth are, in general, affected in
the same way as the upper, and seem to be of even less value in
determining the species. <A detailed discussion of the lower teeth
will therefore be omitted, except in regard to a character given
by Cope, that of the presence or absence of a cup in the external
incisors. An examination of the large amount of material at
Fig. 5. , External lower incisors showing development of cups. <A, B, C,
£. caballus; D, E, F, E. scotti.
104 Bulletin American Museum of Natural History. [Vol. X1V,
hand shows: (1) that the fully developed cup is usually present in
£. caballus, though not always, the writer having examined three
specimens in which it is entirely wanting (Fig. 6, C); (2) inthe two
Fig. 6. Mandibular symphyses. A, E. asinus , B, E. scotti; C, E. caballus.
Jaws of £. asénus in the Museum collection there is no indication of
even a tendency toward cupping of the external incisors; and (3)
the fossil species of Zgwus in which this character can be made out
tgor.] Gidley, The North American Species of Equus, 105,
show all the intermediate stages as well as both extremes of this
character. A fact worthy of mention here is that in the Ameri-
can Museum material, the absence of a cup, or a very much in-
terrupted inner wall, of the external incisor, is the rule in the
species with a short muzzle, while in Aguus scotti, which is
a long-nosed type, the entire absence of a cup is much less.
frequent. Figure 5 shows a series of lower external incisors (4,
B, C,) of £. caballus and another (D, &, /) of £. scoftz, illustrat-
ing the variability of this character. The only other character of
the lower teeth which Cope’ seemed to consider important is the
form of the groove or channel separating the two lobes of the
metaconid-metastylid colurmn, which he designates as (1) a
‘‘sharp entrantangular groove,’’ (2) an ‘‘ open gutter,’’ (3) a
‘* flat channel.’’ An examination of a large number of lower
teeth of Zguus and of the Loup Fork genera of horses soon con-
vinced the writer of the great variability and extreme unreliability
of this character in specific determination. In the Loup Fork
genera it seems to be the rule that when the separation of the
lobes of the metaconid-metastylid column is well marked, the
bottom of the groove is nearly always sharp, but when not well
marked then the groove may be said to be a flat channel. It
seems to be only a matter of individual variation as to which
character is presented. In all the species of the genus Aguus
this separation is usually an open gutter.
SUMMARY.
An application of the foregoing principles of the variability of
tooth characters to the specific definitions used by Owen, Leidy,
and Cope shows the unreliability of most of the characters they
have employed. Thus the degree of complexity of the enamel
folding is seen to be greatly affected by both age and individual
variability —a fact which must be correctly understood before
this character can be of even subordinate value as a specific
determinant; the other characters given by these authors have
been shown to be so affected by either age or individual variabil-
ity, or both, that no dependence whatever can be placed on any
of them; and it has been also shown that size, especially the
transverse diameter of the molars and premolars, although least
1 Report Geol. Surv. Tex., 1893, p. 66.
106 Bulletin American Museum of Natural History. (Vol. XIV,
account has been taken of it, is more constant in a species than
any other tooth character hitherto used.
If the foregoing interpretation of the tooth characters of the
genus Eguus is correct, it seems evident that the synopses for a
determination of the species given by Cope have very little value,
and no dependence can be placed onthem. For convenience of
reference one of these synoptic tables is given below.’
‘I, Long diameter of anterior internal lobe of sup. molars not greater than
one-third the long diameter of the crown.
“ Borders of lakes crenate ; internal anterior lobe notched on the inner side so
as to be bilobate ; crowns a little curved; large.......... E. crenidens.
‘‘TI. Long diameter of anterior internal lobe more than one-third and not
more than one-half the antero-posterior diameter of the crown.
@ Crowns more or less curved.
“« Crowns wider than, or as wide as, long; enamel edges little folded.
EE. curvidens.
@a_ Crowns straight or nearly so.
fi Diastemata longer.
“* Crowns nearly square, enamel not very complex ; no facial fossa ; maxillary
bone produced much beyond m2.............-..-.+2---- E. caballus.
(2 Diastemata shorter.
Y No facial fossa.
‘* Crowns nearly square ; enamel not very complex ; maxillary bone little pro-
duced behind last molar ; smaller.
E.. hemionus ; E. burchelli ; E. quagga ; E. zebra ; and £. asinus.
“« Crowns longer than wide on face ; enamel little complicated ; face and max-
rlMeWmy TMT ANS TENT Ss Go So ooccccemoongsococcccossuss E. occidentalis.
‘* Crowns square; enamel more folded than in other species; face and max-
illany unk Owls Ware e -\erehey<)axelo/olejavel oie s/<inleiersieteieteejare teal E. major.
YT A facial fossa.
“*Crowns nearly square; enamel less complex ; maxillary short posteriorly ;
Smaller’ oj. crete ope oierels\aiaielainyele) o> 6 a's oneness ee .£. andium.
““TII. Long diameter of anterior inner lobe more than half that of crown
of molar teeth.
““ Crowns square; enamel little complex (in Mexican specimens) ; diastemata
and maxillary behind shorter ; no facial fossa; large....... E. excelsus.
““ Crowns square ; enamel little complex ; smallest species..... . E. barcenei,”
The synopsis written by Cope and published in the Journal of
the Acad. Nat. Se1., Phila., 1899, Vol. Xi, pant ep regs.
seems likewise to have no standing.
1 Proc. Am. Phil. Soc., Vol. XXII, 1884, p. ro.
107
of Equus,
125 O
Gidley, The North American Spec
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108 Bulletin American Museum of Natural History, |Vol. XIV,
II.—REVISION AND DISCUSSION OF SPECIES.’
(2) Lguus curvidens Owen.
Type Locality.—Punta Alta in Bahia Blanca, Argentina, S. A.
Horizon.—Found together with remains of Megatherium, Megalonyx, etc.
Type.—Superior m2 ; little worn.
Author's description.—‘‘ A greater relative antero-posterior diameter than in
the recent horse, but especially is it distinguished by the greater degree of in-
curvation of the upper molars.”
Measurements.
Be { Antero-posterior diameter....... Sisliekollee ove akties oho eee 27 mm.
ae
lairansyerse’) |e 8). tdasck. ain, & onaiia wieieeloia aeisio RCL
This species must remain rather indeterminate until better
material reveals its true distinguishing characters. Although
Leidy at one time referred to it some Aguus teeth found in the
United States and, so far as defined above, some of the teeth in
the American Museum collection might be placed in this species,
the probability is that Z. curvidens differs from any of the North
American species, owing to its wide geographical separation from
them.
(3) Zguus major De Kay.
Type Locatities.—Neversink Hills, N. J.; north bank of Susquehanna near
Georgetown, D. C.; North Carolina.
7 ypes.—Indeterminate, cited by author as follows: ‘‘ Some teeth and verte-
brz belonging to the genus Zgzus.”
Author:s description.—‘* They [the teeth and vertebree] resemble those of the
common domestic horse ; but from their size, apparently belong to a larger
animal.”’
£. major is practically a nomen nudum, as no figures or measure-
ments were ever given and it seems impossible now to locate the
original specimens.
(4) Equus americanus Leidy.
Leidy first applied this term to three superior molar teeth (Fig.
7) from Natchez until he ascertained that it was preoccupied by
Gervais for a South American type; he then substituted the term
£,. complicatus.
* For dates, location of types, etc., see Chronological Table, p. 107. The names of valid
species are in heavy-faced type.
1g901.] Gidley, The North American Species of Equus, 109
(6) Equus complicatus (Zezdy).
£, americanus Leidy (non Gervais).
Type Locality.—Near Natchez, Missis-
sippi.
fTorizon.—Tenaceous blue clay under-
lying a diluvial deposit.
Typfe.—A second superior molar of the
left side. Cotypes: Two third superior
molars of the right side.
Author's description. —‘‘ The enamel
folds are one-fourth thicker than in the
recent horse and the isolated enamel folds
of the superior molars are much more pean
plicated, resembling in this respect the I | a
Equus plicidens, Owen.” f! ie { 2}
ean
Measurements. / / / } j ly .
y if 1 Vi aes
j) ;
{ Antero-posterior diameter = 1.2
of an inch — 30 mm. / HH j /
Transverse diameter =1.1 of er |
| an inch = 27 mm. } / i i
When Leidy first described this i / esis |
species he gave it the name & | (ee :
americanus, but, as explained above
(4), he later substituted the name / |
£. complicatus." This name was |
employed by Leidy for some time,’
but in his great Memoir of 1869
(Leidy,. 1869, p. 264) he made £.
complicatus a synonym of £. major,
apparently without any justifiable
reason, stating only that he sus-
pected these two species to be the
same. Thus, although 2. mayor was
employed continuously by Leidy
and by Cope until 1899,° the name
really has no standing and Z. com-
plicatus should be retained. AY
The species 2. complicatus itself Fig. 7. Type of Equus complicatus.
See? : - A, triturating surface ; 41, side view of
Issumpentectly characterized except crown. (After Leidy.)
jee
? Proc. Acad. Nat. Sci. Phila., 1858, p. 11.
2 Holmes’s Post-Pleiocene of So. Carolina, 1860.
3 On p. 259 of the Port Kennedy Cave Memoir (Cope, 99), Cope remarked that having looked
nto De Kay’s account he finds no definite application of the term, £&. #za7or, can be made.
110 Bulletin American Museum of Natural History. [Vol. XIV,
in point of size, and although many teeth from different localities
have been referred to it, there is no assurance as to their refer-
ence.
Among the teeth subsequently referred by Leidy to &. com-
plicatus were those on which Cope founded his species (21) &.
intermedius (= E. eous, Hay) found at Petite Anse, La., not over
100 miles distant from the type locality of Z. complicatus ; they
belonged to an old individual, and the differences are principally
age characters; Leidy was probably correct therefore in his pre-
vious reference of these types to 2. complicatus.
(5) Equus fraternus Lerpy.
Type Locality.—Near Charleston, S. C.
Horizon.—Phosphate Beds.
Type.—Superior p2.
Author's description.—As has been pointed out by Cope,’ when
Leidy first named this species he gave no description by which it
could be distinguished, simply stating that “‘its remains are un-
distinguishable from the corresponding parts of the recent horse,
and are the representative of the £. primigenius of Europe, and
may be distinguished by the name of £. fraternus.’’ He after-
ward figured and described a number of teeth from near Charles-
ton, S. C.,’ referring some of them to this species and others to
E. complicatus. From the teeth of this collection designated by
Leidy as E. fraternus, Cope selected a tooth (No. 6, Pl. xv, H.
P.S.) as the type of &. fraternus. Quoting from the publication
referred to, he says: ‘‘ The superior molar which is first described
is figured on Pl. xv, Fig. 6, of that work (Holmes’s Post-
Pleiocene Fossils of S. C.). Unfortunately the protocone of that
tooth is largely broken off, but enough remains to show that it
had the very small antero-posterior diameter characteristic of the
Floridian teeth, and in other respects it agrees with them, except
that itislargerthan usual, . . . I therefore regard it as the type of
the species as described by Leidy.’’ As there are apparently
two species represented by the figures designated by Leidy as
E.. fraternus it is important that the right tooth be selected as
1 Proc. Am. Phil. Soc., Vol. XXXIV, 1895, p.
2 Holmes’s Post-Pleiocene of S. C., 1860, pp. to a 105, Pl. xv and xvi.
tgor.] Gidley, The North American Species of Equus, Teles
the type of this species; it is evident that Cope made a mistake
in this selection, for the tooth to which he referred was not de-
scribed at all by Leidy but simply mentioned and that not until
after the teeth represented by figures 19-22, Pl. xvi, and figure 8,
Pl. xv, had been described. The teeth represented by figures 19-
22, Pl. xvi, are a composite lot of undistinguishable lower teeth
regarded by Leidy as belonging to Z. complicatus and E. fraternus.
Since he mentioned figures 19 and 21 as exhibiting “‘ a greater
degree of plication in the enamel than is usual in any of the lower
molars of the horse, whether recent or extinct,’’ he evidently re-
garded these two teeth as belonging to Z. complicatus and prob-
ably regarded the other two as belonging to Z. fraternus. But as
there seems to be really nothing in the teeth themselves by which
they can be separated or distinguished, the choice of the type of
£. fraternus seems necessarily to fall on the next described tooth
which is represented by figure 8, Pl. xv, and described on page
102 of the above work.
This tooth, taken by the writer as the type of Z. fraternus (see
Figure 8, 4, No. 9200, Coll. Am. Museum Nat. Hist.), seems to
represent in general the teeth to which Leidy intended to apply
this name, and seems to represent also this species, as understood
by Cope, more nearly than the tooth selected by him as the type.
Cope evidently was led to error in the description of his selection
of the type by the poor representation of this tooth as originally
figured by Leidy, for Figure 8, 2, a careful drawingtaken from the
tooth itself, shows that the protocone has not the *‘ characteristi-
cally small antero-posterior diameter ’’ which he attributed to the
Florida specimens, but is rather large. As there seems to be
nothing then to separate this tooth from 2. complicatus, it should
properly be referred to that species.
L. fraternus (thus clearly separated from the type used for it
by Cope and based upon the first characteristic specimen men-
tioned by Leidy) then represents a rather small species of horse
about intermediate in size between Z. complicatus and £. tau as
described and figured by Owen, with the enamel foldings inclined
to be quite complicated in pattern, as is indicated by two other
teeth, in the American Museum collection, probably belonging to
this species (Nos. 9217 and 9203, Coll. Am. Museum, represented
by figure 8, Cand D). The side views of the crowns show that they
112 Bulletin American Museum of Natural History. (Vol. X1V,
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1go1. | Gidley, The North American Species of Equus. 1 te
are well worn and hence near the stage when the simplest form of
tooth pattern would be presented.
Measurements of type tooth.
2 | ANDRO OSHS? CHEMIN. nso ooaeocageacopoacKoued 32 mm.
Transverse TORO ROIS Ce RPO ELE phy ue
Measurements of Nos. 9203 and 9217.
5 Antero-posterior diameter............. m.
p® (So. Carolina) | bere=p S ; gota:
Transverse seas MD Lg sstevahnrsal nent ait oy
a (ina) ee aca als SR De te cseie ce Cee sae PAG)"
‘Transverse (27) SeariSi ero cca Cr 24.5 °°
(7) Lquus excelsus Leidy.
Type Locality,—Pawnee Loup Branch of the Platte River, Neb.
Type.—A fragment of the right side of an upper jaw, containing p+ - m2
inclusive.
Author's description. — ‘‘ About the size of the largest variety of recent
horse. . . . The teeth do not differ in constitution from those of the recent
horse ; and none of them present a greater degree of complication of the enamel
folds on their triturating surface.”
Leidy gave no further identification of this species in his
original description, but in his publication, ‘ The Extinct Mam-
malia of Dakota and Nebraska,’ 1869, p. 266, he gave as an
additional character, ‘‘ the absence of the little infolding of the
enamel at the bottom of the deep valley between the median and
posterior columns of the upper molar teeth’’; and he figured the
type specimen on Pl. xxi, fig. 31, of that work.
Measurements of Teeth.
A i JENNER NOS CSAC CUEING oto adne ane Oona osc eoad 28.5 mm.
P Transverse EMO Siotegcteynederiits « ialiess, siarecshwutener eters 28.5 ‘
mi J Antero-posterior “ss... eee cere eee eee ees 2075 ee
( Transverse SMES SARS A cl avass er en ai ele a Gites De Pieiats)
m2 PAmterO-POSteriGGs Sou U-lerere sna cen oye ewe oes 20) 5anee
( Transverse eh eran gh eas dan ois ave Sys.i8i ous erst ere Pah ey
av) \ Amiigmosposigmioe °° a bb acoonnao mua oouooDGanT 20.)
( Transverse SME cde rsettcterareresclcl cteleiets ciel oe. a s)he IQ. ee
This species, as it now stands, cannot be clearly defined.
The type specimen is quite simple in tooth pattern, much more
simple than is usual in the teeth of about the same size in the
American Museum collection from the Niobrara River locality.
[May, 1901.] 8
114 Bulletin American Museum of Natural History. |Vol. X1V,
The teeth of the type specimen are little worn and hence the
simplicity of the enamel folding is not due to age. A feature
~ a
Fig. 9. Lguus excelsus Leidy. Type. A, crown
view of upper first, second, and third molars, and
fourth premolar. A?, anterior view of p?.
shown in this specimen which may
distinguish it from the other species
of its size is the extreme anterior
position of the postpalatal foramen
and the anterior extension of the
palatal notch. These are placed
about 15 mm. farther forward than
in any of the Nebraska specimens of
the same size in the American Mu-
seum collection, and indeed their
position is more anterior than in
even very young individuals of the
domestic horse. However, to make
this character a strong distinguishing
feature, it would be desirable to
obtain several other specimens from
the same locality showing this same
character.
(9) Equus occidentalis Zeid).
Type Locality.—Tuolumne Co., California,
7 ype.—Superior p2.
Author's Description.—In describing the teeth from California Leidy said :
‘Two of them are second upper molars of different individuals.
These two upper molars, strongly resembling each other, differ from the more
tgo1.|] Gidley, The North American Spectes of Equus, I15
recent looking specimens, and from the corresponding teeth of the domestic
horse, in the remarkable degree of simplicity of the enamel folding, as seen on
the triturating surface.”
Measurements given.
2 ‘o-posterior diameter — 14 lines (31 mm.).
Second upper molar (? m2) { Antero-pos 4¢ (3
Transverse * — 123 ‘‘ (27 mm.).
L oe 1
Ke ae “ (p8 Type?) { Antero-posterior ye SS (Gy seuhey)
Transverse <a 134 ‘* (28.5 mm.).
This species is apparently undis-
tinguishable from &. excelsus, as the
type specimens agree in size and both
are extremely simple in tooth pattern.
However, the Nebraska specimen shows
a tendency to a more complicated pat-
tern of enamel folding and may be an
extremely simple variation of a species
possessing in general more complicated
teeth (Z. complicatus); while an exam-
ination of some teeth representing sev-
eral individuals from the California
locality, which have been identified by
Leidy as &. occidentalis, shows a uniform
simplicity of enamel folding. While
this is by no means to be considered
sufficient ground for retaining the spe-
cies as separate, there is nothing on the
other hand to prove them synonymous,
and as they have been named from two
widely different localities on opposite
sides of the Rocky Mountain system, it
is perhaps wisest to retain the two spe-
cies as distinct for the present.
In 1869° Dr. Leidy united these two
species on the ground of the absence in :
both, of the little enamel fold, near the Scone eee ar
bottom of the deep valley between the pee oe
protocone and the hypocone. This,
1 This tooth has been chosen as the type inasmuch as it was the one figured.
2 Extinct Mammalian Fauna of Dak. and Neb., p. 267. |. When Leidy combined these two
species he retained the name &. occzdentadis, evidently through a mistake in the date of his
first description of Z. excedsus, which he gives as 1868, but which should be 1858.
116 Bulletin American Museum of Natural History. (Vol. XIV,
however, would seem to be scarcely a sufficient reason for unit-
ing them, as, in addition to this being a somewhat variable char-
acter, the type specimen of Z&. exce/sws shows a deep notch in m®
and a slight though well marked one in m1. (The notch in mt
is not shown in Leidy’s figure of the type.)
Professor Cope again separated these two species in 1884,’ con-
sidering them distinct (as he explained in Rept. Geolog. Surv.
Texas, 1893), on the untenable ground of the smaller size of the
protocone in £&. occidentalis. Moreover the specimens upon
which Cope evidently relied for his characters of &. occidentalis
were taken from an entirely different locality and undoubtedly
belong to a species different from either of the two species under
discussion.
(10) Equus pacificus Ze7dy.
Type Locality.—Martinez, Contra Costa Co., California.
Type.—Superior p2.
Author's description.—‘* The triturating surface in its arrangement of the
enamel presents nothing strikingly different from that of the corresponding
tooth of the recent horse. As in this there is an inflection of the enamel at the
bottom of the principal internal valley, and in this respect and the less simpli-
city of folding of the enamel islets of the triturating surface differs from Zgzzs
excelsus of the Niobrara and of California. The tooth probably represents an
extinct species, upwards of eighteen hands high. Its measurements are as
follows :
Length along the outer median column to the origin of the fangs, 26 lines
56 mm.). Breadth of triturating surface fore and aft, 163 lines (35 mm.).
Thickness independent of cementum, 15 lines (32 mm.).”
Without giving any reason for doing so, Leidy included this
species in his list of synonyms of &. occidentalis given in ‘ Fossil
Vertebrates,’ 1873, p. 322.
It seems altogether likely, however, since Z. pacificus is so
much larger, as is indicated by the tooth measurements, and is
more complicated in enamel folding, that it is distinct from Z.
occidentalis. ‘Vhis view is strengthened greatly by an examination
of the Zguus teeth from the Silver Lake, Oregon, locality, which
were described by Cope and referred to the species Z. occiden-
talts,’ but which agree exactly, both in size and description, with
£. pacificus as given by Leidy and are certainly distinct from the
specimens described and referred by him to the former species.
* Proc. Am, Phil. Soc., Vol. XXII, 1884, p. 12.
tgo1.] Gidley, The North American Species of Equus. Li7
If these Silver Lake specimens can be regarded as representative
of EZ. pacificus as described by Leidy, this species can now be
Fig. 11. Eguus pactficus. A, superior fourth premolar, p* (No. 8602, Coll. A. M. N. H.);
B, superior third preniolsy, p3( No. 8601, Coll. A. M. N. H.); C, superior second molar, m2
(No. 8603, Coll. A. M. N. H.)
pretty clearly defined. eee 11, A, B, C (Nos. 8601, 8602,
and 8603, Coll. Am. Museum Nat. Hist.), are representative teeth
of the Oregon collection. Their measurements are as follows:
Amtero=postenionidiameien..i.c. «eee 35 mm.
(No. 8601) p® - P
l Transverse Gg Resa wernt ss BAe ae
(No. 8602) p4 { Antero-posterior SRY Gioia rotor ance Bulls CC
| Transverse mers ei oreiacet ean ee iste 2 Baie
PN No: 8603) 02 { Antero-posterior ey tte tperereiaterswarsis Bouci
Transverse Se Deca crsatnts, SiG ieoeet sie ie ee
The collection above referred to was briefly but not accurately
described by Cope. Careful measurements of the bones in this
collection show that the astragalus and other bones of the feet
that can be identified as fully adult are not smaller than in £.
caballus, as stated by Cope, but compare in size with the large
draught horse. Also he is probably mistaken in the statement
that ‘‘ the inferior canine issues in direct contact with the last
- incisor, without the diastema seen in the horse.’’ None of the
mandibular symphyses in this collection are well enough preserved
to show the unbroken alveolar borders and only show that these
two teeth were close together or in contact deep down in the jaw.
This is often the case in &. cadal/us, but the much greater curva-
ture of the canine brings it to the surface usually some little dis-
tance behind the last incisor, leaving a short diastema.
It may be added to the description already given by Cope, (1)
that the teeth are actually and proportionally much larger, while
the bones of the skeleton about equal in size the larger varieties
118 Bulletin American Museum of Natural History. (Vol. XIV,
of £. caballus; (2) the occipital region seems to be narrower
transversely, and the muzzle, judging from the front portions of
lower jaws in the collection, seems to be proportionally shorter.
(12) Equus conversidens Owen.
Type Locality.—Valley of Mexico.
Type.—A portion of the upper jaw of the right side containing the full
molar-premolar series. .
Author's description.—In describing the specimens to which he gave the
name £, conversidens, Owen considered the following to be the distinguishing
characters: They correspond closely in ‘‘ size, curvature, and pattern of the
grinding-surface of the upper molar teeth, with Aguwus curvidens. . . .
There is ‘‘ a curved convergence of the two series of upper grinders towards the
fore part of the palate to a degree exceeding that in other Equines. .
The last molar, m®, is relatively smaller than in any old-world Equines.
The first premolar, p2, resembles in the minor production of the anterior lobe
that tooth in Zguus asinus, Equus gquagga, and differs in this respect from Z-
caballus. The grinding-surface, however, retains, as in Hazes aff. caballo, Ld.,
and in £. curvidens, the general conformity of character of enamel-folding so
remarkable in all the modern and in the European Pleistocene Equine species.
hitherto described. . . . But the most distinctive character of the upper
grinders of £. conversidens is their disposition in the jaw, denoted by the
nomen triviale of the extinct Mexican horse; and to the character of the curva-
ture of the molar series of alveoli may be inferentially added a concomitant
modification of the shape of the upper jaw itself, involving that of the lower
one.”
Measurements.
a. § Antero-poesterior diameter... /..-5 a0 sans eee 30 mm.
P ( Transverse i I Se PRE Oy vid Ge Ot
g J Antero-posterior DOA IO R Gian soisGi0-0% bas 2B
P ( Transverse BS hg hid tay! 20% pavaiis Ne ae Cae Se
ar ened ads TO) Saigls edna ey ete eee $22.
Transverse HO OS Beco cepairas tien cated oe ne a 2s
m2 § Antero-posterior Oo) Wye sisieto Rc bibeyas Cee Adore <i) ss
( Transverse OO ” aWals nsec Sila rahe oe ee ifs St
The principal character upon which Owen based this species is
undoubtedly an unnatural feature, not, however, due ‘to distortion,
as suggested by Cope,* but rather to a placing together of upper
jaws of opposite sides of two different individuals. The writer
was led to this conclusion by a careful comparison of the corre-
sponding teeth of the opposite sides as shown in Owen’s figure of
his type specimen, (See Plate XXI, which isa reproduction from
? Proc. Am. Phil. Soc., 1884, p. 13.
tgor1.] Gitadley, The North American Species of Equus. 119
the original plate, Phil. Trans., pl. Ixi, fig. 1.) An examina-
tion of this figure shows that not a single tooth in the one series
corresponds at all closely with its opposite in the other series.
Not only are the comparative measurements different, but the
enamel foldings are so radically different in the corresponding
teeth of the two series as to make it very certain that the speci-
men is a composite of two individuals.
A comparison of the two series in any skull of any species of
Equus will show the reasonableness of this conclusion. ‘These
differences, however, are entirely within the range of individual
variability and of changes due to age. ‘The relatively shorter an-
tero-posterior diameters of the teeth (especially of m+ and m2), the
relatively less length of the entire series and the relatively greater
posterior extension of the maxillary beyond m2, all indicate that
the series of the left side is of an older individual than that of
the right.
Although the principal character assigned to this species by
Owen must be disregarded, there remain still enough other
characters to define it fairly well. Two of the characters
pointed out by Owen, which are shown in both series of his.
type specimen, seem worthy of consideration, namely, the
comparatively small size of m* and of the anterior lobe of
p®. Evidently the comparatively small size of m# is not due
to a little-worn condition of these teeth, as the compara-
tively small antero-posterior diameters of the other molars,
especially in the series of the left side, indicate that it, at
least, is of an old individual with all the teeth much worn.
Cope seems to suggest, in his article above referred to, that
Owen might have taken this character from an imperfectly pro-
portioned photograph of the specimen, for he remarks: ‘‘ This
appearance could be produced by the oblique angle of the aper-
ture of the camera in photographing, due to its too anterior
position.’’’ This view, however, seems scarcely worthy of con-
sideration, as the other teeth seem to be not at all disproportioned,
and had so great a distortion of m® been produced m® at least
would have been affected to a perceptible degree also. Moreover
it is evident, from reading Owen's descriptions of the species £. faz
and £&. conversidens, that, although, as he states, his descriptions
1 Proc. Acad. Sci. Phila., 1884, Vol. XXII, p. 13.
120 Bulletin American Museum of Natural History. [Vol. XIV,
were based on a series of specimens, casts, and photographs sent
to him by the Engineer of Mines, Mexico, he must have had
either the cast or the specimens themselves, upon which he
founded the species Z. conversidens, and from which he took his
characters and measurements; for he mentions at the beginning
of his description (see p. 118) that, among other features, the
curvature of the molar teeth corresponds closely with Z. curvidens,
and this character could not possibly be made out from a photo-
graph showing only the crown view.
Thus, then, the two characters above mentioned, together with
its smaller size and geographical location, seem to determine this
species fairly well.
(13) Equus tau Owen.
Synonym, Laws barcenet Cope.
Type Locality.—Valley of Mexico.
7 ype.—A superior molar-premolar series lacking p2.
Author's description.—‘* They [the teeth] are as much smaller than the cor-
responding teeth of Lgws conversidens as are those of the Asinus fossilis
from the Oreston Cavern (History of British Fossil Mammals, p. 396, figs. 157,
158) compared with the teeth of Aguas plicidens from the same cavern, and
they indicate a species about the size of the common Ass.
‘* As compared with any of the smaller existing kinds of Equines the antero-
posterior diameter of the grinding-surface of the crown, especially in the pre-
molars, is in excess ; and in this character Lguwus tau also differs from Eguus
conversidens, as it does in the greater relative size, especially antero-posterior
breadth, of the last molar m?: £. faz further differs in the greater flattening,
from without inward, of the inner lobe:of most of the molars.”
Unfortunately Owen did not give any measurements with this
description, but if his figure of the type specimen is sufficiently
accurate to be relied upon, as seems quite probable from the
exactness of his other figures, the measurements may be given as
follows:
53 { Antero-posterior diameters <:4c%.ccen dearer 22.5 mm.
( Transverse Tie) sates OS josake ease OeW AOC Ree 19 Gy
p4 \aintero-pesterior: 7 "°°!" 3.3),42 52 ee 234) Pas
' (| Transverse a. Gotha s Rates an See 20 os
au jpAntero-postenon 9.) bik. act sows aa 21 e
Transverse DE Noel ote iaitie, eahe (cree Re MOS
ja) ) Amtero=posterion | {Sabi sis ape Cece ee eee
/ Transverse panne RPE PEE ho Ec G0 c 18 i
g -Amtero-posterion © 8 ele i 22 ame
( Transverse a P55)
ie) (8! [ah 0) caine) ale, ‘aye [elie leviv, jee) e)\eiieier esate
1901.) Gidley, The North American Species of Equus, I2t
This species is much the smallest of any described from
America. Owen considered it about the size of the common
Ass, but it is much smaller than that
species, Z. conversidens and £. asinus
being much more nearly of a size
(see Table I, p. 98).
In. his paper on ‘The Extinct
Mammalia of the Valley of Mexico’?
Cope united £. conversidens with £.
tau, retaining the latter name. Why
he did so, he does not explain, ex-
cept that he says, “‘I could detect
no specific differences between the |
seven or eight specimens I examined.”
As Cope expressed considerable
doubt as to the identity of the type
specimens, especially of £. faz, it
seems quite probable that the true
type of this species was never in his
hands, and his description of £. tau
was probably based upon specimens
that properly belonged to £. conver-
sedens.
There is a superior molar tooth,
m2 (No. g211), in the collection of
the American Museum from Peace
River, Florida, which is much too
small to be referred to any species
reported from the United States. A
second tooth from Lookout Moun-
tain, Tennessee, recently presented
to the American Museum by Mr. E.
S. Hallock, agrees almost exactly in etre ee oe ates:
size and character with the Florida second premolar, p?. (After Owen.)
tooth. These two teeth represent a
very small horse once inhabiting the south-eastern part of the
United States, which in point of size did not differ materially from
£. tau of the Mexican Valley. Hence as there are no characters
1 Proc. Am. Phil. Soc., Vol. XXII, 1884, pp. 12, 13.
122 Bulletin American Museum of Natural History. [Vol. XIV,
in the teeth above mentioned to distinguish them from Z&. ¢au they
should provisionally be referred to this species, although the two
localities are so widely separated. Future discoveries may prove
that the teeth from the southeastern United States belong to a
distinct species.
(14) ZLguus crenidens Cope.
Type Locality.—Tequixquiac, Valley of Mexico.
Type.—Fragment of upper jaw containing the premolars of one side.
Author’s description.—‘‘ Vhis species is primarily distinguished by the close
and strong wrinkling of the enamel border of the lakes of the superior molar
teeth. This wrinkling, or vertical plication, reminds one of what is seen in the
Elephas indicus, This wrinkling is not found in the enamel edges which border
the interior crescents on the inner side, nor in those bordering the internal lobes
or columns. The borders of the lakes are not folded in the complex loops seen
in Equus major Dek., but have the plainer looping seen in Zguas taw Ow
The grinding faces are nearly square. . . . The crimping of the enamel of
the lakes distinguishes this species from the others of the genus.”
Measurements gluen 4
9 | Antero-posterior diameter........-5.2-- censors 43 mm.
_ ( Transverse OS eek sate dgantn te sapere reece Reet 80:5.
pe (‘Antero-posterior 97 0 oes omens nae eater ae 3857
( Transverse ie Mea eroriepe Ot) Ga cmos O80 34
pt ) Antero-posterior EY |) sw sthartyacelainrarsapehcvereersreemreere yt
' ( Transverse Se Na ray caiorstoenyetarae eae eee Be Ue
Although a minute description and measurements have been
given, this species appears to be indeterminate. As seems usually
to have been his custom, Cope has given the transverse measure-
ments of the teeth, including the cement, hence they are not of
much value except in as far as they show that the type of &.
crenidens is of a large horse, apparently about the size of Z. pacif-
cus or the somewhat smaller £. scott#é, The crimping of the
enamel which Cope seemed to consider the distinguishing char-
acter is probably not more than an exaggeration of the same
character often indicated in the premolars of &. caballus. That
Cope himself did not place great reliance on this character is
evidenced by the fact that he afterward described and figured a
large tooth from southwestern Texas,' referring it to this species,
which shows no uncommon tendency to any such wrinkling.
This tooth, however, from its enormous size evidently belongs to
1 Am, Nat., 1885, p. 1208, fig. 4 (No. 8616, Am. Museum Nat. Hist., Cope Coll.).
tgo1.] Gidley, The North American Species of Equus. 123
a species much larger than the Mexican specimen which Cope
called £. crenidens, or even the large Z. pactficus.
(15) Equus barcenei Cope.
Type Locality.—Valley of Mexico.
7 pe.—Two superior molars.
Author's description.—‘‘ This horse is distinguished from all the others here
mentioned or described by its small size. In the characters of its superior
molars it is like the Zgzes excelsus, Vhe anterior internal column is flat, and
its antero-posterior diameter is five-eighths that of the crown of the tooth. The
prism is straight. The lakes have the margin but little looped; the posterior
notch of the anterior lake is trebled or triplex. The grinding face of the crown
of the third superior molar is a little longer than the others.”
Measurements.
: IAMtELO=POStETION. anise Shui vate oe ec 21.5 mm.
Diameters of molar No. 1. P 2
siiramSWense Mierasi1s slcsvs ie cysia a are stomes eit Ly oe
; Ante O=POStEMOLs << ae sae see) os ae ose 22 ah
Diameters of molar No. 2. 5 P
(Malttramsyerserers eve cic vaen enya oe Bay
In 1893’ Cope made this species a synonym of £. faw. It is
difficult to make out whether, in doing so, he referred to 4. tau
Cope (not of Owen),* which has been shown to be the equivalent
of Z. conversidens, or whether he now regarded Z. ¢aw in the light
of the original description given by Owen. It seems probable,
however, that the latter view is the correct one, as in this connec-
tion he referred to Owen’s description of 4. ¢au and not to his
own. ‘Taking into account the fact that the transverse diameters
of the teeth of &. darcenei given by Cope probably include the
cement, there seems to be no distinction between this species and
£. tau Owen, and therefore it may be considered, as Cope has
already placed it, a synonym of this species.
(16) Pliohippus simplicidens Cofz.
Equus simplicidens Cope.
Type Locality,—Mt. Blanco, Texas.
Horizon.—Blanco Beds (Upper Pliocene).
Type.—Superior molar.
Author's description.—‘‘ The size of the teeth is about that of the 2. occtden-
talis and £, caballus. The internal column is of moderate antero-posterior ex-
tent, its posterior border marking the anterior third of the posterior lake. Its
1 Report Geol. Surv. Texas, 1893, p. 7u.
2 Proc. Am. Phil. Soc., 1884, p. 12.
124 Bulletin American Museum of Natural fistory. [Vol. XIV,
long diameter is considerably less than half that of the crown, A peculiarity
found in two of the superior molars, but not in two others, is that the median
dental connection between the external and median crescents is interrupted by
the continuity of the enamel plates bordering the lakes from the one to the
other. This arrangement is frequently seen in the large pm4, in the species of
Equus, but does not occur in the other premolars and molars, It is a reversion
to the condition seen in Anxchitherium. A principal character of the species is
seen in the extreme simplicity of the enamel borders of the lakes.”
Measurements not given.
This species was founded upon exceedingly doubtful characters,
all the characters given in the above description being apparently
due to the little-worn condition of the tooth. (See Fig. 2,
C, D, and discussed p. 95.) However, the validity of the spe-
cies is somewhat strengthened by Cope’s description and figures
of additional specimens from the same locality,’ and from the fact
that it seems to be the only large horse so far discovered in this
geological horizon.
It seems exceedingly doubtful, however, whether this species
can be properly placed in the genus Zguus. Cope did not point
out or suggest any characters that
mark it as belonging to this genus,
nor are there any such characters
shown in any of his figures. On the
contrary, there are in his descriptions
and figures, and also in three teeth
collected by the writer from the same
locality (No. 10624, Am. Mus, Nat.
Tyee Cir ec, Hist; Collec.) and probably, belone=
Superior premolar. (After Cope} ing to the same. species, iseveral
indications of primitive characters
linking this species much more closely with P/iohippus, as this
genus is understood, than with Aguus. The character of the
very small protocone and the tendency to a sharp entrant-
angular groove between the metaconid and metastylid columns,
a distinguishing character given by Cope’ forthe lower teeth, are
both like the Loup Fork genera and unlike any of the Pleistocene
or recent species of Zguus. Besides these characters the Ameri-
can Museum specimens show other characters which seem
* Report Geol, Surv. Texas, 1893, p. 66.
tgo1.| Gidley, The North American Species of Equus, 125
to indicate the close relationship of this species to the more primi-
tive genera of the Loup Fork. The little outward folding of the
enamel at the antero-external border of the protoconid, so gen-
erally observed in all the Loup Fork genera, is plainly indicated
in the two lower teeth by a well-marked groove. This peculiar
fold of enamel, which is entirely wanting in the genus Zguus, seems
to be a primitive character and apparently was derived, in the
evolution of the tooth, from the anterior cingulum of the more
primitive brachydont tooth in passing to the hypsodont form.
Although these teeth are of an old individual and are pretty
well worn, they have the appearance of being of a very short-
crowned pattern.
This species is about the size of Plohippus mirabilis Leidy.
(17) Hipparion eurystylus (Coc).
Equus eurystylus Cope.
Type Locality,—Paloduro Caiion, Texas.
fforizon.—(?) Goodnight Beds.
Type.— Lower molar.
Author's description.—‘' This species belongs with Z£. mznutus Cope, to a
section of the genus characterized by the relatively great width of the metaconid-
metastylid column of the inferior molars, and its close appression to the proto-
conid and hypoconid, and hence by the relatively narrow molar crown ; and
also by the small size.” (For further description see Report Geological Survey
of Texas, 1893, pp. 44, 45.)
Measurements given.
Longitudinal diameter............ 55 mm,
Specimen’ No. 2 (Cope) Antexo-posterior ““ ..-........- 2Al as
( Transverse ay) maynuatisy siakeneekahey = TOs
( Longitudinal Bee Pastries incccesyerey are 405
Specimen No. 5 (Cope) ; Antero-posterior ‘* ............ 22i
Transverse tat Moaoohctrconisc UST
None of the characters given in Cope’s very minute description
of this species, or shown in his figures, prove any relationship to
the genus Zguus ; but they all seemingly show its positive identity
with or at least its very close relationship to the three-toed horses
of the Loup Fork.
An examination of the horse teeth from the true Loup Fork
Beds (in the American Museum collections) indicates that the
126 Bulletin American Museum of Natural History. |Vol. XIV,
character of ‘‘ the relatively great width of the metaconid-metasty-
lid column of the inferior molars ’’ is only a feature of individual
variability, as this column is very variable in size. The charac-
ters that seem to point most strongly to the association of this
species with the genus Hipparion are as follows: (1) The appres-
sion of the metaconid-metastylid column to the protoconid and
wpe ee
Fig. 14. Hipparion eu- Fig. 15. Hipparion eu- Fig. 16. Hipparion eu-
rystylus. Type of Equus rystylus (Equus eurysty- rystylus (Equus eury-
eurystylus Cope. Superior Zus Cope). Lower molar. stylus Cope). Lower pre-
premolar. (After Cope.) (After Cope.) molar. (After Cope.)
hypoconid, which seems to be characteristic of Protohippus and
Hipparion and not of Zguus ; (2) the presence of a well-defined
enamel keel or loop in the anterior border of the protoconid; (3)
the greater extension of the transverse ridge of enamel thrown
out from the parastylid, which extends beyond the middle and
usually to the outer edge of the metaconid, and marks the antero-
internal corner of the tooth in Z. ewrystylus and all the three-toed
horses; and which in Z&. cadal/us and the fossil species of Aguus
seems seldom to extend beyond the middle of the metaconid and
often only as far as its inner margin:
(1) The tendency toward flat external faces of the protoconid
and hypoconid, and (2) toward the formation of the external
median keel, mentioned by Cope, together with its size, seem
definitely to place this species in the genus Hzpparion. (Compare
Cope’s figures of 4. eurystylus with Leidy’s figures given in ‘ Ex-
tinct Mammalia of Dakota and Nebraska,’ Pl. xix, figures 4,
6, 10, 31, 34, 37, and 4o.) The specific distinctions apparently
cannot be made out.
(18) Protohippus cumminsii (Coe).
Equus cumminsii Cope.
Type Locality.—Mt. Blanco, Texas.
Horizon.—Blanco Beds.
7 ype.—Superior molar.
Author's description,—* The enamel borders are entirely simple.
It differs from the Z. stmplicidens and £. tau in the prominent concavity of the
rgot. | Gidley, The North American Species of Equus, 27
internal wall of the paracone and metacone, from which it results that the ex-
terior border of each of the lakes is deeply concave, and the horns appear to be
strongly produced. The protocone has a very short antero-posterior diameter,
in which it resembles the 2. simplicidens, E. crenidens, and E. stenonis, and
differs radically from the 2. faz and the other species of the Equus beds. Itis
not bilobate or grooved on the internal face. . . . The subcylindric char-
acter of the paracone and metacone approach what exists in ‘ Hippidium’ and
some species of the three-toed horses.”
Aleasurements given.
The measurements of the best-preserved crown are: length, 33 mm.; trans-
verse diameter, 24 mm.; antero-posterior, 24 mm.
This species was founded on three superior molars, two of
which were figured by Cope. (See Figure 17.) Both the de-
scription and the figures show some char-
acters that seem to mark its close relation
with the three-toed horses, and like those
given by Cope for the distinction of his Z.
eurystylus and £. stmplicidens they show no
characters that indicate any connection
with the genus Aguus. The very small
size and form of the protocone seems es- Fig. 17. Type of Proto-
pecially to identify this species with the i any Sate ae
Loup Fork genera, and its extreme sim- ™2" “Af Cope.
plicity of enamel folding of the tooth crown seems to place it in
the genus Protohippus.
Une i
SIEtsSTTAt
(19) Protohippus phlegon (//ay).
Lquus phlegon Hay.
Synonym, Laws minutus Cope.
Type Locality.—Mount Blanco, Texas.
fTorizon.—Blanco Beds.
Type.—An inferior molar—(?) mg. :
Author's description.—‘* The inferior molar is narrowly hypsodont, and has
apparently had but a thin cementum investment. . . . The inner side of
the crown shows marked peculiarities. The metaconid and metastylid are ap-
pressed to the hypoconid and protoconid, and are spread widely apart so as to
be connected by a narrow antero-posterior isthmus, and separated on the internal
face of the crown by a wide channel which has a greater antero-posterior diameter
than the metaconid and metastylid respectively, near the apex of the crown,
and an equal width at the base of the crown. The hypostylid (entoconid) is also
appressed to the hypoconid and has a recurved postero-internal angle, which
128 Bulletin American Museum of Natural History, [Vol. XIV,
forms an acute ridge bounding the internal face of the crown posteriorly. This
encloses with the metastylid a flat open gutter as wide as the metastylid. A
sharp ridge marks the internal face of the anterior border, but it is closely ap-
pressed to the metaconid, which it does not equal in elevation.”
Measurements given.
Length of crown preserved... ..- +. .eeee sees eee e eee eee 37. mm.
Antero-posterior diameter at middle............+..-+e0e: ios
Transverse diameter at metastylid.................+-+-- 8 ts
Transverse diameter in front of metastylid.............-. 6 “s
This tooth shows the same characters pointed out in Cope’s £.
eurystylus to distinguish it from the genus Hguus except that the
little fold of enamel at the anterior border of the protoconid is
wanting. A comparison of
Cope’s figure (reproduced in
Cr) QS Figure 18, and Figure 19
; which represents a corres-
oe vO-
Pld, Wit allthis Srope Na es Cas
(Equus minutus Cope). sf. (Loup Fork). te : y , {
Moves ales sites Mus., ) shows the very close
relationship of the species to
which the teeth represented by these figures belong, and were it not
for the little enamel fold mentioned above, in the one, they would
undoubtedly be referred to the same species if their different lo-
calities were not known. In aseries of five lower teeth (No. 10626,
Coll. Am. Mus.) of this species, found in the exact locality
in the Blanco beds from which Cope’s type specimen was taken,
the same general Protohippus-like characters are shown as in
the type tooth, and a rudiment of,the enamel fold of the protoconid
is shown in all of them.
Very unfortunately, an upper molar, found by the writer in
the same locality a few days before the finding of the lower teeth,
was lost. This tooth agreed in size with the lower teeth and
could not have been distinguished from Protohipfpus. The
crown was not much worn and the little oval protocone was still
quite separated from the protoconule.
The species distinctions of this little horse of the Blanco beds
apparently cannot be defined, unless the rudimentary condition
of the anterior outward fold of enamel of the protoconid should
prove to be constant and thus separate it from the species of
Protohippus from the true Loup Fork beds.
tg01,] Gidley, The NM orth American Species of Equus. 129
(20) Equus semiplicatus Cofe.
Type Locality.—Rock Creek, Brisco Co., Texas.
fforizon.—Sheridan Beds.
Type.—Superior molar m2 (?).
Author's description.—‘‘ These teeth indicate a species of about the dimen-
sions of the &. ¢az, but characterized by a greater complexity of the enamel
foldings. . . . In all the Texas specimens the antero-posterior diameter of
the protocone is more than half that of the grinding face of the crown. This places
the Lguus semiplicatus in the same group as the &, excelsus and Z. tau. From
the former its inferior size distinguishes it, as well as the closer plications of its
enamel borders. . . . The characters of the grinding face resemble those
of the Z. conversidens Owen, as far as the latter can be understood from
Owen’s figures. But in this species, according to Owen, the crowns are trans-
versely curved, while in Z. semzplicatus they are straight.”
Measurements given,
Diameters) |supenior \\Antero-postenor.-.1- ee li 25 mm.
mayo eye (Al wike(Cannvern)) ) ewe eKS 504 ao cacbnddcv00bd0God oa
Diameters superior molar § Antero-posterior............... 25a
(sp: from) San Diepo)) ji Transverse. ..c.,c/. > ae w:0'e)e ces ores big] ake
A comparison of the measurements will show that the teeth of
this species are very much larger than
those of Z. fav, and are even larger than
those of Z. conversidens as given by Owen.
The species seems to be about intermediate
in size between £. complicatus and E. con-
versidens, and is about the size of £. fra-
ternus. Its tendency to a very simple tooth
pattern, however, seems to distinguish it
from the latter species.
A skull from southwestern Texas, now in Fig. 20. Eguus semipli-
; : catus Cope (type). Superior
the American Museum collection (No, molar. (After Cope.)
8600), which Cope had previously described,
_and referred to 4. exce/sus,’ and which was again mentioned and
figured in his Rept. Geological Survey of Texas, p. 81, Pl.
xxll, Fig. 3, differs not at all from Cope’s description, measure-
ments, and figures of /. semiplicatus, but does differ in both size
and general character from Leidy’s type of 4. excelsus. Also,
coming from practically the same locality as the type specimen
of L. semiplicatus, and from the exact locality from which two of
the paratypes were collected, it seems to belong more properly to
the latter species than to &. excelsus.
oe
Osrsac=222380ie'
Nika
1 Am. Naturalist, 1891, p. 912.
[May, 1901.] 9
130 Bulletin American Musuem of Natural History, (Vol. XIV,
This skull has been
fully described by Cope as
above cited. A character
mentioned by Cope in
this description, that of
the posterior prolongation
of the vomer over the
presphenoid, (Fig. 21) a
character in which it re-
sembles the ass, seems dis-
tinctly to separate this
species from all the others
of the Pleistocene in
which this character is
known, they being like &.
caballus in this respect.
(21) Lguus intermedius
Cope.
As this term is preoccupied,
Hay substituted for it the term
EZ. cous,
Type Locality,—Petite Anse,
Louisiana.
Type.—Fragment of maxil-
lary containing the three molars
\) and two premolars, p® and p4.
\ Wh, 4 ethor’s description.—Cope’s
:
description of this species is
given at considerable length,
and as much of it is a discus-
sion of characters that have no
standing, only the most import-
ant of the characters mentioned
will be repeated here. *‘‘ The
molar teeth are as large as those
of the Z. major and large forms
of the common horse. The
enamel folds are less complex
than those of the former, but
“c = are more co x than is usual
Fig. 21. Skull of Eguus semiplicatus (No. 8600, Coll... ee . complex t
A. M. N. H., San Diego, Texas). Palate view. inthe &. caballus. . . . A
tgo1.] Grdley, The North American Species of Equus, 131
transverse crest connects the paroccipital processes and the basisphenoid, bound-
ing the precondylar fossa in front, so as to convert it into a basin.
The Z. tntermedius is a shorter-nosed species than the Z. caéallus, the distane
from the anterior lower premolar forwards only equalling the corresponding
length in the quagga, while the parts are in other respects as large as those of
the domestic horse.”
Measurements given for the teeth,
Diameters of p2 WPATitenO=pOStiemOtea-iieeyeloe en irae lei <1 © 30 mm.
Wiiraticverscme:, 20 es inc. ycmindeacee tl ea
WDramere we Ole PATED ETO OSEEGIOLS ooh ich afr ayoreyenie<inqe Sa nie 6 PA
San Mle ATIS CISC ei red sich. ats dicyaseisrarer ses ciavors eee ts .t 30>
Wramiaterwot m2} PAMMEETO=POSLELI Ola aperchelscseyelsitciel creer eis eee gia)
BIRKS WELS Chere syste cies crests. cha eos ierace. one ie PQ)
Length of mandible from pm#® to external border of in-
CISOLSWevere ere shed pavesorste =o eicvetele oi cievorers alesis sel wv ain te wureve Ge aiaie Tkayo)
The specimens upon which Cope founded this species, as above
mentioned (see Z. complicatus, page 110), were considered by him
identical with a certain other lot of specimens from the same
locality which had been referred by Leidy to 2. major (£. com-
plicatus),* but which he thought could not be identified with that
species; separating them on the ground of the less complexity of
the enamel folding in these teeth than in those of 4. complicatus.
One of the specimens, at least, figured by Leidy in the work
above cited is more complicated in tooth pattern than is indi-
cated by the figure of his type specimen of £&. complicatus, and
the specimens described by Cope seem not to differ at all in this
respect from the Z. complicatus type. But even if this lot of teeth
described by Cope are somewhat simpler, a character so variable
as this one seems hardly sufficient grounds for founding a new
species. The specimen figured by Cope (see Cope’s figure *) shows
every indication of belonging to an old individual, which would
account for much of the simplicity of these teeth. Cope at-
tempted no other distinction, nor does there seem to be any. On
the other hand there seem to be good reasons for considering
the two species as identical. There seems to be no difference
in size and practically no difference in tooth pattern; the type
* Trans. Wagner Free Inst. of Sci., EoD p. 38.
2 Proc. Am. Phil. Soc., 1895, Pl. ix, Fig. 8
132 Bulletin American Museum of Natural History. (Vol. XIV,
specimens come from practically the same locality, the type
localities being less than one hundred miles apart; there-
fore as there seems to be absolutely nothing upon which to
separate these species, there is no alternative but to consider
E. eous (E. intermedius Cope) a synonym of £. complicatus, and
the very good characters given by Cope for his species should be
considered additional characters which clearly define the species
£. complicatus.
£. complicatus, as thus understood, seems to be the common
species of Pleistocene times in the United States ; its geographi-
cal distribution extending over a wide area east of the Rocky
_Fig. 22. 2. complicatus. (Tule Cation, Texas.) Superior molar-premolar series of right
side sectioned to show change in teeth due to wear. A, triturating surfaces; 41, sections cut
parallel to the triturating surface about the middle of the crowns. (No. 10611, Coll. A.M.N.H.).
Mountains and south over the Gulf States, and perhaps along the
Atlantic coast as far north as New Jersey; though the specimens
reported from this region cannot be certainly identified, the
material consisting only of single teeth. In the collections from
the Niobrara River there are a great number of jaws, skull frag-
ments and bones, which from their size and general characters
evidently belong to this species. The specimen represented by PI.
XVIII, Fig. B (No. 2725 Am. Mus. Coll.) is typical of the horse
1go1.| Gidley, The North American Species of Equus. 133
of its size from the Niobrara locality. It shows the comparatively
short muzzle and the characters in general pointed out by Cope
in his description of Z. zutermedius. This specimen represents
a young animal with the last molar only just coming into full use,
hence the differences in this region compared with Cope’s
figures. Several symphyses mandibuli containing the incisors,
as well as the lower jaw belonging with the specimen above
figured show the absence or extensive interruption of the internal
wall of the external incisor. Three occiputs in this collection also
show the characters pointed out by Cope for his 4. zntermedius,
and agree very well with the measurements given. A series
of teeth in a portion of the maxillary (Fig. 22) and two fragments
of lower jaws (Nos. 10600, 10601, Coll. Am. Mus.), besides
a few single teeth, represent this species from the Tule Cajion
and Rock Creek, Texas, locality. Leidy reported and figured
some teeth from Hardin Co., Texas, which he referred to this
species. These teeth, now in the American Museum (Nos. 8617-
8618), do not differ in any way from the corresponding teeth
in the specimens from the Staked Plains and Nebraska.
(22) Equus pectinatus (Cofce).
Type Locality.—Port Kennedy, Penn.
fTorizon.—Megalonyx Beds. (Cave deposits).
7ype.—Nine superior molars and premolars probably belonging to one in-
dividual.
Author's description.—In describing some Zguzs teeth from Port Kennedy,
which he considered as represending two varieties of Z. fraternus, Cope gave
the following as distinguishing characters: ‘‘ The superior molar teeth all agree
in the fact that the antero-posterior diameter of the protocone enters the same
‘diameter of the entire crown from two and a third to nearly three times.
In other Plistocene horses of North America, the antero-posterior diameter a
the protocone is from one-half to three-fifths that of the entire crown, excepting
in the premolars of Agus occidentalis Leidy, and in the large 2. crentdens
Cope. . . . One of the individuals differs from the other in the greater
complexity of the enamel plates, especially on the opposed faces of the external
lakes, agreeing in this respect with Egwus complicatus of Leidy. . . . The
superior molars displaying moderate complexity are of smaller size than the
complex ones. . . . Under the circumstances it is necessary to distinguish
two races of Aguus fraternus as already indicated: Z£. f. fraternus and £. f/f.
pectinatus, which may prove to be distinct species.”
34 Bulletin American Museum of Natural History, (Vol, XIV,
Measurements’:
Diameters p* { Antero-posterior.........-.+e+eee00++- 32 mm.
UALTransVeIse ss. weaned ecient 3000 os
, § Antero-posterior........ -+s:++-s+ees aig)
US PransVerses ciocssceoac came neers 2679 ae
Vt mad { Antero-posterior.........--+.+++seseee 26 ve
ARiEIICNeceo Gonabasae Jo wou ODD Oot DOGS py BE
A m2 § Antero-posterior........--+-sesesseeee 27 a
( Pramsvets@us« a.:c'0-0 206 Uelons ene tee QT Be ins
iC ee) Antero-posterior.......- Romenmoncs ees She | oS
(transverse: socio ours ai eee ieee 25 us
The Port Kennedy specimens seem to represent two distinct
species as suggested by Cope; however, it seems highly improb-
able that either bears any close relationship with the South Caro-
lina species, 2. fraternus, which is one of the smallest American
species, while Z. fectinatus is large, comparing in size with Z.
complicatus or £. scotti. The teeth of the type specimen are
much worn, yet they show an unusual degree of complication in
the enamel foldings (see Fig.23 ). As pointed out by Cope the
protocones of the type specimen are unusually small. The skull
characters are not known and until they can be made out this
species cannot be clearly defined. Leidy described and figured
some teeth from Illinois? (see Fig. 24) which he referred to Z.
complicatus, but which seem to agree more closely with ‘4.
pectinatus.
The other Port Kennedy teeth described by Cope and identi-
fied with £. f. fraternus seem not to differ essentially from £Z.
complicatus although they are somewhat smaller.
(25) Equus scotti Gidley.
Type Locality.—Rock Creek, Brisco Co., Texas.
FTorvizon.—Sheridan Beds,
Type.—Part of a skeleton (No. 10,606), including the skull and lower jaws,
all the cervical and the three anterior dorsal vertebrz, both fore limbs and feet
complete, one hind limb and foot complete, and several dorsal and lumbar
vertebrze and ribs probably belonging with it.
Author's description.—Compared with £. caballus: (1) The skull is rela-
tively larger, (2) the teeth are relatively larger compared with the skull, (3) the
neck is shorter, (4) the body is longer, (5) the limbs are shorter and more
1 The measurements given by Cope include cement, hence the writer has carefully remeas~
ured the teeth and substitutes here these measurements for those given by Cope.
s Trans. Wagner Free Inst., Phil. Vol. II, 1889, p. 30.
—s
Gidley, The North American Species of Equus, sy
1901. |
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&
136 Bulletin American Museum of Natural History, [Vol. XIV,
slender than in the larger varieties of Z. caba//us, (6) the lesser curvature of the
ribs near their heads indicates that the back was not nearly so wide, but shaped
more like that of the ass or zebra. (For description of skull characters see
American Museum Bulletin, r900, Vol. XIII, Art. XIII, pp. 114-116.)
Measurements given,
. Antero-posterior.........-.. mm.
Diameters of crown of p# 5 P 33 ie
( Transverses.ia.asle meer eee 31
: Amtero=poSterlOranremceeiette DN
Diameters of crown of m+ S P 3 4
( Pransverse cna cette 30
Wong diameter Of i ISO carole ela) leleral ole trelel elie on Pele thers Pyeye eek
Potal lenethyor miolar-prem olar Series= leit -1-l-ie i isketltetnenete TOC Mme
The type skull as well as the four other associated skulls in
hand when these measurements were given are all of young in-
dividuals that had not yet shed all their milk molars. Very
fortunately the writer has since collected a skull (No. 10628)
from the same deposit or quarry from which the others were taken,
in which all the teeth had come into full use (see Pl. XIX, Fig. 4),
and the measurements can now be given more fully and more
accurately. They are as follows:
Diameters ne {7Atntero-posterionnss-- ee hereeeeeeeee 43 mm.
MEGAN SVEESE viscose aloo rors ae Byoicy we
Diameters p2 NeAmitero-postenioneec ah leilemeee eter SA a as
WEVanSVGPrSer., wautalc ence ee ee CRO Gey RS
Wien ps4 NMS YOMANO do acoogogsneoncogecs ew Me
MGrANSV ELSE) nase see seats yarn tee BB ies
Dinmetacant \PAtIteTro-pOStenl Gree eit ieee SOnaa
g~ (Uitvansverse) o/s2sis.. =, eee BOl nes
Diameters 2 j) > tere-posterior: 0 .), Fee eee Sie ee
{GraANSVEIse acu cameo ere Oe
Dianetercne {, Antero-posterior;. ....o. <eaeraee eee Bee eis
ALTANSV.ELS Cs wieienvesciente eieciete ere Gy
Long dianteter oti ©. i 55, ..ccimals ots ee ee ee je DA,
Total length of molar-premolat series..............4- 204
This species, which is intermediate in size between Z.. compli-
catus and £. pactficus, is distinguished from the former, especially,
by its comparatively longer muzzle, which equals in comparative
length that of Z. cadallus. From £. pacificus it is distinguished
by the comparatively smaller skeleton, the somewhat longer
muzzle, and by the different form of the occiput; that of Z&.
——
;
'
tgo1.] Gidley, The North American Species of Equus. 137
pacificus being much more like Z. caballus. LE. complicatus also.
resembles £. caba/lus in this respect.
Sol
Fig. 26. Occiputs of—A, Eguus scotti; B, Equus caballus.
Equus giganteus, sp. nov.
Type Locaity.—Southwest Texas.
Type.—Superior molar (m2). (No. 8616, Coll. Am. Mus.). This tooth was.
referred by Professor Cope to Z. crenidens.
Description.—T here is a very large tooth, now in the American
Museum (Cope) collection, from southwest Texas which Cope
described and figured, referring it to 2. crenidens, but which
differs in every respect, except the small size of the protocone,
ZF
A.
Fig. 27. Lguus giganteus, sp. noy. Type. Superior second molar m? (No. 8616, Coll.
A.M.N.H.). A, triturating surface; A1, section cut about three-fourths of an inch
below.
from the description of the Mexican tooth upon which this species.
was founded. Its very much larger size; the lack of the charac-
138 Bulletin American Museum of Natural History, [Vol. Ae
ter given by Cope as the one by which &. crenidens could be dis-
tinguished, that of the crimping of the enamel of the lakes; the
comparatively complex folding of the enamel, all prevent the
identification of this tooth with that species. Its grinding-surface
is much weathered and the enamel foldings of the lake borders
are so obliterated that they cannot be made out and appear to be
very simple. This evidently led Cope to the belief, as shown in
his figure and description, that the tooth pattern was very simple.
A section (Fig. 27 A’) made about three-fourths of an inch below
the grinding-surface shows very plainly the true character of the
enamel foldings.
Although in this tooth, as seems characteristic of the genus,
there seem to be no characters which can be said to be of
definite specific value, its great size, which exceeds by at least 5
mm. any other Zguus tooth of which record has been made, and
the relatively small area of the cement lakes or fossettes mark it
as evidently distinct from any species hitherto described; hence
a new name seems necessary. ‘The size suggests the name Z.
giganteus.
The measurements are as follows:
Diameter of srinding-surface { Antero-posterior...... 41.5 mm,
( Transverse.........- 36 a
Diameter where section ( Antero-posterior.......... Moy
was made U Pransverse...0. ce See Bi)
Geno (hvoficrowieyemerervarnieers acer icteric ee TOO Sens
The tooth had evidently been only slightly worn by use, hence
the difference in transverse diameter when the cut was made.
The length of the tooth compared with its other measurements
shows a comparatively short crown.
SUMMARY AND CONCLUSIONS.
The general results of this systematic comparison may be
summed up as follows:
£. curvidens.—A South American horse, indeterminate. North
American specimens referred to this species erroneously.
£. major.—Absolutely indeterminate; a nomen nudum.
£. americanus.—A preoccupied name, replaced by &. complicatus.
E, FRATERNUS.—Southeastern United States, probably a valid
species, but still imperfectly known. It represents a very
tgor.] Gidley, The North American Species of Equus. 139
E.
&
EK.
E.
small horse with teeth scarcely as large as those of the Mexi-
can donkey and of very complex pattern.
COMPLICATUS.—West Southern and middle Western States;
embraces as synonyms, &. ixtermedius, E eows and probably
E. excelsus (which is now indeterminate). Now well charac-
terized, representing a species with teeth about the size of
those of the ordinary draught horse and of moderately com-
plex pattern, but with the bones of the skeleton about the size
of those of the smaller varieties of the western pony. The
species is especially characterized by its short muzzle, in this
respect resembling the ass.
. excelsus,—Nebraska. Very imperfectly known from the teeth
only ; although now indeterminate, may prove to be a valid
species.
. OCCIDENTALIS.—California or Pacific slope. Other species
and specimens have been mistakenly referred to this type.
The characters other than those of the teeth are not now
known, but these indicate a horse about the size of &. com-
plicatus with uniformly simple patterned teeth.
. PACIFICUS.—A large Oregon species, although middle Cali-
fornia is the type locality. A well characterized species.
With the exception of /. giganteus from southwestern Texas,
the largest American horse. The skeleton indicates a horse
about the size of the ordinary draught horse, but the skull is
proportionately larger.
. CONVERSIDENS. Valley of Mexico. Apparently well estab-
lished; of medium size.
. TAU.—Valley of New Mexico, the smallest true horse known
in America, This embraces the Z. darcenei as a synonym.
. crenidens.—Valley of Mexico; indeterminate.
. SEMIPLICATUS.—Western Texas, from the Sheridan beds of
Paloduro Cafion, probably a valid species. The great pos-
terior extension of the vomer over the presphenoid, together
with its size and the proportions of the teeth, seem to show
a close relationship to £&. asinus.
PECTINATUS.—Port Kennedy Bone Cave of eastern Pennsyl-
vania. Probably a valid species.
scoTtTi.—Staked Plains of Texas, Tule Cation. Intermediate
in size between &. complicatus and £. pacificus. A long-faced
type of horse. This species represents a horse about the
140 Bulletin American Museum of Natural flistory, [Vol. XIV,
size of the largest western pony, but with a longer body, a
much larger head, a shorter neck, and a back and steeply
sloping sides shaped very much as in the ass or quagga.
E. GIGANTEUS.—Southwestern Texas. The largest species of
horse hitherto recorded, the teeth exceeding those of the
largest modern draught horses by more than one-third the
diameter of the latter.
Pliocene species wrongly referred to Equus.
. simplicidens = Pliohippus sp. ?
. eurystylus = Hipparion sp. ?
. cumminsii == Protohippus sp. ?
. minutus = Protohippus sp. ?
. phlegon = Protohippus sp. ?
comes imeomico ies)
There are a number of teeth in the Niobrara River collection
which seem to be intermediate between £. complicatus and £.
fraternus, yet they seem not to show characters sufficiently
definite upon which to found a new species. This, however,
may be done when better material from the Nebraska locality has
been found.
BIBLIOGRAPHY.
Corer, E. D:
"84. The Extinct Mammalia of the Valley of Mexico.
(Paleont. Bull. No. 391, 884.) Proc. Am. Phil. Soc., Vol.
XXII (1884), 1885, pp. 1-15.
85. Phocene Horses of Southwest Texas. Am. Naturalist,
Viol, XxX, 18359) p: 1206, (pl. Xxxvil.
’92. Report on Paleontology of the Vertebrata. Zhird Ann.
kept. Geol. Surv. Texas (1891), 1892, pp. 251-252.
‘92. A Contribution to the Vertebrate Paleontology of
Texas. Proc. Am. Phil Soc., Vol. XXX, 1892, pp. 123-125.
93. A Preliminary Report on the Vertebrate Paleontology
of the Llano Estacado. Fourth Ann. Rep. Geol. Surv. Texas,
1893, Pp. 1-137.
‘95. On Some Pleistocene Mammalia from Petite Anse, La,
Proc. Am. Phil. Soc., Vol. XXXIV,"1895, pp. 458-468, pll.
xi and xil.
1g01.]| Gidley, The North American Species of Equus. 141
99. Vertebrate Remains from Port Kennedy Bone Deposit.
Fourn. Acad. Nat. Sct. Phila., Vol. XI, 1899, pp. 193-289.
DE Kay, J. E.
"42. Zodlogy of New York. Nat. Hist. N. Y., Zoél., Part
iG’ (1d42)7 pe tos:
GIDLEY, J. W.
tg00o. A New Species of Pleistocene Horse from the Staked
Plains of Texas. Bull. Am. Mus. Nat. Hist., Vol. XIII,
1900, pp. 111-116.
Hay, O. P:
‘99. On the Nomenclature of Certain American Fossil
Vertebrates. Am. Geologist, Vol. XXIV, 1899, p. 345.
*99. On the Names of Certain North American Fossil Verte-
prates. “Science, N. er., Vol, EX, 1899, p. 503:
LEIDY, JOSEPH.
"47. On the Fossil Horses of America. Proc. Acad. Nat.
en Lia Vole tiles 7. p. 265. pl. ii,
"58. Notice of Remains of Extinct Vertebrata, from the
Valley of the Niobrara River. Proc. Acad. Nat. Sct. Phila.,
Vol. X, 1858, pps11 and 20-26.
*59. Description of Vertebrate Fossils. Post-Pleiocene
Fossils of S. C. (Holmes), Nos. 8, 9, and 10, 1859, pp. 99-
E10; pli. xv and: xvi.
"65. Remarks on Fossil Remains of Horses. Proc. Acad.
Nat. Sct. Phila., Vol. XVII, 1865, p. 94.
*69. Extinct Mammalian Fauna of Dakota and Nebraska,
F ournal Acad. Nat. Sci. Phila., 1869, pp. 257-331 and 399-
4ol, pll. xvii—xxi.
- ?73. Contributions to the Extinct Vertebrate Fauna of the
Western Territories. Report U. S. Geol. Survey of the Ter-
ritortes (F. V. Hayden), Vol. I, Fossil Vertebrates, 1873,
pp. 242-246 and 321-332, pl. xxxiil.
"89. Notice of Some Mammalian Remains from the Salt
Mine of Petite Anse, Louisiana. Zvrans. Wag. Free Institute,
Phila., Vol. II, 1889, pp. 33-40.
MITCHELL, SAMUEL L.
’26. Catalogue Organic Remains, New York, 1826, pp. 7, 8.
142 Bulletin American Museum of Natural History. [Vol. XIV.
Owen, RICHARD. é
’4o. Notice of the Remains of a Species of Equus. Voyage
H. M. S. Beagle, Fossil Mammalia, 1840, pp. 108, 109, pl.
xx
"69. On Fossil Remains of Equines from Central and
South America. P%i/. Trans., London, 1869, pp. 559-573,
pile axa, Toxin.
ISP :
£3):
ay
\ 2079
q *
an
otti; B, Equus complicatus.
A, Equus sc
Vou. XIV, Pu. XIX.
in A. M.N. H.
, TEXAS PONY.
aballus
aballus, DRAUGHT HoRsE; D, Equus ¢
C, Equus c
. a, 7 ,
Sienna Heit eg
ne -
‘
‘SNOLAIAMS OM], JO SLUVd ATION] ‘WAUSAJW NVOINANY AHL NI NOLATANS GHLNAOW ‘(ddA L r) yors snnby
‘HUN ‘WW NI
‘XX “Td ‘ATX "10A
“STVACIAIGNT OM] dO GasOdWOD ATAvdONg ‘(AMAL) NAMO sSuapisvaaucs snnbz
"SIVAGIAIGN] OM] JO GasoOdWOD ATAVHOUd (AaA.L) NAMO sSuapiszzauos snnbz
i all
=
‘HN WY Nicaring
‘IXX “Id “ATX “TOA
:
n Furassic. Stratigraphy in South-
2 eastern Wy Yoming. |
s AUTHOR’S EDITION, extracted from BULLETIN
4 —
_OF THE
: VoL. XIV, ARTICLE XII, pp. 189-197.
: New York, Fune 17, 1901.
~
‘Zhe Tknickerbocker Peces, Rew Work
Article XII.—ON JURASSIC STRATIGRAPHY IN
SOUTHEASTERN WYOMING.
By F. B. Loomis.
PLATES XXVI AND XXVII.
The purpose of this paper is to give some details in regard to
the Jurassic strata of Wyoming, in which, for the last four years,
the American Museum of Natural History has collected Dinosaur
remains. Very little detail geological work has been done in the
region. Prof. W. C. Knight published a map of southeastern
Wyoming showing the Jurassic exposures'; and this is in part
reproduced, with his consent, in Plate XX VI, Fig. 1, to show the
location of the area in which the succeeding maps and sections are
located. Prof. W. N. Logan” has also published a section: made
in the Freezeout Hills. The area studied, lying along the
county line between Albany and Carbon Counties, includes all
-of the quarries worked for the American Museum; and, inciden-
tally, a considerable number of Marsh’s localities, where they
were within the area mapped, are indicated.
Referring to Plate XXVI, Fig. 1, the axis of the Laramie
Mountains is seen in the eastern part of Albany County, extend-
ing nearly north and south; but in Converse County it bends
until it extends nearly west. In the angle thus formed are
situated four short (8-20 miles in length), sharp folds, known
as the Medicine, Como, Prager, and Miser anticlines. Their
axes lie northeast and southwest, and each fold is thrust over be-
yond the vertical on its northern side. These folds are radial
folds formed at the edge of the territory affected when the
Freezeout Hills were thrown up. While the Laramie Mountains
seem to be due to a thrust from the east, there must have been a
force acting from the north in the neighborhood of the Freezeout
Hills. This thrust from the north seems to have affected locally
the axis of the Laramie Mountains, bending it to an east and
west direction. The four short anticlines above named die out
on the eastern boundary of the territory affected by the thrust
from the north. How far west this thrust from the north, to
which the Freezeout Hills are due, acted I cannot say.
It is along the southern exposures of Medicine and Como anti-
? Bull. Geol. Soc. America, Vol. XI, p. 377, 1900.
? Kan. Univ. Quart., Vol. LX, p. 109, 1900.
[189]
190 Bulletin American Museum of Natural History, [Vol. XIV,
clines that the American Museum parties have worked, where the
dip is 14-18°. The north sides of these folds stand too nearly
vertical to be worked. Plate XXVI, Fig. 3, shows a section re-
constructed to show the appearance of these folds had no erosion
taken place. The position of this section is shown on Plate
XXVI, Fig. 2, by a line from H to L running N. 25° W. The
line starts on the Little Medicine River, about one and one half
miles below the mouth of Sheep Creek. For ten rods on either
side of the river all rocks are covered by river deposits. At the
locality c, a collection of fossils’ showed the presence of the
Cretaceous (Fort Pierre); while within three rods to the south
undoubted basal beds of the Jurassic * are found. A fault is
thus indicated. While in the field I did not trace it out, so can-
not speak in regard to its extent. The Jurassic is here inverted
so that its basal member is uppermost. Between the Jurassic and
the typical Triassic occurs a bed of creamy white sandstone, 40
feet thick, the same as is found in the Freezeout Hills, but this
sandstone is lacking on the exposures situated further south.
The Jurassic dips 60° S. (strike N. 53° E.), but proceeding south
the dip quickly becomes vertical, changes to the north, and
gradually diminishes till in the midst of the Triassic it becomes
horizontal. The strata soon dip to the south increasing to about
14°, at which angle most of the rocks, as far as the Fort Pierre
bed, dip.
At G the section changes its course to N. 5° W., crossing
the Jurassic of the south side of the Medicine anticline. The
Jurassic beds are here 274 feet in thickness and are overlaid by
the coarse sandstones of the Dakota to the extent of 275 feet.
Above these in turn lie the Fort Benton shales, the lower 80 feet
of which consist of dense sandy limestone; while the upper 320
feet are thin bedded argillaceous shales, characterized by abun-
dance of teleost fish scales. Above the Fort Benton are about 800
feet of slate-colored clay in which the stratification is obscure.
On account of their soft texture these clays are easily eroded and
the exposure is usually occupied by a long ‘ draw.’ On weather-
ing the clay turns black. As this clay overlies the Fort Benton
* The collection included Jnoceramus, Gryphea, Lucina subundata H. and M., Tedlina,
cf. scztuda M. and H., Anisomyon shumardi W., A. Patella formis W., and Scaphites war-
vent M. and H.,
? The Jurassic layers carried Belesznztes densus M. and B, curtus L.
1901. ] Loomis, Furasste Stratigraphy tn Wyoming. Ig!
and underlies the Fort Pierre, I consider it the representative of
the Niobrara, although it is non-fossiliferous. A bed of gray sand-
stone comes next, making a prairie about three miles wide, so
that to estimate its thickness is difficult, though there can scarcely
be less than 300 feet. The sandstone furnished several badly
preserved fossils among which were /noceramus, Gryphe, and
Scaphites warrent M. & H. While the Ammonites are described
as Fort Benton, inasmuch as I found the same at locality c mixed
with the Fort Pierre Gasteropoda and Lamellibranchiata, and
being strongly influenced by the stratigraphy, I am inclined to con-
sider the sandstone bed Fort Pierre. Across the sandstone the
Niobrara clay reappears, on its northern boundary nearly horizon-
tal, but on the southern side overthrust, so that the Fort Benton
lies inverted over the Niobrara. The Fort Benton dips 60° to the
south (strike N. 48° E.). The change to the vertical occurs in
the Dakota, which at this exposure (Carleton ridge) is only about
one-half its usual thickness. The Jurassic dips 80° N. but as
the Triassic is crossed the dip decreases to horizontal, changes
to south, and gradually increases to about 17° S., at which angle
most of the rocks, as far as the Prager anticline, dip. The
section goes only to Como Bluff but the rocks at Prager anticline
again show overthrust characteristics.
Three sections of the Jurassic are shown in Plate XXVII,
Figs. 1, 2, and 3, giving the details of the series of rocks on the
south side of Medicine anticline, and the north and south sides
of the Como anticline. These sections are also tabulated on
page 192 so that each layer can be traced in its variations across
the two anticlines.
The base of the Jurassic I have located at the beginning of the
series of clays. The Triassic is a great bed of sandstones, usually
alternating red and gray. The first change from this to Jurassic
is a layer of brown clay (No. 2), always cf considerable thickness,
and carrying Belemnites densus M. and B. curtus L. in such abun-
dance that the ground is in many places so strewn with the shells
that one can scarcely step without treading on one. The layer is
widespread, occurring on Sheep Creek,’ Medicine and Como anti-
clines, and is identical with Knight’s No. 14,* and the lower part
of Logan’s No.13.* It is usually designated as the Belemnite layer.
* The Sheep Creek exposures are 12 miles northeast of Bone Cabin Quarries.
2 Bull. Geol. Soc. Amer., Vol. XI, p. 382.
3 Kan. Univ. Quart., Vol. IX, p. rrz2.
192 Bulletin American Museum of Natural History. (Vol. X1V,
PLATE XXVII, FIG. 3. PLATE X£XVIT, BIG, I.) |) PLAGE xox Vile maGa2:
Bed. Section A-B. Thickness.) Section C-D. Thickness.| Section E-F. Thickness.
Feet. Dakota. Feet. Feet
No. 33 Yellowish sand- Straw yellow Sandstones yar 200+
S{ONeG rete ere 249 sandstone..... 120+
‘* 32 Black sandstone 2 /Black sandstone. 3 |Black and _ red
sandstone..... 4
‘© 31 Gray sandstone. 30 | Yellow sandstone 12 |Straw yellowsand-
Stonens tistics 20
Furassic.
‘© 309 Bluish greenclay 21 |Bluish greenclay. 20 |Maroonclay..... I0
Bluish green clay. 15
Sou 2o Mm Greenlclayeryerte 20 |Green clay..... . 40 |Yellow greenclay 13
Bluish green clay, 15
OO “eXero Jali ooo eco WML) vrei amteniaves 4
‘* 28 Gray sandstone. 10 Sandy clay...... 5
sto7 (WoncretionsS .... 2
poonGreensclaya act 3 (Green! clay... tS (Greenclay. rrr meee
‘* 25 Concretions.... 14;Concretions..... 2) |\Coneretions!..7.- I
‘* 24a Green clay..... 8) Greenuclayarreree 9 |Green clay...... a UG
So VE Sandstone....... 4
oO OMe Green) elayn. «00 10
‘© 23 Green clay with Green clay with Green clay with
small concre- small _concre- small concre-
TONS seer aa KG) TIONS! siete Haas bee HONSIA-c eet eave 25
‘© 22 Maroon clay with Maroon clay with Maroon clay with
small concre- small _concre- small concre-
EONS Heal BO) | UtIONS yanertetee AG As HONS! Fass. 20
°8 Oe SENG 9.0 cine 3
So2On Green clayey Sy \Gneenuclayaneriee 20 |Green clay...... 9
*) Sig wandstone’s.. 4) 0h || Sandstonesse one 2
of ids) IRecl GEN co ncaas 5 IRGC ES Aa ace tac 5
OO Fi) SENN Ea o aa} 2
Seton Grecnaclayjejerere: 6) |\Greeny clay? .- g |Maroon clay..... 7
SoS) Sanastonesyatrs. 14)Sandstone....... 2
** 14 Red, green, ma- Green, maroon, RR educl aye mutase 8
ROWE Cocos Me || mal eeieccaouc 26
‘‘ 13 Sandstone...... 2 |Sandstone....... 14 |Gray sandstone.. 28
‘* 72 Maroon, green, Green clay........ 20) |Darklcreen! claynus1O
TEuClayenare 1 LO)
“Tr Sandstone... 1-0 2 |Sandstone......< 2 |Sandstone..... 14
PTO MGreenecl ayer 22)\(Green\clay mnie 60 |Redandgreenclay 10
“© 9 Sandstone ....:. “I |Sandstone...t...! -“Fe|Sanestanees srr - 2
Src (Greenyclayenrrte 20 |Maroon clay. ... 20 |Green clay...... 25
do weddishiclayacms2o
-) JO) Sandstonesceree 1}/Sandstone....... 14 |Sandstone....... 12
‘* 5 Green sand shale 6
‘“ 4 Purple clay with Purple clay with Purple clay with
limestones.... 22] limestones.... 20 limestones.... 15
‘* 3 Nuculalimestone 1
+>) 2) Brown/clay 2... 43 |Brown clay with Gray brown clay. 55
limestone beds. 70
Motallofa\umassics2;7745 | ees series Bi7 BB lisrche chal Acteutoxerevoe . 3324
‘“ 1 Triassic sandstones. | Triassic sandstones. Triassic sandstones.
1901. ] Loomts, Furasste Stratigraphy in Wyoming. 193
On the surface of this, in Bone Cabin Draw, is found a one-foot
bed of limestone (No. 3) made up mostly of Muwculas,' with
occasionally a Zancredia and Ostrea. The band is not continu- °
ous, but similar layers at the same horizon occur both on Sheep
Creek and in the Freezeout Hills.
No. 4 is a purplish clay with large limestone nodules scattered
through it. It is in or on these nodules that the specimens of
Baptanodon are found. That of the American Museum was
found at locality 4, at the foot of Bone Cabin Draw. It seems
best to confine Marsh’s* term, Baptanodon beds, to this layer
instead of designating the whole lower Jurassic by that term.
The bed is universally present in neighboring localities; and is
Knight’s 12 and the upper part of Logan’s 13.
No. 5, green sand shale, is local.
No 6, sandstone. This first sandstone bed seems to be widely
distributed, and marks the beginning of a series of alternations
between sandstone and clay. The clays are variously and
brightly colored and often designated as the ‘ variegated clays.’
The layers from 6 to 20 inclusive form a series of variegated beds
in which there may be eight sandstone layers or only three. Very
few of the clays seem to form wide horizons, Inside of a quar-
ter of a mile I have seen three of the sandstone beds unite into
one thick sandstone, the intervening clays being pinched out.
These indicate a period of shallow water in which there were con-
tinually changing currents, so that deposits from any given source
are laid down first in one place, later in another. In the Bone
Cabin section there are 120 feet of these variegated clays, and
it is somewhere in these layers that the change from marine to
brackish (or freshwater) deposits occurs. In the Como section
there is a 28 foot bed of sandstone, corresponding apparently
with No. 13, in which a Dinosaur quarry (12) is located. This
bed seems to represent the beginning of the freshwater (or brack-
ish) series, and as such is the first member of the Como stage
(Atlantosaurus beds). This sandstone (No. 13) is variable in
+ At the foot of Bone Cabin Draw, locality 4, were found in this layer Mucuda nov. sp.,
Tancredia tnornata, Avicula mucronata, and Ostrea strigilecula. A similar band near
Sheep Creek furnished all of the above with also Cardioceras cordiformis M., and Pseudo-
eta curta ; see also Logan’s No. 13, which furnished some of these and several other
orms.
2 Sixteenth Ann. Rep. U. S. Geol. Surv., 1896, p. 145.
[ Fane, rg07.] 13
194 Bulletin American Museum of Natural History, [Vol. XIV,
thickness but there is no unconformity such as Logan seems to
have found in the Freezeout Hills. é
No. 22 is a bed of maroon colored clay with small limestone
concretions scattered through it, and is a most excellent horizon
indicator, for it is everywhere present, and its brilliant color
makes it a conspicuous feature for miles. At Bone Cabin it is
but ro feet thick but increases to 30 feet in the Como Bluff.
The considerable amount of concretionary limestone would seem
to indicate brackish water.
The maroon clay graduates into a bed of green clay with
similar small limestone concretions (No. 23). This layer like the
maroon bed is widespread and thicker as one goes south.
The green clay with concretions is invariably covered with
smooth green clay (No. 24). In the Como Bluff this layer has
sandwiched into it a 4-foot bed of sandstone (244). The sand-
stone is of interest as marking the horizon at which the few known
Jurassic mammals were found. The mammal layer is the 6
inches of clay immediately underlying this sandstone. Most of
the American Jurassic mammal remains thus far found have
come from one quarry,’ worked most successfully by Marsh and
later by the American Museum. This pocket seems to be
exhausted.
Above 24 is a constant layer of large nodular limestone con-
cretions (No. 25). Single concretions may occur anywhere in
the clay but this layer and No. 27 are definite layers of almost
uninterrupted limestone nodules. The concretion layers are of
considerable importance as, when weathered out, they make a
prominent line on the bluffs just above which the most important
bone bed occurs. In this limestone layer Logan found five
species of fresh or brackish water mollusks.’
No. 27 is a green clay of varying thickness in which the most
numerous quarries are located. On weathering, or having been
dried out, this clay breaks into angular fragments and is there-
fore known as ‘joint clay.’ The ‘Nine Mile Quarry,’ the
‘Aurora 1900 Quarry,’ the ‘ Diplodocus 1897 Quarry,’ and sev-
eral of Marsh’s quarries are in this layer. Most of the fossils
are Diplodocus and Brontosaurus species.
* See Pl. XXVI, the S.E. corner of the map.
= Unga knight? L., U. willistont L., U. baileyi L., Valvata leet L., and Planorbis veter-
MUS L,
1901. | Loomis, Furasstc Stratigraphy in Wyoming. 195
In Bone Cabin Draw this green clay is overlaid by a second
layer (No. 27) of concretions, so that the ‘ Nine Mile Quarry’ is
between two such layers.
No. 28 is a gray sandstone in which the rich Bone Cabin
Quarry is situated, and also the Stegosaurus Quarry. The sand-
stone varies extremely in hardness, being, in the south part of
Bone Cabin Quarry, soft and mixed with considerable clay so that
it is workable with an awl. In the northern part of the quarry,
however, there are bands of the firmest sort of sandstone. In
Como Bluff the layer is clay with merely an admixture of sand.
Bone Cabin Quarry has yielded a great variety of genera: Dip/o-
docus, Morosaurus, Brontosaurus, Allosaurus, Ceratosaurus, Camp-
tosaurus, Stegosaurus, as well as several genera of carnivorous
Dinosaurs; also Compsemys and Gontopholts.
On the surface of this sandstone is usually a 4-inch layer of
flint, No. 28a.
No. 29 is another green clay showing the ‘joint’ structure
similar to No. 26. Several of Marsh's quarries are in this layer.
The series ends with a soft bluish green clay in which I have
never seen fossils. This completes the Jura.
A cap of Dakota sandstone overlies the Jura, near the base of
which is a narrow (2-6 feet) band of soft black, or occasionally
red, sandstone. This band is colored by iron and of interest as
having lately yielded a collection of bones probably crocodilian.
There is some difficulty in comparing the foregoing sections
with those of Logan and Knight, as these authors give fewer
beds. Marsh divided the Jura into three horizons, Hallopus,
Baptanodon, and Atlantosaurus beds. The Hallopus beds do
not occur in southeastern Wyoming. The Baptanodon beds,
according to Marsh, included all of the lower Jurassic, which
has since been subdivided, but the term Baptanodon is now ap-
plied to a single layer (No. 4) in which these reptiles occur.
When this term is restricted to a single layer, the lower strata are
without designation; so for these, Knight has proposed the term
Shirley stage, which shall include all the marine Jura; that is,
beds 2-12 inclusive. The exact boundary between marine and
non-marine is difficult to locate, but bed 13 has furnished some
Dinosaur remains and may therefore be associated with the beds
in which Dinosaurs are abundant. ‘Then beds 13-30, inclusive,
196 Bulletin American Museum of Natural History, |Vol. XIV,
would form the non-marine upper Jura. There are no inverte-
brate forms of decisive character in these beds to determine
between brackish and fresh water, but those found in bed 24
would incline toward fresh water. The change is clearly a
gradual one. These upper beds Marsh calls Atlantosaurus
beds, but as that genus has proved to be a synonym with Cama-
rosaurus, a less misleading term is desirable; and Scott’ has
introduced the name Como stage for exactly these beds. He
suggests that they belong to the Lower Cretaceous; and Logan
also uses the term Atlantosaurus beds as distinct from: Jurassic.
There is no available invertebrate material in these upper beds
for comparisons; but the strikingly close relationships between
the British Purbeck mammalian fauna and the Wyoming mam-
mals cannot be left out of consideration. The considerable
number of British and Wyoming genera which are closely related
will compel us to consider the Como stage of nearly if not just
the same age’ as the Purbeck and retain the Como in the Jurassic
series. It is true there is no apparent unconformity between the
Como and Dakota, as would be expected if this Como stage were
dry land for a period as long as the Lower Cretaceous.
During the Shirley period, the deposits on the Medicine and
Como anticlines were being laid down quite close to shore, as ap-
pears from the fact that within 30 miles to the south the Shirley
is unrepresented, and all of the deposits are strictly shallow water
sediments. However, during and especially at the beginning of
the Como stage, a considerable transgression took place (see
Knight’s map) toward the south, removing the shore line to over
too miles south. The deposits are also of shallow water.
The Dinosaur remains could, however, have travelled such
considerable distances by floating. There could have been no
currents in the Como lake strong enough to transport gigantic
bones, for they were depositing clay. The bones are clearly
floated out to sea by the presence of considerable meat on them.
Most specimens must have started as complete carcasses, which
with decay of the flesh (or its consumption by crocodiles and
fish) have fallen apart, often making series of vertebre, etc.
Doubtless such quarries as the Bone Cabin Quarry mark an eddy,
as there all sorts and sizes of animals are collected together.
1 Ww. Be Scott, Introduction to Geology, p. 4
2 Osborn, Jour. Acad. Nat. Sci. Phil. (2) 1 p. 187, 1888.
gor. | Loomis, Furassic Stratigraphy in Wyoming. 197
In summary it may be said that the Dinosaur remains are
mostly confined to beds from 26 to 29, although occasionally
remains are found lower. The bones are usually nearly perfect,
though in some localities pieces of the more delicate spines are
broken away. The clay usually preserves bones much better
than the sandstone.
EXPLANATION OF PLATE XVE
Fig. 1.—Map of southeastern Wyoming, showing exposures of Jurassic.
The shaded area represents the accompanying map, Fig. 2. This map is a
reproduction of Knight’s (see page 189).
Fig. 2.—Map as indicated on Figure 1, showing position of quarries and
fossil localities on Medicine and Como anticlines; also positions of sections
made through the Jurassic, as follows :
A-B, section of Jurassic on Plate XXVII, Fig. 3.
C-D, section of Jurassic on Plate XXVII, Fig. 1.
E-F, section of Jurassic on Plate XXVII, Fig. 2.
L-H, section across Medicine and Como anticlines, Plate XX VI, Fig. 3.
16, quarry of the American Museum (Dinosaur).
13, Marsh’s Stegosaurus quarry.
12, quarry in bed No, 13, author unknown.
10, II, 14, 15, quarries made by Marsh,
* Quarries of American Museum.
Fig. 3.—Ideal section across Medicine and Como anticlines, as the strata
would have appeared had no erosion taken place.
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EXPLANATION OF PLATE. XXVII.
Fig. 1.—Jurassic section across Carleton ridge, north side of Como anticline ;
C-D on Plate XXVI, Fig. 2.
Fig. 2.—Jurassic section across Como Bluff, south side of Como anticline ;
E-F on Pl. XXVI, Fig. 2.
Fig. 3.—Jurassic section across Bone Cabin Draw, south side of Medicine
anticline ; A-B on PI. XXVI, Fig. 2.
A
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pe Bone Cabin Ouarry.
~ By Henry F. Osporn and Watter GRANGER.
-
AUTHOR'S EDITION, extracted from BULLETIN
OF THE
American Pusenm of Uatural History,
Vou. XIV, Articte XIII, pp. 199-208.
New York, Fuly 9, IQOL.
Article XIII. — FORE AND HIND LIMBS OF SAUROP-
ODA FROM THE BONE CABIN QUARRY.
DINOSAUR CONTRIBUTION No. 6,
By Henry F. Osporn and WALTER GRANGER.
During the early months of 1goo the large collection of limb
bones from the Bone Cabin Quarry was measured and compared
by Mr. Granger in preparation for this paper. Measurements
are chiefly of value in determining proportions ; size, of course,
constantly increasing with age. Further studies during the present
year enable us to establish the following points :
1. The proportions and relations of the radius and ulna in
the Sauropoda are remotely analogous or “parallel with those of
the Proboscidia, owing to the marked extension of the ulna, the
similar weight, and the perfected quadrupedal progression in the
two types.
2. The chief characters of the fore and hind limbs and of the
upper part of the manus of Déplodocus,
The conclusion reached provisionally by Osborn,* that Diplo-
docus was a distinctively long-limbed type, is abundantly con-
firmed. The bones of the fore and hind limbs of Dzplodocus can,
in fact, be readily identified by their relative length and slen-
derness as compared with those of Morosaurus and Brontosaurus
(see Fig. 6).
3. The carpus in the Sauropoda includes the probable coésifi-
cation of the radiale and intermedium into a “ scapho-lunar”’ ;
also a large ulnare, and from two to three small osseous carpalia
in the distal row.
In his numerous and valuable contributions to this group the
late Professor Marsh left the homologies of the carpals undeter-
mined ; his figures (of the Worosaurus and Brontosaurus fore limb)
do not indicate the anterior crossing of the radius and ulna; he
also left the limb structure of Déplodocus practically unknown.
In general, the limbs of the three contemporary Sauropoda of
the Como district can be distinguished as follows :
1 Fore and Hind Limbs of Carnivorous and Herbivorous Dinosaurs from the Jurassic of
Wyoming. Bull. Amer. Mus. Nat. Hist., Vol. XII, 1899, pp. 161-172.
[199]
200 Bulletin American Museum of Natural History, [Vol. XIV,
Morosaurus, scapula relatively short, spreading superiorly ; scapulo-coracoid
plate relatively broad; limbs intermediate in length; tibia, fibula and meta-
podials relatively slender.
Lrontosaurus, scapula long, narrow superiorly ; scapulo-coracoid plate rela-
tively narrow ; limbs long and heavy ; tibia, fibula and metapodials relatively
robust.
Diplodocus, scapula expanding superiorly with intermediate scapulo-coracoid
extension, limbs long and relatively slender, tibia and fibula, radius and ulna,
and metacarpals especially elongated.
Camarasaurus, thus far found only in the Colorado Jurassic, is not related to
Brontosaurus, as Osborn formerly supposed, but is a AZorosaurus of immense
size. The type skeleton will shortly be described.
The distinctive scapulo-coracoid characters are clearly brought
out in the accompanying figures (Fig. 1, A, B, C).
eon ae
© ie
Fig. 1. Scapulo-coracoid of: A. Morvosaurus, Amer. Mus. Coll. No. 332; B. Diplodocus,
Amer. Mus. Coll. No. 221; C. Brontosaurus, Amer. Mus. Coll. No. 222. All from the Bone
Cabin Quarry. 4 nat. size.
Fore Limp or Diptopocus, No. 594.
In the portion of the quarry explored during 1900, within an
area of about 20 feet, were found a scapula and a coracoid which
Fig. 2. Fore limbs of Morosaurus Amer. Mus. Coll, No. 332, and of Diplodocus- Amer. Mus. Coll. No. 594.
wy Nat. size,
[2or]
202 Bulletin American Museum of Natural History, [Vol. XIV,
fits closely to it, a humerus, ulna and radius lying together on the
right side associated with a humerus of the left side ; five meta-
carpals at some little distance associated with metatarsals and a
hind limb of Déplodocus. These bones were considerably scattered
and assigned separate field numbers, but they belong in all
probability to one individual of medium size. They have been
prepared and mounted (Fig. 2) under the direction of Mr. Her-
mann and afford for the first time a fine example of the fore limb
of this stilted Dinosaur. Although the scapula is inclined back-
wards, the upward portion of the blade is ro feet 6 inches, or
3195 mm. from the ground. The scapula is practically identical
in measurement with that of the skeleton described in Osborn’s
Memoir upon Déplodocus.
The principal measurements (in millimeters) are as follows :
Sep MEL, MeN sone ooDoodNoSae SHOMOM BiG oon a dcocoose 1285
3 greatest: bread thhis taster SS avaveyete 500
Scapulo-coracoimucon]OIMedn. miie)-jpefelel-lel tees ele 1650
@oracoid; breadth from) elenoid border... eee 425
Jaliverqliy MenvelN, cou geapnodoObobAassnoUADooe dc Sepa 1080
1 least circumiferenceiay..s. aes nicie ee cee ee eine 470
Wilma demeths «.;csjoren sae cesar ote ainids wale lm th erheanie te epee renee 87
“> least ‘cincumferencepemmce: oee alctintie cee ert 285
Radius; length’. isocacicittecuuuersicteve cate on oiccel velce tee peenene 840
>) © Weasticixeumterencenye mis sss cee chee t eee cence ee 255
Metacarpals......... i JQ QOL. IN, Vv.
Proximal facets, antero-posterior.... 105 e.! 113 76.e. 63 85
- oy “transverses serene 51 64 83 85 43
TEEN GUN 4... sas, 5.5 ieee oarratOeire ee eR se 260 290 315 290 285
The scapula is very similar in form to that already described
by Osborn, The coracotd is short, not yet conjoined to the
scapula, and fortunately retains its natural curvature; on its
postero-inferior surface is the rough facet, observed by Marsh,
for articulation with the sternal plate. The sternals were also
found, but have not been placed in position. The head of the
upper portion of the shaft of the Aumerus is crushed, so that it
extends unnaturally beyond the line of the glenoid facet; the
deltoid or lateral crest (processus lateralis) extends well down
upon the outer side of the shaft ; distally the shaft terminates on
the outer side in a prominent radial condyle which is readily
1 e = estimated.
rgoT. | Osborn and Granger, Limbs of Sauropoda. 203
distinguished by the deep groove upon its anterior face; the
lower articulation of the humerus indicates that the limb was not
greatly flexed at the elbow in the standing position, a fact in
keeping with the great weight of the anterior portion of the body.
In an uncrushed humerus the ulnar condyle is also well defined ;
the ulna, however, extends around and behind the radius.
Proximally the rad/us articulates on the anterior outer side of
the humerus with the radial condyle ; it fits in the broad anterior
groove of the w/a, which
element supports the entire
posterior as wellasthe inner
portion of the humerus,
while the radius supports
the anterior and outer por-
tion only. The analogy of
these elements with the
fore limb of the elephant is
obvious, because in both
the proboscidian and dino-
saurian the ulna is a stouter
element than the radius, as
shown by reference to the
respective circumferences
of the shafts, namely :
Fig. 3. Fore limbs of Sauropoda.
A. Complete fore limb of JZoro-
saurus, Amer. Mus. Coll. No.
332; B. Humerus of Dzplodocus,
Amer. Mus. Coll. No. 342;
Humerus of Broxtosaurus, Amer.
Mus. Coll. No. 280. All from the
Bone Cabin Quarry. 3 nat. size.
Diplodocus. No. 594 No. 588
mm. mm,
Ulna = 285 290
Radius = 255 265
The shaft of the radius i
also crosses that of the ulna
completely, as in the elephant and other ungulate mammals. On
the posterior face of the radius, in its upper fourth, is a stout
tuberosity for the ligaments and muscles connecting this bone with
the ulna, Distally, the radius and ulna present approximately
equal facets for the carpals.
Two carpals were found entirely out of position, making it
difficult to determine their homologies ; the larger of these appa-
rently belongs to the opposite side and, as the evidence is con-
flicting, reference must be made to another fore limb, No. 588,
described below.
204 Bulletin American Museum of Natural History. [Vol. XIV,
There is little question that the metacarpa/s belong to this fore
limb, although they were found mingled with metatarsals of a
hind limb of corresponding size. By comparison with several
series of metatarsals belonging to Morosaurus and Brontosaurus in
the American Museum Collection, their homologies can be de-
termined. Examination of the proximal facets for articulation
with the carpals, shows that Mte. I is
deep and relatively narrow; Mte. II
is somewhat deeper and broader, and
increases in size; Mtc. IIT is less deep,
but much broader in front; Mtc. IV
is still shallower, but of equal front
Fig. 5. Metacarpals of right manus
of undetermined Sauropod, probably
Morosaurus.
Fig. 4. Metacarpals, left manus, of 1. Proximal view of metacarpals.
Diplodocus, with supposed radiale plus 2. Anterior view of metacarpals.
intermedium. Amer. Mus. Coll. No, 3. Distal view of metacarpals.
588. 3 nat. size, Amer. Mus. Coll. No. 462. 4 nat. size.
breadth ; Mtc. V, like Mtc. I, is deep, but narrow in front. The
measurements given above are much affected by crushing, but
indicate that Mtc. III is the largest and stoutest of the meta-
carpals, while Mtc. I and Mtc. V are relatively short.
Fore Limes or Diptopocus, No. 588.
The above characters are supplemented and confirmed by
those of another fore limb (No. 588), consisting of ulna, radius,
supposed scapho-lunar, supposed cuneiform, metacarpals I, I],
III, IV, and a terminal phalanx. The metacarpals correspond
approximately with those figured by Marsh.
Tgot. | Osborn and Granger, Limbs of Sauropoda. 205
Measurements.
mm
INEGI 5 IGREU Ns. 6 po pS Doce od Suse Uae tide ee Sladenoane 870
= leastreincumilerencesemen sates seins eteeiatice nek . 265
Wiltvatgelemcitlirersrsrecsrsi versio iercie ce sieyenssictees cchesehsiay stato hace els gio
pemee LC Astacinclimilenencerere ee ri-ciciom icieiseie sie merrier 290
Supposed scapho-lunary transverse. . sc). ad ete esis selene I51
Metacarpal I, length (vertically crushed)............... 235
ss Oo OO CLs Oy otkd ECO. COIR TR NMC RO ISR OG 280
ss eT lenigithiy sari sens cacies, ociserssie ernisieeoieras 280
af Wem ame Baty chat etevererede ero slcysusissicicve, cvaley aatersvebaters 265
UEnemina)l paelenwe (GOWNS cope sono CEO EeUeOCODOOOME Or 200
Carpals.—There can be little question as to the correctness of
the association of these bones since they were found close together,
and near by was a long series of Dzplodocus caudals. Beneath
the radius was found a large flattened carpal which from its con-
nection with both radius and ulna is believed to represent a
conjoined radiale and intermedium or scapho-lunar bone ; this
bone is much thicker behind than in front ; it has a large radial
facet, slightly concave, and a smaller oblique ulnar facet ; inferi-
orly there is a single convexity with ill-defined areas for the car-
tilaginous trapezium, trapezoid and magnum ; the cuneiform in
this specimen is crushed beyond recognition ; there is a small
ossicle which may represent the unciform or one of the other
carpalia of the second row.
Fore Limes oF DipLopocus, No. 380.
This generic reference is somewhat uncertain. The associated
parts, consisting of radius, ulna, supposed scapho-lunar, cunei-
form and two ossicles probably belong to a young D¢plodocus.
This limb is important, because it includes the terminal phalan-
ges. The position of the latter, however, was not positively
determined. ,
Fore Limes oF Morosaurus, No. 332.
PIGS) Ty 2.3.
The right and left manus found with these limbs have already
been described by Osborn (2. ¢., p. 168) ; the elements were found
in position and therefore yield important information, although
the limb bones are much crushed, As in Déplodocus, the radius
206 Bulletin American Museum of Natural History, (Vol. X1V,
crosses the ulna completely and rests upon the enlarged flattened
scapho-lunar, which also presents a narrower face for the ulna ;
as in Diplodocus this carpal is much thinner in front than behind.
Fig. 6. Hind limbs of Sauropoda, posterior view. A. Mfovosaurus, Amer.
Mus. Coll. No. 461; B. Diplodocus, Amer. Mus. Coll. No. 251; C. Brontosaurus,
Amer. Mus. Coll. No. 353. All from the Bone Cabin Quarry. 2; nat. size.
The cuneiform is a smaller bone, oval and flattened as seen from
above, cyclindrical in anterior view, with an irregular projection
on the lower face. The three median metacarpals are long and
901. | Osborn and Granger, Limbs of Saurofoda, 207
slender, decidedly exceeding in length the outer pair, Mtc. I and
Mtc. V. The doubt expressed in the earlier description of
this manus as to the presence of terminal phalanges and a claw
upon the first digit still remains; the proximal phalanx asso-
ciated with Mtc. I indicates that the terminal phalanx was
missing.
Measurements,
Morosaurus. Camarasaurus, type.
mm mm
Scapulo-coracoid, total length.............. OWOS cele aoenocade 2060
Coracoid, breadth opposite glenoid border...400.............. 760
Eumeruss total lenpthiyess eh 2 <1 ete 770
Wilnarstotalblenothee crys re seers are 560
RAGIN SoMa be Sciccyet pa tere les ie epic, oa 550
Scapho-linaraerans verses ye sea cieteleree oteteve 120
MitGew i mlen other rriardotrseiac +) <laisiar-ts stare -latelc 205
Vite Se eRe! Bet Meryem ater st fet wets lyase ape the 250
Vt Com Ai Seen reeset Posh eae deat cuctopa a chee 250
VEC CHEV ice Rukia Wteceven thet cucisy cacicvesei'es ofa) cake ehente Sirs 225
IMEC S Vid sete ret tiers 2 nr dir creel vee ae Seat e . 205
CARPALS OF SUPPOSED Morosaurus, No. 462.
These are especially interesting and valuable because of the
absence of crushing, which gives the rounded or radial dispos-
ition of these bones as seen from above and in front (Fig. 5).
Measurements.
mm.
VIE Comper ert epee tetrer yar eeet say cte se arcs. tous) ee) eiersyer eisis) suse. IQ
Witte, JO: “Pi gel ayers ie: ea lbkere ree ICI SRO eR ee Par 218
Nylties, WS Coe Bieta arg.chale sets o HCCC eae eC REECE aricxeee 219
IV ICti cas lO i emp RPE IRC e= 1G Slots, Siovs e-aitiesavdigsleucyatapels sos 209
IVC RIV nae See coke Pe crcl an gr ay on)'oiigei oa tae wi'a)'s fate ar 8 av 186
The lateral metacarpals, I, V, have a more flattened section, the
median metacarpals II, III, IV, present a more rounded section.
Portions of the proximal and distal facets are well illustrated in
the figure. As in Diplodocus the proximal facets decrease stead-
ily in antero-posterior diameter as we pass from I to V. The
posterior faces exhibit rugose areas for retractor tendons,
ae pe ONS EA oe eae
208 Bulletin American Museum of Natural History, |Vol. XIV,
Fore Lime oF BRoNTOsSAURUS, No. 276.
In our collection are mounted two fore limbs of Brontosaurus,
neither of which is complete. No. 276 is a young Sronto-
saurus carpus with which have been associated arbitrarily No,
318, ulna and radius, with extensive restoration. The carpals, as
compared with those of Dzplodocus, Nos. 594, 588, or 380, above
described, are shorter and more robust. The manus of No. 268
has already been described (Osborn, 1899, p. 171). The meta-
carpals attain a very considerable length, but also acquire robust
proportions, which enable us to readily distinguish them from
those of Diplodocus ; the placing of these elements is, however,
somewhat conjectural. With the carpus is found the supposed
scapho-lunar and three ossicles, which probably represent the
carpals of the second row.
Dolichocephaly and Brachycephaly
in the Lower Mammats.
By Henry FairFieLp Ospory.
AUTHOR’S EDITION, extracted from BULLETIN
OF THE
American Museum of Matural History,
Vout. XVI, ARTICLE VII, pp. 77-89.
New York, Feb., 3, 1902.
LIST OF ANTHROPOLOGICAL PAPERS FROM THE ,
BULLETIN OF THE AMERICAN MUSEUM OF.
NATURAL HISTORY.
PRICE
A Table of the Geographical Distribution of American Relics
in a Collection exhibited in the American Museum of
Natural History, with Explanatory Text, by A. E. Doug-
dass (VOl VE 1896) i 22rppe. Po 129) ae es, aerial cme $0.75
The Temple of Tepoztlan, Mexico, by M. H. Saville (Vol.
VTE 18 g6)70 Ypps\ Sah Gane eos ete eee et A ea dee eet CAWa5o
A Rock Painting of the Thompson River Indians, British
Columbia, by James Teit, edited by Franz Boas (Vol.
VIL 32 896)0) 4pP sek. Se ise Sp Se ne Seles a 30
The Decorative Art of the Indians of the North Pacific Coast,
by. Franz, Boas(Vol- IX, 1897)2° 184: Dips ststcie pepe ee Bye
An Ancient Figure of Terra Cotta from the Valley of Mexico,
by Mi. Saville: (Vol. ix; 1807) 4p: we pl atic wee -40
The Huichol Indians of Mexico, by Carl Lumholtz (Vol. X,
TSG) = (140 Pree, PUA Re Ae Ca Ii pe eae ati aca mel ema ane -50
Marked Human Bones from a Prehistoric Tarasco Indian
Burial-Place in the State of Michoacan, by Carl Lumholtz
and Ales Hrdli¢ka (Vol..X, 1898).)° 20 pp., 5, pl...) .60
Description of an Ancient Anomalous Skeleton from the Valley |
of Mexico, with Special Reference to Supernumerary and
Bicipital Ribs in Man, by AleS Hrdlicka (Vol. XII, 1899).
Zaps ASupliviesis Oey. ACORN ODER Ler Waking RMT a A A omy a 50
The Eskimo of Smith Sound, by A. L. Kroeber (Vol. XII
1 sXa)ey) AgNO My 2) OAS) 0 | NORE A CY {Mea iran RATE AU oe ELVA SEY .80
Symbolism of the Arapaho Indians, by Alfred L. Kroeber (Vol.
EXAD ET i OO) ip O PAP eS AUNI he easel al aa Reg AMT 2G
A Shell Gorget from the Huasteca, Mexico, by Marshall H.
vaville, (Vols XILT; 2990).1-36 eps, se eee one eae 425
An Onyx Jar from Mexico, in Process of Manufacture, by M. ~
H..Savlle (Volk LIT) 2966)! 4 pp., 1 pling: Neen ere SAE 30
Cruciform Structures near Mitla, by Marshall H. Saville (Vol.
AAG a O00}. 18 ppsa: TOsphes sy GC} SUN een areca hai STS COR
A Bilateral Division of the Parietal Bone in a Chimpanzee, with
a Special Reference to the Oblique Sutures in the Parietat,
by Ale Hrdligka (Vol. XIII, 1r900)..16 pp.20 2... 30
A Bronze Figurine from British Columbia, by Franz Boas (Vol.
XLV TOGE) «| 2 PPay TPES aos oe. ee ee le aes ene are “25
Ac). Stone’s Measurements of Natives of the Northwest Terri-
tories, by Franz Boas (Vol. XIV, 1901). 16 pp., 5 pl.s..-. . 80
The Eskimo of Baffin Land and Hudson Bay, by Franz Boas
(Vol. DOV) qo0r) 370. ps 4. pli ob Re pao genes Rea ee 3.00
Basketry Designs of the Indians of Northern California, by
Roland B. Dixon. (Vol. XVII, 1902).’ 32 pp., 37 pl.:s...\ 1.50
tRnickerbocker Press, few Work
Article VII—DOLICHOCEPHALY AND BRACHYCEPH-
ALY IN THE LOWER MAMMALS:!
By Henry FairFIELD OSBORN.
Skulls are classified according to their cephalic indices into three
groups: dolichocephalic, mesaticephalic, and brachycephalic.—Nature,
>, ©, @ GONE
Dolichocephaly and brachycephaly are familiar terms in
anthropology. The cephalic index, or ratio of breadth to
length, marks a profound distinction between different races
of man; it is one of the most stable of all racial characters,
although no satisfactory theory or explanation of what it
signifies has thus far found general acceptance among anthro-
pologists.*
These facts render it all the more surprising that skull pro-
portion, distinguished from cranial or brain- -case proportion
in man, has not been con-
sidered more generally by
students of the lower mam-
mals as of great value in
the separation of races, as
well as of profound mor-
phological significance. It
is true that certain mam-
mals have been described
as short- or broad-skulled, Fig... lamar, Cris, of dolchocenhalic and
others as long- and narrow- :
skulled. As early as 1873 Kowalevsky demonstrated the
elongation of the face in Ungulates for the accommodation of
long-crowned teeth, but this does not explain the long free
space or diastema in front of these teeth; the studies of Nathu-
suis (1864) on the proportions of the skull in races of pigs,
1 Presented in Abstract before the New York ree ee of Sciences, Nov. 8, r901, and
before the National Academy of Sciences, Nov. 13, 190
2 I make this statement on the authority of Dr. a Eirdliitn of the American Mu-
seum of Natural History.
[77]
78 Bulletin American Museum of Natural History. [Vol. XVI,
are well known; in 1895 Nehring referred to the long- and
short-skulled races of dogs associated with early races of
man; quite recently also Wortman (1899) has distinguished
between short- and long-jawed races of Tertiary dogs, and
Matthew (1901) has distinguished between the long- and
short-skulled races of Oreodonts, as a basis of classification.
These are a few examples, among many which might be
found, of attention directed to such facts; but I am not aware
of any general application of dolichocephaly and brachycephaly
as factors in cranial and dental evolution, and as correlated
with the proportions of the limbs and habits of feeding.
At all events the principle has not found its way into pale-
ontological literature, with which I am fairly familiar, and
was reached by myself independently and purely inductively
while engaged upon the phylogeny of the Rhinoceroses (1900).
After accumulating a great number of facts on the evolution
of this baffling group the correlation of long limbs with long
skulls (dolichopody and dolichocephaly) and short limbs with
short skulls (brachypody and brachycephaly) suddenly ap-
peared as a key, and was expressed in the following statement:
“It [the classification adopted] sets aside several homo-
plastic parallel characters heretofore employed in Rhi-
noceros evolution and attempts to establish a firmer basis 7
the fundamental proportions of the skull, whether dolichocephalic
or brachycephalic, in the correlated proportions of the body, and
in the location of the horn cores. These characters are found
to be more distinctive of phyla than the pattern of the molar
tee ©
The full bearings of the principle were only partly per-
ceived at this time, and singularly enough I turned to the
study of the Titanotheres for the Geological Survey Mono-
graph without reference to my previous work on the Rhi-
noceroses and wholly unbiassed by any theory. Aided by Mr.
W. K. Gregory about eighty-five skulls were measured and
studied, hundreds of facts were noted which seemed to have
no particular significance; finally all these data were put
together and the conclusion was reached again, inductively,
that dolichocephaly and brachycephaly are among the dominat-
1902.] Osborn, Dolichocephaly and Brachycephaly in Mammals. 79
ing factors in the skull of the Titanotheres, and that they are
probably correlated with stmilar proportions in the trunk and
limbs. This result, as in the case of the Rhinoceroses, placed
the whole evolution of the family from its beginning in the
Eocene period in a new light and directly contradicted the
phylogenetic conclusions I had reached in 1896.
Considering the principle, however, as only a working hy-
pothesis I read through various memoirs of Cope, Marsh, Earle,
and others on the structure of the skull in the Rhinoceroses
and Titanotheres and was delighted to find that dolicho-
cephaly and brachycephaly explained a vast number of de-
tailed facts which had been recorded abstractly by these
authors without reference to their significance, not only in
all parts of the skull but in the teeth. In many respects the
teeth were proved to conform to the skull rather than the
skull to the teeth.
In brief, the proportions of the skull were found to involve,
as one might anticipate, every bone in the skull, but more
particularly nasals, horns, zygomatic arches, palate, relations
of the foramina in the base and side of the skull, the occiput,
the mastoid and other bones around the auditory meatus,
the premaxillary and mandibular symphyses, the jaw, the
diastemata between and behind the teeth, the number and
shape of the teeth, the shape, number, and relations of the
cusps, and even, it would appear, the cingulum around the
teeth. In other words all these characters were found cor-
related in many animals with the proportions of the skull,
and consequently with the structure of the limbs and feet,—
a quite unlooked for illustration of Cuvier’s famous law of
correlation. .
This gratifying result suggested a superficial review of the
mammals in general in respect to the same factors. The
conclusions reached in this paper are therefore of a prelim-
inary character.
We may first consider the skull in itself, then the corre-
lation of its proportions with similar proportions in other
parts of the body, the exceptions to such correlation and
special reasons for them, some of the apparent causes of
80 Bulletin American Museum of Natural History. (Vol. XVI,
dolichocephaly and brachycephaly, and finally some of the
facts which await explanation.
In applying these terms to the lower mammals we refer to
the skull as a whole, whereas in man the reference is only to
the cranium.
THE LONG AND THE BROAD SKULL.
The three skulls photographed below from the American
Museum collection (Fig. 2) are three nearly contemporary
species of Eocene Titanotheres which illustrate admirably
dolichocephaly, brachycephaly, and the neutral or inter-
Fig. 2. Eocene Titanotheres. A, Brachycephalic, Paleosyops paludosus. B, Mesati-
cephalic, Limnohyops manteoceras. C, Dolichocephalic, Te/matotherium cornutum.
mediate condition of mesaticephaly. The species are the
classic Pale@osyops paludosus, the extremely long and nar-
row Telmatotherium cornutum, and the moderately broad
Limnohyops manteoceras. The first and second species are
believed to belong to side lines which became extinct; the
third, more generalized, form is now believed to have given
origin to the Oligocene Titanotheres, although this inference
OO
1902.| Osborn, Dolichocephaly and Brachycephaly in Mammals. 81
awaits confirmation. The skulls of these Titanothere species
differ from those of man and of Rhinoceroses in the fact that
the cranium, or skull proper, does not vary in width so widely
as the cheek arches or zygomata. It is chiefly the enormous
expansion of the latter bones in Brontotheritum elatum which
makes the skull actually as broad as it is long. But while
mainly a zygomatic expansion, that there is a very pro-
nounced cranial and facial expansion is attested by the broad
palate, relatively short and crowded dental series, trans-
versely expanded horns, abbreviated nasals, short malar
bridge in front of the orbit, abbreviated mastoid and paroc-
cipital portion behind the external auditory meatus, trans-
versely expanded occiput and occipital condyles, broad
exoccipital and postglenoid processes, short, deep, and thick
lower jaw with less prominent angle. There is no mistaking
a typical brachycephalic for a dolichocephalic jaw, every
contour and proportion is different. Analogous differences
are observed among the Rhinoceroses.
The above are only a few of the correlated effects of skull
proportion. In the comparison of all the Titanotheres from
the beginning to the end of their remarkable history it is
found that the primitive and central types are mesaticephalic,
and the divergence is into brachycephaly and dolichocephaly.
The following table presents the extremes of structure as
observed especially in the Titanotheres.
CORRELATED SKULL CHARACTERS.
BRACHYCEPHALY. DOLICHOCEPHALY.
Teeth.
Wentall series. .-24. Crowded. ..5...-- Elongate.
Diastematameeer aoe Closede.s..:........ lnereased
: ' Persistent and
Anterior premolars, Suppressed. sonora
Pa [IBGE 2 Seo ass ( Two fangs retained.
Intermediate tuber-
cles of molars..... Persistent.......... Reduced.
Opposite dental ser-
LOSHtees oepeterias Somos Convergent or arched More parallel.
[ Fanuary, 1902. |
82 Bulletin American Museum of Natural History. [Vol. XVI,
Grimcino tectitee a
Cingula between
POCO MT wees seein. ss
Canine teeth........
Incisor series.......
Whole skull........
Most of the constitu-
ent bones.
IEA AIEH HED eh epee Ry Sa
INFASRIIS ea Bue 6 6 eae
Malar and maxillary
bridge over infra-
orbital foramen.
Infra-orbital
men.
Lachrymal bone... .
Lachrymal canal.. ..
JOS PROSTUENEN, 5 5 Bo Gobo
Areas of insertion of
masseteric and
temporal muscles.
Mastoid portion of
DEHOLIC.c nr eee Oe
Exoccipital, post-
glenoid, and post-
tympanic processes.
Post - glenoid and
post-tympanic
processes.
fora-
BRACHYCEPHALY,
Teetit.
Shortened and wid-
ened.
SUPP LESse daar
Rounded and broad-
enede einer
Placed transversely. .
Skull.
Shortened and broad-
ened.
Shortened and broad-
ened.
Broadened and flat-
tened.
Shortenedand spread-
ing.
Nanrowedi ea ae eeee
Not seen on side of
face.
Crowded into orbit. .
Crowded into orbit..
Broadened, especially
in) ‘the = buccal
plates’’; in section
broad rather than
deep.
Iincreascdeen eee
Abbreviated........
Broadened........
Approximated, espe-
cially below, en-
closing the exter-
nal auditory mea-
tus.
DOLICHOCEPHALY.
Lengthened and nar-
rowed.
Persistent.
Elongate compressed.
Converging anteriorly.
Lengthened and nar-
rowed.
Lengthened and _ nar-
rowed.
Narrowed and _ trans-
versely arched.
Long with incurving
or straight sides.
Broadened.
Conspicuous on side
of face.
Exposed on side of
face.
Seen on edge of orbit.
Elongate and vertically
deepened; in section
deep rather than
broad.
Balanced or retained.
Exposure persistent.
Deepened and _ nar-
rowed.
External auditory
meatus not closed
below.
1902.| Osborn, Dolichocephaly and Brachycephaly in Mammals. 83
BRACHYCEPHALY. DOLICHOCEPHALY.
Skull.
Tympanic bulla..... Thrust inward...... Exposed laterally.
Foramen ovale and f.
lacerum medius... Approximated...... Separated by a bridge
of bone.
Foramen lacerum
medius and f. lac-
erum posterius.... Approximated...... Separated by periotic.
Alisphenoid canal... Abbreviated........ Elongate.
Presphenoids..< =... Abbreviated... ...... Elongate.
MOMED in A-cae tous.ch Thrust backward.. ..
Premaxillary sym-
DILY.SISas peeecEse oe Abbreviated... .... Elongate.
LORS ls Sects oles Transversely expand-
Cicls Povo oesone Ook ne mae Not so expanded.
Faw.
WE ies eon Gis cat ere Shortened, thick- Elongate, with straight
ened, deepened. lower border and
backwardly pro-
duced angle.
Area of insertion for
femporaluemtiscl eae Neducede ray. on. - Balance maintained.
Coronoid process..:5, (iteducedie . 26s. = ss Lengthened antero-
posteriorly.
Mandibular symphy-
SISsAS ernst ok ee Abbreviated......... Elongate.
The above characters are chiefly observed in the Titan-
otheres, in which the most careful comparison of dolicho-
cephalic and brachycephalic skulls has been made.
Many characters in the first column apply with equal force
to the Primates which are progressively brachycephalic,
marking the passage from the more dolichocephalic Lemurs
and Baboons to the more brachycephalic Lemurs, Monkeys,
and Apes.
On the other hand many characters in the second column
apply also among the Horses, which are progressively dolicho-
cephalic.
Many of these characters also distinguish the brachy-
cephalic from the dolichocephalic Rhinoceroses.
There are, however, notable exceptions, as shown below.
84 Bulletin American Museum of Natural History. [Vol. XVI,
UNEQUAL ELONGATION OF FACE AND CRANIUM.
When we compare a long-skulled with a short-skulled
Rhinoceros the skull of the latter appears compressed antero-
posteriorly, as if composed of india-rubber, all the parts being
affected alike (Fig. 3). But although both the face and the
cranium in the Rhinoceroses and Horses appear to be affected,
this is by no means a general principle. In the Titanotheres
the face is shortened and the cranium greatly elongated, so
af ~~ m.P.Peb..
-
p.0o.
Fig. 3. Influence of progressive brachycephaly upon the ear region of Perissodactyla.
A, Dolichocephalic, Eguus caballus. B, Mesaticephalic, Zapzrus. C, Dolichocephalic,
Ceratorhinus sumatrensis. D, Brachycephalic, Rhinoceros sondaicus. Disappearance of
mastoid portion of periotic, #z. P. Per, and enclosure of auditory meatus, e. a. ., inferiorly.
that the distance between the orbit and the external auditory
meatus is very great, the molar teeth extending back beneath
the orbit. Inthe Horses, on the other hand, the face is greatly
elongated and the cranium only moderately so, and this is
true of by far the greater number of long-skulled Ungulates.
Such unequal elongation of different regions of the skull will
1902.] Osborn, Dolichocephaly and Brachycephaly in Mammals. 85
no doubt be found by examination in every family of mam-
mals.
But every exception has some special adaptive significance.
For example, the nasals in the Tapirs and the Proboscidea
are abbreviated not as an expression of brachycephaly but
in correlation with a prehensile upper lip or proboscis. The
mastoid portion of the periotic, generally exposed in dolicho-
cephalic types such as the Horses, persists also in the
brachycephalic Primates, for the insertion of one of the
most important muscles of the neck. The contrasts of
brachycephalic with dolichocephalic characters, brought out
in the above table, therefore are limited in the various mam-
malian families by special adaptive conditions.
SIGNIFICANCE OF DOLICHOCEPHALY.
The earliest known Ungulates have moderately elongate
or mesaticephalic skulls, from which it follows that brachy-
cephaly and dolichocephaly are for the most part secondary.
Fig. 4. Dolichocephalic skull of Baboon, Cyxocephalus olivaceus.
In Titanotheres and Rhinoceroses they are definitely pro-
gressive characters. The earliest horses (Protorohippus,
Hyracotherium) are already specialized in the direction of
86 Bulletin American Museum of Natural History. [Vol. XVI,
dolichocephaly, which, it is important to note, is here accom-
panied by progressive lengthening of limb.
In fact, dolichopody, in the broad sense of lengthening of
limb, is in general an adaptation to cursorial habits and
speed, associated with life on the plains, cropping front teeth,
absence of defensive weapons.
The lengthening of the limb for the purpose of speed
appears in fact to have been the prime correlate of the lengthen-
ing of the skull. There are numerous cases where the elonga-
tion of the limbs and of the skull have developed part passu,
notably in the case of the long-limbed Rhinoceroses as well
as in the Horses. It is also very characteristic of the long-
limbed Elotheres, which have extraordinarily long skulls, in
contrast with the remotely related Pigs. We reach the con-
clusion that both dolichocephaly and lengthening of neck, in
order to enable grazing animals to reach the ground, may be
primarily due to lengthening of limb.
EXCEPTIONS TO THE CORRELATION OF DOLICHOCEPHALY WITH
DoLICHOPODY AND OF BRACHYCEPHALY WITH
BRACHYPODY.
There are, however, many exceptions to the correlation of
long limbs and long skulls. Among the races of men, although
there are notable cases of such correlation, there are also no-
table exceptions; the bipedal Primates generally offering an
exception to quadrupedal mammals.
Again, the cursorial long-limbed Hyracodonts are a family
of Rhinoceroses with very long limbs and short skulls. Here,
however, brachycephaly is compensated for in a measure by
length of neck, possibly also by a substitution of browsing for
grazing habits. The most remarkable elongation of the
limbs and neck, in connection with an only moderately elon-
gate skull, is in the Giraffes, which are typical tree-browsers.
The opposite combination of long limbs with very long head
and short neck is exemplified in the Moose (Alces), habitually
a browser, which, like the giraffe, extends its mouth to the
ground with great difficulty. Whenever an animal acquires
1902.] Osborn, Dolichocephaly and Brachycephaly in Mammals. 87
the shrub- or tree-browsing habit, therefore, as in the case of
Rhinoceros bicornis, a new factor is introduced. Other fam-
ilies in which the browsing habit appears to have been ac-
quired secondarily are the Chalicotheriidz, Agriochceride, and
Anoplotheriide.
Again, among the Carnivora the Dogs are typically long-
skulled and long-lmbed or cursorial animals. A dog feeds
in a standing position, the food held upon the ground by the
fore feet, the limbs being somewhat flexed. In this family
the skull and limb correlation seems to hold good. Moreover,
the short-faced dogs are generally short limbed. Cats, on the
other hand, present a decided exception, because they are
brachycephalic and dolichopodal, the Cheetah, for example,
having an exceptionally short skull and elongate limbs. We
should recall, however, that cats always feed in the recum-
bent or semirecumbent position, crouching or lying down.
Thus the abbreviation of the Cat skull is correlated with the
functions of the teeth and not with those of the limbs, be-
cause the Cats have a special position in feeding. Similarly
the Proboscidea are extremely brachycephalic and long-
limbed, but the exceptional elongation of the limbs is com-
pensated for by the development of a proboscis.
To sum up, the numerous exceptions to the correlation of
skull and limb proportions are mostly capable of special
adaptive explanations, and, as we shall see below, when cor-
relation does occur it is probably adaptive also. In brief,
there is no innate, invari-
able law of correlation;
skull and limbs may or
may not be dependent
upon each other.
But when such correla-
tion does occur, as in Jel-
matotherium or H yopotamus ==
on the one hand, or in 7 ele- wit. S, pee skull of Monkey,
oceras on the other, it is y sae
likely to affect the whole skeleton: length of the cervical
and dorsal vertebre, form of the scapula and ilium, length
88 Bulletin American Museum of Natural History. [Vol. XVI,
of the metapodials. So complete is the correlation that we
can, for example, immediately distinguish between the ele-
ments of the pes of a long-skulled and of a short-skulled
Rhinoceros.
Eliminating all the exceptions, there appear to have been
two general causes for the elongation of the skull. First, the
elongation of the face for the accommodation of very long
hypsodont grinding teeth in front of the orbit, as observed
by Kowalevsky. Second, the elongation of the skull as a
whole, correlated with the elongation of the limbs, an adap-
tation to grazing and cursorial habit.
No ADEQUATE THEORY OF BRACHYCEPHALY.
It is much more difficult to account for progressive brachy-
cephaly. An adequate theory of its causes is still wanting,
as shown by the following examples:
Among Primates the shortening of the skull takes place
part passu with the increasing use of the manus in conveying
food to the mouth; this is well illustrated by the contrast
between the quadrupedal, long-skulled baboons and the more
bi-pedal short-skulled monkeys.
We are especially at a loss to offer any adequate explana-
tion of the causes of progressive brachycephaly in mammals
which seem to suffer thereby a reduction and compression of
the dental series. In certain Titanotheres and Rhinoceroses
the shortening of the skull seems to crowd and diminish the
usefulness of the teeth, an apparently inadaptive process.
The observations of Nathusius led him to the conclusion
that among the Suide abundant food tended to shorten and
broaden the head and the face. Darwin observes that
domestication tends to shorten the bones of the face in many
animals.
Among Carnivores, and among the long-horned Titan-
otheres, abbreviation of the skull favors the effective-use of
the canine tusks and of the paired horns respectively. But
brachycephaly also develops to an extreme in certain defence-
less types, such as Cyclopidius among the Oreodonts.
1902.] Osborn, Dolichocephaly and Brachycephaly in Mammals. 89
Further investigation and comparison may produce some
general law.
BIBLIOGRAPHY.
1867. Hux iry, THomas H. On Two Widely Contrasted Forms of
the Human Cranium. YFourn. Anat. Phys., Vol. I, 1867, pp.
60-77, Figs. 3-7.
1873. Kowa.Levsky, W. Monographie der Gattung Anthracotherium.
Paleontographica, N. F. II, 3 (XXII).
tgo1. Matruew, W. D. Fossil Mammals of the Tertiary of North-
eastern Colorado. Mem. Amer. Mus. Nat. Hist., Vol. I,
Part V, Nov., 1got.
1864. Natuusius. Vorstudien zur Geschichte und Zucht der Haus-
thiere, zunachst am Schweinschadel. Berlin, 1864.
1897. OsBorn, Henry F. The Cranial Evolution of Titanotherium.
Bull. Amer. Mus. Nat. Hist., Vol. VIII. Art. IX, pp. 157-
197, 1897.
Ig00. OsBorNn, Henry F. Pylogeny of the Rhinoceroses of Europe.
Bull. Amer. Mus. Nat. Htst., Vol. XIV, 1900, p. 229.
1885. Topinarp.—‘‘General Anthropology.” Reviewed in Nature,
1885, Vol. XX XIII, p. 4.
1899. WortMan, J. L., and W. D. MatrHew. The Ancestry of
Certain Members of the Canidz, Viverride, and Procyonide.
Bull. Amer. Mus. Nat. Htst., Vol. XII, Art. VI, pp. 109-138,
1899.
i | The Four Phyla of Oligocene
Litanotheres
By Henry FarrgIELD Osporn.
AUTHOR’S EDITION, extracted from BULLETIN
bes OF THE
A. American DAlusenm of Aatural History,
Sees Vou. XVI, ARTICLE VIII, pp. 91-1009.
ae
New York, February 18, 1902.
BRP DLs AS DIGR coibens mice pee ets ete io reek etal: One ceRe ROE, ASee RTRs
The Eskimo of Smith Sound, by A. L. Kroeber (Vol. XII,
PSQO) O04 Pps ae ple ee ots Ay oe Sen ees ale ipa ae aa eg Neste
Symbolism of the Arapaho Indians, by Alfred L. Kroeber (Vol.
AAT L900) NO. TO PP: tcc hs es | Ue, eee eit a ee
. A Shell Gorget from the Huasteca, Mexico, by Marshall H.
Saville (Volo XILT} ‘1960)i7> 6 2ppiis is) sets unex eos ee :
An Onyx Jar from Mexico, in Process of Manufacture, by M.
Hi Saville c(V.ol) XLF 1900). va ppc. Plc yoo wone eee
Cruciform Structures near Mitla, by Marshall H. Saville (Vol..
MTT, 1900}. TBEpp, Maer hk eh wascnd ote eee como ee
LIST, OF ANTHROPOLOGICAL PAPERS FROM THE
BULLETIN OF: THE AMERICAN MUSEUM OF
NATURAL HISTORY.
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A Table of the Geographical Distribution of American Relics
in a Collection exhibited in the American Museum of
Natural History, with Explanatory Text, by A. E. Doug-
lacs (Vola VIL 1896) 5 22 pp. ane Rees afeiteve helo pnemsenrninns $0.75
The Temple of Tepoztlan, Mexico, by M. H. Saville (Vol.
VATE G6). <6 SP Ps, BY PI Miso oe re anetnrcaeee atc at Eleetateer .50
A Rock Painting of the Thompson River Indians, British
Columbia, by James Teit, edited by Franz Boas (Vol.
WA TAS FS QO) SACD 6 yoo Hts Oe oan Lea coe: Sha nn ce ORS UR fete
The Decorative Art of the Indians of the North Pacific Coast,
byiPranz.Boas*(VoL: TX, \1897)-< S.4 Pps cov cis sheets teemee rans
An Ancient Figure of Terra Cotta from the Valley of Mexico,
by M. H: Saville (Vol. LX, 1897).°“4ipp.; 2 plete
The Huichol Indians of Mexico, by Carl Lumholtz (Vol. X,
TOS). MLA PIP aya oP len va ece Mivady arorene aye a bk het annie a Mapa nee teteatias
Marked Human Bones from a Prehistoric Tarasco Indian
Burial-Place in the State of Michoacan, by Carl Lumholtz
and AleS Hrdlicka (Vol. X, 1898). 20pp.,5pl:..-.......
Description of an Ancient Anomalous Skeleton from the Valley
of Mexico, with Special Reference to Supernumerary and
Bicipital Ribs in Man, by Ales Hrdlicka (Vol. XII, 1899).
A Bilateral Division of the Parietal Bone in a Chimpanzee, with
a Special Reference to the Oblique Sutures in the Parietal,
by AleS Hrdlitka (Vol. XIII; 1900). 16 pp... <.2...2..-
A Bronze Figurine from British Columbia, by Franz Boas (Vol.
XLV; Looa),\ 2 Pp. tpl: ee ee ee eae eae
A. J. Stone’s Measurements of Natives of the Northwest Terri-
tories, by Franz Boas (Vol. XIV, 1901). 16 pp., 5 pl.....
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(Viol, XV argon): 370 pips) acpi 6 AAA ae aeeernye alee tea
Basketry Designs of the Indians of Northern California, by
Roland B. Dixon (Vol. XVII, 1902). 32 pp., 37 pl.....
Knickerbocker Press, Hew Work
a. -s”
v »
Article VIII—THE FOUR PHYLA OF OLIGOCENE
TITANOTHERES.!
TITANOTHERE CONTRIBUTIONS, No. 4.?
By Henry FatIrFIELD OSBORN.
My first review of this group, entitled ‘Cranial Evolution
of Titanotherium’ (Titanotherium Contributions No. 3), was
based upon examination of only part of the original material;
it included a preliminary revision of the species by the dis-
tinction of growth, sexual and variable characters, and by
the method of sections of different parts of the skull.
Two errors invalidated this review. First, the stratigraph-
ical or geological sequence of the types examined was not fully
known, and, second, as a consequence, the group was treated
as more or less monophyletic with certain side branches.
This second review is an abstract of a portion of the results
obtained for the U. 5. Geological Survey monograph, ‘The
Titanotheres,’ now in preparation. It covers practically
all the type material in the Yale, National, American, and
Harvard Museums, and advantage has been taken of the
invaluable field observations by Hatcher of the levels on
which the different skulls in the National Museum collection
were discovered. The section method also has been very
greatly extended and, taken in connection with the teeth and
the detailed structure of the skull, has proved to be a sure
criterion of specific and phyletic character.
. Beginning in January, 1901, the work for the monograph
advanced uninfluenced by any theory as to the evolution of
these mammals until finally, in July, all the data were put
together with most interesting results, which may be briefly
summarized as follows:
1The late Professor O. C. Marsh of Yale University devoted many years to the
collection of materials and preparation of plates and figures for a monograph on the
group treated in this paper. He also completed several valuable papers, but left no
manuscript. Full acknowledgment of this important service to paleontology will be
made by the writer in the final treatise.
2 See contributions 1-3 in Bibliography.
[or]
92 Bulletin American Museum of Natural History. [ Vol. XVI,
I. THE ADAPTIVE RADIATION.
The Oligocene Titanotheres consisted of at least four con-
temporary phyla, to which the prior generic names T1tano-
therium, Megacerops, Symborodon, and Brontotheriwum may be
applied.
They represent an adaptive radiation for different local
habitat, different modes of feeding, fighting, locomotion, etc.,
which took origin, in part at least, in the Middle or Upper
Eocene. Europe and Asia also may have shared in this
radiation, since Titanotheres are now definitely known in the
Balkan region.
The main phyletic characters are analogous to those re-
cently (Osborn, 1900, p. 231) determined among Rhinoceroses;
the great antiquity of the lines leading to the existing species
of Rhinoceroses necessitated the revival of a number of dis-
carded generic names to distinguish them. Similarly the
separateness of four of the Titanothere phyla, throughout the
Oligocene and possibly from the Eocene, render it desirable
to revive certain generic names which in my first review I
considered undefinable.
Radiation involved three main sets of characters, two of
which were correlated:
First, dolichocephaly and brachycephaly, associated with
numerous changes in the skull and teeth, and, in at least two
phyla, with longer and shorter limbs.
Second, four distinct types in the shape and position of
the horns, correlated with the structure of the nasals and
frontals and indicative of different modes of combat among
the males.
Third, canines of different form; and, finally, the presence
of one or two pairs of functional incisor teeth, or the total
degeneration of these teeth.
2. THE FOUR GENERA.
Titanotherium Leidy applies to long-limbed animals with
long skulls, persistently long and broad nasals, short triangu-
1902. | Osborn, Oligocene Titanotheres. 93
lar horns placed slightly in front of the eyes, vestigial in-
cisors, =, large canine teeth. Known from the base to the
summit of the Oligocene.
Megacerops Leidy applies to Titanotheres with broad
skulls, nasals progressively shortening, short horns rounded
or oval in section, shifting anteriorly, one. or two pairs of
incisor teeth, =e medium sized canine teeth. Known from
the base to the summit of the Oligocene.
Probably related to this are the subgenera of the types
named Allops and Diploclonus by Marsh, differing from the
above in horn characters. Known chiefly from the Upper
Beds.
Rises Z Wegacerops, Upper Beds.
6 aon Diploclonus, Upper Beds.
Renews Symborodon, Upper Beds.
YU pry over. Titanotherium, Upper Beds.
Lip aE Brontotherium, Upper Beds.
Fig. 1. Characteristic basal horn sections.
Symborodon Cope includes Titanotheres with skulls of vary-
ing proportion, nasals slender and progressively shortening,
horns elongate and peculiar in being placed above the eyes
instead of shifting forwards, incisors vestigial #=°, canines
small, approximated. Known only from the Middle and
Upper Beds.
94 Bulletin American Museum of Natural History. [Vol. XVI,
Brontotherium Marsh embraces the largest Titanotheres,
with very broad zygomatic arches, nasals shortening while
horns elongate and shift forwards; incisors persistent, $ in
the males, canines stout and obtuse.
Representatives of Titanotherium and Megacerops can now
be continuously traced from the base to the summit of the
Oligocene. Primitive species of Brontotherium also appear
at the base, although the phyletic sequence through the
Middle to the Upper Beds is not so clear. Symborodon sud-
denly appears in the Middle Beds.
3. NEWLY OBSERVED PROGRESSIVE CHARACTERS.
In addition to the progressive and retrogressive charac-
ters previously recorded by various writers (Osborn, 1896,
pp. 162-174), are several of great value in the determination
of species. (1) In the complication of the premolar teeth,
one or more phyla progress in common, although the rate is
unequal; for example, the double internal cones of the upper
premolars are precociously developed in Titanotherium and
very gradually so in Megacerops. (2) The cranial and many
of the dental characters are profoundly affected by the pro-
gressive shortening or lengthening of the skull as recently
explained in some detail by the writer (Osborn, 1902).
(3) The cingulum around the molar and premolar teeth be-
comes progressively stronger in T7itanotherium and Megacerops,
and progressively weaker in Symborodon and Brontotherium.
(4) The persistence or degeneration of incisor teeth are far
more constant systematic characters than appeared to be
the case in my first review. .
In brief, each genus or phylum has its distinctive, persistent,
progressive, and retrogressive characters, of which the above
are a few examples out of many.
4. THE SUCCESSION OF SPECIES.
Phylum I.
Dolichocephalic Titanotheres with short, divergent, triquet-
rous horns placed slightly in advance of the orbits: nasals elon-
1902. | Osborn, Oligocene Titanotheres. 95
gate and square; tnctsors vestigial; canines very long and pointed;
cingulum progressively developing ; premolars 4; buccal pro-
cesses of zygomata deep rather than broad.
Genus TITANOTHERIUM Lenny.
The type of the genus is T. prouti Leidy, represented by
the posterior portion of a jaw fortunately preserved in the
National Museum, recorded from the base of the Titanothe-
rium Beds. The genus is further characterized by a com-
parison of JT. heloceras, T. trigonoceras, and T. ingens. It
may have taken origin from Diplacodon elatus or D. emargi-
natus of the Upper Eocene or Uinta Beds.
Fig. 2. Tzttanothertum heloceras (Cope). (U.S. Nat. Mus. No. 4260, Sk. Q.) X i.
LOWER BEDS.
T. heloceras Cope.—This is probably the earliest and most
primitive representative of the long-skulled series. The doubt
96 ©Bulletin American Museum of Natural History. [Vol. XVI,
arises from the imperfect condition of the type. With this
species, however, is probably associated a skull (Nat. Mus.
No. 4260) definitely recorded by Hatcher as from the Lower
Beds and exhibiting a number of undoubted Tztanotherium
characters.
MIDDLE BEDS.
T. trigonoceras Cope.—Animals of middle size with long
metapodials, represented by numerous specimens in the
American and National Museums, show the state of evolution
of this phylum in the Middle Beds. As compared with T.
heloceras there is a marked progression in size, in the shaping
of the horns, the development of the paired premolar cones,
and of the cingulum on the grinding teeth.
UPPER BEDS.
T. ingens Marsh (Syn., Menops varians Marsh).—These
very large, long-skulled and long-footed Titanotheres, first
described by Marsh as Brontotherium’* ingens, are definitely
recorded by Hatcher from the Upper Beds, and show a cor-
responding increase in size and in all other progressive char-
acters. The incisors are vestigial, the canines very long and
powerful; the cingulum on the grinding teeth is very pro-
nounced in both males and females. There is a marked
difference between the sexes in the‘size of the horns and
canines.
Phylum II.
Brachycephalic Titanotheres with short horns shifting for-
wards, rounded to oval at summits; nasals progressively reduced
in length and broadening at the extremities; one or two pairs of
persistent incisor teeth, above and below; canines pointed, of
medium length; premolars, *=*; zygomata progressively expand-
ing into convex buccal processes; cranium not greatly produced
behind sygomata.
: Ate Soloed below, the name Brontotherium applies to the animals called Titanops
by Marsh.
1902. | ~ Osborn, Oligocene Titanotheres. 97
Genus MEGACEROPS Le rpy.
This genus may have taken origin in the broad-skulled
Limnohyops manteoceras or Paleosyops paludosus of the
Eocene, although no horned type of the latter is known. It
is known from the imperfect type of M. coloradensis and a
very rich variety of species from the Lower, Middle, and
Upper Beds, which have previously been referred to other
genera. In the typical members the horns are slightly diver-
gent and retroverted. There is a wide variation in progres-
sive character between such species as M. brachycephalus
and dispar, M. bicornutus, M. selwynianus, M. tichoceras, M.
robustus, but it is found that the variations of age, growth,
and sex, especially in species such as M. dispar and M.
robustus, bridge over the differences between the types of
said species and present intermediate forms. Furthermore
these species are much more closely united to the general type
of Megacerops than to any other genus.
LOWER BEDS.
Megacerops brachycephalus, sp. nov.
The type of this species is No. 4261, U.S. Nat. Mus. It in-
cludes very small, broad-skulled Titanotheres with very rudi-
mentary sec-
ond internal
cones upon
the dapper
premolars;
nasals elong-
ate, mnarrow-
ing anteri-
Orly,» asin
Pale@osyops.
Horns of an- rig 3. Megacerops brachycephalus. (U.S. Nat. Mus. No. 4258,
. ra v1
teroposterior Sk. F.) Notthetype. X 3.
oval section placed above orbits. It is represented in the
National Museum by numerous skulls besides the type, all
[ February, 7902. | 7
98 Bulletin American Museum of Natural History. [Vol. XVI,
collected and recorded by Hatcher. One of these skulls was
provisionally referred by him to Teleodus avus, from which
this species is quite distinct.
MIDDLE BEDS.
Megacerops dispar Marsh (Syn., Brontops validus Marsh).
—Represented by the type, No. 4941, U. S. Nat. Mus., and
numerous fine male and female skulls in the National Mu-
seum, collected and accurately recorded as to level by
SS
SS CON sy =
4. Megacerops brachycephalus. (U. S. Nat. Mus. No. 4258,Sk. F.) Not the type.
Thiet skull was figured by Marsh (Amer. Jour. Sci., Oct., 1887) as Brontops dispar. X }.
Hatcher. The incisors vary from two to one. Canines
prominent, pointed. Premolars, = with internal cones
more distinct than in M. brachycephalus. Nasals shorter;
horns longer, broader in section at the base; zygomata
more expanded.
1902. | Osborn, Oligocene Titanotheres. 99
LEVEL NOT RECORDED
Megacerops avus Marsh.—This type jaw, distinguished
by the presence of three lower incisors, belongs to a much
larger animal than M. brachycephalus. The presence of
an outer lower incisor may be due to reversion or to the
retention of a milk tooth; all young Titanotheres have three
milk incisors. The formula, I;, Ps, does not appear suffi-
cient to remove this type generically, because the form of the
canine, shape of the jaw, feeble development of the cingulum,
etc., lead to placing it in Wegacerops.
M. coloradensis Leidy.—Horns of medium length, of rela-
tively slender cylindrical section. Nasals elongate, decurved,
and narrow anteriorly. This is the type species.
M. angustigenis Cope.—Type in Ottawa Museum. Horns
as in M. coloradensis, but nasals abbreviated and expanding
anteriorly.
M.? selwynianus Cope.—Level not recorded, probably
Middle Beds. Type in Ottawa Museum. Nasals extremely
narrow, elongate, with highly arched inferior surface as in
the species next to be described.
MIDDLE BEDS.
Megacerops bicornutus, sp. nov. °
Type No. 1476, cotype No. 1081, Amer. Mus. Horns directed an-
teriorly. Hornlets upon the inner and anterior mid-portion of the
horn. Basal section of the horn slightly oval, subtransverse. Nasals
narrow and relatively elongate. Sharp malar bridge in front of orbit.
Orbit large.
This animal stands nearest M. selwyntanus, although dis-
tinguished by the greater size and slightly greater width of
the nasals. The sharp malar bridge is the most absolute
character. The two hornlets are possibly variations.
UPPER BEDS.
Megacerops tichoceras Scott & Osborn.—Type skull in
Harvard Museum. This is distinguished from M. dispar by
100 Bulletin American Museum of Natural History. [Vol. XVI,
the somewhat abbreviated nasals, greater expansion of the
buccal processes, and two distinct internal cones upon the
superior premolars. It is represented also by a number of
Fig. 5. Megacerops bicornutus. (Amer. Mus. Nat. Hist. No. 1476.) Type. 4.
large skulls in the National Museum definitely recorded by
Hatcher from the Upper Beds.
Megacerops marshi, sp. nov.
Type, Skull No. so1, cotype, skull No. 1445, Amer. Mus. Nasals
elongate and square distally, horns short, of oblique oval basal sec-
tion, overhanging the maxilla, or projecting forwards or outwards.
Incisors, ?$. Canines short, tetartocones of premolars moderately
developed.
These skulls were previously confused by the writer with
T. trigonoceras, from which they are readily separated by the
horn section, which relates them to some of the primitive
types of M. brachycephalus and equally to M. robustus. The
canines are more obtuse than in M. dispar, and the superior
1902. ] Osborn, Oligocene Titanotheres. 101
incisors resemble those in Prontotherium rather than in M.
robustus.
Fig. 6. Megacerops marsht, (Amer. Mus. Nat. Hist. No. 501; jaw No. 516.) Type.
This skull was figured by Osborn (1896, p. 181) as 77tanotherium trigonoceras.
Megacerops robustus Marsh.—Type in Yale Museum.
Numerous skulls in the American, Princeton, and National
Museums. Recorded by Hatcher from very summit of the
Upper Beds. In skull development this represents an
extreme evolution of the M. brachycephalus-M. dispar
series. It is, however, distinguished from M. dispar by
the presence of diastemata behind the canines, retarded
development of the tetartocones on the superior premo-
lars, broadly transverse horn-section, procumbent position
of the horns.
The above-described new species, VW. (? Diploclonus) bicor-
nutus, from the Middle Beds is possibly a connecting form
between Megacerops and the aberrant species from the Upper
Beds termed Dziploclonus amplus by Marsh. The latter is
distinguished by short, divergent horns of peculiar triquet-
rous section, with incisors +, and elongate canines flattened
posteriorly. The animals called Allops are certainly more
closely related to Megacerops than to either Titanotherium,
Symborodon, or Brontotherium, but their phyletic position is
uncertain.
102 Bulletin American Museum of Natural History. (Vol. XVI,
Genus ALLOPS Marsz.
Syn. ? Dtploclonus Marsh.
Canines lanceolate, flattened posteriorly, incisors 7. Horns with
greatest diameter progressively transverse, pointed outwards and
forwards.
The distinctness of this genus is doubtful.
UPPER BEDS.
Allops serotinus Marsh.—Superior premolars with small
postero-internal cusps (tetartocones). Represented by two
skulls in the National Museum, recorded from the Upper Beds.
No connecting crest between the horns.
Allops crassicornis Marsh.—Type No. 4289, Nat. Mus.
Superior premolars with large postero-internal cusps; cingula
Fig. 7. Dzploclonus amplus (Marsh). (Yale Museum.) Type. }.
more prominent; zygomata with flattened buccal processes.
No connecting crest between the horns.
Allops amplus Marsh.—Type in Yale Museum. Charac-
ters similar to the above with the exception of a strong
connecting crest between the horns and a pair of rugose
internal hornlets on the horns, hence the term ‘ Diplo-
clonus.’ It is possible that this species descended from M.
1902. | Osborn, Oligocene Titanotheres. 103
bicornutus as a member of a collateral phylum distinguished
by internal hornlets. In such case Diploclonus may prove to
be a valid subgenus.
Phylum ITI.
Medium-sized Titanotheres with long horns placed more di-
rectly above the orbits, of deep anteroposterior diameter at the
base, oval at tip ; connecting crest feeble or wanting. Nasals
thin; skull proportion varying in the species. Incisors degen-
erate. Canines very small, approximated.
Genus SYMBORODON Cope.
Syn., Anisacodon, Diconodon Marsh.
This genus is characterized from the types of S. torvus, S.
acer in the American Museum, and of S. montanus in the
Yale and National Museums. It is known only from the
Middle and Upper Beds, but is entirely distinct. It agrees
with Titanotherium in the disappearance of incisor teeth,
but differs from it absolutely in the horn and nasal struc-
ture. It is also widely separate, both in cutting teeth and’
horn structure, from Megacerops and Brontotheritum.
PROBABLY UPPER BEDS.
Symborodon torvus Cope.— Brachycephalic, incisors %,
nasals thin, buccal processes of zygomata broad and flat,
occipital pillars broad.
Symborodon acer Cope (Syn., S. altirostris Cope).—Dol-
ichocephalic, nasals thicker, abbreviated. Horns of male long,
recurved, low connecting crest, incisors vestigial.
MIDDLE AND UPPER BEDS.
Symborodon montanus Marsh.—Type in Yale Museum.
Nasals thin, of medium length, upper incisors vestigial.
104 Bulletin American Museum of Natural History. [Vol. XVI,
Two skulls in the National Museum (Nos. 4711, 4705) ex-
hibit smaller canines than in Marsh’s type, and possibly repre-
ar sent a fourth species,
because the canines are
exceptionally small, the
nasals excessively thin,
the horns elongate, set
wide apart and very far
back, oval in section at
the top.
Phylum IV.
Titanotheres attaining
the largest size, with vertex
of cranium greatly elong-
ated by extension of horns
in front of orbit and of
° occiput behind zygomata.
Nach ee ee te eee: (U.S. Skull, however, as meas-
ured along the basal line
and across the zygomata progressively brachycephalic. _ Horns
transverse oval in section from base to summit, shifting for-
ward and progressively elongating and flattening. Nasals
abbreviating. Incisors %. Canines stout, blunt, obtuse. Pre-
4-3
molars **, cingulum degenerating.
GENuS BRONTOTHERIUM Marsu.
Syn., 7itanops Marsh.
The type of this genus is the Brontotherium gigas jaw
(Yale Museum), with which the type jaw of Titanops elatus
(Yale Museum) is practically identical. The succeeding
species (B. curtum, B. ramosum, B. dolichoceras, B. platy-
ceras) of very long, flat-horned Titanotheres therefore belong
to Brontotherium.
A discovery of great interest is a very primitive skull of
1902. | Osborn, Oligocene Titanotheres. 105
the same general type, but with rudimentary horns, in the
Lower Beds, to which the name B. leidyi may be given. A
Fig. 9. Brontotherium leidyt, (U.S. Nat. Mus. No. 4249, Sk. R.) Type. x}. The
side view of this skull was figured by Marsh (Amer. Jour. Sci., Oct., 1887) as Wenops varians
successor of this species from the Middle Beds proves to be
identical with Cope’s Symborodon hypoceras.
LOWER BEDS.
Brontotherium leidyi, sp. nov.
Type, Skull No. 4249, U. S. Nat. Mus. Nasals elongate, narrow-
ing anteriorly. Horns very short, slightly recurved, of transverse oval
section. Canines stout and blunt. Premolars non-cingulate, with
rounded contours and well-developed tetartocones, Incisors 2
This rare skull was discovered by Hatcher and is posi-
tively recorded from the Lower Beds, Middle Level. It is
106 Bulletin American Museum of Natural History. (Vol. XVI,
of exceptional importance because it is so clearly distinct
from its contempo-
raries in the Lower
Beds, namely, Tzta-
nothertum heloceras
and Megacerops bra-
chycephalus. It has
numerous. resem-
Fig. 10. Brontotherium leidyi. (U.S. Nat. Mus, blances in the shape
Nose Ser of the nasals, horns,
canines, and back of the occiput to the great Brontotherium
of the Upper Beds.
MIDDLE BEDS.
B. hypoceras Cope.—Cope’s extremely fragmentary type
of this species (No. 6361, Amer. -
Mus., Cope Coll.) is happily sup-
plemented by two skulls found
by Hatcher (Nos. 4702, 4273,
U. S. Nat. Mus.), one of which
is complete though crushed.
These prove that this animal
is undoubtedly a successor of
B. leidyi, but separated by
certain progressive characters, ic an. Bepatliorgie ieee
such as the more anterior posi- (U.S. Naw, Mos No. a7} cotype
tion of the horns, and the entire loss of the median incisors.
Specific Characters——Horns of medium length, transverse oval sec-
tion; nasals shorter than in B. letdyi; zygomata with flattened buccal
processes; incisors reduced to +.
This species, although exhibiting generic resemblances, is
not ancestral to B. gigas.
UPPER BEDS.
The following species, beginning with B. gigas, form a
closely related progressive series which is chiefly confined to
the Upper Beds. One of the most distinctive characters is
1902. | Osborn, Oligocene Titanotheres. 107
the persistence of the large pair of upper incisors in all the
male specimens thus far observed. Certain females appar-
ently lack incisors.
The horns are very ee
long; the nasals oa
gradually disappear.
B. bucco Cope.—
In the writer’s first
review the type of YY
this species was con- /.
fused with the types Be
of Symborodon torvus.
It is actually repre- Z)
sented by the poste-
rior portion of a
cranium in the We
American Museum ae
(No. 6346). The buc-
calesectdom of the
zygomata is more GD Y oN
convex than in the % wee
succeeding species. A. B.
B. gigas Marsh. Fig. 12. Sections of nasals, horns, occiput, zygomata.
3 A, Brontotherium letdy?, type. B, Brontothertum
(Syn., Titanops elatus — kypoceras, cotype.
Marsh).—This is now
one of the best-known species, including the type in the
Yale Museum, and numerous specimens in the American and
National Museums previously referred to B. elatum.
Smaller and somewhat more primitive varieties of B. gigas
are recorded by Hatcher from the Upper Levels of the Middle
Beds.
B. dolichoceras Scott & Osborn. (Syn., Titanops medius
Marsh).—Represented by types in the Harvard and National
Museums. This species is intermediate in many characters
between B. gigas and B. curtum.
B. curtum Marsh. (Syn., Menodus peltoceras Cope).—
The type in the Yale Museum is supplemented by the
female horns named by Cope Menodus peltoceras, also by
108 Bulletin American Museum of Natural History. [Vol. XVI,
several specimens in the American, Harvard, and National
Museums.
B. ramosum Osborn.—There is some question whether this
is really distinguishable from B. curtwm.
Fig. 13. Comparative fronto-nasal sections and horn contours. Showing progressive shift-
ing forward of horns, and abbreviation of free portion of nasals. Brontotherium letdyz, Lower
Beds. Brontothertum hypoceras, Middle Beds. (Nat. Mus. No. 4702.) Brontotherium
hypoceras. (Nat. Mus. No. 4273.) The dotted vertical line shows the position of the orbit.
B. platyceras Scott G Osborn.—Type in the Harvard
Museum.
BIBLIOGRAPHY.
1887.—ScoTt, W. B., and H. F. Osporny Preliminary Account of
the Fossil Mammals from the White River Formation con-
tained in the Museum of Comparative Zodlogy. [Titano-
there Contributions No. 1.] Bull. Mus. Comp. Zodl., Vol.
XIII, No. 5, pp. 151-171, pll. I-II, Aug., 1887.
1893.—HatcHer, J. B. The Titanotherium Beds. Amer. Nat.,
March 1, 1893, pp. 204-221.
1895.—OsBoRN, Henry F. Perissodactyls of the Lower Miocene
White River Beds. [Titanothere Contributions No. 2.] Bull.
Amer. Mus. Nat. Hist., Vol. VII, Art. XII, pp. 343-375,
pill, Vill, EX, Dee, 23;-1805:
1902. | Osborn, Oligocene Titanotheres. 109
1896.—OsBoRN, HENRY F. The Cranial Evolution of Titanotherium.
[Titanothere Contributions, No. 3.] Bull. Amer. Mus. Nat.
Hitst., Vol. VIII, Art. IX, pp. 157-197, July 31, 1896.
1g900.—OsxBorN, Henry F. Phylogeny of the Rhinoceroses of Eu-
rope. Bull. Amer. Mus. Nat. Htst., Vol. XIII, Art. XIX,
Pp. 229-267, Dec. 11, 1900.
1902.—OsBorN, Henry F. Dolichocephaly and Brachycephaly
in the Lower Mammals. Bull. Amer. Mus. Nat. Hist., Vol.
XVI, Art. VII, pp. 77-89.
ea
A Skull of Dinocyon from the Miocene of
Texas.
By W. D. Matruew.
e AUTHOR’S EDITION, extracted from BULLETIN
iu
—— American Biriseum of Aatural History,
VoL. XVI, ARTICLE XI, pp. 129-136.
: New York, April 7, 1902.
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The Tnickerbocker Press,
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Article XI.— A SKULL OF DINOCYON FROM THE
MIOCENE OF TEXAS.
By W. D. MatTTHEw.
Among the valuable specimens brought back by Mr. J. W.
Gidley from his collecting trip for the American Museum last
summer, were the skull and part of the skeleton of an enor-
mous carnivore which on extraction from its matrix proves
to be a Canid of the Amphicyonine group. It appears to be
a very aberrant species of Dinocyon, a genus hitherto known
by teeth and fragments of the jaw of D. thenardi described by
Fig. r. Side View of Skull x 4.
Jourdan in 1862! from the Upper Miocene beds of Grive St.-
Alban.
This specimen is more complete than any Amphicyonine
hitherto described, not only in this country, but in Europe,
1 Comptes Rendus de 1’Institut, LIII; Bull. des Sociétés savantes (1862). Another
species, Amphicyon gériachensis Toula, is referred to this genus by Prof. Deperet. Dr.
Schlosser prefers to place it with Hemicyon, probably identical with H. sansaniensis
Lartet. It would seem to be an intermediate form, like Dinocyon in the carnassial,
like Hemicyon in the tuberculars.
[April, 1902.] [129] :
130 Bulletin American Museum of Natural History. [Vol. XVI,
jaws are complete although crushed, and the first eight ver-
tebre are in place. The succeeding nine vertebre and parts
of the hind limb were found close by. All were enveloped
in hard flinty concretion, which has been removed from one
side only of the specimen. The horizon is the Loup Fork
terrane, which, in the part of Texas in which this specimen was
found, contains a fauna approximately Upper Miocene in age
so far as comparisons have been instituted.
It is only within the past two years that true Amphicyons
have been recognized in this country, the species referred to
that genus by Leidy and Cope being, as Prof. Scott has
shown,’ much more primitive, and nearly or quite in the line
of descent of the modern Canidz, while the true Amphicyons
are an aberrant branch of dogs, related to the Urside, but
not directly ancestral to them, according to Dr. Schlosser’s
recent studies on the group.” In the ‘American Journal of
Science’ for January, rg01, however, Dr. Wortman has de-
scribed a true Amphicyon from the Loup Fork beds of Ne-
braska. Mr. Earl Douglas has recognized the genus in the
same terrane in Montana. The American Museum Expedi-
tions of 1901 obtained remains of Amphicyons both in the
older Loup Fork of Colorado (Middle Miocene) and the newer
Loup Fork of Texas (Upper Miocene). To this group may
also be referred three species of Canidae described some time
since, but whose position has not been recognized. These
are:
Cams (Atlurodon) ursinus Cope, from the Loup Fork of
New Mexico.
Borophagus diversidens Cope,’ from the Blanco of Texas.
4tlurodon m@eandrinus Hatcher,* from the Loup Fork of
Kansas. .
The specimen here described represents a species distinct
from any of those above mentioned, and is named in honor
* Notes on the Canidz of the White River Oligocene, Trans. Amer. Phil. Soc., Vol.
XIX, 1808, p. 326 et seq.
? Ueber die Baren und barenahnlichen Formen des europaischen Tertiars. Palaeon-
tographica, Bd. XLVI, 1800, p. 05 et seq. -
© Ero. Acad. Nat. Sci. Phila., 1875, p. 256; Rep. Wheeler Survey, 1877, p. 304, pl.
seb es ile atk
* American Naturalist, 1892, p. 1028; Rep. Tex. Geol. Sur., 1892, p. 52, pl. xiii, fig. 4.
® American Naturalist, 1894, p. 239 and fig.
1go2. | Matthew, A Skull of Dinocyon from Texas. 131
of Mr. J. W. Gidley, the discoverer of many of the choicest
specimens of fossil mammals in the collections of the Ameri-
can Museum and of Princeton University.
Dinocyon (? Borophagus) gidleyi, sp. nov.
3-1.4.2
SL WY We, ie
Generic and Subfamily Characters. — Dentition Premolars
much reduced, without posterior accessory cusps, one or two of the
inferior series perhaps absent. Carnassials small and low with re-
duced shear, tubercular teeth very large with low cusps. Jaw very
deep and massive, facial part of skull elongated, sagittal and occipital
crests high, brain-case small. Bulle inflated, but smaller than in
typical Canide, mastoid process small, paroccipital process moderately
‘long, coéssified with the bulla. (In the bears the mastoid process is
much enlarged, the paroccipital reduced, and the tympanic bulla not
inflated.)
Subgeneric Characters.—Teeth like those of Amphicyon in form,
especially such species as A. americanus Wortman. Upper molars
much wider transversely, and the first more trigonal in outline than
in Dinocyon thenardzt.
Specific Characters.—Size somewhat greater than in D. thenardi or
D. (Aelurodon’’) meandrinus, premolars more reduced and pre-
molar region of the jaw longer than in the latter species or in Boro-
phagus diversidens.
It is difficult to place this fine species in any of the de-
scribed genera. In form and character the carnassials and
molars (Fig. 2) are like |
those of Amphicyon.
But the third upper
molar, a well developed
tooth in Amphicyon, is
absent from either side
of the Texas skull, as it
isin Dinocyon and Hemui- Figs Crown View of Teeth x ¢.
cyon, and probably in
Pseudamphicyon.. The size is near that of D. thenardt, with
which the proportions of the lower teeth, so far as they can
be seen, agree fairly well. But the first and to some extent
the second upper molars, are trigonal and much extended
transversely, while in Dinocyon, and still more in Hemicyon,
1 Schlosser, J. c.
132 Bulletin American Museum of Natural History. {Vol. XVI,
these teeth approach the round quadrate shape characteristic
of Hyenarctos. D. gidleyi may be considered as an aberrant
member of the genus Dinocyon, but its relationship to the
type species is perhaps rather formal than real.'
The skull (Fig. 1) is the most complete one yet described
of an Amphicyonine dog. Professor Filhol*? has described
and figured a skull of A. Jemanensis, but it was by no means
complete. The Texas skull, although crushed, is quite com-
plete, and the parts of the skeleton preserved enable us to
determine the proportions and general character of this great
carnivore.
The size of the skull equals or exceeds that of any living
carnivore of which | can find record. It is longer, wider, and
deeper than the largest Ursus maritimus skull in our collec-
tion, and its measurements slightly exceed those given for
the Kadiak bear in length and depth (including the jaw).
The crushing of the skull prevents any exact comparison in
width; but the Kadiak skull is probably much wider than
was that of D. gidley1.
The most striking characters of the skull are the size and
depth of the jaw, and the heavy zygomatic arches. The
nasal openings are large and cavernous, as in the polar bear,
unlike the smaller and more slender muzzle of the lesser bears
and of the dogs. The inferior postorbital process is hardly
as prominent as in the wolf, much less than in any of the
bears. The tympanic bulle are inflated, although of pro-
portionately smaller. size than in the wolf. The teeth are
larger in proportion to the size of the skull than in the bears,
especially the molars, which exceed those of U. maritimus in
length and are more than twice as wide. The cranium bears
a high crest, as in Amphicyon, giving attachment for the
powerful jaw-muscles; and the brain is decidedly smaller
than in the modern Urside. Compared with Amphicyon
* Should further study of the American Amphicyons render it advisable to remove
D. gidleyi and the probably nearly allied D. m@eandrinus to a different genus, Cope s
name Borophagus may perhaps be used. But as the type of Borophagus is from the
Blanco beds it would be desirable to know more than we do at present about the Blanco
Amphicyons before making any such change. All that can be said at present is that
there were two or more species of the group in this horizon, of unknown dental formula.
? Arch. Mus. Lyons, [II, 1883, pl. i, figs. 3-5. ;
1902. | Matthew, A Skull of Dinocyon from .Texas. 133
lemanensts of the Upper Oligocene, the premolars are more
reduced, the muzzle larger, the sagittal crest not so high, and
the brain-case fully as large in proportion (although, as the
skull is so much larger, one would expect to see a proportion-
ately smaller brain-case). The arches are heavier and the
jaw much deeper. In the outline sketch of the skull (Fig. 3)
the crushing of the specimen has been corrected, as nearly as
could be estimated.
The cervical vertebre are of the size of those of the polar
bear, but differ rather widely from both bears and modern
Fig. 3. Outlines of skull restored x }.
Canide, suggesting some of the more primitive Canide, such
as Daphenus. The atlas is not very perfectly preserved, and
does not show any very significant characters. The avs is
not unlike that of the bears, the spine ending posteriorly in
a short heavy process directed equally upward and backward.
The anterior prolongation of the spine as a thin high lamina,
carried far forward in the dogs and still further in the cats,
is much reduced, as it is in the Urside. The hemal surface
of the centrum bears a strong median ridge. The remaining
cervicals bear spines of much greater height than in Camzs or
Ursus, and of quite different form; they are flat, slender at
134 Bulletin American Museum of Natural History. (Vol. XVI,
the base, and carry a nearly uniform width to the tip, where
they are slightly enlarged. This form of spine in the cervi-
cals, resembling the usual form of a dorsal spine, is seen to a
less extent in Daphenus, but not in any of the large modern
carnivora, among which the bears offer the nearest approach.
The zygapophyses are considerably smaller than in U. mari-
timus; the transverse processes are as long,
but quite slender,—their inferior lamellz
quite well developed on the one or two
vertebre in which they have not been
broken off.
The centra of probably the first nine
dorsals are preserved, of which the first
was found in position. They are some-
what narrower and a little longer than
in U. maritimus; the first three are
strongly keeled, the others are round in-
feriorly, as in the bear. No part of the
arches or spines is preserved.
The femur (Fig. 4) is smaller than that
of U. maritimus, and resembles much
more that of the wolf in its characters. .
The ball faces more laterally than in the
bears, and is well to one side of the axis
of the shaft; the shaft is somewhat curved,
less so than in Canis, much more than
in Ursus; the condyles project more
posteriorly, the trochlea is narrow and
me: 4. Remue . deep, as in the wolf, not broad and shal-
low as inthe bear. The lesser trochanter
appears to be less prominent than in either Canis or Ursus;
the greater trochanter projects to a level with the top of the
ball, and is considerably more prominent than in Ursus.
The upper end of the tibia is preserved, but considerably
crushed. It appears to have the high cnemial crest and nar-
row proximal facets of the dog, not the lower crest and broad
facets of the bear.
Nothing is preserved of the feet, an unfortunate defect, for
1902. | Matthew, A Skull of Dinocyon from Texas. 135
the best generic distinction between Dinocyon and Hemicyon
lies in the foot characters, the former genus being plantigrade
with short metapodials like the bears, the latter digitigrade
with long metapodials like the dogs.’ If indeed the closer
resemblance to the Canidz which we find in the femur and
tibia of D. gidleyi is equally marked in the distal parts of the
limbs, then our species is not related to Dinocyon, but must
be placed in a distinct genus allied to Hemicyon.
| met eet an all eel
a | 3 | % = 18 g18 4
S| se | 8] 2 lgalss
Measurements. © 5 Oe Weck here ieee
mate eee | asst hecad ae
A ’ i a Ws a 2
See ipo
Length of skull, incisors to occipital
CARES Race ae Ec eae, SOTA CAR Ce ts CO nee ee 450 223)" 2017
Length of upper dentition (i2—m2 in
lOpnociiorp: aya. (OhASPIS)) 5 ao dope se 204 [44] 147
Meneth onuppenp=—ms s4 4. kas. ss 79 52| 64
Fa Fee Ds LOnmoiG es eke 0s 34 Gi 509)
a i sib. adh crate. Si er aa ae 26] 33 15} 19
- a PetMensTI 2 5 tua opty RES a ney. 21| 2090 £3)\| “2%
Width) = 95 sm2 transverse... .est.. Bila 18| 15
S F Otani ss est. 40| 34 Ta@)|| 40g
BOMBA Se easy ily MiG Eee 2 29 Te 8
emigt ine Ole pany annescr ee re att a «aS ow vc 363 252
Wepth, “is. speneath pe he. u. 5... | 982 50] 55 | 69
$ ee ci Pee enat st. mci) LOS 50
Post-canine diastema (c,;—p; exclusive)
ene thst Py sce a ee nee so 2 ws 101 76
ah pen artic 865 oEecinee a cee eee ee 40} 45 2A Ae
ae saat ie. cree lero, bata athe ee RTE Bese Dit 24
TO IBOATS sesso oes Un eer och he ene i072 \) iG) 17
Width of skull (as crushed)........... 270 180] 197
Length of seven cervical vertebre..... 347 330
Width of centra of same, average......| 44 46
Length of ten dorsal vertebre........ 398 345
Average width of centra..............| 39 40
Rena thyar temic rants. gc weaken sala a: 301 438
east diameter ot shatte. 42.5 5255.46. 32 34
Wiameter of distal end >. vo 5.5..'..4:. 87 98
% Sear cll eer cae ee Sree es) s,s 46 56
Although more specialized than Amphicyon, D. gidleyt 1s
apparently not nearer to the bears; the characters of the
1 Filhol, Mammiféres fossiles de Sansan, p. 151.
136 Bulletin American Museum of Natural History. [Vol. XVI.
femur are considerably less bear-like than in A. mayor, judg-
ing from Filhol’s description and figures;* the vertebrz are
bear-like in many respects, but quite peculiar in the length
and form of the spines. The skull and teeth suggest an in-
dependent specialization, parallelling that of the bears in a
few characters, but in most respects peculiar. A further dis-
cussion of its relationship is reserved for a later paper.
The animal must have been of peculiar appearance, not
greatly resembling either bears or dogs. The enormous head
was carried very low (if this is the correct interpretation of
the high cervical spines), more so than in the bears, much
more than in the wolves; the muzzle was long and heavy;
the contour of the head was straight, and continuous with
the neck; the ears of moderate size, jaw very long and deep,
wide gaping, cheeks rather wide. The neck was as massive
as in U. maritimus, the trunk longer and slimmer, the legs
shorter, sharply flexed at the knees, the thigh not as free
from the trunk as in Urside, but much more like the condi-
tion seen in the Canide.
1 Mammiféres fossiles de St. Gerand le Puy.
On the Skull of Bunelurus, a Musteline
~ from the White River Oligocene.
By W. D. MartHew.
AUTHOR’S EDITION, extracted fom BULLETIN
OF THE
American Museum of Natural History,
VoL. XVI, ARTICLE XII, pp. 137-140.
New York, April 7, 1902.
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‘The Tknickerbocker Press, Rew
ay
Article XII.— ON THE SKULL OF BUNALURUS, A
MUSTELINE FROM THE WHITE RIVER
OLIGOCENE.
By W. D. MatTruew.
Bunelurus was described by Prof. Cope in 1873! from
fragments of the lower jaw, which until now have remained
the only representation of the genus. Cope referred it to a
position in the Mustelide near Putorius and Plesiogale; Dr.
Schlosser in his later revision of the European carnivora?
considered it close to or identical with Palgogale (in which
he includes part of Plesiogale). Dr. Wortman has recently
suggested® that it might not improbably prove to be a direct
descendant of certain of the Viverravide.
A finely preserved skull found by Mr. Thomson of the
American Museum Expedition of rgo1, in the Upper Oreodon
TH : ; 4
PE pe PES hy
Fig. 1. Side View of Skull.
beds of East Pawnee Butte, Northeastern Colorado, is the
subject of the present description. No lower jaw is with it,
so that it cannot be positively identified. But the close cor-
respondence in point of size with B. lagophagus, the type of
which is from the same horizon and region, and the corre-
spondence of the teeth of our skull with the upper teeth of
the more carnassial section of the Musteline and of the
teeth of Bunelurus with the lower teeth of the same group,
1 Synopsis New Vert. Col., p. 8; Ann. Rep. Hayd. Sur., 1873 (1874), 507; Tert. Vert.,
p. 946, pl. Ixviia, figs. 13, 14. i
2 Affen Lemuren, u. s. w. d. Europ. Tertiars, p. 386.
%Amer. Jour. Science, 1901, Vol. XII, p. 145, footnote.
[137]
138 Bulletin American Museum of Natural History. (Vol. XVI,
make the identification reasonably safe. No other Musteline
is known from the White River.
The characters of the skull confirm the views expressed by
Cope and Schlosser as to the position of the genus. It is
Paleogale with a minute second molar still retained. It be-
longs to the primitive division of the Musteline, with tri-
angular first molar, no posterior flange on the protocone.
The carnassial is primitive in character, somewhat resem-
bling that of Cynodictis gregarius, the protocone very large, the
shear more oblique than in modern Musteline, less so than in
Cynodictis, the fissure between protocone
and postero-external blade still quite well
p2 marked. There is a small antero-internal
basal cusp and a less marked antero-ex-
ternal one. The second and third pre-
molars are of moderate size without
trittocones, much higher than in Mustela,
higher and proportionately larger than in
Putorius. The first premolar is a single-
rooted tooth of small size; first and second
premolars are spaced. Alveoli of canines,
of moderate size, are preserved.
The bulle are of primitive character,
inflated, short and prominent, instead of
flattened and elongated as in Mustela and
p.3
AIS
= ae if m2 Rea Putorius. The palate extends backward
1g. 2. Town 1ew O A hes
Teeth, only to opposite the anterior edge of the
first molar, while in modern Mustelines
it extends considerably behind the teeth. The shorter bullae
leave a much larger surface of the sphenoids and _ occi-
pitals exposed; the short stout paroccipital process is entirely
free of the bulla. The occipital and sagittal crests have the
same outlines as in Putorius ermineus, but the posterior lobes
of the brain are separated from the cerebral lobes by a strongly
marked depression; the arches are much heavier, muzzle
much longer, resembling that of Mustela more nearly, but
flatter, longer, more slender toward the tip; infraorbital fora-
men smaller, postorbital process of the frontal less prominent.
a
EQO2. 5 Matthew, The Skull of Bunelurus. 139
Postorbital constriction much more narrow than in M.
americana, somewhat more than in P. ermineus. Size slightly
greater than the weasel.
Bunelurus is one of the primitive group of Musteline found
chiefly in the Oligocene of Europe. It belongs to the Putoriine
section, which more nearly ap-
proaches the Felide (through
Proatlurus) in dental reduction
(the typical Musteline section
more nearly approaching Cyno-
dictis and the Viverride), but
shows little indication of the
shortening of the face charac-
teristic of modern Putorius.
Following are the more im-
portant primitive characters:
t. Dentition less reduced, four
premolars and two molars
in upper jaw.
2. Fourth premolar more tri-
angular, shear moretrans-
verse, notch behind pro-
tocone deeper.
3. First molar more viverrine
in shape, protocone not
expanded transversely,
para- and metacones more Fig. 3. Inferior View of Skull.
distinct.
. Second molar present although minute.
Posterior nares not roofed over behind m!.
Bulle short, round, and prominent, instead of long and flat-
tened.
7. Paroccipital process free.
8. Cerebrum small and not extending over cerebellum, transverse
sulcus strongly marked on external surface of skull by a
wide depression.
g. Postorbital processes very rudimentary, postorbital constric-
tion narrow.
10. Infraorbital foramen small.
Ans
The skull is identified with Bunelurus upon the following
evidence: Bunelurus appears to belong to the more carnas-
140 Bulletin American Museum of Natural History. [Vol. XVI,
sial section of the Musteline, typified among modern genera
by Putorius, among ancient ones by Pal@ogale, rather than
to the section with more tubercular teeth, typified among
modern genera by Mustela, among ancient ones by Plesictts,
Stenoplesictis, etc. The latter group retains the metaconid
on m1, and correspondingly the paraconule on mt. The
former group has no metaconid on the lower carnassial, and
on the upper tubercular there is no trace of conules, and the
protocone is smaller in proportion. The primitive members
of this Putoriine group are hardly distinguishable from primi-
tive felines; Proailurus, considered by most writers as a
Felid, is placed by Dr. Schlosser (advisedly as it seems to the
present writer) among the Mustelids of this group. Bbune-
lurus presumably belongs to the primitive division of the
Putoriine group.
The White River skull under discussion belongs unques-
tionably to the Putoriine group, and with the primitive mem-
bers thereof. It has the strongly transverse tubercular, with
reduced protocone and no paraconule. The upper teeth cor-
respond in size and in proportions with the lower teeth on
which bunelurus is based. These were found in the same
formation and horizon as the skull, at a locality about fifty
miles further to the eastward.
— American Eocene Primates, and the
ape Supposed Rodent Family :"s
Mixodectide. Bee
By Henry FAIRFIELD OSBORN.
mn nay un. Oe ee ‘em
__. AUTHOR’S EDITION, extracted from BULLETIN ; Won fo
— American Museum of Datural History,
Vou. XVI, ARTICLE XVII, pp. 169-214. i
New York, June 28, 1902. 3
deh ayes : 4 ANS
Loe
be thnickerbocker ‘Press, Re
Sha ME Pye Veet ae ee Oar he
Article XVII.— AMERICAN EOCENE PRIMATES, AND
THE SUPPOSED RODENT FAMILY MIXODECTID.
By Henry FairFIELD OsBorN.
The only American Primates at present known are those in
the Eocene. The supposed Oligocene Primates, Laopithecus
Marsh and Menotherium Cope, have proved to be identical
with Leptocherus Leidy, an Artiodactyl.
Invariably associated with the discovery and literature of
the Primates is the family Mixodectide, including Mixodectes
and possibly [ndrodon, Cynodontomys, and Microsyops, now
supposed to be very primitive Rodents. In the writings of
Cope, Marsh, and Leidy, the bibliographical relations of these
two groups are so intimate that it is convenient to revise
them together.
Altogether fifty-one species have been named, many of
them based upon defective types; the synonymy is truly
appalling, as shown in the chronological table.
Many years ago I devoted several months to systematic
revision as the basis of the present paper, examining and
comparing the types in the collections made by Leidy, Marsh,
and Cope, now in the Philadelphia Academy (Ph.), Ameri-
can Museum (A. M.) and Yale University (Y.). Unfortu-
nately all the types described by Cope for the Wheeler
‘Survey, and figured in his ‘Extinct Vertebrata of New Mexico’
of 1874 have disappeared. A beginning is made here by
pointing out the synonymous genera but the species require
prolonged and microscopic examination and comparison.
This, however, is intended to supersede and replace all
previous revisions by the author and his staff in the
Museum.
Dr. W. D. Matthew has rendered invaluable aid and advice
both in the morphological and descriptive part. In putting
together these notes and tables | have also been greatly
assisted by Mr. W. K. Gregory. The drawings are chiefly
_ the work of Mr. Weber, Mr. Horsfall, and Mr. Anderson.
[169]
170 Bulletin American Museum of Natural fistory. [Vol. XVI,
Part [. PRIMATES.
I. SUPPOSED BASAL EOCENE (MONTIEN AND THANETIEN)
PRIMATES.
None of the Basal Eocene (Puerco and Torrejon) types
appear to be positively ancestral to the Lower Eocene or
Wasatch Primates; the Primates of the latter stage (Sparna-
cien) thus far appear to represent a new primate fauna like
the new ungulate fauna of horses, tapirs, etc.
The Puerco and Torrejon species include a great number
and variety of small animals whose relationships are still
largely a matter of individual opinion, because the material,
except in the case of one skeleton (No. 823, see below),
affords no absolutely distinctive characters. They have been
referred by different authors, chiefly Cope, Schlosser, Earle,
and Osborn,to such
diverse orders as
the: Creodonta,
Rodentia, Condy-
larthra, Insecti-
vora,and Primates.
I have always in-
clined’ to -reter
many of the smaller
types to: the. Pri:
mates, but without
being able to give
C conclusive grounds
for the opinion, the
main reason being
the general adap-
tive resemblance
which they bear to
Fig. 1. A, Mioclenus acolytus. Amer. Mus. No. 8292. slugs
Lower molar. &, supposed Primate. Am. Mus. No. 823. the existing Le-
B, superior molars, enlarged 2 diam., B', contour of same
still more enlarged. C, lower jaw of same specimen, 2 diam. IMUITS.
Compare Fig. 2. Twice natural size.
Indrodon malaris
(Torrejon stage) was placed by Cope in the Anapto-
morphide; this is an error. The structure of the superior
1902. | Osborn, American Eocene Primates.
n7i
molar teeth relates it
rather to Muixodectes, a
supposed primitive Ro-
dent.
The skeleton (Amer.
Mus. No. 823) originally
associated with ndrodon
by error (Osborn and
Earle, 1895, pp. 16-20)
deserves most careful
examination, for if it
belongs to a Primate
it is by far the most
primitive known. The
astragalus is not like
that of a Rodent. The
figures (Figs. 1, 2) give
the proportions of the
limbs.
Mioclenus acolytus
and M. lemuroides Mat-
thew are two other small
Torrejon species in which
the lower molar teeth
suggest those of Hyop-
sodus, especially in the
reduction of the para-
conid (Fig. 1A).
- Oxyacodon apiculatus
(Puerco) and O. aga petil-
lus (Puerco), also Car-
cinodon filholianus
(Puerco), deserve ex-
amination in this con-
nection.
Fig
size,
. 2. Supposed Primate.
Fore and hind limb bones and tarsals.
Compare Fig. 1.
Am. M
us.
Al
No. 823. .
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[172]
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[173]
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“snupriiags snygsomojgouy i =
fore -d ‘adog snunissads sdotsosnypy
(65)
(85)
(45)
(95)
(SS)
(PS)
(£5)
(8)
(15)
(08)
(6?)
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(4%)
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ps Ut asD gq amasajay pup saquinyy sav2adS
[175]
176 Bulletin American Museum of Natural History. [Vol. XVI,
II. LOWER, MIDDLE, AND UPPER EOCENE PRIMATES.
(SPARNACIEN, YPRESIEN, LUTETIEN, BARTONIEN, LIGURIEN.)
Originating in the lower Eocene or Wasatch (Sparnacien,
Yprésien) of North America are found three phyla of Pri-
mates, quite distinct from those in Europe.
t. The HyopsopoNTID#& are analogous in certain respects
of molar tooth structure to the Microchceride of Europe, but
are distinct in the simple cutting teeth.
2. The NoTHarctip& resemble in size and general form
the contemporaneous Adapide of Europe, but the much more
complex structure of the upper molars constitutes good
grounds for not placing them in the same family.
3. The very specialized ANAPTOMORPHID# so far as known
(skull, teeth and jaws only) remotely resemble the living
Fig. 3. Superior molars. A, Adapis magnus; B, Hyopsodus uin-
tensts, type; C, Notharctus sp.
Tarsiide of the Oriental region, but it seems to be a pre-
mature conclusion to place them in the same family because
they are so widely separated geologically and geographically.
The European Eocene Primates, also placed in three
families, Adapidz, Microchceride, and Plesiadapide, are now
generally (Trouessart, ’97) referred to the Lemuroidea. The
force of Schlosser’s remark (’87, p. 19) uniting them in the
distinct suborder Pseudolemuroidea (‘‘ This suborder includes
only extinct forms, which certainly mark a transition between
the true Monkeys and Lemurs, but stand in direct genetic
relations neither with one nor the other’’) has been vigorously.
disputed by Leche (’96), who, on the ground of resemblances
in milk succession, places certain of these families (Micro-
choeridz) in the Lemuroidea.
It may be possible with the material now in hand to
1902. | Osborn, American Eocene Primates. io avg
positively determine the relationships of some of these forms
to the existing Anthropoidea or Lemuroidea; but it will
require detailed investigation, which I am not able to under-
take at present.’
Three suppositions are possible: First, that these Primates
represent an ancient
and generalized
group (Mesodonta,
Cope) ancestral to
both Lemuroidea
and Anthropoidea;
second, that they
include representa-
tives: of both
Lemuroidea and An-
thropoidea, contem-
poraneous and in-
termingled ;__ third,
that they belong
exclusively to one
Ot the” other order:
There are certain
advantages in the C
revival of the term
Mesodonta Cope, a
suborder (anticipat-
ing the terms
Pseudolemuroidea
and ‘Tarsii) which
would bear some-
CUS foe ie Fig. 4. Jaw outlines. All natural size. A, Pelycodus
lationship to the ge re paulus ; C, Anaptomorphus emulus ;
modern specialized
Monkeys and Lemurs that the Condylarthra bear to the
Ungulata and the Creodonta to the Carnivora. The serious
difficulty with this view is the very considerable separation
of these families.
* Dr. J. L. Wortman is now taking up these problems with the rich materials afforded
‘by the Yale Museum Collections. I therefore omit phylogenetic questions here.
[ Fune, 1902.) 12
IN
178 Bulletin American Museum of Natural History. [Vol. XVI,
The extensive material in the American Museum is at the
disposal of any thoroughly competent investigator who desires
to exhaustively study this group. The present revision,
while not final, will certainly be of service.
GEOLOGICAL DISTRIBUTION.
PARTIAL LIsT OF SPECIES.
Puerco
Torrejon.
Wasatch.
Wind River.
Bridger
Uinta.
HYOPSODONTID2.
H yopsodus MUTCULUS an eel = ek es on
POMLGPNTCNWS a. notre ees om teue eel ee
POWONTOTIUS ic, isto ecole + erates
LU OP EMILIO so ops nite 9 Sse Bes Jo! outa poste lee eae
DUES WR ote asia hie te OR tea seme Pe
WOTUSGUUUS so des canes
GIUSVOMNS Nok cos- hah nea eseiate
UTIOMS TS i, Is, A MAES :
Sarcolemur Baal
furcatus..
xx X
ce
ee
“e
KG OGRE
NOTHARCTID.
Pelycodus FONEOUUD en rty pee eeeah Ace vee 3 eva eee
JUMZUUOTUS. aac sein ¥Sons cerern sw eihcs Me oer ac
TUES opoin's lies soto se TET > patente isle BH plol o oale
ce
5 SOROS
ae
Q
Ss
8
S
nH
XxX KK XK
CHASSIS a etaisieisioa)s) oa
ANAPTOMORPHID#.
OrDER MESODONTA Cope.
Pachylemuriens Filhol, in part; Pseudolemuroidea Schlosser, in part;
Tarsw Gill, in part.
Characters: Primitive Primates. Incisors typical or reduced to 3;
canines typical or enlarged, premolars 4 to 2; molars 2; upper molars
ranging from trituberculy to sexituberculy; lower molars ranging from
quinquetuberculy (tuberculo-sectorial) to quadrituberculy. Lachrymal
1902. |
foramen external or internal to orbit.
fossze, with or without postorbital bar.
foramen.
Osborn, American Eocene Primates.
179
Orbits
opening into temporal
Humerus with entepicondylar
The American forms divide into three contemporaneous
phyla as follows:
1. Hyopsodontide
Schlosser.
1.2, ¢.4,p.4,m: $=44.
Mesaticephalic.
Lachrymal canal mar-
ginal or internal to
orbit. Dental series
not crowded. Pre-
molars slowly re-
duced. Superior mo-
lars becoming sexi-
tubercular, quadrate.
Talonid elevated,
with pointed cusps.
No postorbital bar.
2. Notharctide Os-
born.
ae ==
dete Fan
40,
Dolichocephalic.
Premolars persistent.
Superior molars tri-
angular to quadrate,
tritubercular, pro-
gressively sexituber-
cular. Inferior molars
with depressed, cren-
ulate talonid.
1 4 § —
c.1,p.4,m.3=
3. Anaptomorphide
Cope.
Lia C. +, D-——, ine
36—32.
Brachycephalic.
Lachrymal canal ex-
ternal to orbit. Den-
tal series reduced and
compressed. Premo-
lars rapidly reduced.
Superior molars tri-
tubercular, trans-
versely extended.
Short deep jaw. A
postorbital process.
FAMILY HYOPSODONTID ScuHLosseEr.
Lemuravideé ! Marsh.
Definition.—Dentition with slight or no reduction, incisors and can-
ines normal;
canines slightly enlarged in males;
superior molars
progressive from tri- to sexituberculy, with progressive external
cingulum, but without mesostyle; inferior molars evolved from
quinque- to quadrituberculy by reduction of paraconid; inferior
molars with hypoconulid; external cusps progressively opposite;
pointed cusps both on talonid and trigonid. Lachrymal foramen mar-
ginal or within orbit. Skull without postorbital bar.
Analogous to the Microchceride and Necrolemur in sexitubercular
superior molars; differing in typical incisor and canine teeth. Anal-
ogous to the Adapide in typical incisors and canines; differing in
sexitubercular evolution of molars.
CHRONOLOGICAL LIsT.
Species of Hyopsodus.
(C2) EIpODSORUS PAULUS MELCIDY acco ore swiss «219 dure Wier e siete. Bridger.
(3) wt (Microsus) cuspidatus Leidy..............
(GO) (Stenacodon) varus Marsh. ............++.-
' The genus Lemuravus is a synonym of Hyopsodus.
180 Bulletin American Museum of Natural History. (Vol. XVI,
(ao) Hyopsodus minusculus Leidy: .\ cess see een Bridger.
32) = (Microsyops) vicartus ‘Cope: =. ines. sees “
(34) (Esthonyx) miticulus Cope. >, sn: a: Wasatch.
(36) o (Lemuravus) distans Marsh..............- Bridger.
(48) a lemomsianus Cope... a. ee ea Wasatch.
(50) ? » (Phenacodus, Diacodexis) laticuneus Cope... ‘“
(53) ‘s powellwanus COpPes = 22 otto eee eee id
(55) se worimant- Osborn, 2. .%,: sacha Wind River.
(56) cS marsh? Osborn. vai 5 « snthm os See Bridger.
(57) ve uimionsts ‘Osboriis; ..¢- 20.4522 ee eee Uinta.
Species of Sarcolemur.
(11) Sarcolemur (Hyopsodus) pygm@us Cope............. Bridger.
(31) ce (Auivacodon) jurcatus Cope. sans eee BG
(41) $s (Antiacodon) crassus Cooper... eee ee
0c) era: (Hyopsodus) gracilis, Marsh... 7.022455). - ms
(24) ? ‘1 (Entomodon) comptus Marsh.............. es
This family embraces a great variety of middle-sized Pri-
mates extending from the Wasatch (Suessonien) to the Uinta
(Ligurien) divided into two readily distinguishable genera,
Hyopsodus Leidy and Sarcolemur Cope.
Hyopsodus. Sarcolemur.
Paraconid typically wanting. Paraconid persistent, close to
metaconid.
A single specimen (Am. Mus. No. 4192) of Hyopsodus exhibits a
vestigial paraconid and bridges the gap between these genera.
Genus HYOPSODUS Lume.
Microsus Leidy, Lemuravus Marsh, ? Stenacodon Marsh. Compare
also Diacodexis laticuneus Cope.
Hyopsodus is one of the most abundant and persistent of the
Mesodonta, extending from the Wasatch to the Bridger and
even into the Uinta. It was first described by Leidy in
1870 from a lower jaw found near Fort Bridger; the types
of Microsus Leidy and Lemuravus Marsh, also from
the Bridger, appear to be generically identical with it.
Lemuravus was mistakenly separated by the formula i 3,
a character which is common to all the known species of
Hyopsodus. This is the only Primate with the complete
eutherian dentition known, namely : 3, 4, 4, 3=44. The
1902. | Osborn, American Eocene Primates. 181
narrow form and pointed cusps of the lower molars, both upon
talonid and trigonid, readily distinguish the members of this.
family from the Notharctide. All the known species in-
clude animals of small size.
Hyopsodus does not show a progressive increase in size;
for example, the largest Wasatch species, H. powellianus,
(Fig. 9) is larger than the largest known Bridger species.
Premolar and molar transjormation.—Bridger species of
Ungulates, and in fact of all bunodont mammals, are generally
more complex in dentition than Wind River species, and
invariably far more complex than Wasatch species, so we
cannot agree with Cope in identifying Wasatch specimens
of Hyopsodus with typical Bridger species such as H. paulus.
The species therefore require thorough rearrangement accord-
ing to geological succession and the law of progressive com-
plication of the molar and premolar teeth.
The grinding teeth gradually become more complex, homo-
plastic with those of Ungulates. The upper molars progress
from a triangular, tritubercular condition with a rudimentary
hypocone to a quadrate, sexitubercular condition with a
prominent hypocone (Bridger and Uinta); the external cin-
gulum increases in strength until it entirely extends across.
the outer surface of the crown (H. marsht); it does not
develop a mesostyle. The upper premolars progress by the
addition of internal cusps; the fourth upper premolar has
an internal cusp (deuterocone) in the Wasatch specimens;
the fourth, third, and second have internal cusps in the upper
Bridger specimens. The third and fourth lower premolars
are similarly transformed, but less rapidly.
If we should follow the same principle as that which obtains
among the horses, the species of Hyopsodus in successive
geological stages might well be separated as genera but it
would not subserve clearness to do this.
1. WASATCH (SPARNACIEN, YPRESIEN) STAGE.
Common characters of the Wasatch species —Superior molars
sub-triangular with hypocone depressed and rudimentary or
182 Bulletin American Museum of Natural History. [Vol. XVI,
feebly developed; fourth superior premolar only with a well
developed internal cusp or deuterocone, third superior pre-
molar with a small deuterocone, no trace of paraconid on
Fig. 5. AHyopsodus paulus. Am. Mus. No. 2301. Lateral,
palatal, and superior views of skull. See page 186. Natural size.
lower molars except as an occasional vestige on m,; this is
a remarkably early specialization of these Primates towards
a quadritubercular type and readily distinguishes them from
the contemporary Notharctide, and Anaptomorphide. The
1902. | Osborn, American Eocene Primates, 183
hypoconulid indicated on all the lower molars, is strongly
developed as a posterior spur on m*.
(Sp. 34) Hyopsodus (Esthonyx) ? miticulus Cope. Meas-
urements in the type are: three inferior molars = 12 mm.;
this species may include
the small Wasatch speci-
mens which Cope has
referred to H. vicarius
and H. paulus (both
Bridger species) in which
the molars vary from
ro mm. to1z2mm. The
identification of these
Men remaliegawstwand Gye td) ease Pe
teeth with H. muticulus
is provisional. The best preserved specimen (Am. Mus.
No. 4128) shows an exceptionally reduced third superior
molar (Fig. 6).
(Sp. 48) Hyopsodus §le-
moinianus Cope. — Measure-
ments of three inferior molars
estimated at 15 mm. This
species includes the middle-
= sized individuals collected in
Fig. 7. Hyopsodus lemoinianus. Type: the Wasatch of Wyoming ’
Hen eey laeueeeea ) mamely,, No: \qr39,. which is
the first figured specimen or
. |
. wae im.3,
Fig. 8. Hyopsodus lemoint- . i
anus, Am. Mus. No. 4100. Fig. 7a. Hyopsodus lemoinianus. Am. Mus, No.
Twice natural size. 1. Twice natural size.
type, and Nos. 4140, 4138, as well as many of the specimens
referred by Cope to H. paulus and H. vicarius. No satis-
factory specific distinction can be given atipresent. This may
include specimens ranging as follows: m,—-m,=13 to 15 mm.
184 Bulletin American Museum of Natural History. [Vol. XVI,
(Sp. 53) Hyopsodus powellianus Cope. — Measurements
of three inferior molars in the type: 18 mm. This includes
the robust jaws and teeth, namely, Am. Mus. Coll., the type
No. 4147, and Nos. 4148, 4150, 4151, 4152. These are as
large as, or larger than, the largest species known from the
Bridger; the m, - m,; = 16 to 18 mm.
(Sp. 50) ?Hyopsodus (Diacodexis) laticuneus resembles
NO
G ST)
Fig.9. Hyopsodus powellianus. Am, Fig. 10. Déacodexis laticuneus.
Mus. No. 4147. Slightly less than twice Type. Am. Mus. No, 4202. Twice
natural size. natural size.
this species in size but differs in the presence of a paraconid.
This type is, however, of uncertain reference.
Dr. Matthew has observed that these species may be ar-
ranged in three groups, including larger and smaller jaws in
each group, which may represent successive stages of develop-
ment in a long geological age; these differences of size may
however represent males and females respectively.
2. WunpD RIVER (LUTETIEN) STAGE.
Common characters of the Wind River species.—Hyopsodus
is represented by a large number of specimens from the Wind
River beds. which Cope mistakenly referred to the Bridger
species H. paulus and H. vicarius. The superior molar teeth
1902. | Osborn, American Eocene Primates. 185
are somewhat more primitive than those found in the Bridger
in the more triangular shape and less prominent development
of the hypocone. The hypocone is stronger than in the Was-
atch specimens and both third and fourth premolars exhibit
deuterocones. It is therefore probable that there are valid
specific differences between these animals and the overlying
Bridger species.
(Sp. 55) Hyopsodus wortmani, sp. nov.
Type No. 4716, Am. Mus. (figured by Cope as H. vicarius), Fig 11.
is valuable because it shows a
complete maxilla and premaxilla,
the latter with three incisors of
which the median pair is possibly
slightly enlarged; the crown of
the second incisor preserved is
pointed. Third superior premolar
narrow with a deuterocone. Su-
perior ms =10 mm. Inferior pms—
ms = 21 mm.
Fig. rr. AHyopsodus wortman?. Type.
Am. Mus. Cope, No. 4716. Superior and in-
The best specimens in the ferior teeth. ‘Twice natural size.
American Museum collection are
Nos. 4716, 4712. Inferior true molars measure 11 to 13 mm.
A slightly larger and somewhat different variety is represented by
Nos. 4701, 4715, 4732.
pe
3. BRIDGER (BARTONIEN) STAGE.
As shown in the chronological list above, six species
have been named from this formation and the synonomy can
only be cleared up by exact comparison of the types. Deuter-
ocones begin to appear on the second as well as the third and
fourth upper premolars; hypocones are still more prominent.
The Bridger specimens show a more or less decided external
cingulum in the upper molars.
(Sp. 2) Hyopsodus paulus.— This includes the species of
middle size, inferior molars = 14 mm. With this should be
compared the type of (3) H. (Microsus) cuspidatus Leidy,
also of (16) Stenacodon rarus Marsh; also of (32) H. (Micro-
syops) vicarius Cope.
186 Bulletin American Museum of Natural History. [Vol. XVI,
Skull of Hyopsodus.—The most beautiful specimen in the
collection consists of a skull and jaws of Hyopsodus (Fig. 5),
collected during the American Museum expedition of 1895
by Dr. J. L. Wortman. Unfortunately the anterior region of
the orbit is fractured, but on the left side the lachrymal
foramen is seen to be marginal or internal as in the Anthro-
poidea and not external as in many of the Lemuroidea; the
infraorbital foramen is placed above the interval between the
third and fourth premolars; the temporal fossa is surmounted
by a thin sagittal crest, which is broken away in this specimen.
The external auditory meatus is widely open inferiorly, and
there is no trace of a tympanic tube (the absence of a tym-
panic tube distinguishes the South American from the
European monkeys); the bulla is also broken away, exposing
a portion of the semicircular canals. Behind the auditory
meatus is apparently a narrow exposure of the mastoid, per-
forated by the mastoid foramen; the posterior nares open
just behind the last molar.
Dentition.— The last molar is a small tooth with a
small hypocone, the first and second molars have the
hypocone better developed. Unlike H. marsht, the third and
fourth premolars only have internal cusps (deuterocones),
the second premolar has an internal basal cingulum, the first
is a Simple conical tooth; close in front of this is the canine,
a much larger tooth. The grinding series do not converge
anteriorly, being nearly parallel; this is an important point.
There is a faint external cingulum.
The lower jaw. — The inferior molars measure 14 mm;
they cannot be distinguished from those of H. paulus ; the
animal was thus a small one. The fourth lower premolar
presents a deuteroconid, the third premolar presents a rudi-
ment of the same, the second and first are absolutely simple.
The jaw has a well marked masseteric fossa, the condyle is
raised somewhat above the level of the molar series, the chin
is not very distinctly defined, the mandibular symphysis is
coossified.
(Sp. 29) Hyopsodus minusculus Leidy.— This appears to be
1902. | Osborn, American Eocene Primates. 187
the smallest representative of the genus in this geological
stage.
(Sp. 32) Hyopsodus vicarius Cope.—The type of this species
(Amer. Mus. 5003)
is a single worn
molar tooth, very
uncharacteristic.
We have _ provi-
sionally associated
with it a finely pre-
served .jaw, Am.
Nas. “Coll., ~ :No:
7304 o. £2!
(Sp. 36) Hyop-
a eel sige es een ences ee
distans Marsh.— A
small animal; inferior true molars = 12.5;mm. In separating
this genus Marsh mistakenly supposed that the true Hyopsodus
had but two superior in-
cisors. The codssification
of the symphysis is pos-
sibly a valid specific dis-
tinction of H.distans from
H. paulus in which the
jaws are usually found
with the symphyses im-
perfect ; the symphysis
is lacking. in ‘the 2:
paulus type but codssified
in the skull described
above, so that the present
character of this species
Fig. 13. Hyopsodus marshi. Am. Mus. No. c 5
17o6a. Slightly less than twice natural size. 1S entirely a matter of
Dental series too convergent in figure.
conjecture.
(Sp. 56) Hyopsodus marshi, sp. nov.
Among the Bridger specimens in the American Museum
is a pair of upper jaws (No. 1706a) with a very perfectly
188 Bulletin American Museum of Natural History. [Vol. XVI,
preserved molar series which may be taken as the type of a
new species. It is probably from the upper Bridger (Fig. 13).
The second, third, and fourth superior premolars show well
developed internal cusps, thus differing widely from the
Wasatch and Wind River specimens. The superior molars
(m1—m* = 13 mm.) are quadrate with a well developed hypo-
cone now almost as prominent as the protocone; the para- and
metacones are conic and there is no trace of a mesostyle.
4. Uinta (LIGURIEN) STAGE.
The genus in this stage is represented by three specimens:
two parts of jaws, Nos. 2078, 2078a, also the molar series
No. 2079, which unmistakably belongs to Hyopsodus, but
indicates a new species.
Fig. 14. Hyopsodus uinten-
szs. Am. Mus. No. 2079. Twice
natural size. Type.
Fig. 15. Hyopsodus uin- Fig. 16. Sarcolemur furcatus. Type. Am. Mus. Cope,
tensis. Am, Mus. No, 20782. No. 5008. A, superior, A’, internal, A?, external views of
Twice natural size. teeth and jaw. Twice natural size.
(Sp. 57) Hyopsodus uintensis, sp. nov.
Type No. 2079, Am. Mus. A right maxilla containing p*—m}3.
Superior molars with broad, well defined external cingulum, but no
mesostyle; m'—m* = 12.5 mm. Hypocone feeble or wanting on m°.
The level is Horizon C or the upper true Uinta beds of Utah.
=
1902. | Osborn, American Eocene Primates, 189
Genus SARCOLEMUR Cope.
Entomodon Marsh, Antiacodon Cope.
1. BRIDGER (BARTONIEN) STAGE.
The type of the genus is the jaw of the species S. (Antia-
codon) furcatus Cope, belonging to an animal about the same
size as Hyopsodus paulus but well distinguished by the
presence of a prominent paraconid which is closely connate
with the metaconid, also by the more crescentic form of the
external cusps (protoconid and hypoconid), and by the more
elongate form of the fourth premolar (Fig. 16). Three in-
ferior molars = 15 mm.
S. pygmaeus Cope. S. furcatus Cope.
P, simple, lacking internal cusp. P, complex, elongate, with promi-
nent internal cusp.
(Sp. 11) Sarcolemur pygmzus Cope.—This species, from the
Bridger basin of Wyoming, was originally referred by Cope to
Hyopsodus. It bears a su-
perficial resemblance to M1- cKkBaB
crosyops but the _ simple Ee ee © ae
structure of p, and the more :
elevated and connate para-
and metaconids readily dis-
tinguish it; the hypoconulid
onm, andm, is quite distinct 4... sarcolemur pygmaeus. Coe
as in Hyopsodus, enabling us Am. Mus. No. soon fat ramuy germ
to readily distinguish this
type of molar from that of Anaptomorphus.
(Sp. 31) Sarcolemur furcatus Cope.— Represented by the
type lower jaw in the American Museum (No. 5008) con-
taining the fourth premolar and three molars. With this
should be compared the prior type of (Sp. 24) Entomodon
comptus Marsh, a fourth lower premolar closely resembling
that of Sarcolemur; if identical Entomodon has priority.
190 Bulletin American Museum of Natural History. (Vol. XVI,
Famity NOTHARCTIDZ Oszorwn.
Limnotheride Marsh.
The type genus, Limnotherium, is apparently preoccupied
by Notharctus Leidy; if not, Limnotheride takes precedence.
Definition.—Upper incisors early reduced to 3. Premolars persis-
tent, but with reduced fangs. Molars relatively low crowned and low
cusped; lower molars especially like those of many true Monkeys in
the broad depressed and early crenulate or tuberculate talonid. Para-
conid gradually reduced. External cusps of upper molars more cres-
centic than in Hyopsodus or Adapts, consequently developing a
meso-style; also showing external cingulum.
Unlike the Hyopsodontide these animals show a pro-
gressive increase in Size in as-
cending levels. This phylum
certainly includes the numerous
Wasatch and Wind River species
referred to Pelycodus ;by Cope,
the Bridger species referred to
Notharctus and Htpposyus by
Leidy; to Hyopsodus (in part),
Limnotherium, Thinolestes, and
Telmatolestes by Marsh; to Pro-
totomus and Tomitheritum by Cope.
Fig. 18. Stxopa(Prosinopa)eximtia. : . °
Type. Phila. Acad. Twice natural The species Sinopa (Prosinopa)
yeas eximia Leidy was based upon a
jaw containing ps3, p,, which should be compared with
Notharctus.
To be compared with this family are the small animals,
Omomys cartert, Leidy (7i. s,
c. 7, p.2); and Hemiacodon
gracilis Marsh (5,7,3, 3-);
true molars=11 mm., molars
ss}
i
1 ma) men emi pan po. mz ce
Fig. 19. Omoneyscarterz. Type. Superior
and premolars Sap 105 peed acahealy view of right ramus, traced from a photograph.
Twi atural size.
Also H. nanus Marsh, H. “\.
pucillus Marsh ; the two species last named, however, should
also be compared with Anaptomorphus.
1902. | Osborn, American Eocene Primates. IgI
CHRONOLOGICAL LIST.
Species of Pelycodus.
(35) Pelycodus (Proisiomus) sarrovit Cope...4,..220008 6. 0a 8 Wasatch
(38) (Tomitherium) frugivorus Cope. . Se relates 4s
i ie LU SEC OD CER et a ny ee ahs ace ets ee
(45) - MUTUCCIL US: CONG a Sere Ge bo oR Aste ahha Sate Ste Wind River
Species of Notharctus and Allied Types.
(4) Notharctus LENCO OSUSMIECIGN Aken Ron oe crore soso hea ee Bridger
(6) (Limnotherium) DMGnnuseMiarshl tes eee cieiaie a
(7) ss CLES AVSIM ASI ooh ye gaat se Ha re
(xo) a (Hip posyus) formosus Leidy.. Se eas OOe
(12) TooUstior Welding wae eee te ty
(73) ‘i (Thinolestes) anceps Marsh.................. *
(14) a (Lelmatolestes) crassus Marsh................. “
(15) a (Limnotherium) affints Marsh................. Pe ite:
(27) a (Tomuthertum) rostratus Cope................- SEN,
PELYCODUS. NOTHARCTUS.
Jaw elongate. Mandibular sym- Jaw stout. Symphysis typically
physis uncodssified. Superior coéssified. Superior molars
molars triangular with rudi- quadrate, with pronounced
mentary hypocone; no meso- hypocone; a mesostyle.
style.
Rpt pig pi4 mb m2 m8
Fig. 20. Evolution of molars in Notharctide. <A, Pelycodus
Srugivorus, Wasatch; B, Notharctus nunienus, Wasatch; C,
Notharctus sp. indet. Bridger.
192 Bulletin American Museum of Natural History. [Vol. XVI,
Genus PELYCODUS Cope.
Pelycodus is distinguished generically from the later mem-
bers of the family by the more tritubercular upper molars,
which exhibit the hypocone in all stages of development
(Fig. 20) and the quinquetubercular lower molars which
correspondingly show the paraconid in various stages of de-
generation (Fig. 22). P. frugivorus is more tritubercular;
P. tutus is more sexitubercular; a constant distinction from
the contemporary Hyopsodus in the upper molars is the
presence of an internal cingulum in Pelycodus which is wanting
in Hyopsodus. In the Wind River specimens we also note
the rise of the external intermediate column or mesostyle in
a manner precisely analogous (homoplastic) to its develop-
ment in the equine Perissodactyla. In other words the
Lower Eocene Pelycodus is in a lower stage of evolution from
the tritubercular (tuberculo-sectorial) type than its successors
in the Middle Eocene (or Bridger) stage (Fig. 20).
Dentition.—In examining the rich Cope collection, now in the
American Museum, the incisors are apparently 3; the upper
pairs are conical and not spaced; the canines are slightly
enlarged and erect; the dental series is somewhat spaced,
that is, the first and second premolars are not crowded (Fig.
21). The lower molars show traces of a hypoconulid; the
first premolar is usually single fanged, and exceptionally bi-
fanged; the crowns of pms 1-2 are simple. The third and
fourth upper premolars show a single external cusp (proto-
cone) and an internal cusp (deuterocone) while the fourth
lower premolar is also slowly transforming into the molar
pattern by the addition of a tritoconid. The upper molars
may be clearly distinguished from those of the Microsyops line
by the stronger development of the intermediate tubercles or
conules (Fig. 20), which are exceptionally progressive, also
by the more rounded or quadrate contour.
In the following descriptions dependence is placed largely
upon the specific determinations made by Cope himself. The
species undoubtedly require careful reéxamination.
Skeleton.—In the Wasatch species the jaw (P. tutus) ‘is
1902. | Osborn, American Eocene Primates. 193
stout but not very deep with a well rounded border and an
uncodssified symphysis ; the jaw increases in depth in the
Wind River species (P. nunienus, Fig. 22). Many portions
of the skeleton have been described by Cope, including meta-
carpals and digits, also a clawed terminal phalanx (as in the
second digit of the lemuroid pes); unfortunately we must
consider this association as somewhat doubtful. The femur
(P. tutus) has a pit for the ligamentum teres and a long crest
below the great trochanter. The radius has an oval head.
The scapula has a prominent coracoid process. The head
of the astragalus (P. jarrovit, Coll. U.S. Nat. Mus.) is convex
and prolonged beyond the caleaneum. The caudals are long
and slender.
1. WASATCH (SPARNACIEN, YPRESIEN) STAGE.
Common Characters.—Superior molars more or less triangu-
lar, with rudimentary hypocone, without mesostyle.
(Sp. 35) Pelycodus jarrovii Cope.— This, the first species
described, is represented by a rather imperfect type in the
National Museum.
Fig. 21. Pelycodus frugivorus. Am. Mus. No. 65. Slightly less than 2 diam.
(Sp. 38) Pelycodus frugivorus Cope.— Includes the smaller
and more primitive Wasatch specimens in which there is a
variable rudiment of the hypocone in the upper molars (see
{ Fune, 7902.] 13
194 Bulletin American Museum of Natural History. [Vol. XVI,
especially Nos. 55, 65, 4174, 4182, Amer. Mus.); the upper
molars are strictly tritubercular, with little or no indication
of a mesostyle. This is certainly a very primitive species.
and it probably comes from the lower levels of the Wasatch
beds: Inf mis; = 35.5:
(Sp. 43) Pelycodus tutus Cope.— This, on the contrary, 1s.
the largest (inf. m. series=17.5 mm.), most progressive, and
most abundant species. The superior molars are triangular
in form but show a well developed hypocone forming a double
internal lobe, but no mesostyle (see No. 4162, Am. Mus.).
The paraconid, which is always the first primitive element to
disappear among the Primates, shows every stage of position
and development; it sometimes appears on m,—m,, but is.
always distinct on m,.
Fig. 22. Pelycodus nunienus. Type.
Am. Mus. Cope, No. 4734.
Genus NOTHARCTUS Leipy.
2. WIND RIVER (LUTETIEN) STAGE.
As we might expect, in the Wind River specimens the tirst.
lower premolar is always single fanged, while in the superior
molars the hypocone is decidedly more prominent so that in
some cases they might be described as quadrate and sexi-
cuspidate; a very conspicuous difference is the presence of
the mesostyle (Fig. 20). The Wind River species are also
generally distinguished by the more advanced transformation
of the posterior premolars. As observed by Matthew, the Wind.
River species show closer affinities to those of the Bridger. In
fact the Wind River specimens may well be referred to the
Bridger genus Notharctus as characterized below. Cope was
entirely mistaken in identifying the progressive Wind River
species with the older Wasatch species (P. jarrovit and P.
tutus).
a a
EEE
1902. | Osborn, American Eocene Primates. 195
(Sp. 45) Notharctus nunienus Cope.— Includes the smaller
Wind River forms (m,-m,=15mm.). We find a strong
deuterocone (internal cusp) on pm,.
(Sp. 58) Notharctus venticolus, Sp. nov.
A much larger monkey (No. 4715, Am. Mus.) was referred
by Cope to P. tutus but is clearly distinguished from this
older Wasatch species by the presence of a mesostyle in the
upper molars, and by the more progressive character of the
grinding teeth throughout m—m, = 17. The type (No.
4715 b) was figured by Cope, ‘ Tertiary Vertebrata,’ Pl. xxv,
figs. 1, 2. Other specimens are No. 4726 (op. cit. fig. 3), Nos.
4728, 4738.
3. BRIDGER (BARTONIEN) STAGE.
The gradual steps toward sexituberculy in the upper molars
and quadrituberculy in the lower, begun in Pelycodus, lead
directly into a number of Middle Eocene (Bridger) forms,
mostly of larger size and on a higher plane of general develop-
ment, baptized by Leidy, Cope, and Marsh with an unusual
number and variety of names, as shown in the chronological
table of species above. The identification of all these genera,
however, needs confirmation by further comparison of types.
Unfortunately we have an incomplete record of the levels
at which the types of these species and genera were found,
but it is important to remember that the Bridger was a
very long period, with time for the marked progression in
dental structure observed in various specimens which may
provisionally be referred to the single genus Notharctus. While
there was considerable range of progression from the lower
to the higher forms we cannot at present specify any single
generic character which will enable us to clearly subdivide
the Bridger species into different genera, because the pro-
gression although on a higher scale, is precisely analogous to
that observed in the transition from Pelycodus frugivorus to
P. tutus.
1960 Bulletin American Museum of Natural History. [Vol. XVI,
Notharctus has the same dental formula as Pelycodus but is
readily distinguished by the codssified mandibular symphysis,
the chisel-shaped incisors, the usually single fang of the first
and sometimes of the second lower premolar, the compara-
tively well developed hypocone and the subquadrate shape of
the upper molars; the usually marked reduction or absence
of the paraconid in the lower molars.
£8 pd om Ae m.3
Fig. 23. Notharctus sp. indet. Princeton Mus, No. 10,020,
Thanks to the excellent type selected by Leidy, N. tene-
brosus, to the beautiful specimen of 7. rostratum (Am. Mus.
No. 5009), and to the very complete series belonging to the
Yale Museum, this genus is by far the best represented of all
the American Primates, or in fact of any sub-Pliocene form
excepting possibly Adapis. It is most interesting to observe
the exact homoplasy between the variations in the cheek
teeth with those seen in the early Ungulates.
(Sp. 4) N. tenebrosus Leidy.— The type (Phila. Acad.) is a
relatively primitive species in which the second premolar is
still bifanged, and there are traces of the paraconid on all the
true molars; the third lower molar has a well developed heel
or hypoconulid, the fourth premolar is sub-molariform. On
the other hand its progressive specialization is marked by
the low uniform wearing surface of the molars, the primitive
trigonid being almost as depressed as the talonid; also by
the large erect canine (probably indicating a male individual),
by the well defined chin, by the unbroken dental series, and
by the elevated condyle.
Furst Stage? Lower Bridger.
A stage beyond N. tenebrosus is the species or variety
1902. | Osborn, American Eocene Primates. 1Q7
(Sp. 13) N. (Thinolestes) anceps Marsh, in the type of which
the second lower premolar is bifanged but the paraconid has
disappeared upon the second and third molars, which are now
truly quadritubercular. The mandibular dentition is other-
wise closely similar to that of N. tenebrosus; the lower jaws
are codssified, with the suture visible externally. In the upper
molars of this important specimen we find the crown sub-
triangular, the primitive triangle with distinct intermediate
tubercles, but the hypocone is prominent and well separated;
there is also an external intermediate cusp or mesostyle; the
first upper premolar is small orrudimentary. The third upper
premolar has a broad internal cingulum, the fourth is submo-
lariform.
Second Stage.
(Sp. 6) N. (Limnotherium) tyrannus Marsh.— (Type,
Yale Mus.). The second specimen named in 1871 was also
founded upon a lower jaw. Marsh described this as a “‘ pachy-
derm’’ and distinguished the genus Limnotherium from
Notharctus by the single fangs of the first and second lower
premolars, by the quadritubercular lower molars ‘‘with a
rudimentary double tubercle on the anterior margin’’ (para-
conid). This type was probably found upon a somewhat
higher level than N. tenebrosus. It marks perhaps the next
higher stage of evolution in which the first and second pre-
molars have single fangs; the paraconid is a vestigial tubercle
seen on all three lower molars; the third upper molar is.
tritubercular. Close to this stage is the type of
(Sp. 27) N. (Tomitherium) rostratum Cope, with small,
spaced, first and second lower premolars; in the latter the
fang is still grooved.
Related to these are the more slender jaws forming the
type of
(Sp. 15) Limnotherium affine Marsh. In this beautiful
specimen, belonging to a young individual, we note a slight
progression in the cheek teeth, the fourth upper premolar
differs from the first molar only in the absence of the
198 Bulletin American Museum of Natural History. (Vol. XVI,
hypocone, and sexituberculy is slightly more marked in the
true molars than in N. anceps; the first and second lower pre-
molars are single fanged, and, correlated with the development
of the hypocone above, the paraconid has degenerated and
disappeared upon the first and second molars below.
(Sp. 7) N. (Limnotherium) elegans, according to Marsh, is
a much smaller but related species.
(Sp. 5) N. (Hyopsodus) gracilis Marsh exhibits a paraconid
on the lower molars and thus either antedates Notharctus
(Limnotherium) elegans or is possibly referable to Sarcolemur.
Third Stage.
It is evident from the study of the foregoing series that
this line would inevitably terminate in sexituberculy above
and quadrituberculy below. This condition is fulfilled in the
type of
(Sp. 14) N. (Telmatolestes) crassus Marsh, a large species
in which the first and second upper molars are nearly quad-
rate in form and bear six tubercles, the hypocone being almost
as large as the protocone on m!' and m?; the hypocone how-
ever is wanting on m*%. The lower molars still retain a
faintly developed paraconid. Near this stage is
(Sp. 10) Hipposyus formosus Lezdy, founded upon a single
upper molar (Leidy, ’73, plate vi, fig. 41).
General Characteristics of the Teeth.— Notharctus was very
abundant in the Bridger period and as seen in the above
analysis presented progressive variations which are certainly
due to the passage from lower to higher geological levels.
The two pairs of incisors are compactly placed, with
chisel-edges as contrasted with the rounded incisors of
Pelycodus; the opposite pairs are spaced, that is there is a
slight interval between them. The unworn lower molars are
elongate (Fig. 23); they exhibit a transverse anterior crest
(metalophid) between the protoconid and metaconid, in front
of which is an oval valley bounded internally by the para-
conid in all stages of degeneration; behind this ridge is the
1902. | Osborn, American Eocene Primates. 199
posterior basin or talonid in which the enamel is waving or
crenulate; the hypoconulid has disappeared excepting on the
broad heel of m,; the lower molars are thus very similar to
those of monkeys and it is a complete surprise to find the
upper molars with greatest diameter transverse and almost
indistinguishable in pattern from those of the contemporary
horses such as Orohippus. The protocone forms a low trans-
verse crest with the protoconules (this is a rudimentary
protoloph mechanically correlated with the metalophid
below), while the hypocone and metaconule are isolated; the
external cusps (paracone and metacone) are compressed
with apical ridges running into an external intermediate
mesostyle; the anterior cingule, or parastyle, is also developed.
The fourth upper premolar is submolariform, with three
large cusps (protocone, tritocone, deuterocone); it also ex-
hibits traces of the conules (Fig. 20). The fourth lower pre-
molar has an elevated protoconid connected by a low crest
with a tritoconid and a deuteroconid.
This genus is finely represented in the American Museum
by the skeleton and teeth, No. 1727; the upper molars are in
a very progressive (Telmatolestes, Hipposyus) stage.
Famity ANAPTOMORPHIDZ Cope.
Definition.—Skull brachycephalic. Post-orbital process. Facial por-
tion of lachrymal greater than orbital; fossa lacrymalis in front of crista.
Premolars reduced, ;=,. Grinding teeth arched, molars compressed
antero-posteriorly, extended transversely, tritubercular, rudimentary
hypocone. Short, deep, lower jaw. Lower molars with elevated
trigonid region, reduced paraconid, no hypoconulid.
- This family is represented by the type jaw of A. emulus
from the Bridger, the famous skull of A. homunculus from the
Wasatch, together with portions of four isolated jaws; also
possibly by a larger Upper Eocene species Microsyops uin-
tensts.
Prior to Cope’s description of the Bridger jaw are a num-
ber of specimens named by Marsh, as shown in the following
chronological list, which possibly are related to this family or
to the genus Omomys,
200 Bulletin American Museum of Natural History. [Vol. XVI,
CHRONOLOGICAL LIST OF SPECIES.
Sp. (22) ? Hemtacodon gracilis: Marshi. 5230-57 seine Bridger.
(a2)in2 i nanus |) nalvatyes Blache eae tenn eara eae x
(23) ? “ DUCTUS ON Sapiens Gee ss
(26) ? (Paleacodon) vagus Marsh............... of
(28) ? Anaptomorphus emulus Cope.............0.+.2e0005
(39) ? Pelycodus angulatus Cope.. ....Wasatch (Indeterminate).
(47) Anaptomorphus homunculus Cope. ..............- Wasatch.
(54) " (Microsyops) uintensts Osborn....... Uinta.
Other animals to be
considered in this con-
nection are Leidy’s
Washakius insignis, Fig.
23a, ‘Microsyops’ speiri-
anus Cope, Fig. 37,
Nig. 24. Anapromordies* and Paleacodom™ wee
homunculus. * ss m.
Mus. Cope. Skull, No. 4104;
partly reconstructed from the Marsh.
cad. External.su- two sides. Jaw from another
perior, and internal specimen, Am. Mus, No. 43. te ae BEY Us:
views. whestoel ous: born was mistaken (Os-
born and Wortman1892,
p. 102) in referring to this family the genus and species
Omomys cartert Leidy.
Fig. 23a. Wa-
shakius insignis.
Type. Phila. A-
Genus ANAPTOMORPHUS Cope.
Paraconid reduced, hypoconulid absent except on m,, canines of
medium size.
rt. WASATCH (SPARNACIEN, YPRESIEN) STAGE.
(Sp. 47) Anaptomorphus homunculus Cope.—Type skull,
Amer. Mus. Cope Coll. No. 4194.
Definttion.—i.2, c.}, p-?, m.2. M ,.-m,, with reduced paraconid,
P. with very slight rudiment of deuterocone.
The species is represented by the famous type skull, also
by specimens Nos. 41, 44. Paraconid on m, somewhat larger
than in the Bridger A. emulus; the deuteroconid is barely
visible on p,; there is a very small alveolus for the root of
the second lower premolar.
The skull has been refigured with care (Fig. 24) to exhibit
1902. | Osborn, American Eocene Primates. 201
its principal characters. In reference to Forsyth-Major’s
(1901) very precise examination of the lachrymal in the Lem-
uroidea and Anthropoidea it is important to note that this
bone in Anaptomorphus resembles that in the. Lemurs, espe-
cially such a form as Opolemur (op. cit., p. 139, text fig. 37),
much more closely than.it does the lachrymal of Adapis or
of any of the Anthropoidea, in the following respects: (1)
the pars facialis is broader than the pars orbitalis; (2) the
lachrymal fossa is extra-orbital, being bounded posteriorly by
the crista posterior lacrymalis which forms the anterior rim of
the orbit. In the words of Forsyth-Major: ‘“‘In Lemurs, as
a rule the crista lacrymalis posterior rides on the lower orbital
margin, of which therefore it forms a portion . . . the
anterior part of the lachrymal thus becoming the pars jaczalis,
the posterior part the pars orbitalis . . . Asa result, we
have the lachrymal fossa outside the orbit. . . .” (op. cit.
p. 134). Anaptomorphus re-
sembles Chrysothrix in the re-
duplication of the infra-orbital
foramen.
This sustains Cope’s state-
ment (2884, p. 250) and
definitely proves that in the
structure of its lachrymal
Anaptomorphus is lemuroid; it
does not, however, prove posi-
tively that it is a Lemur.
The transversely extended
form of the upper molars and
premolars is correlated with
the brachycephaly of the skull;
the molar pattern being best
indicated in Fig.25. Observe
especially the depression and gy 4. a paptomor Maslin cae
transversely oval form of the Am. Mus No. 4! meet Dae
superior teeth, the relatively
broad short crowns of the inferior teeth, the trigonid and
talonid being of approximately the same width.
202 Bulletin American Museum of Natural History. {Vol. XVI,
The species Pelycodus angulatus Cope was based upon a
type (Nat. Mus.), now unfortunately lost, containing a single
lower molar which resembles that of Anaptomorphus, also that
of Cynodontomys.
indeterminate.
2.
Fig. 26. Axnafto-
morphus amulus,
ype) —eAm™ Mis.
Cope, No. soto. A,
from side; A‘, recon-
structed from above.
(Sp. 54) ?‘* Microsyops’’ uintensis
In the absence of the type this species is
BRIDGER (BARTONIEN) STAGE.
(Sp. 28) Anaptomorphus emulus Cope.—
Type, No. 5010, Amer. Mus. Cope Coll.
This famous little jaw (Fig. 26) is the type
of the genus. It exhibits progression on the
Wasatch species in the loss of the second
premolar, ‘the formula; beingey 3. ee.
P. 5, M. 5; paraconid especially on mg is
also slightly more reduced, while the deu-
teroconid on p; 1s slightly more pronounced,
but still not separate.
3. UINnTA (LIGURIEN) STAGE.
INCERT SEDIS.
Osborn.—A reéxamination of the type
of Microsyops uintensis (Amer. Mus.
No. 1899) demonstrates that the
reference of this type to Microsyops
was an error, because the fourth lower
premolar is totally unlike the molars.
Its nearer reference is either to the
Anaptomorphide or to some member
of the Notharctide.
Fig. 27. ‘* Microsyops”’ uin-
tensts, Type. Am. Mus. No.
1902. | Osborn, American Eocene Primates. 20
» GQ
Part II. RopDeENTIA.
SUBORDER PROGLIRES, subordo nov.
A primitive suborder of Rodents distinguished by the presence of
rooted incisors, and canine teeth, and by the absence of any consider-
able diastemata and of antero-posterior motion of the jaw. Types:
Mixodectes, Olbodotes, Microsyops.
It is obvious that these animals are far too primitive to
be classed with the Protrogomorpha of Zittel which was framed
to include all those modernized fossil and living rodents which
do not naturally enter either of the four great divisions of
Brandt.
Famity MIXODECTIDZ Cope.
Characters.—Median lower incisors close to symphysis, enlarged and
elongating (unlike Tillodontia, in which second incisor is enlarged),
lateral incisors early reduced; canines persistent (unlike Rodentia) ;
no diastemata (unlike Rodentia), first and second premolars rapidly
reduced; third premolar slowly reduced, fourth premolar progressively
molariform (as in Tillodontia and Rodentia); lower molars with nar-
row, slightly elevated trigonid, but early reduced paraconid; talonid
broad, hypoconulid small, except in third lower molar; superior molars
tritubercular. A feature of the jaw is the sharp definition of a ridge
descending from the coronoid and defining the masseteric insertion
anteriorly (Fig. 3).
This phylum specialized very early.’ The little animals
which represent it are rare in the Torrejon and Wasatch,
more abundant in the Wind River, and very common in the
Bridger; not as yet reported in the Uinta. The specific
forms range greatly in size but the essential progressive
characters of the lower teeth are the same throughout this
long geological period.
Ordinal, position. — Cope placed Mixodectes among the
Primates. Matthew (’97, p. 265) was the first to point out
that the enlarged median tooth was probably an incisor and
that the astragalus was exactly similar to that of a Rodent.
He therefore took the important step of transferring this
1In 1892, Schlosser (Neues Jahrb. f. Min. Geol. u. Pal., Bd. II, s. 238) referred the
contemporary Cernaysian Plesiadapide, Plesiadapis and Protoadapis Lemoine, to the
Rodentia, removing them from the Insectivora.
204 Bulletin American Museum of Natural History. (Vol. XVI,
genus to the Rodentia, leaving the position of the more recent
members of the family undetermined.
Relationship to the Rodentia is now found to be indicated by:
(1) progressive elongation of median incisor; (2) disappear-
ance of lateral incisors; (3) reduction
of canines; (4) disappearance of two
anterior premolars and reduction of
third premolar; (5) transformation of
fourth premolar into molar form, thus
foreshadowing a homodont molar-
premolar series; (6) width and ex-
tension of talonid (as in Eocene
Paramys); (7) rodent form of as-
tragalus.
Against the Rodent relationshtp are:
(1) Persistence’ ei the ‘canine; (2)
absence of diastemata; (3) absence
of any evidence (except the levelling
An Me No eaets greens of the premolars). of adaptation, \for
Nao ee eee antero-posterior or oftial motion of
or distal aspect. Natural'size. the jaw. Pendine the tinal demon.
stration of this problem the Mixo-
dectidea may be placed in the new
primitive suborder Proglires, defined above.
A careful reéxamination of all the material belonging to
Mixodectes, Cynodontomys, and Muicrosyops has confirmed
Matthew’s observation that the enlarged median tooth is an
incisor and has convinced us that these animals represent
three successive stages in the same family.
A still more primitive stage is represented by a new genus
to which the name Olbodotes (oABodérns) may be given, in
reference to the happy solution it affords of the problem of
the homology of the enlarged incisor teeth.
SYNOPSIS OF GENERA.
TORREJON.
Olbodotes.— 5, 3, s, 3 One enlarged and two reduced incisors; two
premolars, fourth premolar pointed; depressed paraconid on the molars.
1902. | Osborn, American Eocene Primates. 205
Mixodectes.— ;, 7, y-z, y- One enlarged incisor tooth only; a canine,
three to two premolars, fourth premolar pointed; depressed paraconid
on the molars, a rudimentary hypoconulid.
WASATCH.
Cynodontomys.— ;, 7, 5, y- One enlarged incisor only, two pre-
molars, fourth premolar submolariform; a small paraconid and hypo-
conulid on the molars.
WIND RIVER AND BRIDGER.
Microsyops.— ;, 7, 5, y- Greatly enlarged incisor; two premolars,
third premolar further reduced, fourth premolar molariform; a small
paraconid on the molars.
CHRONOLOGICAL LIST OF SPECIES. PARTLY INCERT# SEDIS.
Sp: Gn) iewodectes pungens Cope =: . 5.0600 0s cea. es ss. oe LOrrejon.
(52) = crasstusculus sale FE NS AMR Ss TP a
(8) Mucrosyops gracilis Leidy. eae ode <c0 oo 5 ELC:
(9) os (Paleacodon) verus = leeidiye Cty dao ie «oe nea o
(18) i CE aiirodorn) ety pusaNiats tases a eee te eee
(19) i (Bathrodon) annectens Marsh............ “
(20) Ms (Mesacodon) speciosus Marsh........... oS
(46) of SCOUMAN USE CODE. an ek sy eae ee Wind River.
(49) a ens. latidens aie ... Wasatch.
(59) Olbodotes copet Osborn. A PUR At noaeha ate agdieedt ROLE CTOs
Compare also Indrodon malaris — OC mes :
rs Chriacus angulatus Cope................. Wasatch.
1. TORREJON (THANETIEN) BASAL EOCENE
(Sp. 59) Olbodotes copei, gen. et spec. nov.
Type, No. 2385, Amer. Mus., left lower jaw.
. Dentition.— x, ~, 3, x» An enlarged median incisor, two smaller in-
cisors on the alveolar border behind it; canine small; third premolar
reduced; fourth premolar high, simple, pointed, as in Mzxodectes.
This specimen had previously been referred to Mzixodectes
but it differs in the retention of three incisor teeth and the
loss of the second premolar tooth, which is represented by an
alveolus in the type of Mixodectes pungens. There is little
question about the presence of three incisors, the median
one of which, although not preserved, has been much enlarged
206 Bulletin American Museum of Natural History. [Vol. XVI,
so far as we can judge by its alveolus. The second and third
incisors are equal sized with laterally compressed fangs (Fig.
mi] tO
Fig. 29. Olbodotes copez. Type. Am. Mus. No. 2385. A, lateral view, 4’, anterior
view, of left ramus.
29A'). This further enables us to determine the single-fanged
tooth at the edge of the jaw as a canine, an interpretation
which is supported by the condition of this long
single-fanged tooth, heretofore described as an
anterior premolar, in Cynodontomys and Micro-
\ syops.
Hig aga) NOE With the type of Olbodotes is associated an
Uscet melas. upper molar tooth (Fig. 29a) resembling that
tpn £4 ¥Hh Of Indrodon malaris in the possession of a
prominent mesostyle, and suggesting that
Indrodon is probably a member of the Mixodectide. (See
below.)
(Sp. 51) Mixodectes pungens Cope.— The well known
type of this species (No. 3081 Amer. Mus.) shows no evidence
wed \ ¢
Fig.30. Mzzxodectes pungens. Type. Am. Mus. Cope, No. 3081.
of the existence of the reduced lateral incisors seen in Olbodotes
and is further distinguished by the variable presence of the
second premolar.
1902. | Osborn, American Eocene Primates. 207
Another specimen (No. 2557 b, Amer. Mus.) shows that
the enlarged incisor is still a spatulate tooth with the enamel
Fig. 31. MWixodectes pungens. Am. Mus. No. 25574. Left lower jaw. Median
incisor partly displaced.
completely surrounding the crown and a persistent fang
(Fig. 31). Still another specimen (No. 3083 Amer. Mus.)
gives a better view of
the molar teeth.
(Sp. 52) Mixodectes
crassiusculus Cope.—
The type, No. 3087
Amer. Mus., consists
of the posterior por-
tion of the right and
left rami of the lower
jaws containing molar
teeth. This specimen
gives a perfect view
of the structure of
the molars, showing
that they are even
more specialized than
those of Cynodonto- —=
rye ancl WMCTOSVOPS) ic 52, | arszodectes crassivecutus). Arm: Mus, Cope:
faethe depeneration | N°: 27 Pars of right and left rami.
of the paraconid.
208 Bulletin American Museum of Natural History. \Vol. XVI,
INCERTA SEDIS.
Genus INDRODON Cope.
As stated above (p. 170) this does not belong near the
Anaptomorphide, the molar structure being entirely different.
In previous articles, owing to the incorrect association of
another specimen (No. 823), there has also been much con-
fusion, which Matthew has partly cleared up (1897, p. 265).
Indrodon malaris Cope.— The type skull (Amer. Mus. No.
3080) is carefully redrawn in Fig. 33. Its conspicuous
characteristics are:
sightly enlarged
median incisors,
three premolars
well spaced, fourth
premolar with deu-
terocone ; molars
Ne noes Le ee ew ithaelbsoaclanen
ternal cingulum,
crescentic para- and metacones and prominent mesostyle,
rudimentary hypocone. The molar teeth resemble those of
Olbodotes.
Specimen No. 833 also belongs to
Indrodon but is more progressive than
I. malaris, being distinguished by the
breadth.of the ectoloph and additional
cusps on the fourth superior premolar.
Specimen No. 823 (Figs. 1, 2) was as-
sociated by Osborn and Earle (1895,
p- 17) and believed to give us the
- Skeletal characters of this animal; this eee ee.
19t1 1 Mus. No. 833. Fourth pre-
association appears very questionable TaGlE ees ealecoearletrraes
(see page 172).
2. WASATCH (SPARNACIEN, YPRESIEN) STAGE.
The genus Cynodontomys is only by courtesy and for want
of better knowledge separated from Microsyops.
(Sp. 49) Cynodontomys latidens Cope. ?Syn.M. (Chriacus)
angulatus Cope.—Type : the two rami of a lower jaw (Amer. Mus.
1902. | Osborn, American Eocene Primates. 209
No. 4195), with molar teeth (ms.=11.5 mm.) in nearly parallel series;
anterior pair apparently well developed and procumbent; lower pre-
molars spaced, p, with paired fangs; p, a sub-quadritubercular tooth,
namely with protoconid, tritoconid, deuteroconid, and tetartoconid;
molars with narrow trigonid, paraconid small but distinct and median
in position, broad talonid with a small hypoconulid. The ramus is
long and rather slender, and the angle is produced posteriorly.
Cynodontomys is barely distinguished from its successors in
the Wind River and Bridger by the less complete transforma-
tion of the fourth
premolar and by
the smaller single
incisor. It is repre-
sented also by the « '
juvenile jaw (No.
65), by the speci-
men (No. 4184) re-
ferred to Chriacus Fig. 35. Cynodontomys latidens. Type. Cope Coll-
angulatus by Cope. No. 4195 Wasatch. Big Horn Valley, Wyoming. Twice
natural size.
Cope erroneously
referred to this species a number of specimens from the
Wasatch and Wind River horizons, with a short deep
mandibular ramus, small heels upon the third lower molars,
and general structure more similar to the Bridger series
described below; ms. =13.5 mm.
3. WIND RIVER (LUTETIEN) STAGE.
Genus MICROSYOPS LE rpy.
Paleacodon Leidy, Bathrodon Marsh, Mesacodon Marsh.
(Sp. 46) Microsyops scottianus Cope.—Type: A long, shallow
mandible (Amer. Mus. No. 4748), large semi-procumbent tooth with
narrow diastema behind it; ms.=14 mm.; formula: j;, 7, 3, 3-
Fig. 36. MWicrosyops scottianus. Am. Mus. No. 4748. Slightly less than 2 diam.
[ Fune, 1902. | 1
210 Bulletin American Museum of Natural History. [Vol. XVI,
The eight specimens from the Wind River formation (Amer.
Mus. Nos. 4743-4748 inclusive), referred by Cope (Tertiary Ver-
tebrata, p. 217) to M. elegans Marsh (or
M. gracilis Leidy), represent a larger animal
than C. latidens of the Wasatch; and agree
closely with M. scottianus in size of the
teeth, but certain of. them differ in the
greater depth of the mandible and coal-
Bip yn i cerosy- escence of the fangs of p, (see No. 4743).
eeavrmes Tee (Sp. 44)“ Microsyops” speirianus (Fig.
4190. Portion of right = 34). The type certainly does not belong
to this genus. It resembles Anaptomorphus
slightly.
4. BRIDGER (BARTONIEN) STAGE.
(Sp. 8) Microsyops gracilis— Leidy’s type of the genus
(Microsyops gracilis of the Bridger) was a small lower jaw in
which he mistook the homologies and erroneously described
six molars (ms. and pms.) and enlarged “‘canines,”’ remarking
that the number of incisors was indeterminate. Unlike those
of the Anaptomorphide the lower molars are readily recog-
nized by the narrow trigonid, now depressed to the level of the
talonid, 7. ¢., more bunodont, depressed paraconid, behind
which is the broad talonid bearing a hypoconulid; m, has a
small cuspidate hypoconulid, unlike that in the Notharctide.
So far as reported, upper molars have not been found as-
sociated, but it is probable that they are rightly identified
in the broadly triangular (as distinguished from the more
transversely oval form of the molars in the Anaptomorphide
and the more quadrate form in the Hyopsodontide) tri-
tubercular teeth, with a small cingule representing the hypo-
cone, with intermediate spaces on the palatal side, as in all
forms in which the trigonid is present; rudimentary conules
and para-, meso-, and metastyles; the type of Paleacodon
verus Leidy, described immediately after that of Muicrosyops,
1s such a tooth with small conules and a rudimentary hypo-
cone! (Leidy,) 73, pl, vix fie, 46),
1902. ]
Osborn, American Eocene Primates. 211
It is probable that the types of Paleacodon verus Leidy,
Mesacodon speciosus Marsh, Bathrodon annectens Marsh, also
belong to this genus. The species
Paleacodon vagus Marsh apparently
belongs with the Anaptomorphide.
Leidy chose a rather uncharacteris-
tic specimen' as the type and first
adopted the specific name M. (Hy-
opsodus) gracilis Marsh;* but as the
type of the latter species (H. gracilis)
has four premolars it is probably
related to Notharctus (?Limnotherium)
p.4 mf m2
Fig. 38. Jzcrosyops. Princ.
Mus. Superior molars, iso-
lated.
elegans, as in fact suggested by Leidy himself (1873, p. 84).
We are unable at present to straighten out the names of the
Bridger formation species; they appear to be numerous.
Principal Characters of Microsyops.— 7, 7, x, x; median incisors very
large, semi-procumbent, laterally compressed; p, submolariform;
m, with small third lobe: symphysis not codssified.
The fourth premolar of Microsyops presents an advance
Am. Mus. No.
Fig. 39. Microsyops.
1732. A, external, 4’, internal aspect of
right ramus.
upon that of Cynodontomys
in the presence of a ridge
uniting the two anterior cusps;
the incisors were still larger
and more procumbent, ex-
tending well back, below and
inside of the fangs of the
canines (tig: 39), The ex=
ternal cingulum of the upper
molars is not constant, some-
times faint; the valleys are
smooth or slightly ridged. The
upper molars (Fig. 38) are
tritubercular, the primitive
cusps (protocone, paracone,
metacone), when unworn, be-
ing sharp and prominent; the
conules faintly developed on m' and m?; the hypocone is a
4 Leidy ’73 pl. vi, figs. 14, 17.
2 Amer, Jour. Sci., July, 1871, p. £0.
212 Bulletin American Museum of Natural History. [Vol. XVI,
mere cingule on the postero-internal slope of the protocone.
The posterior heel (hypoconulid) of the lower molars is always
very small, but sometimes forms a sharp narrow heel upon
m,; these teeth are further distinguished by the narrow
primitive triangle; the paraconid is distinct on m,, but de-
creases on m,,m,. The lower premolars are reduced. |
(Sp. 20) Microsyops (Mesacodon) speciosus Marsh.—Type,
a complete lower jaw (Yale Museum), lacking the incisive
border and the articular portion. The measurements corre-
spond with those of the other Bridger small species. The
Fig. 40. MWicrosyops? annectens. Princ. Mus. Portions of rami of two indi-
viduals. Lower figure slightly exceeding 2 diam.
identification with Microsyops is due to the sub-molariform
pattern of p,. As observed by Marsh the anterior tooth 1s
large, compressed, almost in contact with the symphysis; sym-
physis not codssified; lower border of jaw produced posteriorly
angle slightly inflected; trigonid of lower molars very distinct.
(Sp. 18) Microsyops (Bathrodon) typus Marsh.— Type, a
lower jaw (Yale Museum), containing the molar teeth (m,-—
m,;=12 mm.) resembling that of Microsyops and Mesacodon,
1902. | Osborn, American Eocene Primates. 213
coinciding in measurements and description, although the
crucial tooth, p,, is wanting.
(Sp. 19) Microsyops (Bathrodon) annectens Marsh.—Type,
Yale Museum. This corresponds with the large specimens at
the Princeton Museum (Fig. 40), but the paraconid is less
elevated; the trigonid is narrow and the talonid broad, with
three distinct cusps. The Princeton specimens have a deep
jaw, with an unusually large procumbent incisor; the canine has
a long single fang; p, is possibly represented by a rudimentary
socket; the trigonid narrow and slightly elevated; the para-
conid more or less distinct on m,—m,; hypoconulid faint on
m,, m,, strong on m,. The upper molars, which are pro-
visionally associated with this species, bear a low, external,
intermediate cusp or mesostyle, minute conules, and a low
cingule representing a hypocone.
BIBLIOGRAPHY.
(The references to the descriptions of the types will be found in the
table, pp. 172-175.)
1884. Cope, E. D. Tertiary Vertebrata. 4to. Washington, 1884.
1897. EARLE, CHARLES. On the Affinities of Tarsius: A Contribu-
tion to the Phylogeny of the Primates. American Natural-
ast, Vol. XXXI, No. 367, July, 1897, pp. 569-575; zbid.,
August, 1897, pp. 680-689.
1896. Lrcue, W. Untersuchungen ueber das Zahnsystem lebender
und fossiler Halbaffen. Festschrift fir Carl Gegenbaur, pp.
128-166, Leipzig, 1896.
1901. Major, C. I. ForsytH. On Some Characters of the Skull in
the Lemurs and Monkeys. Proc. Zodl. Soc. London, Feb. 19,
1900, pp. 129-153, pll. xi—xiil.
1897. MattHew, W. D. A Revision of the Puerco Fauna. Bull.
Amer. Mus. Nat. Hist., Vol. 1X, Art. xxii, Nov. 16, 1897,
PP. 259-323.
1895. Ossporn, Henry F., and CHARLES EaruteE. Fossil Mammals of
the Puerco Beds. Collection of 1892. Bull. Amer. Mus.
Nat. Hist., Vol. VII, Art. i, March 8, 1895, pp. 1-70.
1892. Osporn, Henry F., and J. L. Wortman. Fossil Mammals of
the Wasatch and Wind River Beds. Collection of 18g91.
Bull. Amer. Mus. Nat. Hist., Vol. IV, No. 1, Art. xi, Oct. 20,
1892, pp. 81-148.
214 Bulletin American Museum of Natural History. (Vol. XV1.]
Ossporn, Henry F. Fossil Mammals of the Uinta Basin.
Expedition of 1894. Bull. Amer. Mus. Nat. Hist., Vol.
Wil, Arta, May 18; 1805, pp. 71-05;
1887. ScHLossER, Max. Die Affen, Lemuren, Chiropteren, Insectiv-
oren, Marsupialier, Creodonten und Carnivoren des Euro-
paischen Tertiars und deren Beziehungen zu ihren lebenden
und fossilen aussereuropaischen Verwandten. Separat-
Abdr. aus: “ Beitrage zur Paldontologie Osterreich-ungarns,”’
VI. Band, Theil I, s. 1-224. 4to. Wien, 1887.
1901. WortTMAN, J. L. The Probable Successors of Certain North
American Primates. Science, N. S., Vol. XIII, No. 319,
Feb. 8, 1901, pp. 209-211.
1895.
F =4
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— Mew C anida from the Miocene of ae
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By W. D. Marrnew. —
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_ AUTHOR'S EDITION, extracted from BULLETIN __
OF THE
mevican Museum of Hatnval History,
: VoL. XVL ARTICLE XXI, pp. 281-290.
2 New York, Sept. 18, IQO2.
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The Knickerbocker P
Article XXI.—NEW CANIDA FROM THE MIOCENE
OF COLORADO.
By W. D. Matruew.
Cynarctus, new genus.
Family Canide, Subfamily Amphicyonine.
Dentition 34.4.3. Carnassials reduced and molars enlarged, talonids
bicuspid in the type species, and two accessory cusps on the trigonids.
Jaw long and slender as in the dogs, premolars cynoid.
Cynarctus saxatilis, n. sp.
Size of the Coyote. Saw somewhat more slender anteriorly, in-
ferior border more convex, angular process longer, curving more up-
ward and inward. Coronoid process more triangular, the tip narrow,
Lower jaw, outer side, x 2, and inside view of teeth.
Fig. 1. Cynarctus saxatilis.
Type, No. 9453. Loup Fork (Pawnee Creek Beds), Colorado.
the anterior border with slight uniform backward curvation, while in
the coyote it is straight until near the tip and then curves suddenly
backward. Masseteric fossa deeper and wider, its inferior border
much more marked.
Teeth. Incisors not preserved. Canine and premolars little worn,
molars much worn. Premolars somewhat smaller than in C. latrans,
carnassial nearly one-fourth smaller, molars over one-fourth larger.
The trigonid of the carnassial is low, and reduced in size, and two
[281]
282 Bulletin American Museum of Natural History. |Vol. XVI,
accessory cusps are added to it, one external to the protoconid, one
behind the metaconid. The protoconid is central in position, greatly
reduced in proportionate size compared with Canis or even with Am-
phicyon, and the shearing edges of pr? and pa‘ are reduced and little
used. The accessory cusps and heel are nearly as high as the pa‘.
The heel consists of a larger external and smaller internal cusp, both
greatly worn, but apparently low and rounded. The external cin-
gulum is strong and crenulate.
The second molar has the same composition as the first, except that
the paraconid is small and connate with the protoconid, which is of the
same size as the well-separated metaconid. The external cingulum is
very broad in the anterior half of the tooth and bears one well-defined
cusp external to the protoconid. The heel is nearly as long as the
trigonid.
The third molar is obovate with shallow basin heel, and larger trig-
onid too much worn for distinction of cusps. The cusps of the heel
are mostly obsolete, the surface wrinkled.
The premolars are shorter than in C. latrans, all except the first bear-
ing the posterior accessory cusp, characteristic of the dogs, but absent
in the bears. The first premolar is single rooted, spaced equally be-
tween the canine and second premolar. (In the dogs it approaches
the other premolars, in the bears usually the canine.)
Canine slightly more slender than in C. /atrans, more curved at base,
less curved toward tip.
The deep masseteric fossa, long angular process, and strong
metaconid suggest Daphenus, which, however, has the normal
canine proportion of carnassial and molar teeth, and, like all
the more ancient genera, has the shear more oblique to the
tooth-line than in the later Canide.
From the more ancient genus Cephalogale it differs in the
presence of the accessory premolar cusps, slender jaw, larger
molars, the posterior molars less unlike to the carnassial, and
in the presence on the carnassial of two accessory cusps. All
the modern microdont Canide except Otocyon have a more
typical proportion of carnassial and tubercular teeth, and lack
the accessory carnassial cusps. Their premolars are nar-
rower. Cynarctus is near to Haplocyon Schlosser, founded on
the jaw-fragment with pms 2 to 4 from St. Gérand-le-Puy,
described by the late Prof. Filhol under the name of Amphi-
cyon crucians. But the premolars are more cynoid, not so,
high, and the posterior accessory cusp is present on p,.. The
1902. | Matthew, New Miocene Canidae. 283
horizon of the two is different, and I hardly think that they
are really allied, although the distinctions on known parts
may appear rather slight.
From Pseudarctos it differs in the presence of the two ac-
cessory cusps on the trigonid of the carnassial, in the larger
premolars with well-marked deuteroconid, the slender jaw
and small third molar.
From Amphicyon the genus differs in the bicuspid heels of
the molars, greater reduction of pr*, and presence of accessory
external and internal cusps, long slender jaw, and cynoid
premolars.
From Ursavus it differs in the less reduction of the pre-
molars, presence of accessory cusps on-p,, p,, and p,, and the
much more cynoid character of molars 2 and 3. In Ursavus,
judging from Dr. Schlosser’s figures and description, the
cusps on m, are nearly obsolete, and the surface flat and
wrinkled, while m, is a round, peg-like tooth with flat, wrinkled
crown. The jaw of Ursavus is deep and short like that of the
bears, and the coronoid directed nearly upward as in the
Urside.
The foregoing description is based on a nearly perfect pair
of lower jaws found in the Loup Fork (Pawnee Creek beds) of
Cedar Creek, Colorado, by Mr. Brown of the American Mu-
seum Expedition of rgor1. A single lower carnassial in the
Cope Collection from the Colorado Loup Fork probably rep-
resents the same species. No upper teeth are known, and
the position of the genus is therefore uncertain. Judging
from the characters of the lower teeth it would seem probable
-that it must be placed with the Canide, and cannot be con-
sidered as near to Ursavus, which is unmistakably a bear. As
far as can be determined from the lower jaw characters, it
seems to be partly intermediate between Ursavus and Cants,
with some primitive characters retained, no doubt, from its
Oligocene ancestors. If this be borne out by the characters
of the upper teeth, Cynarctus will help to bridge the most
serious gap in the series of extinct genera connecting the
Urside and Canide. Amphicyon, as Dr. Schlosser has shown,
does not fulfil the requirements for a direct ancestor of the
284 Bulletin American Museum of Natural History. [Vol. XVI,
bears, but must be considered as a side branch paralleling
them. Cynarctus would seem somewhat more but by no
means exactly in the line of descent. The slender jaw ex-
cludes it from direct relationship.
In a previous paper the writer has discussed an alternate
hypothesis of the origin of the Urside which derives them
from the Creodont family Arctocyonide, instead of from the
Canide. It was then stated that the apparent chain of ex-
tinct types connecting the Canide and Urside formed a most
serious objection to considering any other hypothesis as pos-
sible, but it was pointed out that there was a wide gap in the
series between such genera as Amphicyon and Dinocyon—un-
mistakable dogs although bear-like—and Hyenarctus and
Ursavus, unmistakable bears, although with the primitive
carnivore formula lost by the modern bears. Dr. Schlosser
has shown that Amphicyon is a side branch of the Canide and
Hyenarctus of the Urside and that the wide gap between the
primitive Oligocene dogs, such as Cephalogale and the earliest
true Urside cannot be filled by any genera hitherto known.
The genus here described reduces this gap, as its close re-
semblance to Ursavus in the composition of the teeth seems
hardly explicable except on the ground of a near relationship;
while in most characters it is as unquestionably a dog as Ursa-
vusisabear. It does not seem, however, to point especially to
Cephalogale as an ancestor. Its relationship to the direct line
of descent is uncertain.
Nevertheless this additional evidence in favor of the deriva-
tion of the bears from early Canidz seems to render untenable
any other hypothesis. The Arctocyonide must then be con-
sidered as a case of parallelism not confined to the general
characters of teeth and feet, but extending to the detailed
structure of both, the rather exceptional cusp composition of
carnassial and molar teeth, the relative proportions of the
digits, even certain details in the character of the carpals
and tarsals being common to both, besides the more
general characters of large quadrate, flattened molars, re-
duced premolars, slender canines, plantigrade, large-clawed
feet.
1902. | Matthew, New Miocene Canide. 285
?? Ursavus sp.
Another small Amphicyonoid of about the same size as C.
saxatilis is indicated by No. 9454, a lower carnassial, and a few
fragments from Pawnee Buttes. The tooth is composed of
very low trigonid of three cusps, paraconid nearly as large
as protoconid, me* well developed, more internal than pos-
terior, long basin heel completely enclosed by a well-marked
ridge, which begins at the metaconid and swings around the
margin nearly to the protoconid, the heel-cusps being scarcely
seen. Trigonid is proportioned much as in Ursavus, but no
accessory cusps, and heel without well-marked cusps. Trig-
onid and especially the protoconid lower than in Amphicyon,
and marked basin heel.
? Cyon or Icticyon sp.
A palate and a ramus of the lower jaw, both young indi-
viduals showing the milk dentition, are referred here. The
permanent sectorials are formed within the jaw, but not ex-
truded. The character of the milk dentition proves that the
specimens belong to the Canide, but to the division of the
family with most highly secant teeth. This is confirmed by
the metaconid; in the superior one the deuterocone is minute,
and there is no anteroexternal cusp. I am unable to make
comparisons with the milk dentition of either Cyon or Icticyon;
the permanent upper carnassials differ from the figures of
Icticyon and from specimens of Cyon alpinus in the greater
reduction of the anterointernal cusp and presence of two
- strong ridges on the anterior slope of the protocone, diverging
from the point, one running to the anteroexternal corner of
the base, the other to the base of the anterior internal cusp.
The permanent incisors are trifid, the lateral cusps being
stronger than in any Canid that I have seen, equal almost to
the median cusp. The external temporary incisor has but
one strong lateral cusp, the external one minute; a posterior
cusp is also present. The temporary canine is short and
small with prominent posterior ridge. The permanent first
premolar is one-rooted, with anterior and posterior cusps and
286 Bulletin American Museum of Natural History. [Vol. XVI,
small posterior cingular cusp. The second temporary pre-
molar is smaller, more compressed, two-rooted, with rudi-
mentary posterior cusp. The third is the carnassial, and is
three-rooted, composed of large protocone and strong posterior
blade, with a minute anterointernal basal cusp situate be-
tween the anterior and internal roots. The fourth milk pre-
molar is molariform, with no protoconule, strong hypocone (in
reality probably a metaconule), besides the three main cusps.
The second lower milk premolar is two-rooted, set obliquely
in the jaw, and has a small posterior cusp. The fourth (car-
nassial) has the trigonid of shearing protoconid and paraconid
blades, small metaconid, and three-cusped basin heel. Com-
pared with the corresponding teeth in the Coyote these teeth
differ in larger size, greater robustness, more sectorial char-
acter in the carnassials, proportionately smaller and narrower
heel on dp,, dp* of less transverse and greater longitudinal
width, the inner cusps less marginal, reducing the size of the
basin enclosed by them. The anterointernal cusp on dp? is
smaller and situated much more anteriorly; the anteroexter-
nal cingular cusp is hardly noticeable. The jaw is very much
shorter and deeper, ‘the premaxilla is carried much farther
back between maxilla and nasal. The cusp composition is
the same in both.
No Canid has been described with which this can well be
identified. It is of the size of Avlurodon, but differs in ab-.
sence of anteroexternal cusp on p* (and other characters). It
is much more modernized than any of the John Day dogs, and
the size is too great for C. brachypus Cope, temerarius or vafer
Leidy, anceps Scott, all of which, moreover, seem more
typically cynoid.
In a previous article Dr. Wortman and the writer attempted
to trace a line of descent from the Eocene Uintacyon and
Prodaphenus through the Oligocene Daphenus and Tem-
nocyon to the modern Cyon. It seems not unlikely that the
Canid here described may nearly represent the Upper Miocene
stage of evolution of this race. Jcticyon seems also to be more
or less nearly connected with it—and if this hypothesis of
descent be correct, this group parallels the Camels in their
1902. | Matthew, New Miocene Canide. 237
present and past distribution, originating in North America,
spreading to South America and Asia, and becoming extinct in
their old home while still surviving in the two widely separated
districts to which they had wandered.
In this as in other cases the writer desires to guard against
expressing any belief that the evolutionary series worked out
in various lines represent the actual species through which
descent has occurred. They represent indeed the history of
the evolution of certain parts; they may in some cases be not
far from the direct line of descent. But it appears probable
that each ‘stage’ represents in most cases a migration rather
than a mutation of species. Believing that the principal
causes of the evolutionary changes among the Tertiary mam-
malia lay in the secular world-wide alteration in climatic and
geographic conditions, it seems improbable that in any given
locality a change in the fauna occurred directly without a
change in the area over which the species flourished. It
seems much more likely that most of the changes in fauna in
a locality were due to successive waves of migration, setting
out from the region in which the new climatic conditions first
appeared. This would involve in general a succession of
waves of migration spreading from the north into America on
one hand, Europe, southern Asia, and Africa on the other,
differentiating to some extent as the separation increased, and
driving the older faunas southward before them. Hence the
Tertiary aspect of so large a part of the South American and
African faunas, and hence the primitive aspect of forest faunas
in general, the new conditions of cold and arid climate which
_ culminated in the Glacial Epoch involving the spread of open
plains, and diminution of the forest areas.
Amphicyon americanus Wortman.
Amphicyon americanus WortMAN, Amer. Journ. Sci., Vol. XI, 201,
Sep. Jan. 25, March, rgor.
This species is of moderate size in the genus and of rather
primitive character, in some respects approaching the species
of the European Oligocene (A. lemanensts, etc.). Dr. Wort-
man gives the following measurements:
288 Bulletin American Museum of Natural History. [Vol. XVI,
Length of superior molar series, including canine... 134 mm.
Anteroposterior diameter of canine at base........ 24
enethot true molar Series: jit. ever teers 46
Transverse diameter of first superior molar........ 27
Anteroposterior diameter of superior sectorial..... 27
Width of palate at first molar, including crowns... 98
Amphicyon sinapius, n. sp.
? Canis, sp. incerta, Cope, Rep. Vert. Pal. Col. U.S. G. S. Terrs.
Ann. Rep. 1873 (1874), 519.
A larger American species of Amphicyon is represented by
a number of fragmentary specimens from the Colorado Loup
Fork.
Am. Mus. No. 9358. Jaw fragment with broken carnassial and com-
plete first tubercular molar. Type.
9357. Carnassial tooth, unworn. Co-type.
?9356. Twenty-three vertebre, ribs, humerus, and ulna.
?9355. Astragalus, parts of tibia, humerus, radius, and
several metapodials, of uncertain association.
“28248. (Cope Coll.). Astragalus, and anterior part ofa
lower jaw, without teeth.
Our material unfortunately does not enable us to determine
the dental formula, but the characters of the teeth agree best
with those of Amphi-
cyon, and are more pri-
mitive than those of
Dinocyon. It is much
larger than A. lemanen-
sis, but resembles rather
nearly the figures of that
genus given by Dr.
Schlosser. It somewhat
exceeds A. major and A.
giganteus in size, the heel
: of m, is broader, m? is
: Fig. 2. Amphicyon sinapius. Part of lower much larger and broader
jaw, inner side, x 3. Type, No. 9358. Loup Fork i i
(Pawnee Creek Beds), Colorado. comparing it with de
Blainville’s figures, and on both molars the entoconid is
represented only by a broad cingular ridge. Both in size
1902. | Matthew, New Miocene Canide. 289
and characters the second lower molar is very like that
figured by Dr. Schlosser in ‘Paleontographica’ and referred
Fig. 4. Amphicyon eee
Lower carnassial x.3 No.
Fig. 3. Amphicyon sinapius. Crown view of Loup Fork (Pawnee Creek Beis,
m, injaw, x3. Type, No. 9358. Colorado.
doubtfully to A. major. Without more complete material
the position of this species cannot certainly be determined.
? Dinocyon (Borophagus) mzandrinus (Hatcher).
4Elurodon meandrinus HaTcuHEeR, Amer. Nat. 1893, 240.
The type consists of part of a lower jaw, with the second and
third premolars greatly worn, and roots of fourth premolar
and sectorial. It differs from other Alurodons, according to
Mr. Hatcher’s description and figure, in the much greater size,
extremely short jaw, and reduced premolars, large posterior
root to the sectorial. All these characters point to the Am-
phicyonine rather than to the true dogs; the second molar,
whose proportionate size would make the position of the
species certain, is unfortunately not indicated in the type.
To this species may be referred provisionally Am. Mus. No.
10583, a fragmentary lower jaw with roots of the teeth, asso-
ciated with parts of tibia, etc., found by Mr. Gidley of the
American Museum Expedition of 1899 in the Loup Fork forma-
tion of Donley Co., Texas.
Measurements.
Type. 10583
Post-canine;diastemany.. 444. 4 oo 19 14
Premolar Gestition..t.c:s =... acu. «226 io 60 62
Carnassialmlengthinn. 2+ sles cere ee one 47 47
Severo; 08 1710] Fk ans eR, ee ae aoe — 24
Carnassial, width ant. root........-..... — 15
a AE DOS bam a tarereyoe ns cusia sc — 20
Depthbvot jaw behind pig .iednis Seed coos, 55 69
[ September, 1902] 19
290 Bulletin American Museum of Natural History. [Vol. XVI,
? Dinocyon (Borophagus) diversidens (Cope).
Borophagus diversidens Corre, Amer. Nat., 1892, 1028; Vert. Pal.
Llano Estac. (4th Ann. Rep. Geol. Surv. Tex., 1892), 54, pl. xiii, fig. 4.
Blanco horizon (Upper Pliocene). Referred to the Hyz-
nide by Professor Cope. The type is a fragment of a lower
jaw, with two premolars preserved and the root of athird. It
agrees with Amphicyon more nearly than with Hyena in the
form of the individual teeth, as well as in their proportion one
to another; the second premolar is smaller, apparently, than
in the Loup Fork species.
? Dinocyon (? Borophagus) gidleyi Matthew.
Dinocyon (? Borophagus) gidleyi MATTHEW, Bull. Am. Mus. Nat.
Hist., Vol. XVI, 1902, 129-136.
? Amphicyon ursinus Cope.
Cants ursinus Cope, Proc. Phila. Acad. Nat. Sci. 1875, 275; Rep.
Wheeler Survey, Vol. IV, pl. ii, p. 304, pl. Ixix, fig. 1.
The reduction of the premolars, proportionately large tu-
bercular teeth, deep, massive jaw with comparatively straight
inferior margin, large heel on the lower sectorial, etc., place
this species with the Amphicyons. Professor Cope remarks
on the probability that C. ursinus is very close to C. hayden.
Leidy’s species is, however, much more like the wolf in pro-
portion of sectorial to tubercular teeth, and the heel of the
sectorial is comparatively small, as in C. lupus or in the
/Elurodons.
A. ursinus is about the size of A. americanus, and is perhaps
synonymous with it.
1 ey egal From the Colorado Mio- oe
ee ath a Revision of the Myla
A merican TLertiary.
By W. D. MartrtHew.
AUTHOR'S EDITION, extracted from BULLETIN
~ OF THE
Article XXII. A HORNED RODENT FROM THE COLO-
RADO MIOCENE. WITH A REVISION OF THE
MYLAGAULI, BEAVERS, AND HARES OF THE
AMERICAN TERTIARY.
By W. D. MatrHew.
Ceratogaulus rhinocerus, n. g. et sp.
The writer has recently described part of the skull of a
Mylagaulus from the Colorado Loup Fork beds, found in
1898. A nearly complete skull, with one ramus of the lower
jaw, found by Mr. Brown of the Expedition of 1901, indicates
a new genus of this family, distinguished by the unique
character (for a rodent) of a pair of large connate processes on
the nasals resembling the horn-cores of some Ungulata, and
giving the skull a profile absurdly like that of a miniature
rhinoceros.
The skull is a little larger than that of Mylagaulus, and dis-
plays considerable modifications, chiefly conditioned by the
development of the horn-like processes on the nasals. The
muzzle is much wider and tapers forward; the nasals are
much wider throughout, and especially in the middle, where
they bear the horn-cores. The postorbital processes of the
frontal and jugal bones are considerably less prominent and
placed farther back, making the orbit larger and more ex-
tended anteroposteriorly. The zygomata are deeper. The
enlarged molar in the upper jaw differs a little in form, and
considerably in the pattern of the crown. The penultimate
upper molar appears to be considerably larger in proportion,
but is so much damaged in the Mylagaulus skull that it cannot
be closely compared. The enlarged molar of the lower jaw
displays a crown pattern with the usual lakes in three longi-
tudinal rows, instead of four as in Mylagaulus. The alveoli
of the second and third molars are of nearly equal size, while
in Mylagaulus the penultimate alveolus is much larger. The
type specimen No. 9456, is of nearly the same age as the My-
lagaulus skull with which it has been compared, the wear of
[291]
292 Bulletin American Museum of Natural History. (Vol. XVI,
Fig. 1. Ceratogaulus rhinocerus. Skull and jaw, natural size. Superior, lateral and
anterior views. No, 9456 (type). Loup Fork, (Pawnee Creek Beds) Colorado,
1902.] Matthew, A Horned Rodent from Colorado Miocene. 293
the teeth being slightly less advanced. The distinctions,
therefore, cannot be due to age. The horn-like processes
might be sexual, although I can find no parallel among the
rodents for such a wide divergence between male and female;
but the marked distinctions in the teeth and other characters
are not likely to be sexual, although in themselves they are
not of generic importance.
The height of the horn-cores is about one-fifth the length
of the skull, their length a little more, and their conjoined
width about one-fourth the skull-length. The longer axis of
each process is diagonal, posteroexternal and anterointernal;
the conjoined process is subtrigonal, the angles posteroexter-
nal and anterior.
A character so marked as this would seem a good basis for a
separate genus. Nevertheless, the resemblance to the skull
of Mylagaulus obtained in 1898 is considerable in most char-
acters, except in the horn-cores, the position of the postor-
bital processes, and the pattern of the enlarged grinding teeth.
.The occiput is extraordinarily wide and low, its width
equalling the entire length of the skull. The postorbital
crests do not unite behind; the top of the skull is flat trans-
versely, concave anteroposteriorly, and the occipital surface
slopes 30° forward from the condyles to the top of the crest.
The zygomatic arches are stout, deepest in front, somewhat
wider than the occiput, both postorbital processes (on the
frontal and jugal) moderately strong. In both upper and
lower jaws the alveoli of two smaller molars are preserved,
but no clear indication of a third, behind the enlarged tooth.
Measurements.
Length of skull (condyle estimated)................... 68 mm.
Which GH AGhOSSralLGhesm runt y emt = foie Rina), | ar, Gandia ets 64
UNDIES) SEA. oh SP SU ARI Ane ie Oe Ao 65
PAO OI NORM M Ola. isi fein: Setagt se Gates Saree 6 Rito wos oye 13
Length “ SRM gee Nee ets ek EAs aca dc Sstte au gst ois 17
Conjomedsavidthomhorn-COTES. wm... oes ge oes sls ere 19
Width across postorbital processes of frontals........... 32
Width across postorbital constriction. ......0......06. 18
Least depth of zygomatic arch beneath orbit............ 9
294 Bulletin American Museum of Natural History. [ Vol. XVI,
Bengthyotdiashenia na. ciheisy. cys cre elo elie ier ets Oana ene 20 mm.
“-) = “three upper smolars:\(? p2—m")ia.).) pee areal 15
‘ty ten genlarced .<o mmolar (P94) )s Mice shan ieeerenees eee 8
Width of - uy e Ss Was aioe eats aes 6.5
Height ofsame (rootandcrowm)),2-- ss er eee 12
Bstimated length of dower jaw tsi. oe knee eek ees 58
Depth*of jaw beneath) molars... .i0.2 sles eee nel as
Height ‘‘ “ (angle to tip of coronoid process)......... 41
Length of enlarged lower molar (? p,)...:..........2.. 10
Width of o i “ ROA og: apes to os
This remarkable skull has no parallel among the Rodentia.
Haplodontia most nearly approaches it in width, but the horn-
cores and the specialized teeth are unique.
HIND LIMB AND FEET OF MYLAGAULID&A.
We know but little as yet of the skeleton of this family.
The pelvis was very massive, and beaver-like on a smaller
scale, the tail probably not flattened, and the scaphoid and
lunar were united. A metacarpal associated with two teeth
of Mylagaulus shows some remarkable characters. It is
nearly as large as the metacarpals of Castor canadensis and
much stouter. The distal facet is strongly keeled on the
inferior surface, in a manner recalling the distal ends of meta-
podials of Chalicotherium or metacarpals of Dasypus. The
facet is limited superiorly, so that it is not at all reflexed over
the superior surface of the metacarpal, but faces entirely in-
feriorly and distally. The characters of the bone may be
taken to indicate the presence of large digging claws on the
manus.
I refer also to this family a remarkable specimen in the Cope
Collection from the Loup Fork beds of the Republican River,
Nebraska, consisting of a nearly complete hind limb and foot,
with an ungual phalanx of the fore foot, not associated with
any parts of the skull. This specimen combines characters of
a Castoromorph rodent, with a resemblance to the modern ar-
madillo so striking that Iam unable to state positively that it
is not edentate.t As nearly as I can judge, however, the ar-
* Professor W. B. Scott, whose recent extensive and thorough studies of the Santa
Cruzian fauna give especial weight to his authority, has examined this specimen and
pronounces it probably rodent and certainly not edentate.
1902.] Matthew, A Horned Rodent from Colorado Miocene. 295
madillo resemblances may be explained as all associated with
the development of digging claws, chiefly on the fore foot, and
walking on one side
of the foot in conse-
quence. The Casto-
romorph characters,
on the other hand,
appear to be such as
would indicate real
relationship, al-
though not close re-
lationship to any
living form. From
the phalanx just de-
scribed we have rea-
son to infer that
Mylagaulus devel-
oped large digging
claws on the fore
foot, and this speci-
men is of appropri-
ate size and propor-
tions to belong to
the Mylagaulus. It
is found in the same
horizon, and could
not belong to any
other rodent known
_ from those beds, for
all the others are
quite nearly allied
to still existing gen-
era. Edentates have
not been found in
the Loup Fork,! and
our specimen shows
no resemblance to
Fig. 2. Mylagaulid, indet. Hind limb bones and claw,
natural size: 7, anterior view of femur; /%. 7. posterior view
of tibia and fibula; superior view of pes; A, A,’ superior and
lateral views of ungual phalange of fore foot. No. 8336. Loup
Fork, (Republican R. Beds) Kansas.
1 The supposed exception, Caryoderma snovianum, is, according to Williston, a turtle.
206 Bulletin American Museum of Natural History. |Vol. XVI,
any known fossil edentate, the resemblance being only to
Dasypus, and to a much less extent to Tatusia. From this it
seems reasonably safe to infer that it is a Mylagaulid, and that
this family paralleled the Armadillos in the structure of their
feet.
The femur (Fig. 2, fF) is nearly complete; it differs from
either beavers or armadillos in the position of the very large
and powerful third trochanter, which is placed high up on the
shaft, nearly opposite the second trochanter, instead of in the
middle of the shaft as is usually the case. The greater and
lesser trochanter are much as in Castor; the distal condyles
are wide and low, and the trochlea short, broad, and shallow.
Most of the tzbia and fibula (Fig. 2, Fb., T.) are preserved;
the tibia is short and stout, beaver-like at the lower end; the
fibula as strong as in Castor, separate from the tibia, with a
vertical internal facet for the astragalus,
but no distal facet, and no contact with
the calcaneum. The astragalus (Fig.
2), is quite rodent-like, with moderately
broad trochlea defined by sharp keels
internally and externally, rather small
neck, and broad, flattened head. Meta-
tarsals II to V are present, but the
hallux was rudimentary or absent. The
second metatarsal is much more slender
than the third and fourth, but of about
the same length. Mt. V is only three-
fifths as long as the others, but fully as
Ee Tathicnee or stout asmt.IIIandIV. The phalanges
eae pa See of the first row are rather short, their
(Pawnee ‘Greek Beds), Colo- distal facets wide, not deep, moderately
concave from side to side. The size
and strength of the metatarsals and proximal phalanges
does not appear adequate to bear the very large claw
(Fig. 2, A, At) with which they are associated, and I therefore
suppose that it belongs to the fore foot, where the much
stouter and more specialized metacarpal, such as has been de-
scribed as occurring with teeth of Mylagaulus, could very ap-
1902.] Matthew, A Horned Rodent from Colorado Miocene. 297
propriately bear it. This claw phalange is long, compressed,
the proximal facet very little keeled, the distal end slightly
fissured but not symmetrically so, and shows no hood at the
base.
The hind foot when set in position on wax shows an un-
mistakable twist, the external side being bent down distally
as if the animal walked on the outside edge of the foot, turning
the claws inward underneath. This may serve to explain the
short, stout fifth digit, as contrasted with the long, slender
second; a proportion seen also in the armadillo and some
other modern Edentates.
REVISION OF THE SPECIES.
On comparing the two skulls and three other more frag-
mentary specimens from Colorado with five specimens of
Mylagauli from Nebraska in the Cope Collection, 1t appears
that a considerable number of species are represented. On
reviewing the description of Mesogaulus ballensis Riggs I find
that I was in error in identifying it with Mylagaulus monodon,
but its position can hardly be determined until more is known
of the milk dentition and the history of the changes in tooth
pattern in this curious family. As far as at present deter-
minable the characters of the known species are:
Mylagaulus monodon Cope. Type, a jaw with the enlarged
molar and two alveoli posterior to it, from the Loup Fork of
the Republican River Valley. A second specimen, a lower
tooth from the same locality, referred to it by Professor Cope,
is considerably larger. Enamel lakes in four rows. There
are seven lakes in the type; nine in the associated specimen.
No cement outside external enamel ring.
No. 8327 (type). No. 8328
Extreme anteroposterior diam. of molar ?ir.5mm. 13.5 mm,
transverse Re, ch — Gms
Anteroposterior diam. of grinding surface 10.5 II
Transverse ‘> yaad i Fs 6 6
Mylagaulus sesquipedalis Cope. Type, an upper molar, No.
8329. Referred specimen a lower molar and incisor anda
metacarpal, No. 8330. Both from the same locality as M.
298 Bulletin American Museum of Natural History. [ Vol. XVI,
monodon. They are much smaller, and the pattern of the
enamel lakes simpler and more irregular. Those of the lower
tooth are six in number, corresponding in position to the lakes
in M. monodon, but less elongated, fewer in number, and less
regularly arranged. Grinding surface of upper molar regu-
larly oval, with six lakes irregularly arranged.
No. 8329 (type). No. 8330
Upper molar, extreme anteroposterior diameter... 10 mm.
ie i CranSVersess. «ares eres Bak
anteroposterior diam. of grinding
SUITPACes cisee cen athe bias eee 8
transverse diam. of grinding surface. 5.5
Lower molar, anteroposterior diam. (estimated) . . 8.5
i tTANSVETSe) 0 Se. acy ener es
“eé
Mylagaulus (Mesogaulus) ballensis Kiggs. Type, a lower jaw
containing three teeth, from the Deep River beds of Montana.
Size of M. sesquipedalis, but with only four enamel lakes and,
according to Mr. Riggs’s drawing, a heavy band of cement
surrounding the grinder. Our specimens show nothing like
this; several have a thin layer of cement over parts of the
outside, but never at the grinding surface, except in a sup-
posed milk-tooth in which the enamel ring does not come up
to the grinding surface.
Dimensions, from Riggs’s Description.
Anteroposterior diameter of grinder.......5............. 9 mm.
Greatest lateral breadth of SO ce ae eee 4.2
Mylagaulus levis, n. sp.
M.monodon MattHew, Mem. Am. Mus. Nat. Hist. I, LO OMe sive
Not M. monodon Cope.
Type, front half of skull and jaw, pelvis, and other frag-
ments from the Loup Fork of Colorado. Smaller and less
robust than MM. monodon, pattern of lower molar similar, with
seven lakes arranged in four rows. Upper molar flattened ex-
ternally, not regularly oval like that of M. sesqutpedalis, lakes
longer, narrower, and lying more regularly parallel. Nasals
smooth.
1902.] Matthew, A Horned Rodent from Colorado Miocene. 299
Dimensions.
Lower jaw, diameter of enlarged molar, anteroposterior(grind-
LER GE SUL CS Pa etae Murer c os ceexac co cauat ee meretepshetecs aor etext eren ee 9 mm.
Lower jaw, diameter of enlarged molar, transverse (grinding
SUEPACE)! sce Veer cwicntederots os Sica o wea. smiere ate wolatons 5
Upper jaw, diam. of enlarged molar, anteroposterior (grind-
AOS SUPTACE i asks poectt ch ones SS TAs, Mel eae a ale” een se aaa ee 9
Upper jaw, diam. of enlarged molar, transverse (grinding sur-
FACE) Norctc ch eet creer ate sick <) eet onchekAtets: srever Pawencheuobaleuale sinetewars 5.8
Ceratogaulus rhinocerus, n. sp.
Generic characters: Nasals bearing a large pair of horn-like
processes, closely twinned. Specific characters: Size some-
what larger than that of M. levis, less than that of M. mono-
don. Enamel lakes of lower molar simpler, seven in number,
arranged in three rows, less regular than those in MZ. monodon
or M. levis. Alveoli of last two molars of subequal size. Ex-
ternal side of upper grinder flat, internal strongly convex.
Second upper grinder larger than third. Muzzle broader
than in M. levis, postorbital processes shorter and more
posterior.
Dimensions of the Enlarged Grinders.
Upper. Lower.
Anteroposterior diameter. 8.7 mm. 9.7mm.
Transverse i Oi. 5 535
Mylagaulus paniensis, n. sp.
A small and simple species indicated by
‘half a lower jaw and a few fragments.
There are five enamel lakes, of which the
three interior ones are arranged in a row as
in M. ballensis; and external to these are a
large anda small lake, the latter correspond-
ing to the fourth lake of ballensis. The ee eee
tooth is worn well down, while that of eee Nore (Penne
ballensis appears to be a comparatively Cee Beds), Colorado.
young individual, so that the less number of lakes in Mr.
Riggs’s species can hardly be due to greater age; and on
300 Bulletin American Museum of Natural History. [Vol. XVI,
our specimen is no trace of external cement. This specimen
comes from the base of the Loup Fork beds at Courthouse
Butte, near Pawnee Buttes, Colorado.
Dimensions, No. 9361.
Anteroposterior diameter of lower molar............... 7.4mm.
Transverse i y ct Sed a odo, Bae et eRe ae 4
pensth) of dsastemiae 3, sc cisyat o attaeis ae ane ae 7
Mransviersenwidth Okan cisor- essen te tae eee ree 3
Anteroposterior diameter Of incisons areas eee 4
CASTORIDA.
Steneofiber Geoffroy.
The pattern of the molars in this genus is so evanescent
that it is almost impossible to make satisfactory comparisons
of species on the limited number of specimens available. The
essential pattern of the upper teeth consists of a deep internal
enamel inflection and three external ones, the anterior and
posterior of which quickly become fossettes. In the lower
teeth the pattern consists of a simple external and three in-
ternal inflections, the anterior and posterior internal inflec-
tions soon becoming fossettes. With moderate wear the
crown is divided transversely, by the internal and external in-
flections, into an anterior and a posterior column, united at the
base, each column containing a fossette, sometimes more than
one. The teeth become broader transversely with wear, and
much less in their anteroposterior diameter; the anterior col-
umn increases and the posterior one diminishes in size in the
upper teeth, while the converse holds true in the lower teeth.
The enamel folds become closer, and their direction changes with
wear; the internal one in the upper teeth, the external one in
the lower, becoming more nearly anteroposterior in direction.
The principal inflections in old individuals have become fos-
settes, while the minor inflections have disappeared. The last
molar in young individuals appears small in proportion, because
of the small area of the grinding surface; in older individuals
it appears larger in proportion. The fourth premolar of
1902.] Matthew, A Horned Rodent from Colorado Miocene. 301
young individuals likewise appears small in proportion to the
first and second molars, but reaches its maximum of size
earlier than does the third molar.
These observations are based on the American species,
which are a rather closely allied group, characteristic of the
Oligocene, and more primitive than the European assemblage
of species, which are characteristic of the Miocene epoch, and
in general larger, longer-toothed, and nearer to the modern
branches. The S. viciacensis of the Upper Oligocene (St.
Gérand-le-Puy) appears to be the nearest to the American
group. The distinctions between our species have been based,
unfortunately, chiefly on the evanescent pattern of the teeth,
not sufficiently considering the great alteration due to a com-
paratively slight difference in their wear. Seven species
have been described, as follows:
1. S. nebrascensis Leidy. White River, 5. Dakota, skull and jaws.
2. S. pansus Cope. Loup Fork, N. Mexico, upper and lower jaws, etc.
3. S. peninsulatus Cope. John Day, Oregon, skull.
4. S. gradatus Cope. John Day, Oregon, skull.
5. S. montanus Scott. White River, Montana, teeth and skeleton
fragments.
. S. hesperus Douglas. White River, Montana, lower jaw.
7. S. complexus Douglas. White River, Montana, part of skull and
jaws.
an
I have at hand for comparison the types of all the species
except the first two, besides three other skulls and some less
complete material. I am indebted to the courtesy of Pro-
fessor Scott and Mr. Douglas for the loan of the type speci-
mens of the species described by
them. In revising these species
it has been necessary to reject a
large part of the distinctions made
by their authors, as being merely
a matter of different age in the fig.s. Stencofiber nebrascensis.
as Upper molars x ?. No. 1428. White
type specimens. River (Protoceras Beds), South Da-
1. §. nebrascensis. I refer here ‘**
a skull, No. 1428, which, like Leidy’s type, comes from the
Protoceras beds, and part of a lower jaw, No. 10288, in our
302 Bulletin American Museum of Natural History. [Vol. XVI,
collections. The former is an older animal than Leidy’s
type, the latter younger. The species appears to be dis-
tinguishable by the long, narrow muzzle, small bulle, sharp
sagittal crest, and small brain-case. The postorbital constric-
tion is moderate, the pattern of the teeth rather complicated,
two deep fossettes anterior to the external inflection on p+
remaining in the well-worn teeth of No. 1428.
2. S. peninsulatus. Besides the type skull, another skull,
less crushed, a skull and jaws, and several parts of jaws, etc.,
referable to this species, are
in the Cope Collection in this
museum. It isa more robust
species than the last, distin-
guishable by the large bullz
and probably by the broader
muzzle, wide occiput, larger
brain-case, and wider sagittal
Za crest. The postorbital con-
Hig. 6. Steneofiber peninsulatus. Upper SbTictionds very nanowem nae
an De Coe hdiud Eis Nore” type, but not’ in the second
Specimen: - “Thea steetnrreane
much like those of S. nebrascensis, but the second anterior
fossette of p* apparently remains longer as a branch from
the medial external enamel inflection. The fossette remaining
from the posterior external enamel inflection of p* has almost
disappeared in the type and another specimen, but shows no
signs of breaking up into three little fossettes, as it apparently
has done in the type of S. nebrascensis. The specific validity
of these distinctions in the tooth pattern is very ques-
tionable; the two sides of a single skull seldom agree at
all closely, and this may well be an advanced mutation or
subspecies of S. nebrascensis, the large bulle being the
clearest distinction.
3. S. gradatus. The type skull, and the palate of a some-
what older individual are in the Cope Collection. It is a
smaller animal than the two preceding species, with short,
wide muzzle, postorbital constriction moderate, brain-case
short and rounded, temporal crests not uniting to form a single
1902.] Matthew, A Horned Rodent from Colorado Miocene. 3093
sagittal crest for some distance back of the postorbital con-
striction. Bulle of moderate size; grinding series of teeth
near together anteriorly, divergent posteriorly. The teeth
decrease in size from p* to m* more
than they do in S. nebrascensts or
S. peninsulatus; there is but one
fossette anterior to the external
enamel inflection on the type, while
our referred specimen of S. nebra- _ Fig.7. Steneofiber gradatus. Up-
; : per molarsx%. Type. No. 7007.
scensts shows two, neither near ex- Jou Dex (? Diceratherium Beds),
tinction, although the teeth have
attained the same stage of wear. The external enamel in-
flection has given off a small fossette on the left premolar of
the type, but not on the right one; in both type and referred
specimen the fossette of the posterior enamel inflection has ,
disappeared.
4. S. pansus. The upper and lower jaws are the only parts
of the head known, and I have no certainly referable material
to asisst in determination of the characters. Professor Cope’s
figures indicate an old individual, and apparently that the
posterior enamel inflection of the upper
teeth was more deeply impressed than in S.
gradatus, which it resembled in the presence
of but one fossette anterior to the median
enamel inflection. The form and propor-
tions of the molars, on which Professor Cope
relied to distinguish it from other species,
are merely a matter of age, as far as any
separation from S. peninsulatus and nebra-
scensis is concerned. The bulle are very
large, as in peninsulatus, which it most
nearly approaches as far as known, although
presumably distinct, as it is recorded as
found at a much higher horizon.
ee ppeee adie: s. S.montanus. AlliedtoS. nebrascensis,
molars x ?. Type. Lower
Beds of Smith Creek, but larger, with somewhat longer teeth,
Montana (Lower Mio- ; ‘
cene). and enamel inflections deeper and more
complex. The type is an old individual, whence the antero-
304 Bulletin American Museum of Natural History. [Vol. XVI,
posterior direction of the internal upper and external lower
enamel inflections, on which Professor Scott largely relies to
distinguish the species. Two anterior fossettes are preserved
on p*, while the posterior fossette has already disappeared.
6. S. hesperus. Founded on the lower jaw of a young in-
dividual which is certainly close to S. montanus if not identical;
the difference in age prevents any accurate comparison. The
size is the same at similar
points of wear; the upper
incisors are more rounded
externally, but the value of
; this character is doubtful.
Fig. 9. Steneofiber hesperus. Lower molars x}. The depth and complexity
Type. White River, Montana. BRS ees So folds, Ae
_ nearly as I can judge on the specimens, correspond fairly well.
7. §. complexus. Founded on the anterior half of the skull
and jaws of an animal
younger than any of the
preceding types, still re-
taining the milk premolars.
The skull has the long,
slender muzzle of S. ne-
brascensis; the postorbital
constriction is moderate,
and the temporal crests do
not unite into a sagittal
towel Ratan meegeer omplenus. Upper and crest, but are separate, as
cen aes in S. gradatus. The differ-
ence in wear precludes comparison of the teeth with those
of the remaining species; the dimensions of the masseteric
scar and coronoid process given by Mr. Douglas as distinguish-
ing characters likewise change with age so much as to be un-
safe specific distinctions. The separate temporal crests may
constitute a valid specific distinction.
Eucastor (Leidy) Allen.
This genus is represented by a single species found as yet
only in the Nebraska Loup Fork. The teeth are considerably
1902.] Matthew, A Horned Rodent from Colorado Miocene. 305
more hypsodont than in the American Steneofibers; the
molars have but one internal and one external enamel inflec-
tion (fossettes in the type
specimen). The upper pre-
molar has three external and
the lower premolar three in-
ternal inflections, asin Steneo-
mnie oy cesta cores hai. Lanet fiber and Castor. The position
(Republican River Beds), Nebraska. of Eucastor is very doubtful ;
if, indeed, it is a true Castorid at all, it cannot be very nearly
allied to either Castor, or Steneofiber. To this genus and
species probably belong, besides the type, parts of two lower
jaws in the Cope Collection.
OTHER CASTORIDAZ FROM THE LOUP FORK.
The two isolated teeth which form the type of Leidy’s
Hystrix venustus, if they are really from the Loup Fork, are
much more likely to be Castorid than Hy-
stricid, the latter not being likely to occur
= as far down in the Tertiary of the Western
i] has States asthe Miocene. Possibly with these
I Ce
species should be associ-
e ated a broken tooth from
ee Uc eh the Colorado Loup Fork,
nae Srdek Baty) Cok of the same size and gen-
ee eral proportions and pat-
tern as H. venustus. Another isolated tooth
Fig. 13. ? Castorid, indet.
from the same beds, No. 9364, represents a x2. Loup Fork (Pawnee
é Z : Creek Beds), Colorado.
species probably undescribed, a little larger
than S. montanus, more hypsodont and of more complicated
d pattern. A third specimen is part of a lower
1 . . . .
EAP jaw with m,-,, the latter just emerging.
aS QZ iC This is much more brachydont than Steneo-
Teg i Coste tuade fiber, although of somewhat the same gen-
Last two lower molars x 3. eral pattern, and somewhat smaller than
Loup Fork (Pawnee Creek ; p
ee area the described species. The style of the
teeth suggests Spalax rather than any other genus of Rodentia
with which I am acquainted.
[August, 1902. | 20
306 Bulletin American Museum of Natural History. [Vol. XVI,
LEPORID&.
Paleolagus Leidy.
The genus was based on the division of the anterior lower
premolar into two columns, instead of three asin Lepus. The
dentition and characters of the front of the skull and of parts
of the skeleton were fully figured and described at length by
Professor Cope in his ‘Tertiary Vertebrata,’ and have been
further discussed and revised in recent papers by Dr. Forsyth-
Major on the Lagomorpha. We are now enabled to add cer-
tain skull characters, not hitherto known.
Pal@olagus has a well-developed postfrontal process in all
four species, scarcely less than in Lepus ennisianus, but much
less than in modern Lepores. The angle between basicranial
and basifacial axes varies considerably in the different species,
but in none is it as great as in the modern species of Lepus.
The brain is relatively smaller than in Lepus. The tooth
pattern varies greatly during life; in the young it approxi-
mates that of Lepus, especially in such species as L. ennisianus;
in the old animal it becomes much simpler. P. agapetillus is
the most advanced in tooth, but least in skull, characters; P.
intermedius most nearly approaches the John Day Lepus.
The pattern of the teeth changes greatly in Pal@olagus, -ap-
parently from the superposition of a new pattern (that of
Lepus) on an older and simpler one. The young individuals
show an internal inflec-
tion on the upper molars,
whose depth and persist -
7 fj ence varies in the differ-
iJ ent species; there is also
an external inflection,
never deep, but rather
persistent, and a median
crescent which origin-
ates as an anterior mar-
_ Fig. 15. Lepus ennisianus. Cope. Skulland lower. 2
jaw natural size. Type specimen No. 7190. John Day, ginal inflection on Dey
Oregon. Muzzle supplied from another individual.
an amnteroexternal one
on p* (apparently median-external on p* and m'), a postero-
1902.] Matthew, A Horned Rodent from Colorado Miocene. 307
external one on m®, and a posterior one on m3. The median
crescent is somewhat less persistent than the internal inflec-
tion. Enamel is lacking on the external side of the teeth
except in very young animals.
The internal inflection is that which has become deeper and
more persistent in Lepus; the crescent has disappeared in
modern species of Lepus, and in the John Day L. ennisianus
apparently does not persist as long as in Paleolagus. The
enamel fails on the external side of the upper teeth of Lepus
except for a little while after they are protruded.
Dr. Forsyth-Major has suggested that the second upper in-
cisor of the Lagomorpha has originated from the posterior cusp
of such a tooth as that of Plesiadapis; the bifanged tooth
splitting in two and the large anterior cusp giving rise to the
large incisor of the modern lagomorph. Pal@olagus, he thinks,
might help to verify this theory. But the incisors of Paleolagus
are quite of modern type. It is to the lower Eocene Rodentia,
including the Mixodectide, or to the as yet little-known fauna
of the Cretaceous, that we must look for light on the method
of evolution of their teeth. But Eocene Lagomorpha have
not yet been discovered.
Pal@olagus approaches most nearly to Forsyth-Major’s
Caprolagus group among modern Leporide. These species
are less specialized for speed, and in consequence the head is
carried lower and more forward, and the basicranial and basi-
facial axes are at a smaller angle.
Angle between bastfacial and basicranial axes in different
species of Palgolagus and Lepus.
PP ORG PEN US Rena he bot eyo feat blave, E<CRAD as, be Ue
DERE HE) SSG AE SE i Ra Rc ig, Soot 20°
Lp SIEEVILE II the, Seleg a BRE NOOR Arete aia abies ooo < thn Soi a20
TEINS AIS EOS hg ace Sa ose OGG 33°
ECO MEV ESC S EARLS SO Ree. Fa Ro ee 47°
Paleolagus ? agapetillus Cope.
Professor Cope held this species as distinct in 1874, but in
1884 united it with P. haydeni. A skull found by our party
308 Bulletin American Museum of Natural History. [Vol. XVI,
in 1898 in the upper levels of the White River beds appears
distinct from P. haydent. It is smaller, more narrow and
elongated, the muzzle more slender. The anterior half of
another skull and several jaws, chiefly or all from the same
upper horizon, may be referred here, although they approach
P. haydeni somewhat more nearly than does the complete
skull. The distinctive
characters are: slender and
narrow muzzle, small teeth,
internal median fold of
enamel on upper molars
Fig. 6. Paleolagus ? agapetillus, Skull, nat- MOTe deeply incised and
ee White River (Martin Cafion yy yore persistent than in P.
haydent. The species ap-
pears to be good, on the evidence of some half-dozen speci-
mens referred to it and compared with the very numerous
P. haydent specimens in our collections. Whether the type
of P. agapetillus is properly referred to it, I am unable to
decide; but leave it provisionally.
I, p23 4 m1 23 7
Palezolagus haydeni Cope.
Trictum anne Cope; Tricium avunculus Cope; Tricium leporinum
Cope.
The additional specimens collected by American Museum
parties include no complete skull of this species. The best
one, No. 9327, shows a skull shorter and wider than P. aga-
petillus or intermedius, brain-case rounder than in either, basi-
facial axis bent down about as in P. intermedius, arch heavier
than in Lepus ennistanus, the jugal a band of uniform width,
thickened at the upper and lower margins. Teeth larger than
in P. agapetillus, less transversely broadened, medial internal
furrow less deeply incised or persistent. A scapula, vertebra,
and metapodial, associated with this specimen, agree in size
with Cope’s skeleton material of P. haydent.
Paleolagus intermedius Matthew.
Type, a nearly complete skull from the upper levels of the
White River, at Castle Rock, Cedar Creek, Col. Associated
1902.] Matthew, A Horned Rodent from Colorado Miocene. 309
type, upper and lower jaws and fragments of skeleton from
same level and region.
This skull is much more depressed on the basicranial axis
than P. agapetillus; it has a long and heavy muzzle, unlike
the slender, sharp muzzles of the two preceding species; the
teeth are larger than |
those of P. haydent,
butresemble themin
pattern; the length
of the diastema is
equal to that in P.
turgidus, but the a ;
teeth are much i Mae ue 1
smaller,andofmore
. Fig. 17. Paleolagus intermedius. Skull, natural size.
Lepus-like pattern; Type. No. 8722. White River (Martin Cafion Beds), Colorado
the skeleton appears
to be as large as that of P. turgidus, and of similar propor-
tions. The postfrontal process is nearly as large as in Lepus
ennistanus,; the muzzle is of the same length as in that species,
but heavier; the basifacial axis 1s somewhat less depressed,
and the brain-case is distinctly smaller.
Paleolagus turgidus Cope.
P. triplex Cope; ? Tricium paniense Cope.
Young jaws of this species show a third lobe on the p,—m,
and a third lobe on p,, both of which disappear in the old ani-
mal. The jaw on which P. triplex was founded appears to be
a juvenile stage of P. turgidus, in which these characters are
“very marked. We have no other jaws of the same age; but
if the twelve or fourteen examples of lower jaws be arranged
according to age (determined by wear on énd of p,) they form
a perfect series from P. triplex to the type of P. turgidus.
Cope, in comparing the series, came to the conclusion that
the difference between P. turgidus and P. triplex could not
be entirely explained as a matter of age; but the present
writer is unable to see sufficient difference to warrant the
retention of the species.
310 Bulletin American Museum of Natural History. (Vol. XVI.
Part of a skull, No. 1429a, is referred here; it belongs to a
very old individual, and the tip of the muzzle and brain-case
are missing. It appears to be a short-skulled species, con-
siderably broader but not much longer than P. intermedius;
the angle of the basifacial axis cannot be determined, and the
teeth are very much worn, so that the internal inflection of
the enamel has disappeared on the molars, although it persists
on pm,.
Very little additional material of this species has been found
by our party, and none that throws any new light on the
younger stages of tooth-change, so that Professor Cope’s pro-
visional reference of Tvicium paniense (juvenile P. turgidus)
cannot be confirmed.
Paleolagus temnodon Dovweglas.
Allied to P. haydeni but probably distinct, as it comes from
a lower horizon and a widely separate locality.
eae ) ; ; y
-AUTHOR’S EDITION, extracted from BULLETIN ~
r “:
OF THE
mevican Museum of Hatural History,
Vou, XVI, ARTICLE XXIII, pp. 311-316.
‘ae . x “
New York, Sept. 25, 1902. ;
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Article XXIII.— THE SKULL OF HYPISODUS, THE
SMALLEST OF THE ARTIODACTYLA, WITH A
REVISION OF THE HYPERTRAGULID/.
By W. D. MatTrHew.
A very well-preserved skull of this tiny Artiodactyl was
found by the writer at Pawnee Buttes, northeastern Colorado.
In connection with the fragments of the skeleton already de-
scribed from the same region, it gives a fairly complete idea of
the characters.
The animal was not larger than a ‘cottontail’ rabbit. The
orbits are remarkably large, as are likewise the tympanic
bulla. The tip of the muzzle is unfortunately missing, but
enough is preserved to show that it was slender and short.
The whole skull is distinctly more brachycephalic than in any
other White River selenodont—more so, indeed, than in any
of the modern Cervide or Antilopidz with which I have com-
pared it. The molar dentition consists of five teeth in each
jaw, p3} — mé; p§ is present in young individuals and repre-
Fig. 1. Hypisodus minimus Cope. Skull, x 3. No. 9354. White River (Cedar
. Creek Beds), Pawnee Buttes, Colorado.
sented by an alveolus in the older animals. A considerable
diastema is in front of this alveolus in the upper jaw, and
[311]
312 Bulletin American Museum of Natural History. [Vol. XVI,
anteriorly to this the muzzle is broken off on both sides (or is
not calcified). A longer diastema precedes the alveolus of p,
in the lower jaw, and in front of that, on evidence of other
specimens, it is known that there are five small subequal teeth,
probably incisiform.
The antorbital foramen is double, opening above the dia-
stema in front of the molar series. There appears to be a
small prelachrymal vacuity. The postorbital bar is com-
plete. The orbit is surrounded by a thin prominent ring, of
which the inferior and anterior parts (jugal and lachrymal)
are more prominent, and the superior part (frontal) less
prominent than in Hypertragulus or Leptomeryx. The eye,
therefore, faced much more upward than in these genera; it
was more prominent and much larger in proportion. The
basifacial axis is much more bent down on the basicranial
axis than in other Hypertragulide. The bulle are very large,
connate anteriorly, with a long, prominent, enclosed meatus
opening behind the origin of the zygoma. The occiput
projects much more backward than in Hypertragulus or Lepto-
meryx. The paroccipital processes are slender, and are co-
ossified with the bulle except just at the tip.
The lower jaw is slender, its condyle set high up, and the
long coronoid process is slightly curved. It does not possess
the angular hook seen in certain of the Camelide.
The limbs and feet I have described in a previous article.
The ulna and radius are codssified, and the distal end of the
fibula is codssified with the tibia. The cuboid and navicular
were coossified, the median metatarsals distinct though ap-
pressed, the laterals thread-like but still complete.
Hypisodus was much less Tragulus-like than Hypertragulus
or Leptomeryx, and superficially resembled rather the dwarf
antelope Madoqua. Its real relationships are more nearly
with Hypertragulus than anything else, but it is a remarkably
modernized animal for the formation in which it is found.
The resemblance to Madoqua is a striking instance of parallel
adaptation.
1902. ] Matthew, The Skull of Hypisodus. 313
Famity HYPERTRAGULID Cope.
(LEPTOMERYCID# Scott.)
Leptomeryx Leidy.
Dentition, If C®°71 P? M3. Lower canine incisiform, first premolar
small, caniniform, with a long diastema in front of it and a somewhat
shorter one behind. Upper canine small or absent. Second, third,
and fourth premolars in series with molars, p* with two crescents, p*~%
with anterior, posterior, and internal accessory cusps. Mesostyle
prominent on upper molars. Second, third, and fourth lower pre-
molars in series with molars, trenchant, with anterior basal cusp,
basin heel, and accessory ridges from the protoconid. Last molar
with heel composed of a large posteroexternal crescent and a small
anterointernal flattened cusp (as in Palgomeryx).
Skull of moderate length, muzzle slender, elongated, orbits not very
prominent. Prelachrymal vacuity diamond-shaped, situated rather
above than in front of the
lachrymal. Bulle small, not
filled with cancellous tissue.
Ulna and radius separate;
tibia and fibula separate; navi-
cular and cuboid united. Man-
us of four usable digits, the
lateral pair smaller, no cannon-
bone. Pes with cannon-bone
and no lateral digits (? the
small proximal splints of me-
tatarsals II and V fused to the cannon-bone). Hoofs small, pointed.
Fig. 2. Dentition of Leptomeryx.
1. L. evansi Leidy. White River (Oreodon Beds). 5S. Dak.
Leripy, Ext. Mam. Dak. and Neb., p. 16s, pl. xiv, figs. 1-8.— ScorrT,
jour. Morphs,) V,, Dec: rso07. Scott, Trans. Wagn. ‘Inst. Sci.,
Nis OOs 05
2. L. mammifer Cope. White River (? Titanotherium
Beds), Swift Current Creek, Canada.
Core, Rep. Geol. and Nat. Hist. Surv. Canada, I, 1885 (1886), 84.
One and a half times the linear dimensions of L. evanst.
Prof. Cope distinguished it by the presence of a separate
cusp between the entoconid and the heel of m3; this appears,
however, to be only the anterointernal cusp common to all
species of Leptomeryx.
314 Bulletin American Museum of Natural History. (Vol. XVI,
3. “L.” esulcatus Cope. White River (? Titanotherium
Beds), Swift Current Creek, Canada.
Corre, Mem. Geol. Sur. Can. III, 1891, 22, pl. xiv fig. 5.
Based on a single upper molar, which, judging from Cope’s
figure, is not Leptomeryx, suggesting rather an ally of Pal-
@omeryx, so far as such slight evidence is worth consider-
ation.
4. “L.”? semicinctus Cope. White River (? Titanotherium
Beds), Swift Current Creek, Canada.
COPE 8G. Py 23 nips Suvi wll One.
This species is also known by a single upper molar. It has
twice the linear dimensions of L. evans1; tooth broader
transversely, with heavy cingular ridge (protostyle) internal
to protocone and strong cusp (hypostyle) anterointernal to
hypocone. This is clearly not Leptomeryx, and is distinct
from any described White River genus (except possibly Cal-
ops, with which I am unable to compare it; Prof. Marsh
states, however, that the molars of Calops are like those
of Protoceras, in which case it is not “L.” semuicinctus).
Two upper molars in the Am. Mus. Collection from the
Protoceras Beds of South Dakota probably represent this
species.
5. Leptomeryx sp. indesc.
A smaller species occurs in the Leptauchenia clays in Col-
orado and in the Protoceras Beds of South Dakota distin-
guished by simpler premolars, narrower, more hypsodont
molars. The deuteroconid of p, is not distinct as in L.
evanst, but represented only by a ridge descending ante-
rointernally from the point of the protoconid; p, is smaller
and more trenchant; the internal faces of the molar cusps
are more convex. This may prove varietal when L. evanst
is examined from more localities and regions.
6. Leptomeryx sp. indesc.
A larger, somewhat more brachydont species or variety,
characteristic of the Protoceras sandstones. Premolars pro-
1902. | Matthew, The Skull of Hypisodus. 315
portionately larger, longer, and more complicated; antero-
internal cusp of p, less prominent, posterointernal (hyposty-
lid) more so; hypoconid ridge bifid posteriorly, into a
posterior branch which passes backward to the posterior
margin, and a posterointernal branch projecting into the
basin of the heel. P, more anterior in position than in L.
evanst, probably near anterior end of diastema, while in
L. evanst it is nearer to p, than to c,.
Hypertragulus Cope.
Dentition, It Ct P+ M$. Lower canine incisiform, first premolar
large, fully caniniform, with a short diastema in front and a long one
behind. Upper canine enlarged, first upper premolar two-rooted, with
diastema before and behind. Second lower premolar spaced, simple,
without accessory cusps; third with heel; fourth with anterior cusp
and heel; but all though equally trenchant simpler than the corre-
sponding teeth in Lepto-
meryx. Second upper pre-
molar simple, two-rooted;
third with internal cusp;
fourth with two crescents.
No mesostyle on upper mo-
lars. Heel of last lower
molar composed of two
equal opposite crescents.
Molars somewhat more hyp- -
sodont than those of Lepto- Fig. 3. Dentition of Hypertragulus.
meryx.
Skull much like that of Leptomeryx, but somewhat shorter, muzzle
slenderer, supraorbital ridges much more prominent. Prelachrymal
vacuity as in Leptomeryx, bulla somewhat larger.”
Ulna and radius codssified; tibia codssified with distal end of fibula;
navicular and cuboid united. Manus of four separate usable digits;
pes with two separate digits and splints representing the lateral meta-
tarsals.
1In his recent discussion of this genus Prof. Scott interprets the dentition as
I? C} P$ M3, the first premolar absent and lower canine caniniform. It would appear
rather that, as in most other selenodonts, the canine is incisiform. The number of in-
cisors is not certainly known; but it seems certain that both in the John Day skulls on
which Prof. Scott's description and figures were based, and in the White River skulls in
the Amer. Mus. collection, the inferior tooth, which Prof. Scott considers as a canine,
closes behind, not in front of, the upper canine; his drawing does not agree with the
specimen in this respect. The one skull from the White River in which this part is
preserved shows the lower caniniform tooth shutting unmistakably behind the upper
canine; it is, therefore, a premolar, and the canine is incisiform as in Leptomeryx.
316 Bulletin American Museum of Natural History. [Vol. XV1.]
1. Hypertragulus calcaratus Cope. White River, Oreodon
Beds.
”
H. tricostatus Corr.t Not H. calcaratus ‘“‘Cope”’ Scott, Trans.
Wagn. Inst. Sci., 1899.
2. Hypertragulus sp. indesc. John Day.
Hypertragulus calcaratus Scott, Trans. Wagn. Inst., 1899, pl. i,
figs. 3-4.
The John Day specimens referred by Profs. Cope and Scott
to H. calcaratus are a larger, more brachydont species, with
heavier muzzle, etc.
3. ‘‘Hypertragulus’’? transversus Cope. White River,
Titanotherium Beds (?), Swift Current Creek.
Cope, Mem. Geol. Sur. Can. III, 1891, 22.
Twice the linear size of H. calcaratus, para- and meta-
cones uniformly convex externally, small para- and metastyles
and prominent hypostyle anterointernal to hypocone. It is
not at all probable that this species is Hypertragulus or re-
lated thereto.
Hypisodus Cope.
Dentition, I; Ci p*$-# M8. Lower canine incisiform, first pre-
molar small, probably incisiform. Second upper and lower premolars
a decadent at anterior end of grind-
ing series. Teeth hypsodont, skull
very brachycephalic, but muzzle
rather long and slender. Pre-
lachrymal vacuity irregular, orbits
very prominent, bullz very large.
Lateral digits of pes extremely
slender although still entire; no
cannon-bone; ulna and radius united.
Fig. 4. Dentition of Hypisodus.
1. Hypisodus minimus Cope. White River (Oreodon clays).
yrs I know but the one species, the H. calcaratus Cope.’’—Cope, Proc. Amer.
Phil. Soc., 1884, Vol. XXII, p. 24. This statement invalidates H. tricostatus, and quite
correctly so.
aS prings, Nebraska.
By W. D. Martuew.
ENOL. NG, ARTICLE XXIV, pp. 317-322.
New York, Sept.’ 25, 1902.
Article XXIV.—LIST OF THE PLEISTOCENE FAUNA
FROM HAY SPRINGS, NEBRASKA.
By W. D. MatTTHeEw.
In 1893 and 1897 field-parties from the American Museum
were sent out by Professor Osborn to collect in the Pleistocene
at this locality, a bone-bed near the Niobrara River, not far
from Hay Springs. A large collection of horse and camel re-
mains was obtained, and a few specimens of other animals.
The horses have been carefully studied by Mr. Gidley, the
camels by Dr. Wortman; the remainder of the fauna has been
revised and partially studied by the writer, but no results
have hitherto been published. A list of the fauna may be of
some interest to compare with that found in the Sheridan or
Equus Beds at other localities. The-appended lists, when
based on materials in the Cope Collection, are from deter-
minations by Professor Cope, revised in a few cases by the
writer.
Hay SPRINGS, AMERICAN MusEvUM COLLECTION, 1893 AND 1897.
Canis ? latrans. Lower jaw.
? Dinocyon or large Ursid. Metacarpal. This bone ap-
pears to exceed in size the corresponding parts in any living
species of bear. The character of the diaclasts excludes it
from the Felide or typical Canide, but the Amphicyonine
Dogs are somewhat similar, and if not a bear it might repre-
sent a very large species of Dinocyon (Borophagus), a genus
known to occur in the late Pliocene (Blanco).
? Felidze indet. Several species are represented by foot-
bones and fragments of limb-bones.
Fiber zibethicus. A skull and several jaws, all of which
come within the limits of variation of the modern muskrat.
Arvicola, cf. amphibius. Upper and lower jaws.
Cynomys, cf. ludovicianus. Palate.
Thomomys sp. indet. Upper and lower jaws.
Castoroides sp. indet. Teeth, limb-bones, and astragali.
Mylodon sp. indesc. A complete skull, jaw, and large part
[317]
318 Bulletin American Museum of Natural History. [Vol. XVI,
of the skeleton of a sloth allied to but distinct from Mylodon
harlani. Description is reserved for the present.
Equus complicatus Leidy. This, the first-described and
most characteristic of American fossil horses, is by far the
most abundant fossil at the Hay Springs locality. Numerous
bones of all parts of the skeleton were obtained.
Equus fraternus. Smaller and not nearly as abundant as
E. complicatus.
Equus ? scotti Gidley. Upper and lower jaws.
Elephas primigenius columbi. Tusk, several foot-bones,
and grinders.
Platygonus vetus Leidy. Palate.
Platygonus compressus Leconte. Upper and lower jaws.
Undescribed Porcine, cf. ? Leptocherus. Upper premolar.
(obtained in digging a well near Hay Springs).
Camelops kansanus Leidy. Parts of jaws, teeth, vertebre,
limb- and foot-bones. More than one species is quite probably
represented, but the material does not warrant attempt at
separation.
Camelops vitakerianus Leidy. Teeth, jaw fragments, etc.
Camelus americanus Wortman. Lower jaw.
Antilocapra, cf. americana. Parts of jaws, limb- and foot-
bones.
Capromeryx furcifer, n. g. et sp.
A small jaw containing p,—m, indicates an animal allied to
Antilocapra and somewhat more nearly to Merycodus, but
generically distinct from either. The premolars are most
nearly like those of Merycodus; p, is long, trenchant, simple,
lacking the deuteroconid; p, and especially p, are more com-
plicated. In Antilocapra p, is shorter, with prominent deu-
teroconid, and the anterior premolars are short and simple,
thus approaching the premolar characters of goats and sheep.
The molars in Capromeryx are fully as hypsodont as in Antzlo-
capra, much more so than in Merycodus, which more nearly
approach the brachydont teeth of the deer.
The species was of about two-thirds the (lineal) dimensions
of the American antelope.
1902.] Matthew, Pletstocene Fauna from Hay Springs. 319
This unexpected addition to the short list of Ruminants of
the American Pleistocene is rather interesting despite its
fragmentary character. It is, much more certainly than An-
tilocapra, descended from the little group of antelopine deer of
the American Miocene, of which Blastomeryx, Cosoryx, and
Merycodus are the known forms. This group is characterized
by the combination of antlers approaching those of the deer
and teeth approaching those of the antelopes. The antlers
are forked or several times branched, provided usually (if not
always) with a burr,—hence, in Professor Cope’s opinion,
deciduous; smooth surfaced,— hence probably covered per-
manently with ‘velvet.’ The teeth are more hypsodont
than in any of the deer, less so than in Antilocapra, and the
premolars have preserved somewhat of that primitive, long,
trenchant character seen among modern genera only in Trag-
ulus, but generally present among the older Tertiary seleno-
donts. Blastomeryx (B. gemmzfer) is the oldest, smallest, and
most brachydont genus; Merycodus is more hypsodont.
Cosoryx has a simple forked antler, just above the eyes, like
the horn of Antilocapra, with a burr at base, and quite hypso-
dont teeth.
Blastomeryx has antlers like those of the Virginia deer, but
I have not seen more than four tines; the face is very short and
the antlers more nearly over the eyes than in modern deer,
less so than in Antilocapra. The position of Merycodus is un-
certain; present evidence indicates that it may be distinct
from Cosoryx. The large, completely brachydont species re-
ferred by Cope and Scott provisionally to Blastomeryx and by
Douglas to Palgomeryx are distinct from any of these antelo-
pine deer (and from any European genus as well), forming a
transition between them and the contemporary and later true
deer of Europe and America. The new genus, Capromeryx,
may, when better known, prove to be a transition between
the antelopine deer and Antilocapra.
The above list is obviously a plains fauna. Horses and
camels are the most abundant. There are antelopes, but no
deer; Canide are found, but few if any Felide. Platygonus
320 Bulletin American Museum of Natural History. [Vol. XVI,
is much more swift-footed and more advanced in dentition
than is the modern peccary, and may be supposed to have
lived more in the open. Prairie-dogs, gophers, and field-mice
are now to be found on the plains in the same region, and
muskrats along the streams. Of the habits of Mylodon we
know little; perhaps it, ike the mammoth, frequented the
watercourses and valleys of open country, rather than the
denser forest regions which were the home of the contem-
porary mastodons. The Castoroides remains are too fragmen-
tary to tell whether it was the same as the eastern species.
SILVER LAKE (LAKE Co.), OREGON.!
Canis ? latrans, jaws, limb- and foot-bones.
Canis, cf. occidentalis, limb- and foot-bones.
Vulpes, cf. pennsylvanicus, femur and tibia.
Lutra canadensis, front of skull, jaws, limb-bones.
Fiber zibethicus, jaws, limb-bones.
Arvicola sp. div., jaws, limb-bones.
Thomomys sp., skeleton nearly complete.
Geomys sp., jaws, limb-bones.
Castor sp., one molar.
Castoroides sp., teeth.
Lepus sp. (cf. campestris), parts of jaws, limb-bones, etc.
‘Mylodon sodalis Cope (? = M. harlant), phalanges.
Equus pacificus, numerous bones from all parts of skeleton.
Elephas primigenius ? columbi, teeth, foot-bones, vertebre,
etc.
Platygonus, cf. vetus, teeth.
Platygonus sp. minor, teeth.
Eschatius conidens, parts of jaws.
Camelops kansanus, parts of jaws, limb- and foot-bones.
Camelops vitakerianus, upper jaw, ? foot-bones.
? Camelops sp. max., teeth, foot-bones, etc.
Antilocapra, fragments of feet.
With the above mammalia were found numerous bird re-
mains, which have been studied by Dr. Shufeldt.
1 Revised from Prof. Cope’s list.
1902.] Matthew, Pleistocene Fauna from Hay Springs. 321
This is equally a plains fauna, with two aquatic mammals,
Castor and Lutra, not found at Hay Springs. Otherwise the
list is very similar to that of Hay Springs, and, like it, is
characterized by the absence of the forest types found in the
Pleistocene cave deposits, river-gravels, and peat-bogs of the
East.
OREGON DESERT.
A collection made by Geo. C. Duncan at some point or
points of which I can find no exact record. The collection
was considered as of Pliocene age by Professor Cope, because
it contained Equus, Hipparion, and Teleoceras, along with less
characteristic remains referred to Holomeniscus (= Camelops)
and Elephas. The Equus and Elephas bones, however, are
from a different matrix from the other bones, and the Holo-
meniscus is more probably Pliauchenia or Procamelus. They
are, therefore, probably from two distinct formations, the
older one of the age of the later Loup Fork, the newer one of
the age of, if not identical with, the Silver Lake Equus Beds.
WASHTUCKNA LAKE, WASHINGTON.
Taxidea sulcata (=americana), parts of skulls, jaws,
limb- and foot-bones.
Felis, cf. ampertalis, parts of limb-bones.
Felis, cf. concolor, parts of limb-bones.
Felis, cf. canadensts, parts of limb-bones.
Mylodon sp., astragali and foot-bones.
Equus sp., bones of feet and some teeth.
? Camelops, cf. kansanus, foot-bones.
? Camelops, cf. vitakerianus, foot-bones.
? Camelops, sp. max, foot-bones.
Alces brevitrabalis, parts of antlers, foot-bones, etc.
Alces semipalmatus, parts of antlers, foot-bones, etc.
Cariacus enstfer, parts of antlers, foot-bones, etc.
Oreamnus, parts of horn.
This fauna shows a large proportion of forest and mountain
types, and no aquatic mammals. It is a very inadequate list,
322 Bulletin American Museum of Natural History. (Vol. XV1.]
and I have seen no description of the locality where the speci-
mens were found, but it is probable that the physical condi-
tions were quite different from those prevalent in the Silver
Lake and Hay Springs localities.
_ AUTHOR’S EDITION, extracted from BULLETIN
OF THE
ae a ARTICLE XKKE: pp. 401-407, Pll. tiv-Lyv.
Mew Voee: Nov. 3, 1902.
ra Press, ew Work
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Article XXXI.—ON JURASSIC STRATIGRAPHY ON
THE WEST SIDE OF THE BLACK HILLS—SEC-
OND PAPER ON AMERICAN JURASSIC STRATIG-
RAPEDY
By F. B. Loomis.
PiLaTes LIV anv LV.
During June, rgo1, a party, sent out by Professor Osborn,
from the American Museum of Natural History, prospected
the Jurassic exposures of the west side of the Black Hills for
Dinosaur remains. In connection with this work sections
were made by the writer at six of the best and most character-
istic exposures, to show the stratigraphy of these deposits.
These were made at Professor Osborn’s request for com-
; q
parison with the sections previously made by the writer in the
Como district of Wyoming.
The area covered extends from a little north of Hulett to
fifteen miles south of Newcastle; the exposures stretching
some 125 miles in length, and varying from a fourth of a mile
to ten or twelve in width. The distribution of the Jurassic
in the region is shown in the map accompanying the paper,
as is also the location of the individual sections. The Juras-
sic is exposed mostly in escarpments, capped by the heavy
Dakota sandstone, which makes the ‘rim.’ These escarp-
ments generally face toward the centre of the Hills and con-
tinue on around the north and east sides as well as on the
west side. The soft clays which predominate in the Upper
Jurassic are the cause of a considerable valley all around, just
inside the ‘rim.’ The strata dip in varying degrees away
from the centre of the Hills, but are in the best exposures
nearly horizontal.
The Jurassic is divisible into two parts: a lower marine,
corresponding to Knight’s Shirley‘; and an upper fresh or
brackish water corresponding to Scott’s Como.?_ On the east
1 W. C. Knight, ‘Jurassic Rocks of Southeastern Wyoming,’ Bull. U. S. Geol. Surv.,
Vol. GF ee 377-388.
2 W. B. Scott, intoanction to Geology, p. 447 (footnote).
[4or]
402 Bulletin American Museum of Natural History. |Vol. XVI,
side of the Hills, a thick bed of sandstone immediately over-
lies the Triassic; but this bed is everywhere lacking on
the west side of the Hills} the transition being uninterrupted,
and seen only in the change from sandy red clay to sandy
green clay.
On comparing the sections made on the east side of the
Hills,t these on the west side, and similar sections made in
central and eastern Wyoming,? it becomes clear that the
Jurassic of the west side corresponds more closely with that in
central Wyoming (especially on the Medicine and Como anti-
clines) than the west side corresponds to the Jurassic on the
east side of the Black Hills. However, the upper or fresh-
water beds of the east and west sides correspond with one
another better than the marine layers. Many of the most
striking layers of the west side can be detected in the centre
of Wyoming almost exactly as they occur in the Hills. To
bring this out strongly, in the table where the sections are
set side by side, the same series of numbers is used as in the
table 3 of sections from central Wyoming, and such layers as
are recognized as being equivalent are printed in heavy type.
The Triassic of the west side of the Black Hills is made up of
barren red sandy clay. This grades into the green sandy clay
of the base of the Shirley, which for some eight to ten feet is
also barren. Here, however, Belemnites densus begins to ap-
pear in great abundance. This clay is then the equivalent of
and like in texture to the Belemnites layer further west (No. 2).
The upper part of the layer may carry thin beds of limestone
as in the Inyan Kara Peak section. This is overlaid by a
layer of green clay with large limestone nodules (No. 4),
which vary in size from six inches to a couple of feet in di-
ameter. It is in and on just such nodules that Baptanodon
remains are found in the centre of the State. The layer was
everywhere 4 present both on the west and east sides of the
Hills. However, no trace of Baptanodon was found in any of
the exposures; but in the stone-pile of a yard on Miller Creek,
* The east side sections are in manuscript; they were made by G. R. Wieland, and
verified by myself.
? Loomis, Bull. Amer. Museum. Nat. Hist., Vol. XIV, pp. 189-107.
3 Last cit., p. 192
* Except in the Salt Creek section, which is an unusual one.
1902.| Loomis, American Jurassic Stratigraphy. 403
two or three Baptanodon vertebre were discovered, and
probably came from this horizon in some of the nearby ex-
posures. In the centre of Wyoming this layer has a purple
hue, but is otherwise similar.
A sandy limestone or shell sandstone (No. 5) usually fol-
lows, which carries several invertebrate marine forms. At
the Bellé Fourche station, this layer held Amaltheus cardi-
jormis, Afgoceras tumidus v. Buch, Ostrea strigulecula W..,
Tancredia tnornata M. & H., Pseudomonotis curta W., P. orbic-
ulata W., Dosinia jurassica W., Trigonia sp., and Pholadomya
sp. This fauna with some variations is widely distributed at
this horizon.
From this horizon to the top of the marine Jurassic there
is no uniformity in the character of the beds, usually, as in
‘the Belle Fourche section, the clays alternating rapidly with
thin beds of sandstone. The Salt Creek section has soft sand-
stones alternating with denser ones. The Beaver Creek sec-
tion has nothing to represent the alternations. In central
Wyoming there are several clays and sandstones. There is
no uniformity except in the rapidly changing character of the
deposits.
The top of the marine beds is a green clay (No. 12) of vary-
ing texture. The change to fresh- or brackish-water is a
gradual one, so that a distinct boundary is difficult to find, but
this bed is the highest in which any trace of marine life was
found, and is, therefore, used here as a convenient separating
horizon.
The base of the freshwater deposits is a bed of sandstone
(No. 13), varying greatly in thickness and in character.
No. 14 isa layer of green clay, the lowest in which any traces
of Dinosaurs were found. On the north side of Inyan Kara
Peak a few fragments of sauropod limb bones occurred; and
near the Sheldon P. O. section a few foot bones were found at
this level. At the Belle Fourche station traces of Dinosaurs
also occurred. In all cases the bones were uniformly hard, but
very scarce.
1 At the Sheldon P. O. station there occurred Camponectes platissijormis W., Tan-
credia warrent M.& H., T. bulbosa M. & H., Avicula mucronata W., and Ostrea strigu-
leculaM.& H. The Kara Peak section had Amaltheus cardiformis M. & H., A®goceras
tumidus v. Buch, and Pseudomonotis curta W
404 Bulletin American Museum of Natural History.[Vol. XVI,
The Inyan Kara Peak section is remarkable for its thick-
ness. This is most marked in the No. 15 bed of sandstone
which is here 75 feet thick. The layer is very generally a
fairly heavy one and occurs in all the sections.
Next follow several thin layers of variable character, con-
sisting of clays, sandstones and limestone concretions. There
is no constant bed till No. 22 is reached, which is a band of
maroon clay filled with tiny concretions. It is present in
three of the sections and occupies the same position as a simi-
lar band in the centre of Wyoming. It is avery good horizon
marker, being so distinctive in texture and conspicuous in
color. Just above a very constant layer of limestone nodules
(No. 23) occurs.
The layers from 24 to 28 are a series of brilliantly colored
clays, red, purple, and green in color, and popularly called
the ‘‘varigated clays.’’ These clays have occasional beds of
limestone nodules. In the variegated beds, especially toward
the top, Dinosaur remains are not infrequent, but without
exception the bones are in a wretched state of preservation,
the iron in the colored beds having eaten into them till great
spots are mere powder. In all the sections some traces of
Dinosaurs were found at this horizon, but they were especially
abundant along the lower reaches of the Inyan Kara Creek.
On Inyan Kara Creek in these same beds Mr. Thompson found
several specimens of Unto baileyi L. and Valvata leet L. similar
to those found by Logan in the Freezeout Mountains. These
variegated clays occur on the east side of the Hills, as well as
on the west side, also in the Freezeout Mountains, and in the
southeastern part of Wyoming.
The top of the freshwater series is everywhere a bed of olive-
green clay of considerable thickness, in which Dinosaur re-
mains are extremely rare, but do occur; and where present,
the bones are in a good state of preservation. Fragments
were found at the Belle Fourche station and near Inyan Kara
Peak. The whole is capped by the heavy bedded Dakota
sandstone. The sections are often complicated by this sand-
stone faulting and slipping part way down the slope of the
escarpment. In fact the greatest care is required to find ex-
1902.] Loomis, American Jurassic Stratigraphy. 405
posures where more or less of the face has not faulted and
slipped to some extent. The majority of all the Dinosaur
prospects were in clays which were faulted out of place.
Of all the sections the Inyan Kara Peak one is the thickest.
This is due in great part to the frequency and extra thickness
of the sandstones. The Beaver and Salt Creek sections in the
same general neighborhood also have larger quantities of
sandstone, which fact I take to mean that in the neighbor-
hood there was some land mass during a part at least of the
Jurassic period.
EXPLANATION OF THE PLATES.
Fig. I, Map of the country on the west side of the Black Hills, cov-
ered by the prospecting party of the American Museum of Natural
History in June, gor.
Line A-B, Belle Fourche Section.
C-D, Inyan Kara Creek Section.
E-F, Sheldon P. O. Section.
G-H, Inyan Kara Peak Section.
I-J, Salt Creek Section.
K-L, Beaver Creek Section.
The map is modified from Scott’s Mineral and Geological Map of the
Black Hills.
Fig. II, Belle Fourche Section.
The vertical and longitudinal enlargements are the same in all the
sections.
Fig. III, Inyan Kara Creek Section.
Fig. IV, Sheldon P. O. Section.
Fig. V, Inyan Kara Peak Section.
Fig. VI, Salt Creek Section.
Fig. VII, Beaver Creek Section.
BELLE FOURCHE SECTION.
Ft,
INYAN KARA CREEK/SHELDON P. O. SEC-
SECTION. TION.
31 Dakota sandstone. ..180+
Olive-green clay.... 70
30 | & MA i
20
f
Oa)
o % | Yellow-green clay
NA with small concre-
vs HONS oe ee ees r2
| 24
3
a 8
a l Maroon clay......... 10
Ss | Green clay......... 6
Limestone concre-
23 ; TOUS WorieiLieiials I
Greeniclayscc..c. 6
22 Maroon clay with
small concretions. 6
. 21 Greenclay with small
2 concretions....... 10
B20 ;
<
%
5
ey Limestone _concre-
ao ; HONS epee cote I
& (\Green| clay. .-~ 4. 0-- 4
= 18
m 17 2
2 Limestone _concre-
~% 16 IONS ee creer el I
ied eee 6
15 Soft yellow sand-
StOGC rr ioe ieee 2
14 Greenclay......... 7
13. Soft yellow sand-
STOME meas eucwevenersies 2
12a
(Green clay weather-
| ing brown........ 25
12 4 Limestone _concre-
3 LOS rao ao odtados I
- | Green clay weather-
B inp brow. 1-20
~ It Gray sandstone.... 2
fe OF Green /ClaVs 5a. 10
a (Gray sandstone..... 2
z Green-clay: . tic. ce: U |
5 Gray sandstone..... 1
<
= 7 4Greenclay......... 4
Gray sandstone..... 1
Green clay......... 6
Gray sandstone..... 1
Ol Sand vaiclayemeeeeoes I4
5 Shell limestone..... 3
fiSandy clays cic IO
|
(Green clay with
Belemnites densus 20+
2
Ft. Ft.
Dakota sandstone... 80+} Dakota sandstone... 50+
Olive-green clay.... 40 |Olive-green clay.... 30
Light green clay.... 12
Maroon clay with
small concretions. 10
Green clay with
smallconcretions. 5
Red clays. ans ae 5 '
Limestone _concre-
HL 556 GO EC O00 I
Green clay......... 5 |Greenclay....... Io
Maroon clay....... 6. |Maroon clay....... 5
Green clay......... 20 | Yellow-green clay... 9
: Rediclaye serie 5
Limestone concre- Limestone concre-
IONS arohaeeeterte ee I GionS Fe syste
Greenvelay ate enn
Limestone concre-
Vator ees Dt oho ooo I
Rediclay=-asenner 3
Yellow sandstone... 9
Dense gray sand-
SCONCE iene cisteieiotcas 8
Green‘clay.. «2... 12
Limestone concre-
+1OMS' ashes eect I
Green clay; eo. mene
Yellow sandstone... 5
Limestone concre-
CLOTS vevetelaraietare lated I
Yellow sandstone... 12 |Gray sandstone..... 2
Limestone _concre-
EIQUS sie tse eee I
Green clay.......... 8 | Yellow-green clay... 2
Gray sandstone..... 2
Green clay with
Modwlesmaane cares 6
Soft sandstone..... 1o |Soft sandstone..... 10
Gray sandstone..... 4
Olivetclayy-i4-ch- mo
Sandstone with
shells? nearer 2
Olive clay:. 0. <2. re Ss
Sandstone with
Shells erertel eters 2
Oliveiclay....+5.-«1 5
Gray sandstone..... 2
Olivetclayz ae cioe 1+| Olive-green clay
with nodules..... 35
Concretions aac aee
Olive-green clay with
MOdtUeSmerr tenets 12
Era allevateus laqetadous ay eeeveyers LOT sroeneene ates Ont
Norr.—Numbers in heavy type indicate that the layer can be recognized in the
centre of Wyoming
[406]
KARA PEAK SECTION
Ft.
31 Dakota sandstone. . 15-4
304
20
2 |
a
oO <
a4
a Red clays ste .e. 3
ei
2
ee Green clay.. ase ke
a & | Cream sandstone. . 8
Bs Blue-green clay.. 6
iS leed™ Claymecrite- tater I
Limestone concre-
23 TIONS teers I
Purple’ clay sass ase: 3
22) ) Reditclayae-es- ee 2
321 Cream sandstone... 2
a
B20
Pe)
5
aS
% 19
E oe clay. . aS
“<1 Cream sandstone... 2
E 17. White sandstone.... 11
z Black iclayaerrre seis 2
16
fe White sandstone.... 8
1s Yellow sandstone... 75
14 Slate-green clay.... 50
13. Gray sandstone..... 15
12a
( Yellow-green clay.. 6
\
Pelz
2 | Oli ive-green clay.... 6
2 aa
(o} LE
P 8
= (Sandstone ......- 5a
B Green clay with
5 MOUUMIES eee iecersues 3
<
em a
White sandstone.... 12
i
6
5 :
(Green clay with
nodules aaa). sce 12
4 4Soft gray sandstone. 8
|Green clay with
L eOsKEIES. oa ooo pe 82
(Belemnites densus
(DEdINER A has 2
(Geese, EN 6 on conne 20
2 4 Belemnites densus
Aseel coe onccusedon Io
Limestone layer.... 3}
(Sandy green clay... 16
ra
Motel nay: mectemte 4s 373%
t Red Triassic sandy
ClaVitid ste sys a eeu 50+
SALT CREEK SEC-
TION.
Ft,
Dark red sandstone 20
Olive-green clay with
Belemnites densus 12
Pink sandstone..... 8
Buff sandstone..... 4
Soft pink sandstone 4
Dense buff sandstone 15
Soft buff sandstone. 8
Buff sandstone..... 8
Olive-green clay ae SYS
Gypsumiyer aceite 6 3
Voter eben aclaoes <= Tay,
Red Triassic sandy
EK nos Fou D oe eeee 90+
BEAVER CREEK SEC-
TION.
Ft.
Dakota sandstone.. 20+
Brown-green clay.. 15
Olive-green clay.... 66
Green clay with small
concretions...... 15
Maroon clay with
small concretions 6
Green! clay... - 4: I5
Maroon clay with
small concretions. 12
Green clay.........
Limestone concre-
TIONS Poh ae eile I
Olive-green clay.... 8
Buff sandstone..... 12
Green clay with gyp-
SUMeeeias. cesiaee 20
Gray sandstone.... 1
Green clay with
MLOGUIES! -felete wiesete 30
Olive clay with
Belemnites densus 60
Sandy greenclay... 10
TAI COE eee 277
Red Triassic sandy
GlAYe cin iaie aioe 40
eee
i.
ae
- _,
j
rere
us
4
2.
—
2
a
oe
*
OPED sry
( Be/70 fourche.Fiver
I}
il
n ;
~ > Little Beaver CF
‘Ohive Green chay,
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ae re 4
On Certain Genera and Species of North
American Cretaceous Actinopterous Fishes.
By Ov P. HAY;
AUTHOR’S EDITION, extracted fom BULLETIN
OF THE
American Bluseun of Hatuval History, -
VoL. XIX, ARTICLE J, pp. I-95.
PLATES I-V, AND 72 TEXT FIGURES.
New York, March 4, 1903.
“
The tknlickerbocker ‘Press, Rew :
BULLETIN 4
AMERICAN MUSEUM OF NATURAL HISTORY. 4
Votume XIX, 1908. ee
Article I— ON CERTAIN GENERA AND SPECIES OF 2
NORTH AMERICAN CRETACEOUS ACTINOP-
TEROUS FISHES.
By: PEAY.
PiaTes I-V, AND 72 TEXT FIGURES.
The present paper has resulted from observations made by
the author on Cretaceous fishes in the course of his work of
identifying, cataloguing, and arranging the Cope Collection of
fishes and reptiles, now the property of the American Museum ee
of Natural History. In this collection are most of the types ah
of the fishes which Professor Cope described from the Creta-
ceous deposits of Kansas, South Dakota, and New Jersey; and
there are likewise many other specimens which had not been
carefully identified and studied. A comparison of these ma-
terials with the types, and of the types with one another, and t
‘an estimation of the value of the proposed species in the light
of work done by more recent investigators, have resulted in
the reduction of a considerable number of nominal species to
_ the position ofsynonyms. Many of Cope’s types have hitherto
never been figured and the opportunity offered by the liber-
ality of the Museum authorities has been employed to furnish
many drawings and photographic reproductions of interest-
ing specimens. My thanks are especially due to Prof. Henry
F. Osborn for the opportunity to prepare and present this 4
[ fanuary, 1903.] {r] it
2 Bulletin American Museum of Natural History. [Vol. X1X,
paper. Of the drawings, Mr. R. Weber has made numbers
27,50, 50,00, OL, 62, and 65% -All ithe others except genau
72, have been prepared by Mrs. L. M. Sterling. The photo-
graphs for the plates have been made by Mr. A. E. Anderson,
photographer of the Paleontological Department.
PROTOSPHYRANIDA.
The genus Protosphyrena is referred by Dr. A. S. Wood-
ward to the Pachycormide; and in this procedure he is fol-
lowed by Loomis (Palzeontogr., XLVI, 1900, p. 221), by
Stewart (Univ. Geol. Surv. Kan., VI, 1900, p. 362), and by
the present writer (Bibliog. and Cat. Foss. Vert. N. A., 1902,
p- 378). A reconsideration of the subject and the study of
the materials at hand in this Museum have led me to a differ-
ent view.
If we refer the genus to this family we must assume that
the vertebral column was not at all ossified or only feebly so.
That it was composed of well ossified vertebra cannot yet be
proved. However, accompanying the type of P. dimidiata
there is a single vertebra which bélonged close to the skull.
It is figured on page 19. This vertebra may be an intrusion
from some other fish, but there is nothing in its appearance
to suggest this.
So far as the writer can gather from the literature, neither
the Isopholide, hitherto called Eugnathide, nor the Pachy-
cormidé, possess ossified scapulz and coracoids. Even the
members of the more advanced family Amiide, with well
developed vertebre, have scapule and coracoids cartilagi-
nous. Protosphyrena, on the other hand, has the elements
of the shoulder girdle developed as in the modern Isospondyli.
While there may be no necessary connection between an
ossified shoulder girdle and ossified vertebre, it seems logical
to believe that, when the shoulder girdle is so advanced in
its development as it is in Protosphyrena, there were probably
also well defined vertebre. It is remarkable that vertebre
have not been certainly collected, but neither has the tail fin
been obtained, nor the anal, nor the dorsal fin.
It is proper to add to the above paragraph the statement
1903. | Hay, North American Cretaceous Fishes. a
that in a specimen of Hypsocormus from Solenhofen, which
the writer has been able to examine, there are evidences of
the presence of ossified scapula and coracoid. If this shall
prove to be the case, the fact that these elements are ossified
in Protosphyrena will have no bearing on the question re-
garding the presence of vertebre.
While there are many interesting and important characters
common to Hypsocormus and Protosphyrena, there are also
many striking differences. I regard the deeply socketed
teeth of Protosphyrena as furnishing a character of family
value. Woodward indeed states that the teeth of Hypso-
cormus are in incomplete sockets, those of the dentary of H.
tenutrostris (Cat. Foss. Fishes, IV, p. 397) being fused with
the bone in sockets which are incomplete on the inner side.
The condition of such teeth is certainly very different from
that of the teeth of Protosphyrena, which are in complete and
very deep sockets. It evidently signifies a great departure
from the primitive condition in fishes, when teeth become so
deeply implanted in the bone and are replaced, not by new
teeth developing in the mucous membrane of the mouth, but
from germs lodged deep in sockets.
The pectoral fin of Protosphyrena is quite different from
that of Hypsocormus, as may be seen by comparing the
description and figures of P. perniciosa presented in this
paper with the description of the fin given by Dr. Wood-
ward on page 398 of the work cited above. In Protosphyrena
there is no such intimate fusion of rays and the foremost rays
are extremely short.
The condition of the shoulder girdle of Protosphyrena
would appear to relegate the genus to the Isospondyli.
Whether the possession of a splenial and a slight excess of
baseosts is sufficient to exclude it from this order may have
to be determined hereafter. At any rate, the genus is close
to the border line between the Halecomorphi and the Isos-
pondyli.
Protosphyrena nitida (Cope).
Erisichthe nitida Core (E.D.), Proc. Acad. Nat. Sci. Phila. 1872, p.
280; Bull. U.S. Geol. and Geog. Surv. Terrs. I, No. 2, 1874, p. 42;
4 Bulletin American Museum of Natural History. (Vol. XIX,
Vert. Cret. Form. West, 1875, pp. 217, 275, pl. xlviii, figs. 3-8;
Bull. U. S. Geol. and Geog. Surv. Terrs. III, 1877, p. 821 (in part).
Protos phyrena nitida NewTon (E. T.), Quart. Jour. Geol. Soc. XXXIV,
1878, p. 794.— FeLix (J.), Zeitschr. deutsch. geol. Gesellsch.
XLII, 1890, p. 278 (in part).— Woopwarp (A. S.), Cat. Foss.
Fishes Brit. Mus. III, 1895, p. 409. — Loomis (F. B.), Palezontogr.
XLVI, 1900, p. 227 (in part only). — Hay (O. P.), Bibliog. and
Gat. Bosse Vert. Nea t902."p370:
Pelecopterus chirurgus Cope (E. D.), Vert. Cret. Form. West, 1875,
PP. 2448, 273, pl. xlviti, fig. 1; pl. liv, fig. 9:
Protosphyrena chirurgus Hay (O. P.), Bibliog. and Cat. Foss. Vert.
NEMA £902, )..370:
Erisichthe penetrans Corr (E. D.), Bull. U. S. Geol. and Geog. Surv.
errs ENS 7/7 ap o2 2)
Protosphyrena penetrans Newton (E. T.), Quart. Jour. Geol. Soc.
XXXIV, 1878, p. 795. — FELIX (J.), Zeitschr. deutsch. geol. Ge-
sellsch. XLII, 1890, p. 297, pl. xiv, fig. 1.— Crook (A. R.), Pala-
ontogr. XX XIX, 1892, p. 1o9. — Woopwarp (A. S.), Cat. Foss.
Fishes Brit. Mus. III, 1895, p. 409. —StTEWwartT (A.), Kan. Univ.
Quart. VII, A. 1898, p. 192; Univ. Geol. Surv. Kansas, VI, 1g00,
p- 369, pl. xiii, fig. 4. — Loomis (F. B.), Paleontogr. XLVI, 1900,
p. 227, pl. xix, figs. r-;.—Hay (O. P.) Bibliog. and Cat. Foss. Vert.
Nig dels HOO, 1a S/O):
Protosphyrena obliquidens Loomts (F. B.), Paleontogr. XLVI, 1900,
p. 225, pl. xx, figs. 1-4.— Hay (O. P.), Bibliog. and Cat. Foss.
Werte NeeAG To O25 pans zor
The type of Professor Cope’s Erisichthe nitida is now in the
American Museum of Natural History. It consists of both
premaxillze, a portion of the left dentary, and the adhering
postsplenial, the supposed hyomandibular, and a bone (Vert.
Cret. Form. West, pl. xlviii, fig. 7) which is as yet unidentified.
Of the same specimen Cope possessed also a fragment of the
fin, which lacked the anterior edge and was therefore not
susceptible of comparison with other species based on parts of
fins. This fragment is now missing. Dr. Loomis (Palzontogr.,
XLVI, p. 228) has questioned that the fragment of the den-
tary figured by Cope belongs with the premaxilla; but there
is no reason to doubt that all the parts of the type belong to
the same individual. As regards the premaxilla it may be
remarked that the anterior fang is probably directed more
horizontally forward than is natural, a fact due to pressure.
1903. | fay, North American Cretaceous Fishes. 5
In 1877, Cope, as cited, described from materials collected
in Gove County, Kansas, by Mr. Russell Hill, a species which
he called Erisichthe penetrans. The type, never figured
hitherto, is now in the American Museum. Its number is
2105. Views of this type, seen from below and from above,
and three cross-
sections are here
presented (Figs.
mand 2). - The
specimen con-
sists of the snout
from the front
of the orbits
tO, the tip of
the rostrum, but
no other parts.
On the lower
side the surface
of the bone has
been damaged,
so that the or-
namentation is
removed over a
considerable
area; but where
preserved, it is
not especially — Fig. 1._ Protosphyrena
different from Aah pier ears pe
trans Cope. Rostrum seen Fig. 2. Protosphyrena nitida
that of the Uup- from below. X 3. eth., (Cope). No. 2105. Type of Evés-
ethmoid; Jay., parasphe- zchthe penetrans Cope. Seen from
per side. It 18 noid; von, vomer; vom.?¢., above. % 4%. With cross-sections.
yvomerine tooth. eth., ethmoid ; /~., frontal bone.
somewhat coars-
er, and for a short distance in front of the vomerine fangs
forms areole resembling those of the shell of 7rionyx.
The specimen has suffered some crushing, and this affects
especially the region from the orbits to the vomerine fangs,
but also to some extent nearly the whole length of the rostrum.
Cope has given the vertical diameter of the base of the ros-
trum as 20 mm., but this is not correct. The diameter is
6 Bulletin American Museum of Natural History. (Vol. XIX,
24 mm., and was in life probably somewhat more. The same
author has also stated that the superior surface of the skull
is swollen above the base of the great vomerine tooth, while
no such enlargement marks the position of its young com-
panion. This is readily explained. The downward crushing
during fossilization has been resisted by the base of the great
fang, while on the other side there has been nothing to resist
crushing.
The vomerine fang referred to (Fig. 1, vom. t.) has an
antero-posterior diameter of 17 mm. and a transverse diam-
eter of 8 mm. The crown is mostly missing. It has been
directed strongly forward. In the alveolus of the other side
is seen the tip of the fang which was to have come into func-
tion on the shedding of the large one now present.
In 1890, Felix, as cited, identified correctly, as it appears,
and figured a beak as that of P. penetrans.
Various other specimens which were collected for Cope in
1877, by Sternberg and Hill, are regarded as belonging to the
same species as P. penetrans and serve to throw light on its
relations to P. nitida. One of these, No. 1871, has been less
affected by pressure than any others of the collection. The
cross-sections of the rostrum are oval, with the transverse
axis the longer, until near the insertions of the vomerine
fangs, where the two axes are about equal. The ornamenta-
tion of the lower side is coarser than that of the upper, but
the pattern is the same. In this beak the left vomerine fang
is functional, while the right alveolus is a cavity 9 mm. deep.
The distance from the fangs to the tip of the rostrum is 125
mm.; the transverse diameter, 27 mm.; the vertical, 26 mm.
Halfway from the fangs to the tip the transverse diameter is
18 mm.,the vertical; 15 mm:
No. 2121 of this Museum furnishes a complete beak, with
the base of the right tooth; the anterior end of the splenial,
freed from the dentary; the tip of the left dentary with three
large teeth; some other fragments of the jaws and skull; and
three sections of the pectoral fin blade, measuring all together
250 mm. The beak is rather slenderer than the type of P.
penetrans and is smoother near the base. Sections of the
1903. | Hay, North American Cretaceous Fishes. 7
beak are broader than high; but the posterior portion has
suffered some distortion. The right vomerine tooth has been
functional, but there is hardly a vestige of even the alveolus
of the other fang.
Reference must be made here to a species of this genus
which has been described by Dr. Loomis (op. cit., p. 225, pl.
xx, figs. 1-4) under the name P. obliquidens. The beak of
this is described as being compressed. The author has fig-
ured a portion of the pectoral fin; and this appears to agree
in every respect with that of our No. 2121. There is the
same front edge, without serrations or undulations; and a
band along this edge is represented as being ornamented with
fine enamel ridges which run at right angles with the edge.
It appears to the present writer that these pectoral fins must
be identified as belonging to the same species. Too much
importance must not be attributed to the compressed or de-
pressed form of the beak. Many of them have been modified
by pressure, and there was probably a good deal of individual
variation.
The premaxilla of No. 2121 is missing. The anterior end of
the dentary and that of the splenial are well preserved and
are figured (Fig. 3). These
are attached to the frag-
ment of the fin. {Between
these bones and the cor-
responding ones of P.
obliquidens I find no dif-
ferences that appear to be
important. Dr. Loomis
states that his species
has three rows of small
teeth on the splenial; but
in some cases there may iY.
be one row on a portion ae pa ects ea rer
5 sfl., splenial.
of the splenial and more
than one row in another part. The number of rows of these
small teeth is probably not a constant character. The small
teeth of the dentary are directed forward, as they are in P.
8 Bulletin American Museum of Natural History, |Vol. XIX,
obliquidens. The possession of five fangs in the premaxilla,
as in P. obliquidens, is unusual in Protosphyrena, but when
we consider that the number of great fang-like teeth in the
jaws of Portheus varies, we shall not give too great value to
this character. In short, it appears to the writer that P.
obliquidens is identical with P. penetrans. Furthermore, it
seems impossible to distinguish these two nominal species
from Cope’s P. nitida.
No. 1634 is a part of the Cope Collection, and consists of a
considerable part of a crushed skull, with the basal half of
the rostrum. The axis of the skull is presented from the
vomerine fangs to the basioccipital articulation. A section
of the rostrum 52 mm. in front of the vomerine teeth is oval,
with the long axis transverse. At the vomerine teeth the
section is quadrate, but this is evidently due to lateral pres-
sure. The ornamentation is quite like that of the type of
P. penetrans, the base of the beak not being so smooth as in
No. 2121. Seen from below, it much resembles that of P.
obliquidens. There is present a part of one of the pectoral
fins, presenting about 95 mm. of the edge, and this is identical
with that of No. 2121, and, so far as we may judge from the
figure, with that of P. obliquidens. But the latter has a com-
pressed section; No. 2121, a depressed section. It appears to
the writer that the evidence furnished by the fins outweighs
that to be derived from the cross-sections of the rostra.
Pelecopterus chirurgus was based on a part of a pectoral fin,
with the characteristic front edge broken away. Judging
from the part remaining, which comes up close to the edge,
and from the angle made by the rays with the edge, there
seems little doubt that this fin is identical with P. obliquidens,
and, therefore, with P. nitida. The specimen bears the Ameri-
can Museum of Natural History’s number 1894.
Under the name of Pelecopterus chirurgus Cope has de-
scribed (Vert. Cret. Form. West, p. 2448, pl. liv, fig. 9) the
articular portion of the shoulder girdle of another specimen.
There is no evidence that the latter belongs to the same
species as the type of P. chirurgus; and the specimen, which
is now in the American Museum, No. 1609, cannot be distin-
1903. | Hay, North American Cretaceous Fishes. 9
guished from the corresponding part of P. perniciosa. In-
deed, this part is probably much the same in all the species.
Impelled by the evidences furnished by the materials before
me, I am compelled to regard Cope’s Erisichthe nitida, E.
penetrans, and Pelecopterus chirurgus, and Loomis’s P. obli-
quidens as belonging to a single species, to which the name
Protosphyrena nitida must be applied.
Protosphyrena perniciosa (Cope).
Pra eG ser
Ichthyodectes perniciosus Cope (E. D.), Bull. U. S. Geol. and Geog,
Surv. Terrs: I, No, 2, 1374, p: 41; Vert. Cret. Form. West, 1875,
Pp. 275.
Pelecopterus perniciosus CopE (E. D.), Vert. Cret. Form. West, 1875,
PP. 244D, 273, pl. xlviii, fig. 2; pl. lii, fig. 2.
Protosphyrena perniciosa Woopwarb (A. S.), Cat. Foss. Fishes Brit.
Mus. III, 1895, p. 414.— ? Loomis (F. B.), Paleontogr. XLVI,
Ig00, p. 221, text fig. 2.— Hay (O. P.), Bibliog. and Cat. Foss.
Werte Nee Aci OO2 pane 70:
This species was established in 1874, as cited, on a frag-
ment of a fin which was afterwards figured on Plate li of
Cope’s ‘ Vertebrata of the Cretaceous Formations of the West.’
In this work there were figured also two other fragments of
fins which were referred to this species. Of these the one
which furnished Fig. 2 of Plate xlviii quite certainly belongs
to P. perniciosa; the one which is represented by Fig. 13,
Plate xliv, appears to belong with those fins which have been
referred to P. tenuis. It will be observed that the undula-
tions of the edge of this specimen, instead of increasing in
-height from the base toward the tip, seem to be subsiding.
In the Cope Collection of fishes and reptiles are fragments
of several pectoral fins of P. perniciosa; but one specimen is
especially worthy of description and illustration. This bears
the Museum’s number 1go1. The record accompanying the
specimen shows that it was collected by Mr. R. Hill, in 1877,
in the Niobrara beds along the South Fork of Solomon River,
Kansas. The shoulder girdle accompanies the fin.
The present length of the fin blade (Pl. I, Fig. 1) is 838 mm.,
but it has doubtless been originally somewhat longer. It is
10 Bulletin American Museum of Natural History. |Vol. XIX,
curved saber-like, and the greater part of the front edge is
provided with coarse serrations. From the posterior border
some rays are probably missing, and a few of those present
have been floated away somewhat from the body of the fin.
A count of the rays at the base of the fin shows the presence
of 45. Even the most posterior of these may be traced to
near the anterior border of the fin at the distal end. As in
the specimen of P. tenuis, described in this paper, there must
have been a posterior fringe of soft flexible rays.
The breadth of the base of the fin is about 105 mm. Here
the rays midway between the two borders are very slender,
the exposed edges of four of them occupying only 5 mm.;
but they grow broader toward their distal ends, so that the
four referred to occupy a breadth of 22 mm. Near the distal
ends of those rays which outcrop in the anterior border of the
fin, at the middle of its length, we find signs of a separation of
each into two portions, as in P. tenuis. In the distal end of
the fin the two components are as distinctly separated from
each other as they are from the components of contiguous
fins. The thickness of the fin at the middle is 9 mm. Asin
the case of the fin rays of fishes in general, each ray is com-
posed of an upper and a lower half. At the base of the fin
these become broader perpendicularly to the surface of the
fin, so as to form plates. Finally these diverge, so as to re-
ceive between them the baseosts. Near the anterior border
of the base there is a large acetabular cavity for the reception
of the rounded head of the scapula.
The tooth-like projections on the front of the fin vary in
distance apart from 10 to 15 millimeters. Those of the most
distal half of the fin protrude beyond their bases as much as
5 mm. and are retrorse; those of the proximal half are shorter
and are dentate in form. In the proximal half of the an-
terior border there is a tooth at the end of each fin ray; in
the distal half, a tooth for each of the two subdivisions of the
ray. From the tip of each tooth a rounded ridge runs back-
ward on the surface of the fin at right angles with the course
of the rays. This appears to be for the purpose of strengthen-
ing the tooth. Each ridge soon divides into two diverging
1903. | Hay, North American Cretaceous Fishes. II
smaller ridges and at length disappears. Those of the distal
portion of the fin may be traced nearly across the fin.
A considerable part of the shoulder girdle of the fin above
described is present (Figs. 4, 5), and it enables me to make
some corrections in Cope’s account of this part of the anatomy
(Vert. Cret. Form. West, p. 244A). This author affirmed
that all the basilar bones, which support the fin, articulated
with the scapula; and on this character he founded the order
Actinochiri.
OS
é . d yy
Fig. 4. Protosphyrena perniciosa (Cope), _ Fig. 5. Protosphyrena perniciosa (Cope).
No. 1901. X 4. cé., cleithrum;corv., coracoid; No. rgor. X 4. cé., cleithrum; cor., cora-
Sc., scapula. coid ; g. cor., precoracoid,
The conclusion reached by Cope is obviously erroneous; for
it is certain that the greater portion of the bone which he
identified as the scapula is the coracoid; while the bone sup-
posed by him to be the coracoid is the precoracoid. The
result of his erroneous determinations was that he had the
shoulder girdle turned wrong end up. All this is satisfac-
torily proved by comparing these bones with the correspond-
ing ones of Tarpon or Salmo. Unfortunately, most of the
sutures in this complex of bones are obsolete, and Cope him-
self was unable to make them out in his specimens. At the
base of the precoracoid of No. rgor there is a dislocation of
the latter which may indicate the position of the suture. In
the tarpon the enlarged anterior fin ray and two baseosts
articulate with the scapula. In the specimens of Proto-
sphyrena there are no satisfactory indications of the suture
that once existed between the scapula and coracoid.
12 Bulletin American Museum of Natural History. [Vol. X1X,
The scapula (Figs. 4, 5, sc.) is applied to the inner side of
the cleithrum, while the precoracoid, p. cor., appears to be
applied to the inner side of the scapula. There may, how-
ever, be some distortion here. In 7arpon the precoracoid is
applied to the inside of the cleithrum in front of the scapula,
but it also articulates with the latter.
On the united scapula and coracoid are borne the surfaces
for articulation with the fin and its supports. Nearest the
cleithrum there is a rounded head (Fig. 4), which fits into
a cavity at the base of the fin. Beyond this are two surfaces
for the paired baseosts which Cope mentions; and still fur-
ther out is a row of six grooves for the reception of six un-
paired baseosts. Between this shoulder girdle and that of
Tarpon I see no important differences, except in the presence
of the paired baseosts. Of these the outer one corresponds,
no doubt, to the articulatory surface for the first baseost
of Tarpon. For the other, situated below the rounded head
on the scapula and mesiad of the surfaces for the other
baseosts, I find no equivalent in Tarpon. Possibly we must
credit to Protosphyrena the possession of eight baseosts.
Of these the first has possibly been crowded out of its place
to a position below the rounded head on the scapula,
while the second has been crowded to a position above the
head.
The disposition of the paired baseosts with reference to the
rounded head and the row of unpaired baseosts must limit
greatly the movements of the fin. In fishes there is generally
a free movement of the fin at right angles to its plane; but it
seems that in Protosphyrena there could have been only very
restricted motion perpendicular to the plane of the fin. The
action of the paired baseosts would have had the effect of
steadying the motion in the plane. Such motion would have
had as its end the employment of the fin as a weapon, with
which its possessor could slash an enemy or a victim of its
appetite. The position of the paired and the unpaired base-
osts may be determined from Cope’s figures (Vert. Cret. Form.
West, pl. liv, fig. 9), from the figures already cited of the
present paper, and from Fig. 6. In the latter figure the
1903. | Hay, North American Cretaceous Fishes. 13
front border of the fin is above, and the upper (bas.) rests on
one of the paired baseosts.
No. 2009 of this Museum furnishes both cleithra of
Protosphyrena perniciosa, which may be conveniently de-
scribed here. One of these is represented by Fig. 7. It is
possible that a small part of the bone is missing from the
upper end of the element. Compared with a specimen of
Tarpon atlanticus, 5 feet 6 inches long (1.67 m.), the cleithrum
is narrower. The portion below the curve is of the same
length, but the upper end of that of Protosphyrena is about
50 mm. shorter. How much of this deficiency is due to in-
jury cannot be determined. The outside of the cleithrum
presents no features worthy of mention; the surface probably
Fig. 6. Protosphyrena sp.
No. 1646. X43. 4das., baseosts;
cor., coracoid ; Act., pectoral fin ; No.7. Protosphyrena perniciosa (Cope). No.2009. X 3.
sc., scapula. Cleithrum seen from inner surface.
has been convex in cross-section. On the inside we see, near
the upper border of the lower, or anterior, end, a deep groove
which is bounded both above and below by a sharp ridge.
On the hinder border of the upper, or posterior, portion there
is another groove. This appears to have started at the upper
end of the bone and to have run downward as far as the
articulation of the fin. It is possible that, as in Tarpon, some
accessory bone fitted in this groove.
It is remarkable that, although this is perhaps the com-
monest species of Protosphyrena, except perhaps P. nitida,
it is represented by remains of fins and shoulder girdle bones
alone. None of these have been found in association with
remains of the head. It appears to be possible that P. tenuis
Loomis and P. perniciosa are identical. The fins appear to
differ only in the character of the anterior edge, the angle
14 Bulletin American Museum of Natural History. |Vol. XIX,
which the rays make with the anterior border being the same
in both species. Under P. tenuis mention is made of fins
whose edges are not repand, but have the summits of the
waves sharp. No. 2044 is a specimen which is evidently to
be referred to P. perniciosa, since it has the serrations well
developed on many parts of the edge. And yet, well out
toward the distal end these serrations subside and the edge
is nearly straight. It seems possible that there was con-
siderable variation in the degree of development of the
serrations in different individuals. Only more and better
materials will enable us to settle this point.
Protosphyrena tenuis Looms.
PLATE I, FIGS. 2 AND 3.
Pelecopterus perniciosus Cope (E. D.), Vert. Cret. Form. West, 1875,
pl. xliv, fig. 13 (erroneous identification).
Protosphyrena tenuis Loomis (F. B.), Paleontogr. XLVI, 1900, p. 226,
pl. xx, figs. 5-7.—Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A.
Igo2, p. 379.
Protosphyrena penetrans STEWART (A.), Univ. Geol. Surv. Kansas, VI,
1900, pp. 370, 402, pl. lxiu, figs. 1-3.
This species is said to be based on three individuals which
were collected by Mr. C. H. Sternberg in the Niobrara de-
posits of Kansas and are now in the Museum at Munich,
Germany. The parts figured belong to one individual and
consist of a rostrum, premaxilla and maxilla of the left side,
and a part of one pectoral fin. This appears to be a well
founded species. The rostrum is, relatively to the length of
the maxilla, very long and slender. The maxilla enters into
the length of the rostrum two times, while in a specimen of
P. nitida the maxilla is contained in the rostrum only one
and a half times. The teeth of the maxilla appear to be
smaller than they are in P. nitida. The most certain evi-
dence that this species does not belong to P. nitida is to be
found in the pectoral fin blade. On comparing it with the
figure of Dr. Loomis’s P. obliquidens (= P. nitida), figured on
the same plate, we find that in the latter the edge of the fin
is devoid of any undulations and that the rays make an angle
of nearly 10° with the edge; while in P. tenuis the edge is
1903. | Hay, North American Cretaceous Fishes. 15
wavy, especially near the base, and the rays make an angle of
15° with the edge. I regard these differences as important.
In the American Museum there are several specimens of
fins which I refer to P. tenuis. One of these, No. 205 (Pl. I,
Fig. 2), is 528 mm. long and is accompanied by a portion of
the baseosts and a part of the cleithrum. It presents appar-
ently 36 rays, including the shortest one at the base in front.
As a result probably of maceration and some disturbance be-
fore burial, some of the hindermost rays are separated from
one another, except immediately at the base. It is probable
that others of the hindermost have been wholly removed.
At its base, as now found, the fin is 62 mm. wide. The front
edge is gently repand in the basal half; but in the distal half
the edge has a uniform curve. The edge is everywhere thin
and sharp, and is strengthened by a iayer of enamel, as in the
other species. This layer is disposed more or less in ridges
at right angles to the edge; but these do not have the regu-
larity and fineness which they present in P. mitida. As in
other species of the genus, the anterior edge of the fin is
formed by the ends of the rays which successively outcrop at
their distal ends. All the rays, except a few of the first,
become broader as they proceed outward. The greatest in-
crease in width is found in the most posterior rays. At the
base they are only about 1.5 mm. in diameter, but distally
they may be as much as 5 mm.in diameter. At about the 20th
ray we find at its distal end evidences of a division into an
anterior and a posterior portion. This separation becomes
still more distinct in the succeeding rays. In another speci-
men, No. 215, traces of the cleft condition may be found as
far forward as the 12th ray from the front, and is indicated
by a narrow furrow, or line of pits.
In the front of the fin in No. 215 are two holes which are
made entirely through the rays, and these, with some fractures,
must have been produced before the specimen was covered
with the matrix. It is easy to imagine that this fish had
been seized and destroyed by some Portheus or some mosasaur.
The fins of this species resemble those of P. perniciosa in
the angle which the rays make with the edge of the fin. As
16 Bulletin American Museum of Natural History. [Vol. XIX,
in the latter species, the base of the fin is undulated for a few
centimeters, but beyond this the character of the edge in the
two species is very different. In P. tenwis the undulations
subside and the edge is continuous; in P. perniciosa, the
elevations increase in height and soon take the form of hooked
teeEh:
No. 1620 of this Museum, a part of the Cope Collection,
probably belongs to this species. It appears to differ only
in having the edge resemble a series of waves whose summits
are not rounded but sharp (like Figs. 1, 2, Pl. lxiii, of Stewart’s
memoir) and in having them continued well out toward the
distal end of the fin. Such fins possibly belong to a distinct
species. This specimen displays the distal end of the fin
apparently to within a few centimeters of the tip (Pl. I,
Fig. 3). Behind the rays which are consolidated to-
gether are several others which evidently have been only
loosely connected and which have been subdivided into very
fine filaments. Evidently, too, the hindermost of these loose
rays did not reach quite to the tip of the fin. It is quite
probable, therefore, that a considerable number of soft flexi-
ble rays occupied the hinder border of these remarkable fins.
No cross-segmentation of these rays can be observed. As
stated under P. perniciosa, there is some reason to doubt
that P. tenuis is distinct from the species just mentioned.
Protosphyrena dimidiata (Cope).
Erisichthe nitida Cope (E. D.), Bull. U.S. Geol. and Geog. Surv. Terrs.
ITI, 1877, p. 822.
Protosphyrena nitida Loomis (F. B.), Paleontogr. XLVI, 1900, p.
227 Gn part), pl. xix, fies: 6,4.
Erisichthe dimidiata Corr (E. D.), Proc. Amer. Assoc. Adv. Sci. XXVI,
1878, p. 300.
Protosphyrena dimidiata Hay (O. P.), Bibliog. and Cat. Foss. Vert.
N; A.1902; ps 370:
This species, the type of which is now in the American
Museum of Natural History, having the number 1635, was
originally described as belonging to Erisichthe nitida. How-
ever, within the next year, Prof. Cope came to the conclusion
1903. | flay, North American Cretaceous Fishes. 17
that it represented a distinct species and to this he gave the
name Erisichthe dimidiata (Proc. Amer. Assoc. Adv. Sci., X XVI,
1878, p. 300). Of this change of opinion and this new
name neither Felix, writing in 1890, nor Loomis, writing in
1900, was apprised.
The most obvious characters of the species are those
presented by Cope in his original description, the upward
curvature of the rostrum and the flattening of the upper
surface of its distal half. Felix, having before him a speci-
men which he regarded as belonging to the same species,
denies the presence of the angular ridges on the sides of the
rostrum. They are present, nevertheless. The rostrum (Fig.
8) has been broken across just in front of the vomerine fangs
and a portion of
the upper surface
nas serumbled
away, perhaps for
a distance of 25
mm.; but there is
little or nothing
missing on _ the
lower side. About _ Fig.8. Protosphyrena dimidiata (Cope). No. 1635. X }.
Oe erin) Gran hraiy edna stacoccsccans, cok) ctkmeidl, pemiza vomeuad
: teeth.
of this break there
has been another break and a slight faulting in the bone.
From this point there is a gradual upward curve to the
tip of the beak. Just in front of the last-mentioned break
the vertical diameter is 23 mm., the transverse, 26 mm.,
the section being oval. Halfway to the tip, the upper sur-
face has become decidedly flat, the vertical diameter being
16 mm., the transverse, 22 mm. Beyond this section the
upper surface is somewhat concave, with a sharp ridge bound-
ing the concavity on each side. Near the tip the upper sur-
face again becomes convex. Both the upward curvature of
the beak and the concavity of the upper surface appear to
be wholly natural.
There appears to be less difference in the sculpture of the
upper and lower sides than in the case of P. mitida.
[ fanuary, 1903.] 2
18 Bulletin American Museum of Natural History. (Vol. X1X,
The premaxilla (Fig. 9, pmx.) possessed four fangs. Its
length has been about 75 mm.; the elevation of the hinder
end about 43
mm. The max-
illa (Fig. 9, mx.)
is 137 mm. long,
with a depth of
only, 17a sami
Fig. 9. Protosphyrena dimidiata (Cope). No. 1635. X 3.
Type of Erisichthe dimidiata Cope. Upper jaw, side view. where mnarrow-
mx., maxilla; faZ., portion of palatine ; Awzx., premaxilla.
est, then again
expanding to near 30 mm. Another bone, probably a jugal,
has overlapped the hinder end of the maxilla on the
upper side. There is space for about 33 teeth on the maxilla.
They are lancet-shaped and are not so strongly inclined for-
ward as they are in P. nitida. The same is true of the teeth
offthe mandible. Outside of the row of cutting teeth is a
row of much smaller teeth.
The lower jaws are represented by 68 mm. of the anterior
end of the left mandible and by nearly the whole length of the
right. The tip of the latter and a section of about 30 mm.
are wanting, as well as most of the lower border of the bone.
In the anterior end of the dentary we find the usual three
lancet-shaped fangs. These are succeeded by an interval in
which the teeth are very small or wanting. Then follows a
single row of cutting teeth, of which those in front and those
behind are small. The presplenial is occupied, as in the
other species, by two strong fangs. Laterad of these and
running nearly the full length of the presplenial is a band of
small teeth. This consists of three rows where narrowest.
A short distance behind the posterior presplenial fang the
band of small teeth is interrupted by what may be a suture
between the presplenial and the postsplenial. It is not un-
likely, however, that it is only a fracture, since the whole jaw
has suffered flexure at this point. In specimens of P. nitida
no suture can be seen; although at this point the band of
teeth becomes very narrow and thereafter widens rapidly.
Felix was unable to find a suture between the supposed two
bones. If they have ever been distinct at any time of the
1903. | Hay, North American Cretaceous Fishes. 19
animal’s life, they are now probably consolidated. The “‘pre-
dentary” of Felix was evidently the result of erroneous
observation.
Lying against the inner surface of the posterior end of the
premaxilla (Fig. 10) is a bone whose edge bears a band of
three rows of teeth which resemble those of the presplenial,
some being two-edged, but most of them conical. This bone,
now 30 mm. long, but with its anterior yy
and posterior ends missing, I take to be 4]
the palatine. Lying against the median
surface of the anterior end of this palatine
and extending forward nearly to the an-
terior end of the premaxilla is another
bone which bears a large patch of granular
teeth. Itis possibly a part of the palatine.
The writer sees little reason to doubt fig 1c. Protosphyrena
that the specimens referred to P. nitida by Uiate (Rope). No.
1635. X 3 Type of Zrzs-
Drasloomis, and) feured on Plate xix oe ore
Part of upper jaw from be-
Figs. 6, 7, of his paper here frequently Cent 3e7 reread oF
quoted, really belong to P. dimidiata, palatine: Ame. premax-
There is the same narrow maxilla; but,
especially, the dentary teeth are only slightly inclined for-
ward.
Among the remains belonging to the type of P. dimidiata
there is present a single vertebra, apparently one belonging
close to the head. It
is possible that this is
an intrusion, either at
the time of burial or
ete etabiocase Se (lao Me alter collection, but the
1
1635, Be imidion of Erhitithe dimidina matrix is the same and
Sas = a Sere Cap alaci tthe bone is: atemilanly
fossilized. The verte-
bra is represented in Figs. 11 and 12. It is 12 mm. long and
Ig mm. in the transverse and vertical diameters. The ends
(Fig. 11) are deeply cupped. The arches have dropped
away, not having been coéssified. The lower surface (Fig.
12) is ornamented with a network of low ridges. The verte-
20 Bulletin American Museum of Natural History. [Vol. X1X,
bra agrees in size with the basioccipital of some skulls of
Protosphyrena. It is possible that it belongs to either
Pachyrhizodus or Anogmius, but it appears to be different.
Cope believed (Proc. Amer. Assoc. Adv. Sci., XXVI, 1878,
p- 299) that this genus possessed well developed vertebre, but
whether or not he based his opinion on this particular verte-
bra is not known. It is very desirable that collectors shall
give attention to the finding of such parts of the body of the
fishes of this genus as have not yet been described; and such
parts include practically all parts behind the shoulder girdle
and pectoral fins.
Protosphyrzena sequax, sp. nov.
Protosphyrena nitida FEwtx (J.), Zeitschr. deutsch. geol. Gesellsch.
XLII, 1890, p. 278, pl.xii, figs: 1—9;, pli xine figs:-1—25; plo aaa,
figs. 2-7.—Loomis (F. B.), Paleontogr. XLVI, 1900, p. 227 (in
part).—Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A. 1902,
Pp. 379 (in part).
Dr. Loomis has already concluded that, on account of the
position of the teeth of the dentary bone, Felix’s specimen
does not belong to the same species as the dentary figured by
Cope as part of P. nitida; but Loomis’s explanation is that
Cope’s dentary does not belong with the premaxillary, and
is, therefore, not a part of P. nitida, an idea already sug-
gested by Felix (op. cit.). As elsewhere remarked, there is
no reason for questioning the unity of Cope’s type. The
more reasonable explanation is that Felix’s specimen does
not belong to Cope’s species, P. nitida. And one of the
grounds for this conclusion is found in the fact that the
teeth of the maxille and those of the dentary, except
the three anterior fangs, are, in Felix’s specimen, inserted
at nearly right angles to the containing bones. Even those
of the premaxillea emerge nearly perpendicular to the border
of the latter bone.
Nor can Felix’s specimen belong to P. dimidiata, The
posterior teeth of the maxilla of the type of the latter have a
decided slant forward. The median and anterior teeth are
nearly perpendicular. All the teeth of the premaxilla appear
1903. | Hay, North American Cretaceous Fishes. 24
to have inclined forward. The cutting teeth of the middle
of the dentary are only slightly inclined forward, not greatly
different from the corresponding ones of the skull described
by Felix. The maxilla of the latter is quite different from
that of P. dimidiata and that of P. nitida, as represented by
the specimens described under that species. In the type of
P. dimidiata the maxilla has a length of 137 mm., a width
where widest of 17 mm. and where broadest, near the hinder
extremity, of 30 mm. The maxilla of Felix’s specimen is
probably little, if any, longer. The figure gives evidence
that little of that of the left side is missing. Its width, where
narrowest, is 20 mm.; where widest, at least 32 mm. Of
the right maxilla of Felix’s specimen perhaps nothing is want-
ing and it measures only 130 mm. This indicates that the
bone was of considerably heavier construction than in P.
dimidiata. If it be contended that the maxilla of Felix’s in-
dividual belongs to a larger animal and was both longer and
broader, it may be shown that it must have contained a con-
siderably larger number of teeth. On measuring backward
from a point 25 mm. behind the anterior end of the maxilla
of P. dimidiata, there are found 9g teeth or alveoli for them, in
32 mm. In the same distance on the left maxilla of Felix’s
specimen are g or 10 teeth, or spaces for them. This indi-
cates either that the maxilla was no longer or that the teeth
were relatively smaller. Indeed, in the portion of the left
maxilla represented by Felix, 103 mm. long, there is room
for as many of its teeth as are found in the 122 mm. of tooth
line of P. dimidiata. Furthermore, the rostrum of the speci-
men described by Felix is very different from that described
by Cope, as Felix himself has pointed out.
Protosphyrena sequax differs from the specimens which are
here referred to P. nitida in most of the respects in which it
differs from P. dimidiata, viz., in having teeth perpendicular,
or nearly so, to the supporting bones and in having a broad
heavy maxilla.
It is, of course, impossible to say that the skull here de-
scribed does not belong to some species which has already
received a name based on a fin blade. It may, for example,
22 Bulletin American Museum of Natural History, |Vol. X1X,
be the skull of P. perniciosa. In such case P. sequax will
become a synonym.
Protosphyrena ziphioides (Cope).
Erisichthe ziphioides (Cope E. D.), Bull. U.S. Geol. and Geog. Surv.
errs.) LL, 28775\p- 823:
Protosphyrena ztphioides Newton (E. T.), Quart. Jour. Geol. Soc.
XXXIV, 1878, p. 795. — FELIx (J.), Zeitschr. deutsch. geol. Ge-
sellsch. XLII, 1890, p. 297.—- Woopwarp (A. S.), Cat. Foss.
Fishes Brit. Mus. 1895, p.413.— Hay (O. P.), Bibliog. and Cat.
Boss: Vert. Ne A. 1902) pp. 370:
Erisichthe xitphioides Loomis (F. B.), Paleontogr. XLVI, 1900, p. 222.
This species was described by Cope in 1877, as above cited.
The description was quoted by Felix and accompanied with
brief remarks. The type specimen is now in the American
Museum of Natural History, and has the number 2131. Cope
states that the specimen is the muzzle of an old individual
which has lost a good deal of its apex by attrition. It is
probable that he meant that this attrition had been suffered
during the life of the animal. The present writer, however,
finds no evidences of any attrition. It appears improbable
that the beak could have been worn to any considerable
amount without revealing it either in the character of the
surface or in some asymmetry. Where the wear must have
been suffered, the bone is dense and smooth and the form
wholly symmetrical. The specimen appears to have belonged
to a species having a short and blunt snout.
The following measurements are given.:
Length-of the specimen. 70:7.) it. oe eee 102 mm.
Width of the hinder-end. 2: ..<:2f:ks . cise eee eS ean
Transverse diameter at vomerine alveoli.......... 22 mm.
Vertical - " oi ey gee Ben aeraener see 19 mm.
Transverse “ 15 mm. from tip of snouts---.... 2eu0uL
Vertical = ik ma hip Oe ees 14 mm.
On the under side of the snout (Fig. 13, vom. t.) is a pair of
longitudinal depressions. These Cope regarded as alveoli for
vomerine fangs, from which the teeth had fallen and which
1903.| Hay, North American Cretaceous Fishes. - 23
had become filled up with bone. His explanation is probably
the correct one. At the hinder end of the specimen, on the
under side, is a triangular depression. This appears to be
produced by the :
parting of the hinder
ends of the vomers,
so as to .expose the
parasphenoid (Fig.
£3, pars), but the
bones are somewhat
eroded. The apex
of this depression is
considerably farther
behind the alveoli
of the vomerine
fangs than in any
other described B;
species of the genus, 2s
being about og Fig. 13. Protosphyrena ziphiotdes (Cope). No. 2131.
i i f x4. Type of Erisichthe ziphioides Cope, seen from below.
while in the type 2 eth., ethmoid ; Jar., parasphenoid ; vow. ¢., position of vom-
P. penetrans the in- «rine teeth.
: Fig. 14. Same rostrum as Fig. 13, viewed from above, with
tervalis only I5 mM. three cross-sections and a longitudinal section of tip. e¢h.,
Oar Pomin ele ae ethmoid ; /., frontal; ~a., nasals.
the basal half of the specimen and extending forward to a line
crossing just in front of the vomerine alveoli is a longitudinal
excavation. Cope regarded these as probably intended for
the reception of the forward prolongations of the premaxille.
It is more probable that they receive the anterior ends of
those bones which Felix has called ‘‘ethmoidea lateralia.”’
There is some reason to think that a portion of these bones
is yet in these excavations. Possibly if the whole of these
lateral ethmoids were present the lower surface of the beak
- would not be so constricted as it is.
The hinder end of the upper surface is occupied by a tri-
angular smooth depression (Fig. 14, jr.). The writer takes
this to mark the anterior termination of the frontals, united
into one bone. In front of this depression the surface is
rough, as if from erosion. On each side of this rough area
ED
. eX SS
= = ees = = = aS =
SS SS =
Lizz
aE
SZ
= 5s
24 Bulletin American Museum of Natural History. [Vol. X1X,
there is a moderate longitudinal excavation, which appears
to be partially filled by the remains of a distinct bone (Fig.
14, na.). These two bones appear to have overlapped the
anterior ends of the frontals and to have met in the midline.
They are probably the nasals.
The supposed nasals on the upper side and the supposed
“lateral ethmoids”’ on the lower side are separated by right
and left ridges of bone which evidently form a part of the
ethmoid. These are continued forward to the tip of the
beak and form the boundary lines which separate the convex
lower surface of the beak from the flatter upper surface. As
these ridges pass forward they descend on the sides of the beak,
until at its tip they meet at the level of the lower surface. The
convexity of the upper surface increases as we move toward
the tip of the beak and that of the lower surface diminishes,
until at length, 15 mm. behind the tip, the upper surface
becomes more convex than the lower. In Figure 14 are pre-
sented cross-sections of the beak at three points. The outline
figure above the sections represents a longitudinal section at
the end of the beak.
The specimen was found in the Niobrara deposits of Gove
County, Kansas. It seems not unlikely that it represents a
distinct genus.
Protosphyrena gladius (Cope).
Portheus gladius Corr (E. D.), Proc. Acad. Nat. Sci. Phila. 1873, p.
337; Bull. U.S. Geol. and Geog. Surv. Terrs. I, No. 2, 1874, p. 40.
Pelecopterus gladius Cope (E. D.), Vert. Cret. Form. West, 1875, pp.
244E, 273, pl. xliv, fig. 12; pl. li. fig. 3.— Loomis (F. B.), Pale-
ontogr. XLVI, 1900, p. 221, text figure 3, tail.
Protosphyrena gladius Hay (O. P.), Bibliog. Cat. Foss. Vert. N. A.
TGO2,'D. 370:
The type of this species is now in the American Museum of
Natural History, and bears the number 1849. It measures
728 mm.in length, Prof. Cope’s statement making it equal
to 31 inches being slightly erroneous. In his description of
the specimen Cope says that Prof. Mudge, the collector, re-
1903. | Hay, North American Cretaceous Fishes. 25
ported that the original length was 41 inches. This state-
ment is not at allimprobable. The second specimen described
by Cope (Vert. Cret. Form. West, p. 244F, pl. xliv, fig. 12) is
also in the Museum’s collection and is numbered 2064.
The type fin blade has been somewhat distorted by pressure,
so that the surface, instead of being flat, as it undoubtedly
was originally, is, as Cope says, trough-like. That the sur-
face was flat is shown by other specimens in the collection
which were obtained for Cope by Sternberg and Hill. The
enamelled edge is sharp, sharper than the edge of the regula-
tion table knife. The edge is not straight, but has the ap-
pearance of having been nicked here and there, as is shown by
Cope’s figure. This author thought that these irregularities
had been produced by the rough uses which the fish had
made of the fin; but, since the bottoms of the notches are as
sharp as the edge elsewhere, it is not probable that there
have been any injuries done to the edge.
At the base of the fin the anterior rays have been so thor-
oughly consolidated that they cannot be counted; but, after
making proper allowances, there appear to be about 50 rays
entering into the portion of the fin represented by the speci-
men. The rays, beyond the most anterior ones at the base,
grow wider as they are followed toward the distal end; where,
one after the other, they emerge at the anterior border, becom-
ing consolidated with those in contact with them. Those
which reach the distal half of the fin become divided each
into an anterior and a posterior portion, as in the case of P.
pernictiosa.
In the case that the fin was originally about rogo mm. long,
that is about 312 mm. longer than it is now, there must have
been about 13 more spines entering into its construction,
‘that is 63 altogether; for the last 312 mm. of the length is
now occupied by 26 outcropping ends, and these represent
13 tays at the base. If this estimate is correct, the fin must
have been about 200 mm. wide at the base.
For the greater part of its length the fin is 20 mm. thick,
measured at a distance of 50 mm. behind the edge. Near the
base the thickness is still greater. Other specimens in the
26 Bulletin American Museum of Natural History. (Vol. X1X,
collection show that the front border is bevelled off on both
sides, as a board may be bevelled off by a carpenter’s plane.
At the middle of the length of the fin the component rays
make an angle of about 22° with the edge. At the distal end
the angle is somewhat smaller.
This fin must have belonged to a large and powerful fish,
of which no other parts are known.
Dr. Loomis in his paper on Kansas fishes holds that this fin
formed one lobe of the caudal fin of some species of Proto-
Sphyrena; and in his restoration of Protosphyrena he recon-
structs the caudal fin from this specimen. This is, however,
manifestly an error. In the caudal fin of fishes the right and
left halves of the constituent rays diverge slightly at their
proximal ends, so as to receive between them the hypural
bones. They are also each drawn out to a point. In the
pectoral fins the two portions of the ray not only diverge
strongly, but each half is broadened so as to form two pro-
cesses. One of these is directed toward the corresponding
surface of the fin, while the other is brought into close con-
tact with the small bones at the distal ends of the baseosts.
The fin known as Protosphyrena gladius has the same struc-
ture as that of the pectoral fin of ordinary fishes and of other
species of Protosphyrena.
PLETHODIDA.
Anogmius Cope.
This genus was erected by Prof. Cope in 1871 (Proc. Amer.
Philos. Soc., XII, p. 170), the type species being A. contractus,
and the type specimen consisted of a large number of vertebrae
representing a fish believed to be about four feet in length.
The vertebre were in the Agricultural College, at Manhattan,
Kansas, and had been collected by Prof. B. F. Mudge. These
vertebre are further described on page 354 of the volume re-
ferred to. This description is repeated on page 241 of the
same author’s ‘Vertebrata of the Cretaceous Formations of
the West’; but on page 220A, evidently written later, he
records his conclusion that the genus in question was really
1903. | flay, North American Cretaceous Fishes. 27,
identical with Pachyrhizodus. In 1877 (Bull. U. S. Geol. and
Geog. Surv. Terrs., III, p. 584) Prof. Cope again restored his
genus Anogmius to favor, and described the new species A.
aratus, based on a nearly complete individual. This per-
mitted him to define more fully the characters of the genus.
In the same year he described two additional species, A.
javirostris and A. evolutus, collected for him in Kansas, by
Sternberg’s party.
Dr. A. S. Woodward (Cat. Foss. Fishes, IV, 1901, p. 71)
apparently takes the position that Cope, when he described
Anogmius aratus, intended to employ the generic name in a
new sense, and to make A. aratus the type of the new genus.
Prof. Cope’s language may give some justification to this
conclusion; but it is evident that he intended to include the
original species, since he cites the original description. He
had evidently again changed his mind regarding the generic
position of the type vertebre. The vertebre of the species
assigned to Anogmius and those of Pachyrhizodus resemble
one another closely, and Cope’s vacillation is not to be won-
dered at as long as he possessed no other parts for comparison.
But in his second description (Proc. Amer. Philos. Soc., XII,
Pp. 354) he mentions characters which appear to separate the
two genera. One of these is found in the crowded condition
of the vertebre at the base of the caudal fin of Anogmius;
the other, in the failure of the upper and lower arches in this
region to become codssified
with their centra. Figures
15 and 16 of this paper
represent the condition of
this part of the vertebral Ze
columns slachpriuzodus. yi, 23 Anopnius Wigs a6)! Amepwieas
Poctemicw apparently Tessviyocrcae (8 (Pine ae
crowding of the vertebre,
and the arches become codssified with the centra. On
this point see Dr. Loomis’s figure (op. cit., pl. xxvi, fig. 9).
Stewart (Univ. Geol. Surv. Kansas, VI, pl. lxvi) presents
a tail of Anogmius in which the shortening and crowding
are less than in any that I have seen.
28 Bulletin American Museum of Natural History. (Vol. XIX,
Just what Cope’s Anogmius contractus was we shall prob-
ably never know. Prof. E. A. Popenoe, who has charge of
the collection of the Agricultural College at Manhattan, Kan-
sas, informs me that he is unable to find any traces of such
vertebre as Cope described. The type being lost, it seems
proper to accept Cope’s Anogmius on the definition given of
it. When it becomes necessary to divide the genus as thus
defined, it will be time to consider whether or not Anogmius
is indeterminable.
Recently Dr. Loomis (op. cit., p. 254) has made Cope’s
Anogmius (spelled, however, Agnomius) a synonym of Os-
meroides Agassiz. Mr. Woodward properly, as it seems to
me, does not follow this identification. So far as we know, the
species of Anogmius have an elongated dorsal fin, while
Osmeroides (Holcolepis) has a short dorsal. There exist un-
doubtedly many other distinctive characters.
Dr. Loomis (op. cit., pp. 229, 235, 252) has described the
new genera Thryptodus, Pseudothryptodus, and Syntegmodus.
Stewart in his work referred to, p. 391, has expressed the
opinion that the first two genera mentioned are synonymous
with Anogmius. Dr. A. S. Woodward (Cat. Foss. Fishes, IV,
pp. 84, 85) regards all three as closely related to Plethodus.
There are so many structures common to the genera
Plethodus, Anogmius, Thryptodus, Pseudothryptodus, and
Syntegmodus, that it appears evident that they are all closely
related, and some of them are quite certainly identical with
others. All appear to have an upper grinding plate developed
on the parasphenoid and one or more lower plates developed
on the median bones in the floor of the mouth. These plates
are composed apparently of dense bone, and are often, if not
always, furnished with pits, sometimes shallow, sometimes
deep. The maxilla forms a considerable part of the boundary of
the mouth, the bones about the mouth are similarly sculptured,
and, so far as we know, the rear of the skull is similar in all.
Plethodus appears to be characterized by coéssification of
the premaxille and ethmoid and the possession of a lower
dental plate composed of a single bone. The upper and
lower plates are smooth or furnished with shallow pits.
1903. | fay, North American Cretaceous Fishes. 29
Anogmuius, as represented by A. aratus and A. javirostris,
described on succeeding pages, has the premaxille free from
the ethmoid, and possibly from each other, and the dental
plates are deeply pitted. The lower plate is composed of
two and possibly of three median bones.
Syntegmodus is described as having the parasphenoid and
some other bones of the mouth covered with a thick mass of
osteodentine. This is penetrated by large canals which pass
from the surface to the bone on which the supposed osteo-
dentine mass rests. This mass is probably composed of
dense bone. On comparing Dr. Loomis’s Syntegmodus, and
the known remains of A. aratus and A. favirostris and a
specimen resembling Syntegmodus altus, among one another,
it is difficult to see how they may be distinguished generically.
Thryptodus is regarded by Dr. Loomis as having the
premaxille not free, but probably consolidated with the eth-
moid. The parasphenoid has developed a great oval, con-
cave, dental plate, which is furnished with pits; while the
median bones of the floor of the mouth support plates which
work against the parasphenoidal plate. A study of the type
of Cope’s Anogmius aratus, figures of which are presented in
this paper, have convinced me that the relation of the pre-
maxille to the ethmoid has been misunderstood by both Prof.
Cope and Dr. Loomis, and that Thryptodus is identical with
the type of Anogmius aratus. In this form, as illustrated by
the types of A. aratus and Thryptodus, the premaxille appear
to be united, while in A. javirostris they are possibly, but not
certainly, free from each other. If this difference is confirmed
by other specimens it may require that two genera shall be
recognized; but for the present it seems to the author best
to unite all under Anogmius. Pseudothryptodus, with free
premaxille, will be included.
Reference has already been made to the differences sup-
posed to exist between the vertebre of Pachyrhizodus and
Anogmius. However much they may resemble, there is great
difference in the structure of the tails of the two genera.
There is apparently little difference in the form of the tail
fins, both being deeply forked. In Pachyrhizodus, as shown
30 «Bulletin American Museum of Natural History, (Vol. X1X,
by Pl. III, Figs. 1 and 2, the rays constituting each lobe
are comparatively few, and these are large, and cross-seg-
mented. In Anogmius, on the other hand,
the rays (Fig. 17, a fragment of the caudal
fin) appear to have been in greater number
and to have shown no segmentation, unless
this may have appeared toward the distal
ends. It will likewise probably prove true
Fig. 17. Saeaees that in Pachyrhizodus the neural and hemal
Fragment tor eeaudl arches of the caudal region are always con-
solidated with the centra, while in Anogmius
they remain distinct.
Anogmius favirostris (Cope).
Anogmius favirostris Cope (E. D.), Proc. Amer. Philos. Soc. XVII,
1877, p. 178—Woopwarp (A. S.), Cat. Foss. Fishes Brit. Mus.
IV, 1901, p. 73.—Hay (O. P.), Bibliog. Cat. Foss. Vert. N. A.
1902, Pp. 393- ;
Osmerotdes favirostris Loomis (F. B.), Paleontogr. XLVI, 1900, p. 256.
This species was based on the skulls of two individuals, one
of them accompanied by a number of vertebrae. These speci-
mens are now in the American Museum of Natural History,
but they appear to be somewhat less complete than when
they were described by Prof. Cope. “Onelofithem, so penis
which must be regarded as the type, since from it the descrip-
tion was mostly drawn, consists of the rear of the skull and
some fragments of its upper surface, the premaxille and
maxille, the anterior portions of both dentaries, the anterior
half of the left palatine and the whole of the right, and a
considerable part of the parasphenoid. With these skull parts
are 11 vertebre.
The rear of the skull shows that the parietals are broadly
joined. Behind these is a narrow area occupied by the supra-
occipital and the epiotics, but the exact extent of these can-
not be determined. The midline of the rear of the skull is
occupied by a valley, deepest and widest between the parie-
tals, where its width is about ro mm.
Figure 18 presents a view of the front of the head, seen
1903. ] Flay, North American Cretaceous Fishes. 31
from below. The outer surface of the right dentary is ob-
served; also the tooth-bearing and triturating surfaces of the
premaxille, the maxille,
the palatines, the vomer,
and the parasphenoid.
The premaxille are sculp-
tured with grooves su-
periorly and with pits
nearer the tooth line.
Here we find a band of
5 or 6 rows of small teeth.
The outer rows of teeth
become very small and
some of them are found
standing on the narrow
walls ‘surrounding the
pits. A similar band is ae 18. Anogmius favirostris Cope. No, 2111,
z- Type. den., dentary; mx., maxilla; fad.,
palatine; far., parasphenoid; fyza., premaxilla ;
found on seach maxilla, Vom. vomer.
and on each of the den-
taries. What I regard as the parasphenoid Cope has described
asthe vomer. It is, however, too elongated; and besides, it
seems to correspond with what is certainly the parasphenoid
in other specimens of the genus figuredin this paper. More-
over, there is a patch of teeth farther forward which is sup-
posed to belong to the vomer. Cope has spoken of this
parasphenoid as if it were covered with small teeth. I find
traces of teeth around its border and a considerable patch in
front; elsewhere, the bone is occupied by pits and is devoid
of teeth. Where the teeth are very small, they occupy the
summit of the bone surrounding the pits. This bone, as
well as the palatines, is thin, only about 2 mm., but this is
probably due to pressure. The pits have the appearance
of passing deeply down into the mass of the parasphenoid.
The surface of the dentine-like layer of the bone presents
evidences of wear. The lower surface of the palatines re-
sembles the surface of the parasphenoid. A few small
teeth are to be seen around the borders of the bone.
The patches of teeth between the maxilla and palatine
32 Bulletin American Museum of Natural History. [Vol. X1X,
in Fig. 18, probably, but not certainly, belong to the
palatine.
The hinder end of the parasphenoid is missing; but it is
not likely to have been much wider than the part present.
The species, therefore, appears to be characterized by a nar-
row parasphenoid. In one important respect this parasphe-
noid differs from that of A. aratus, figured in this paper. In
the latter species the parasphenoid extends forward nearly as
far as the palatines do. In A. javirostris the parasphenoidal
dental plate, at least, falls far short of the anterior end of the
palatines. I am not able to see that this is due to displace-
tment.
The vertebre (Fig. 19) are little constricted, devoid of
conspicuous lateral grooves, and provided with fine longi-
tudinal Tridees-
There appear to be
no lateral processes
such as are found
in some related
species.
The paratype. of
the. «species, No:
2109, consists of
the anterior half
of the skull, and
is of most interest
seen from below
(Fig. 20, natural
par.
Fig.19. Anogmius favi-
vrostris Cope. No. 2011. Xr. Fig. 20. Anogmius favirostris Cope. No. 2109. Xt.
2 ype: Three caudal ver- Paratype. dex., dentary ; mzx., maxilla; far., parasphe-
tebrze noid; fyzx., premaxille.
size). The specimen has evidently had pressure applied to
the snout, so as to drive it backward against the other bones.
The anterior ends of the dentaries overlap. Just behind and
1903. | fay, North American Cretaceous Fishes. 33
between these overlapping ends are seen the edges of two
toothed bones, onein front of the other. Probably the one in
front is the vomer, the other possibly a part of the glossohyal.
In front of the symphysis of the dentaries (den.) are seen
the premaxille (pmx.) evidently pressed downward and back-
ward. They are ornamented as in the type. As shown in
the figure, the maxilla (mx.) join the premaxille and form
the greater part of the border of the-mouth. The lower
border is pitted, and some or all of these pits enclosed the
bases of teeth. There is also a pitted band on the outer sur-
face of the bone, above the tooth line.
Through a splitting of the specimen a view is afforded of
the inside of the mouth. Fig. 21 is a view of the right-hand
side of the floor of the mouth seen from above; while Fig.
22 shows the right side of the roof of the mouth turned up-
side down. If this piece is supposed to be rolled to the left
it will fall into its place on Fig. 21, the points marked a in the
two figures coinciding. Both
these figures are of the size
of the objects.- In-Fig. 21
the whole of the surface cov-
ered with little rings, which
represent pits, except a nar-
row strip occupying a part
of the right-hand side, ap-
pears to be made up of one
or more dental plates. This
is flat along the middle of
the figure, but shelves off
quite steeply on the right.
Near the anterior end is a_ Fig. 21. Anogmius favirostris Cope. No,
5 é 2109, X 1. Paratype. Floor of mouth, a, point
fold which runs obliquely coinciding with a of Fig. 22; 4. é7., basi-
~_ branchials; dex., dentary ; g?. 4., glossohyal.
outward and backward; and pig. 22. Same as Fig. 21. Roof of mouth, a,
there is a similar fold near prac (pec Sabgpmetel ae agate b
the hinder end of the dental
plate. These folds may be artificial, but they probably
represent sutures separating distinct bones. If so, the most
anterior one is probably the glossohyal (gl. h.); the others
2
[ Fanuary, 1903.] 3
34 Bulletin American Museum of Natural History. [Vol. XIX,
are median bones behind the glossohyal, baso-branchials (b.
br.). Certainly, the antero-posterior extent of this mass is
too great to belong to any one bone in the floor of the mouth.
Along the right-hand border of Fig. 21 are seen the broken
edges of the dentary bone and of at least one, perhaps two
bones, between the latter and the tritoral surfaces described
above. Probably the hyoid and possibly a branchial arch
are thus represented.
Fig. 22 presents, on the left-hand side, the right maxilla; on
the right, the tritoral surface of the parasphenoid. Between
them there is a toothed and pitted bone (pal.) which appears to
have been folded longitudinally through pressure. This bone
is taken to be the palatine. The parasphenoid, as seen, prob-
ably does not represent the whole width of this bone, but
posteriorly it was at least 12 mm. wide. The bones, both upper
and lower, of this triturating apparatus appear to be masses of
dense osseous tissue penetrated by deep pits. On the outer
border of the palatine are numerous sharpteeth. Noteeth are
observable on the border of the maxilla, but doubtless the shal-
low pits there seen lodged small teeth, as in the type specimen.
In both of the specimens described here there is present a
broad thin bone which lies on the snout occupying the region
' between the anterior ends of the frontals and the premaxille.
No median suture can be made out. Laterally the borders
extend outward as far as the outer ends of the premaxille.
This bone appears to be separated from the premaxille, but
the evidence is not satisfactory. The bodies of the premaxille
are perfectly distinct from each other, but the bone referred
to may be their codssified ascending plates. In No. 2111 the
bone extends backward about 15 mm. The large area occu-
pied by this bone and its scale-like appearance indicate that
it is not the ethmoid.
This species differs from A. evolutus in having a much nar-
rower band of teeth on the dentary.
Anogmius aratus (Cope).
Puate II.
Anogmius aratus Cope (E. D.), Bull. U. S. Geol. and Geog. Surv.
Terrs. III, 1877, p. 585.—StEwart (A.), Univ. Geol. Surv. Kan-
1903. | Hay, North American Cretaceous Fishes. a5
sas, VI, 1900, p. 340.—Woopwarp (A. S8.), Cat. Foss. Fishes Brit.
Maus: IV; 1901p. .72:—Hay (©. P.),, Bibliog. and Cat: Foss. Vert.
IN; AL 1902; Pp. 303.
The following description is based on the type of the species.
The matrix containing this fish has apparently formed a large
flattened concretion. Of this there are now in the posses-
sion of the American Museum four pieces. The largest block
presents the head and the anterior half of the fin. Another
has been split from the right side and presents the pectoral
fin and some ribs and the imprint of some of the opercular
bones. <A third block shows the body in the region of the
anal fin. This piece does not join accurately the largest
block, but Cope did not believe that any considerable part was
wanting. The fourth block contains the base of the caudal
fin and about 10 vertebre. Between this section and the
third there is missing an unknown number of vertebre,
probably about 8. From a specimen of A. polymicrodus,
studied at the University of Kansas, Dr. A. S. Woodward
came to the conclusion that there were altogether about 80
vertebre, of which nearly 40 were in front of the pelvic fins.
This indicates that at least 10 vertebre are missing in the
abdominal region of our specimen and several others from
the caudal region. The number of the type of A. aratus is
2403. With regard to the dorsal fin Professor Cope’s state-
ments are not as positive as the specimen seems to justify.
The fin begins just above the hinder border of the operculum.
Its first ray is supported by a strong interneural bone (PI.
II, z.”.). The succeeding interneurals diminish rapidly in
size, so that after the sixth they are slender. Cope says that
the fin is ‘“‘continuous as far as the specimen is preserved in
this region, viz., to the fifteenth vertebra behind the scapula.”
But there are 22 vertebre preserved on the block and behind
these are the imprints of 8 more; and there are impressions
of interneurals as far as over the 28th vertebra. There is
also the base of a dorsal ray over the 21st vertebra, so that
the dorsal fin must have extended to this vertebra, at least.
On the succeeding block there are interneurals and fin rays
(i..,d.7r.). Of the latter several must have had their origin
36 §Bulletin American Museum of Natural History. |Vol. X1X,
considerably farther forward, possibly on the anterior block.
On the hindermost block again there are remains of 4 or 5
Tays. If the latter represent a distinct fin the one in front
must have ended very abruptly. To the writer it appears
quite certain that there was a single dorsal fin and that this
extended from near the head to near the root of the tail.
This must have resembled considerably the fin of Coryphena
hippuris (Jordan and Evermann, Fishes of North and Middle
Amer, pl. exlix, fig. 402).
Unfortunately, throughout most of its length only the bases
of the fin rays remain, so that we have little idea regarding
the height of the fin. On the second block there are re-
mains of about 9 rays and some of these were at least 75 mm.
high.
As stated by Cope some portions of the anal fin are present
(a.r.). It was certainly short, but its exact length cannot
be ascertained. A part of the anterior ray is seen, and about
6 interhemal supports (7. h.) may be counted; but weathering
has removed most of the bones.
There are present the distal ends of about 5 rays of one
ventral fin (v. 7.). Their tips have reached the front of the
anal. They are cross-segmented.
One pectoral fin (pet.) is well preserved. It has lain in
the matrix with the distal end directed across the vertebral
column and with the convex anterior border directed up-
ward. It is represented, for the sake. of convenience, in a
different position in the figure, the figure of it being drawn
from the block split from the one bearing the body of the fish.
The fin is 220 mm. long and seems to have consisted of about
20 rays.
Only the base of the caudal fin is present. The rays are
supported by a fan-shaped hypural bone. In front of this
are seen the crowded terminal vertebre. The fin rays appear
to have been slenderer and more numerous than in Pachy-
rhizodus. No evidences of cross segmentation appear.
The ribs are long and slender, and they have apparently
been connected with the vertebral centra by means of dis-
tinct pieces of bone, parapophyses, as in Tarpon, Portheus,
1903. | flay, North American Cretaceous Fishes. 37
and some other fishes. Intermuscular bones are well de-
veloped, those arising just behind the head being especially
long and slender.
Some scales are present. One is 18 mm. long and 5 mm.
wide. The exposed portion is marked with lines radiating
from the centre of growth.
Cope has described the form of the upper surface of the
skull. It presents three planes, a median and two lateral.
The median is narrow at the supraoccipital, but increases in
width to the snout. The lateral planes are widest behind
and narrow anteriorly. They slope off at an angle of about
30° with the median
plane. The sides of the
head are about vertical.
The bones of the upper
surface of the skull in
front of the orbits have
\!
been damaged. A por- , \ ny \y elf
tion of the frontals is JCA 4 f JA! i\ if
but the imprint of MW wa (ZY 7 2
gone, bu e impri IIXINSA WZ \\ We ft
SSR FE aie
these and some splinters
ofthe bones remain. Their
anterior border cannot
be determined exactly.
The anterior extremity
of the ethmoid is present
and is thick and broad,
as seen at the upper end
oniiig.. 23. Cope’ re=
garded this bone as the
consolidated premaxil-
laries and thought that ~Me7j,47emrius arate Cope. No. s4oa.. x 4
little if any of the ethmoid bry Bei ser em supraoccipital ; s. 07., supra-
was exposed. The im-
print of the premaxille and splinters of these bones still
remain on the upper surface of the snout on a portion of the
matrix which overlies the ethmoid. Again, a smooth articular
surface is found at the proximal end of each of the maxille
38 Bulletin American Museum of Natural History. (Vol. X1X,
*.
(Pl. II, mx.), and these smooth surfaces must have been
applied to the premaxillz, unless the structure of this region
was entirely different from that of A. favirostris. The pre-
maxille have quite certainly been similar to those of Loomis’s
Thryptodus zitteli, and when present have nearly concealed
the ethmoid. Onthe other hand, Loomis is probably mistaken
in thinking that the premaxille of his species have coalesced
with the ethmoid. His specimen and the type of A. aratus
explain each other, and the explanation is confirmed by the
types of A. favirostris. Whether or not the premaxille of
A. aratus were codssified, we cannot be sure, No evidences
of a median suture appear. Nor is it absolutely certain that
the ascending processes of those bones were not united in A.
javirostris.
The nasal cavities are situated above the level of the eyes
and a little in front of them.
The bones of the upper surface of the head (Fig. 23) are in
general like those figured by Loomis in his Thryptodus zztttelt.
The supraoccipital (s. oc.) does not separate the small and
square parietals (pa.). Laterad of the last named bones are
the large squamosals (sq.). The eye has been surrounded
by a complete ring of bones. Over the eye are two sculptured
supraorbitals (Fig. 23, s. or.). Behind the eye is the smooth
postorbital (pt. or.); while in front is the prefrontal (p77.).
This bone is sculptured above, but it sends downward in front
of the orbit a long smooth process. Another elongated bone,
apparently the preorbital, occupies the area between the pre-
frontal and the maxilla. The number of the bones covering
the cheeks cannot be determined. They extend to the pre-
opercular. The preoperculum (p. op.) and operculum (0p.)
are as represented in the plate. The other opercular bones
were doubtless present, but are not preserved. The post-
temporal (pt.) and the supracleithrum (su. cl.) are present.
Of the cleithrum (cl.) only fragments remain on the block.
An imprint of the coracoid is seen on the matrix below the
throat. The articulation of the pectoral fin is high, being
just below the vertebral column. If the precoracoid was
present, it must have been short.
1903. | Hay, North American Cretaceous Fishes. 39
The articulation of the lower jaw is below the hinder half
of the orbit. The articular sends up a strong hook-like pro-
cess behind the quadrate. The maxilla is curved and does
" not extend back as far as to the quadrate. Neither the
maxilla nor the mandible appears to have differed much from
those of Thryptodus zitteli. The mouth has been relatively
small and quite oblique.
The structure of the greater portion of the dental apparatus
was unknown to Cope. He states that teeth are found on
the ethmoid bone, his united premaxille; but the present
writer has been able to find there only a slight roughness.
When the fish died, the mouth was left in a gaping position.
Recently the matrix has been removed from it to a depth of
98 mm. from the edge of the vomer. This reveals the fact
that the mouth is armed with large bony plates which closely
resemble those described by Loomis
as belonging to Thryptodus. From a
gelatine mold, plaster casts have been
made of the upper surface of the
mouth and of the floor. These have
given much assistance to the artist
in making drawings ofthe parts. The
excavation of the cavity of the mouth
did not extend quite to the hinder
end of the plates, but must have ap-
proached them closely. In the front
of the mouth we find a short, broad
vomer (Fig. 24, vom.) which is covered
with villiform teeth. Behind this is
found a dental plate (par.) at least ono Agno eS tig 93 OA,
S2umelonsiand about. 40 mm. wide. Rot eee ee ea
it fassnearty parallel sides, and. the °?7*>Y°™<
lower surface is concave. The concavity is greatest just
behind the middle. No doubt, this plate rests on the par-
asphenoidal bone, as in A. favirostris. On each side of this
parasphenoidal plate and articulating closely with it is another
plate (pal.), long, narrow in front, broadening behind, and
convex in cross-section. These plates represent the palatine
40 Bulletin American Museum of Natural History, [Vol. XIX,
bones. The surfaces present the same structure as we have
found in the case of the parasphenoid, being furnished with
numerous pits. In places, all of these bones present evi-
dences of attrition. Near the hinder end of the palatine is
seen a portion of the ectopterygoid.
The floor of the mouth is occupied by two great convex
plates which are joined by a transverse suture. The anterior
(Fig. 25, gl. h.) has the form of the plate
figured by Loomis (op. cit., pl. xxi, fig. 4a)
as the entoglossal, although it is smaller
than the one figured by this writer. The
length and breadth are each about 34 mm.
It is strongly convex from side to side. The
posterior plate (b. br.) is still more convex
than the anterior, especially behind. The
hinder border has not been exposed, so that
ee. we do not know its form and whether or
Fig. 25. Anogmius NOt it was followed by a third plate. Both
aratus Cope. Same
specimen as Fig. 23. these plates are everywhere pitted, and the
3, Floor of th : 5 :
mouth, & dr. base Posterior one, which has fitted into the con-
wheal 8% 481 Cavity of the parasphenoidal plate, is worn
smooth over a considerable part of its sur-
face. It is quite certain that these two plates have been
developed on the glossohyal and the basi-branchials. Doubt-
less the structure of these plates is the same as that of the
plates described under A. altus. Fig-
ure 26 illustrates a section across the
mouth at about the middle of the
basi-branchial plate (Fig. 25, b. br.)
and some distance in front of the ; ec pt:
hinder end of the parasphenoidal plate fig 26. Anvemtus aratus
(Fig. 24, par.). The line above b. br. Foe ge ie Ugeaea
represents the upper surface of the Sctign sFss the cavity of ats
basi-branchial plate. The upper line adie One ale anal
shows the vaulted roof of the mouth
as formed by the parasphenoidal plate (par.) and the right
and left palatines (pal.).
It is quite certain that the specimen described by Dr.
1903. | Hay, North American Cretaceous Fishes. 4I
Loomis as Thryptodus zitteli belongs to the same genus as
the one here described, but it is also quite as certain that it
represents a different species. It appears to have had a
flatter skull and probably a blunter snout. Furthermore, the
upper and the lower dental plates were all proportionally
shorter and broader than in A. aratus.
Anogmius altus (Loomis).
Syntegmodus altus Loomis (F. B.), Paleontogr. XLVI, 1900, p. 253,
pl. xxii, fig. 9 —Woopwarp (A. S.), Cat. Foss. Fishes Brit. Mus.
IV, rg01, p. 84.—Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A.
1g02, p. 390.
Dr. Loomis’s type of this species included the hinder portion
of the skull and the parasphenoid. The specimen is figured
by him so as to present a lateral view.
No. 2112 of the American Museum of Natural History is
a part of the Cope Collection, and was collected by C. H.
Sternberg in 1877, probably in Gove County, Kansas. It
furnishes about the same parts as does Loomis’s specimen;
but it is crushed obliquely downward. Figure 27 presents a
Fig. 27. Anogmius altus ? (Loomis). Fig. 28. Same as Fig. 27. X 3. Base of
No. 2112. X 4. Upper hinder part of skull. ads., alisphenoid ; ex. oc., exoccipital ;
skull. 4. 0c., basioccipital; ef. 0., epi- op. 0., opisthotic ; 0. s., orbitosphenoid ; par.,
otic; /*., frontal; of.0., opisthotic ; parasphenoid; fvo., prodtic; f. sp., pre-
pa., parietal ; sg., squamosal. sphenoid ; sg., squamosal,
view from above; Figure 28 from below. ‘The specimen can-
not be identified with certainty as A. altus; since, as will
42 Bulletin American Museum of Natural History. [Vol. XIX,
be observed, in the type of the species the anterior edge of
the presphenoid is but little in front of the middle of the para-
sphenoidal triturating surface; while in the specimen here
described the presphenoid comes much farther forward.
These differences may be individual, however.
As will be observed, the supraoccipital lies some 25 mm.
to the right of the basioccipital, having been crushed to the
right and downward. This shows that the skull must have
been elevated at least that many millimeters above the fora-
men magnum, a conclusion which is confirmed by the skull
described by Dr. Loomis. About 20 mm. in front of the
hinder extremity of the supraoccipital begins an abrupt de-
pression. It is about ro mm. wide, and extends well forward
on the frontals. From Fig. 28 the position and boundaries
of the various bones may be seen. In general, these agree
with those shown in the specimen described by Loomis as
Thryptodus zittelr (op. cit., pl. xxi, fig. 1). On the left the
opisthotic, sphenotic, and hinder part of the frontal are in
their natural positions and form the border of the skull. On
the right the squamosal has been flexed downward at right
angles with the surface of the skull. The border of the fron-
tal on the left has probably been bent downward somewhat
more than in life.
Figure 28 presents the skull as seen from below, only the
outlines of the presphenoid (p. sp.) and parasphenoid (par.)
being drawn. This bone will be described below. The ob-
server must imagine himself as turning it to the left until it
stands at right angles with the paper, and has the triturating
surface (par.) facing him.
In this figure the prodtics are drawn as meeting in the mid-
line. This cannot be certainly affirmed; but the appearances
are to that effect. There appears to have been a considerable
excavation of some sort beneath the projecting edge of the
Squamosal and outside of the hyomandibular articulation.
The alisphenoids are large, agreeing with those of Dr. Loomis’s
figure of A. altus. We come now to the bone called by Dr.
Loomis the orbitosphenoid. It appears to agree with the
bone so-called by Parker (Philos. Trans. Roy. Soc., CLXIII,
1903. | Hay, North American Cretaceous Fishes. 43
1874, pl. vii.). There is, however, satisfactory evidence of
a pair of bones, or more probably of an unpaired bone with
right and left wings, which is placed in front of the alisphe-
noids. This bone, marked o. s.in Figure 28, is certainly not
the lower surface of the frontal, and there is a plain suture for
union with the alisphenoid. The lateral edge of this bone is
not so certainly determined, but seems to be where drawn in
the figure. The surface for articulation with the large median
bone (Figs. 28, 30, p. sp.), lies about 10 mm. to the left (right
in the figure) of the midline between the frontals. This dis-
tortion could not occur if the bone (p. sp.) were articulated
to the under surface of the frontals. Hence, I hold that the
bones o. s. are the distinct, or more probably united, orbito-
sphenoids, while the bone (p. sp.) is the presphenoid. The
latter would occupy the position of the presphenoidal carti-
lage shown in Fig. 10 of Pl. v and Fig. 10 of Pl. vii of Parker’s
paper just quoted. Dr. Loomis’s figure of A. altus also shows
an arch of bone running upward and outward from the upper
edge of the bone called by him orbitosphenoid. It is above
these bones that the anterior portion of the brain and the
olfactory nerve must have lain.
The parasphenoid of this specimen must have had a median
process behind, like that represented in Loomis’s drawing
(Histo, Pilly xxi): At the anterior end or the
lower surface of the supposed prodtics, in the
midline, there is an excavation in which this pro-
cess must have been lodged. As already stated,
the parasphenoidal grind-
ing plate is deeply pitted
(Fig. 29). Where a small
fracture occurs it is seen
frig, a0 man: that these pits pass down
wee“! to tae: one OL the ordin=
(Loomis). Same
individual as ; Fig. 30. <Anogmius altus ?
Figs. 27, 28. X 3. any) sort on which the (Loomis). Same individual as
Paras phenoid, no. Figs. 27-29. X 4. Lateral view of
palatal Figkace’ denser mass TEpOses. Ac seats i a and presphencid.
companying this plate is
another bone which is represented by Fig. 31. This was
broken, and the injured end has been ground down to show
44 Bulletin American Museum of Natural History. [Vol. X1X,
the structure. Fig. 32 shows a small portion magnified two
times. The pits and cavities in the bone are shown by the
black. It will be observed that the pits pen-
etrate nearly the whole thickness of the bony
mass. Loomis has described the dense mass
which forms the bulk of these triturating plates
Fig.3r. Ang as “‘dentine’”’ and “osteodentine.” I do not
(Loomis). Sane discover the peculiar structure of dentine in
27-30. X 4, im any of them. The canals which Dr. Loomis
plate. "© calls ‘‘Haversian’’ appear to be the pits which
open on the surface. To me the plates appear
to be merely a kind of dense bone, similar to
that found on the maxilla and premaxilla out-
side of the mouth cavity; and these bones dis-
play not very dissimilar pits. Dr. Woodward
has mentioned the presence of dentinal tubules
(Loomis). Section in Plethodus (Ann. and Mag. Nat. Hist., Ser.
ee Dae 7, Vol. III, p. 355). Unfortunately he has not
furnished figures of these microscopical ele-
ments, or given us their dimensions. What is
the function of these pits, and how they have
been produced, the present writer does not
attempt to explain. It would be interesting
ie tem to know how these bony masses increased in
ineeaie dative ee
Seen from the lower side this bone is orna-
mented with fine ridges, which run, for the most part, longi-
tudinally and anastomose, but which, in some parts, run in
other and often irregular directions. In cross-section these
ridges form narrow perpendicular plates, as appears at
the bottom of Fig. 32. They appear to be similar to the
fine ridges found by Dr. Woodward in similar situations in
Plethodus.
The convex surface of this plate fits well the concave sur-
face of the upper plate. Fig. 33 presents a view of the grind-
ing surface of a lower dental plate collected for the author
in the region of Butte Creek, Kansas, but which is now the
property of the American Museum. It is strongly convex
1903. | Hay, North American Cretaceous Fishes. 45
above, concave below. The ends have evidently been sutur-
ally joined to other bones; so that it has apparently been the
middle one of a series of at least three bones forming a tritu-
rating plate. In this bone, asin No. 2112, the pitted surface
gives evidences of polishing through use. No
evidences of teeth are to be found on the cen-
tral portions of this plate, but around the
borders, especially in front, many small, sand-
like teeth are observed. These are clustered ;
on the ridges of dense bone surrounding the ee Heda
pits, as seen in Fig. 34, which represents an es vel
enlarged view of the surface of Fig. 32 bounded
by the two fractures on the upper left-hand
border. The pits are shown in black; the teeth
bythe small citcles, This bone closely re-
sembles the one figured by Stewart (Univ.
Geol. Surv. Kansas, VI, 1900, pl. lxvii) as a
pharyngeal of Anogmius polymicrodus. eg
Fig. 35 represents an upper view of another Fig. 35. Anog-
lower dental plate which evidently belongs to eae
some species of Anogmius. The upper surface aa ae
is in general convex, but the central and
hinder part is somewhat concave. On each
side of the concavity a ridge runs forward to
about the middle of the length. Here it te et
divides, one branch running outward to the X3- Caudal verte-
border of its side. The other unites with the
corresponding ridge of the other side, and the single ridge
thus formed continues to the anterior end of the triturating
surface. In front of the pitted mass of dense bone there is a
thin expansion of ordinary bone. On the right-hand side of
the figure this does not appear, but this is because it has
been crushed downward and to the left beneath the part
seen. It seems most probable that this bone is the glos-
sohyal. It belongs to the same specimen as the piece of tail
represented by Fig. 16. Fig. 36 gives a view from above of
a caudal vertebra of the same specimen, No. 1116. There
are seen the pits for the reception of the neural arches.
46 Bulletin American Museum of Natural History. [Vol. XIX,
There is also presented on each side an outstanding process
which is found on some of the vertebre of some members of
the genus. They occupy a position about the middle of the
height of the vertebral centrum and near the hinder end.
Anogmius evolutus Cope.
Anogmius evolutus Corer (E. D.), Proc. Amer. Philos. Soc. XVII, 1877,
p. 179. — Stewart (A.), Univ. Geol. Surv. Kansas, VI, 1900, p.
347, pl. Ixv, fig. 7; pl. Ixvii.— Woopwarp (A. S.), Cat. Foss.
Fishes, IV, 1901, p. 72. —Hay (O. P.), Bibliog. and Cat. Foss.
Wetter Ne eae rOO2n D303.
Osmeroides evolutus Loomis (F. B.), Paleontogr. XLVI, 1900, p. 257,
pl. xxv, figs? 556.
Beryx multidentatus Stewart (A.), Kansas Univ. Quart. VII, 1898, p.
196.
The type of this species is here figured (Fig. 37). The
original description given by Professor Cope seems to be
sufficiently clear and accurate. Dr. Loomis has figured a
mandible and premaxilla of what appears to be this species.
His specimen is different from that described by Cope in
having the band of teeth wholly on the inside of the dentary,
instead of having it about equally distributed on the inside
and the outside of the bone. Fig. 37 shows how much of the
Fig. 37. Anogmitus evolutus Cope. No. 2101. X 3. Type. Lower jaw,
Ce ee figure, a portion of outer surface. avrt.,
band of teeth was on the inside of the jaw in the type of the
species; the detached part of the figure represents the teeth
on the outer side of the jaw. It is possible that the differ-
ence in the two specimens is due to the crushing of one or
the other; but of which, it is now impossible to say.
A comparison of the figure of the premaxilla given by Dr.
1903. | Flay, North American Cretaceous Fishes. 47
Loomis shows that this bone must have been quite like that
of A. javirostris, in both form and sculpture.
Cope’s type of A. evolutus was, according to Mr. C. H.
Sternberg’s diary of his expedition of 1877, found near the
line between Lane and Gove counties, Kansas. It is now
No. 21o1 of the American Museum of Natural History.
ELOPIDA.
Spaniodon simus Cope.
PLATE IV, FIGS. r AND 2.
Spaniodon simus Cort (E. D.), Bull. U. S. Geol. and Geog. Surv. Terrs.
IV, 1878, p. 69. — Woopwarp (A.S.), Cat. Foss. Fishes, IV, 1901,
p- 53.— Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A. 1902, p.
391.
Illustrations are herewith presented of the specimens from
which Professor Cope drew his description of this species.
He has mentioned the possession of three specimens, of which
one, said to be nearly entire, served as his type. These speci-
mens are now in this Museum. Fig. 1, Pl. IV, is taken
from No. 2508; Fig. 2 of the same plate, from No. 2509.
On the block containing the latter specimen are remains of
one or more fishes lying behind and above the fish represented
on the plate, but it is doubtful whether or not they belong to
the latter. In any case the vertebre are all wanting. Yet it is
from this fish that most of the description of the head is derived.
Both specimens are‘labelled by Cope as being his types.
Little criticism can be made on Cope’s description. To the
writer it appears evident that the lower portion of the body was
scaled. The dorsal and anal seem to have had each about two
rays fewer than the numbers given in the original description.
The number of vertebre in this fish is less than in any of
the other described species. S. latus (Agassiz) is stated by
Dr. A. S. Woodward (Cat. Foss. Fishes, IV, p. 53) to have
50 vertebra, whereas the present species has only 45.
The depth is contained in the length to the end of the
vertebral column three and one-half times; the length of
the head in the same distance about three and one-third
48 Bulletin American Museum of Natural History, [Vol. XIX,
times. Three other species, S. blondeli Pictet, S. elongatus
Pictet, and S. latus (Agassiz) are found in the upper Creta-
ceous of Sahel Alma, Mt. Lebanon.
On the block bearing No. 2509 are found written in pencil
the words ‘Yankton, Neb.’’ Professor Cope merely stated
that the specimens came from Dakota. The formation and
locality are therefore as follows: Niobrara Cretaceous,’
Yankton, South Dakota.
ICHTHYODECTIDA.
Saurocephalus Harlan.
So far as has yet been shown the only difference between
Saurocephalus Harlan and Saurodon Hays is found in the
presence in the former of a row of foramina, one foramen for
each tooth, placed some distance from the dental border of
the jaws, upper and lower, while in Saurodon there is at
the base of each tooth a deep notch. In the latter genus the
notches are often converted into foramina by the growth of
bone across the notch. There can be little doubt that the
foramina of Saurocephalus lanciformts originated from notches
like those of Sawrodon leanus. Whether or not this difference
shall be regarded as sufficient to indicate distinct genera may
be a matter of individual judgment. The writer has pre-
ferred to retain all the species under Saurocephalus.
These openings in the bones of the jaws have been called
nutritive foramina, and as such Dr. Woodward speaks of
them in the final volume of his work on fossil fishes. There
can be little doubt that it is through these foramina that the
young teeth enter the sockets. The writer has ground down
a small piece of a jaw of this genus and found the very young
tooth at the bottom of one of these foramina, lying against
the functional tooth. As growth occurs, the root of the
tooth pushes itself above the foramen, while the blade grows
toward the dental border. It is very improbable that any
nutrient vessels enter the sockets through these foramina.
From Dr. Loomis’s memoir I gather that the view here pre-
sented is also that of Dr. Rése.
1903. | fay, North American Cretaceous Fishes. - 49
Saurocephalus phlebotomus Cope.
Saurocephalus phlebotomus Corr (E. D.), Proc. Amer. Philos. Soc. XI,
1870, p. 530; U. S. Geol! Surv. Wyomung, etc. 1875, p. 426;
Proc. Amer.- Philos. Soc. XII, 1871, p. 343.— Hay (O. P.),
Bibliog. and Cat. Foss. Vert. N. A. 1902, p. 386.
Daptinus phlebotomus Corr (E. D.), Proc. Acad. Nat. Sci. Phila. 1873,
pescos ullbiUeomGeolsand Geos sun. lly Nome S74.eps cn:
Vert. Cret. Form. West, 1875, pp. 213, 275, pl. xlvii, figs. 3, 4, 6;
pl. xlix, figs. 1-4. —- Newton (E. T.), Quart. Jour. Geol. Soc.
XXIV, 1878, p. 440. — ZitteL (K. A.), Handbuch Paleont. III,
1890, p. 264.
Saurodon phlebotomus Corer (E. D.), Bull. U. 5S. Geol. Surv. III, 1877,
p. 588. — Stewart (A.), Kansas Univ. Quart. VII, A. 1808, p.
186; Univ. Geol. Surv. Kansas, VI, 1900, p. 312, pl. lvu, figs.
4, 5.-— Loomis (F. B.), Paleontogr. XLVI, rg00, p. 248, pl.
xxiv, figs. 1-5. — Woopwarb (A. S.), Cat. Foss. Fishes, IV, rgor,
Dp: 422:
Daptinus phlebotonum Crook (A. J.), Paleontogr. XX XIX, 1892, p.
m2ae
The type of this species is in the American Museum and
has the number 1906. There are present all of the left max-
illa, except the distal end; the alveolar border of the right
maxilla; both palatine malleoli; and considerable parts of
both dentaries, including the symphysis and surface for articu-
lation of the predentary. All the parts are more or less frag-
mentary; but it is believed that they furnish a correct idea
of the tooth lines, both of the maxilla and the dentaries. The
conclusion is reached that there were not more than 4o teeth
in each dentary and about 30 in each
maxilla.
~The specimen which Stewart has de-
scribed and figured as Saurodon phileboto-
mus (Univ. Geol. Surv. of Kansas, VI,
Pp. 312, pl. lvii, figs. 4,5)is quite certainly _ ERE.
such. On the other hand, the specimens Ji etblomuntone Mokena
described and figured by Loomis (Pale- * * Premarin inner view.
ontogr., XLVI, p. 248, pl. xxiv, figs. 1-5) are probably not
of this species. In these there are 47 teeth in the dentary.
Likewise, the premaxilla does not resemble that of one of
[ Fanuary, 1903.) 4
50 Bulletin American Museum of Natural History. [Vol. XIX,
the specimens which Cope has (and, so far as I can see, cor-
rectly) identified as S. phlebotomus. This specimen, No. 1907,
is here figured (Fig. 38) and it will be seen that it is much
more pointed than is the one figured by Dr. Loomis. The
latter is probably S. x:phirostris Stew., the premaxilla of
which is shown here by Fig. 39.
There seems to be only one objection to Stewart’s identifica-
tion of his specimen, and that is found in the relative lengths
of the dentary and maxilla. Cope’s specimens indicate that
the dentary projected farther in front of the premaxillaries
than Stewart’s figures would suggest. The tooth line of the
dentary of Cope’s type must have had a length of 98 mm.;
the maxillary tooth line a length of 60 mm. If now we add
to the latter 20 mm. for the premaxilla, we have 18 mm. for
the distance which the dentary projected beyond the pre-
maxillary. How the discrepancy is to be explained is now
uncertain.
Saurocephalus lanciformis Harlan.
Saurocephalus lanctformis HARLAN (R.), Jour. Acad. Nat. Sci. Phila.
(1), IMI, 1824, p: 337, pl. xii; Trans! Geol: Soc: Pean: I,-uaan-
pt. 1, p. 83; Med. and Phys. Res. 1835, pp. 286, 289, 366. —
? Morton (S. G.), Amer. Jour. Sci. XXVIII, 1835, p. 277. —
OweEN (R.), Odontog. 1845, p. 130, pl. lv.— GiEBEL (C. G),
Fauna Vorwelt, I, pt. iii, 1848, p. 89. — Lripy (J.), Proc. Acad.
Nat. Sci. Phila. 1856, p. 302; Trans. Amer. Philos. Soc. XI, 1857,
p- 87, pl. vi, figs. 8-11. — Pictet (F. J.), Traité Paléont. 1854,
ed. 2, p. 93. — ?? SPILLMAN (W.), Hilgard’s Rep’t on Geol. Miss.
1860, pp. 142, 389. — Core (E. D.), Proc. Amer. Philos. Soc. XI,
1870, p. 530; U.S. Geol. Surv. Wyom. 1871, p. 415; Vert. Cret.
Form. West, 1875, pp. 216, 275. — Daviess (W.), Geol. Mag. (2),
V, 1878, p. 260.— Newton (E. T.), Quart. Jour. Geol. Soc.
XXXIII, 1878, p. 786. — Stewart (A.), Kansas Univ. Quart.
VII, 1898, p. 186. — Hay (O. P.), Amer. Jour. Sci. (4), VII, 1899,
p. 299, figs. 1-4. — Loomis (F. B.), Palzeontogr. XLVI, 1900, p.
251, pl. xxv, figs. 2-5.— Stewart (A.), Univ. Geol. Surv.
Kansas, VI, 1900, p. 392. Woopwarp (A. §S.), Cat. Foss.
Fishes, IV, rgo1, p. 113. — Hay (O. P.), Bibliog. and Cat. Foss,
Vert. N. A. 1902, p. 386.
Saurodon lanciformis Hays (I.), Trans. Amer. Philos. Soc. (2), III.
£S30, Pi 4775 ple XViy ew rT:
1903. | Hay, North American Cretaceous Fishes. 51
Saurocephalus arapahovius Corer (E. D.), Proc. Amer. Philos. Soc. XII,
1872, p. 343; U.S. Geol. Surv. Mont. 1872, pp. 344, 348; Bull.
WU. S: Geol. and Geos. Surv. I; No. 2, 1874, p. 41; Vert. Cret.
Form. West, 1875, pp. 216, 275, pl. xlix, fig. 5. — Woopwarpb
(A. S.), Cat. Foss. Fishes IV, r901, p. 114.—— Hay (O. P.), Bib-
liog. and Cat. Foss. Vert. N. A. 1902, p. 385.
Saurocephalus arapalovius Loomis (F. B.), Paleontogr. XLVI, 1900,
p. 251 (syn. of S. lanciformis).
The type of Cope’s Saurocephalus arapahovius is in the
Cope Collection, now belonging to the American Museum of
Natural History, and has the number 2073. It is a fragment
of the maxilla. Cope distinguished his species from S. lanci-
jormis on the ground that the facets shown by Leidy to exist
on the roots of the teeth of the latter were absent in the for-
mer. Loomis unites the two species because he regarded the
presence of facets to be variable. I have exposed the root of
one of the teeth of the type of S. arapahovius and find that
there are very distinct facets. There appears, therefore, to
be no reason for retaining it as a distinct species.
Saurocephalus xiphirostris (Stewart).
Saurodon xiphirostris STEWART (A.), Kansas Univ. Quart. VII, 1898,
p. 178, pl. xiv; Univ. Geol. Surv. Kansas, VI, rgo0, p. 314, pl. lv.
— Loomis (F. B.), Palzontogr. XLVI, 1900, p. 247. — Woop-
WARD (A. S.), Cat. Foss. Fishes, IV, 1901, p. 113.
Saurocephalus xtphirostris Hay (O. P.), Bibliog. and Cat. Foss. Vert.
N. A. 1902, p. 386.
Saurodon phlebotomus Loomis (F. B.), Paleontogr. XLVI, 1900, p. 248,
pl. xxiv, figs. 1-5.
I find difficulty in estimating the value of Stewart’s species,
Saurodon broadheadi, S. ferox, and S. xiphirostris. There is
at least one good species among these; if only one, it must
bear the earliest name, broadheadi. S. jferox appears to differ
from S. broadheadi in having the maxilla proportionally a
little higher. It has also an unusually large number of teeth
in the maxilla, 40; an unusually small number, 46, in the
dentary; and only ro in the premaxilla. The number is
variable, however; a specimen, No. 1614, referred to S.
52 Bulletin American Museum of Natural History. [Vol. XIX,
xiphirostris, having 32 in one maxilla, 35 in the other; 50
teeth in the dentary; and 11 in the premaxilla. Another
has 32 and 34 teeth in the maxillz, 50 in the dentary, and 13
in the premaxilla. However, it is probably better for the
present to regard the three species as distinct.
No. 1614 (Figs. 39, 40) was collected in 1877, in Gove
County, Kansas, by Mr. R. Hill, for Professor Cope. It con-
sists of both upper jaws complete; the
left palatine complete, or nearly so; the
greater part of both lower jaws; one quad-
rate; and the predentary. The number
of the teeth has been stated. The pre-
dentary has a length of 55 mm., a vertical
diameter of 27 mm. at the base, and a
Bich ay) pene ire transverse diameter of 16 mm. It does
ee evar ie, Not appear to have suffered any crushing.
Bee: The premaxilla is represented by Figure 39
for comparison with that of S. phlebotomus.
The palatine has the form represented in Fig. 40, seen
from the outside. The mesial sur-
face is concave in cross-section.
On this surface are seen two con-
siderable patches of small teeth,
and probably the whole surface was
originally furnished with teeth.
Attached to the hinder end of the
upper border of the maxilla is a
supramaxilla.
No. 2012 has 35 teeth in the
maxilla. The predentary is 66 mm. long and 28 mm. high.
Fig. 40. Saurocephalus xiphiros-
tris (Stewart). No. 1614. X 4. Pala-
tine, outer view.
Saurocephalus goodeanus (Cope).
Ichthyodectes goodeanus Corr (E. D.), Proc. Amer. Philos. Soc. XVII,
1877, p. 176. — Hay (O. P.), Amer. Jour. Sci. (4), VI, 1898, p.
227; Bibliog. and Cat. Foss. Vert. N. A. 1902, p. 385. — Woop-
WARD, (A. S8.), Cat. Foss. Fishes, IV. 1901, p. 107.
In the Cope Collection of fossil fishes has been found Cope’s
type of his Ichthyodectes goodeanus. It proves to be a species
a1
oo
1903. | Hay, North American Cretaceous Fishes. 53
of Saurocephalus, belonging to the section or subgenus Sauro-
don. Its number is 2110. It is difficult to understand how
Professor Cope came to refer the species to Ichthyodectes,
since the forms of the maxillary, premaxillary, and palatine,
RVC
Fig. 41. Saurocephalus goodeanus (Cope). No. 2110. X 3. Type.
Upper jaw, outer view. mx., maxilla; Aa/., palatine; fyzx., pre-
maxilla; s#zz., supramaxilla,
and the presence of notches at the bases of the teeth plainly
indicate its relationship with Saurocephalus phlebotomus (Fig.
41).
Cope’s description agrees in almost every respect with the
specimen, but in one or two cases there is some apparent lack
of agreement, due evidently to a misuse of terms. He states
that “‘the maxillary border is incurved at its anterior ex-
tremity,’ etc. This is exact, if instead of maxillary we read
premaxillary. That the latter is meant is indicated by the
statements “‘the middle part of the border being most
prominent,” and “‘the anterior border is sigmoidally curved,”
statements true of the premaxilla, but having little or no
significance when applied to the maxilla. The statements
as to the number and character of the premaxillary teeth
are correct. As Cope says, the maxillary teeth are round
in section; but there can be little doubt that if we had the
crowns of these teeth they would be found to be two-
edged, as are those of the premaxilla. Cope’s measurements
are correct.
This species differs from all other described species of
54 Bulletin American Museum of Natural History. [Vol. XIX,
Saurocephalus in its heavy structure. The bones are much
thicker than those of specimens which I refer to S. xiphiros-
tris, as the following measurements indicate.
S. xiph. S. good.
Thickness of premaxilla 20 mm. above alveolar
DOrders. cavern seca 8 mm. 12 mm:
7 ‘* maxilla 10 mm. below condyle.. 8 mm. 12 mm.
os 1omm. above alveolar
border at middle of
fehieth. ee fs. ene 5.5 Man. 725 ain.
The exact length of the maxilla cannot be determined, but
it is quite evident from the way in which the alveolar border
is curved upward posteriorly and the small size of the teeth
that the bone did not extend much farther backward. Alve-
oli for 31 teeth are counted; and there are six alveoliin 20 mm.
The palatine malleolus, measuring from the articular surface
for the maxilla to that for the prefrontal, is high; not low, as
we find it in Ichthyodectes.
The supramaxilla (Fig. 41, smx.) is present. In the draw-
ing this bone is lifted somewhat above its natural position.
It is flat on the outside. On the mesial side it is traversed
longitudinally by a sharp ridge.
This species was collected by C. H. Sternberg’s party during
the year 1877, probably in Gove County, Kansas.
The maxilla figured by Stewart as the type of Saurodon
broadheadi resembles in outline and proportions that of Sauro-
cephalus goodeanus, but Dr. S. W. Williston has kindly given
me measurements which show that Stewart’s species is every-
where much thinner. At 10 mm. below the condyle the
thickness is only 7 mm.; at middle of length, 10 mm.; above
the lower border, only 4.5 mm.
Ichthyodectes anaides Cope.
Ichthyodectes anaides Corr (E. D.), Proc. Amer. Philos. Soc. XII,
1872, p. 339; 5th Ann. Rep. U. S. Geol. Surv. Mont. etc. 1872,
Pp. 343; Bull. U. S. Geol. and Geog. Surv. Terrs. I, No. 2, 1874,
p. 40; Vert. Cret. Form. West, 1875, pp. 206, 274, pl. xliv, figs.
14,15; pl. xlv, figs. 1-8. — Crook (A. R.), Paleontogr. XX XIX,
1892, pp. 111, 123, pl. xv. — Hay (O. P.), Amer. Jour. Sci. (4),
1903. | Hay, North American Cretaceous Fishes. 55
VI, 1898, p. 226, fig. 2; Bibliog. and Cat. Foss. Vert. N. A. 1902,
Pp. 384. — Loomis (F. B.), Palezontogr. XLVI, 1900, p. 244. —
STEwaRT (A.), Univ. Geol. Surv. Kansas, VI, 1900, p. 296, pl.
xlix, figs. 1-3. — Woopwarbp (A. S.), Cat. Foss. Fishes, IV, rgor,
Pp. 100.
Portheus arcuatus Corr (E. D.), Vert. Cret. Form. West, 1875, p. 274,
pl. xlvii, figs. 7-9.
This is one of the commoner species found in the Niobrara
beds of Kansas, and it has been figured by Cope, Crook, Hay,
and Stewart. As shown by some of the published figures and
by specimens in the American Museum, the mouth has been
very oblique. The eyeball is large, as shown by the pre-
served sclerotic bone.
Crook has figured portions of the shoulder girdle and the
first pectoral ray. As in the case of Portheus, he has mis-
taken the upper end of the clavicle for the lower, and conse-
quently the right for the left fin. The first pectoral ray has
the same structure as that of Portheus, but has evidently been
proportionately shorter and more curved distally.
An examination of the original of Cope’s figures 7-9, pl.
xlvii, ‘ Vertebrata of the Cretaceous Formations of the West,’
has convinced me that they belong to this species and not to
Gillicus arcuatus. Crook has already surmised this to be true
(Paleontogr., XX XIX, p. 112). The skull is shorter and
broader and of heavier construction.
After examining skulls of Portheus, Saurocephalus, and
Gillicus I am confirmed in my opinion that the parietals are
united in the mid-line and lie in front of the supraoccipital.
They form the base of the great crest at the back of the
head, and extend slightly backward on each side like the
horns of a crescent. Posteriorly these horns join the an-
terior prolongations of the epiotics. I have not been able in
any case to discover sutures between the parietals and the
epiotics, but doubtless these exist.
No. 2005 of the American Museum was collected in 1877,
in Gove County, Kansas, by Russell Hill. It furnishes a
complete head, 19 anterior vertebre, the shoulder girdle, and
the first rays of both pectoral fins. Unfortunately, the head
560 Bulletin American Museum of Natural History, [Vol. XIX,
is somewhat distorted and crushed, and the greater part of
the shoulder girdle hidden. Above each orbit are two supra-
orbitals, one behind the other and each about 25 mm. wide.
There are remains of a supramaxilla, but its limits are not
definable. A portion of the palatine behind the malleolus
has been exposed in life. Below and behind the eye the
bones of the palatopterygoid arch have been wholly hidden
by the suborbitals. The boundaries between these latter
bones cannot be made out, the bones themselves having
probably been very thin. If correctly identified, the supra-
cleithrum is large, about 160 mm. long and 50 mm. or more
wide. The preopercular resembles that of Portheus. The
opercle is large. All the opercular bones are roughened, as
if there had been here and there bony nodules. The cleith-
rum appears to have a backwardly extending flap behind the
articulation of the jaw.
The head of this specimen has been shortened by distortion,
but must have been, from snout to gill clefts, about 300 mm.
long. The length of the whole fish must have been about
5 feet (1.64 m.).
The vertebre of this species resemble those of Portheus.
In his description of these (Vert. Cret. Form. West, p. 207)
Cope states that the ribs are not articulated directly to the
centra, but by means of free elements which were inserted
into the lateral grooves. Had I been aware of this fact when
writing my observations on the vertebral column of Portheus
(Zool. Bull., II, 1898, pp. 25-54) I might have been saved
from the blunder which I there made, that of calling the
upper side of the vertebral column the lower. The sec-
tions of the column there studied had been crushed so
that the ribs of opposite sides had been brought into close
contact and so as to resemble neural arches. In Tarpon,
with which they were being compared, there are also free
parapophyses, but posteriorly these diminish and disap-
pear. In Portheus, on the contrary, as is now realized,
these parapophyses increase in size toward the tail region,
and the same is probably true in the cases of other members.
of the family.
1903. ] Hay, North American Cretaceous Fishes. 57
Ichthyodectes multidentatus Cope.
Ichthyodectes multidentatus CopE (E. D.), Proc. Amer. Philos. Soc.
XII, 1872, pp. 339, 342; 5th Ann. Rep. U.S. Geol. Surv. Montana,
etc. 1872, p. 343; Bull. U. S. Geol. and Geog. Surv. Terrs. I, No.
2, 1874, p. 41; Vert. Cret. Form. West, 1875, pp. 212, 275, pl. 1,
figs. 6, 7. — Crook (A. R.), Palzontogr. XX XIX, 1892, p. 123.
— Hay (O. P.), Amer. Jour. Sci. (4), VI, 1898, p. 227; Bibliog.
and Cat. Foss. Vert. N. A. 1902, p. 385.— Loomis (F. B.),
Paleontogr. XLVI, 1900, pp. 243, 245, text figs. 8, 9; pl. xxiii,
fig. 9. — Woopwarp (A. 5.), Cat. Foss. Fishes, IV, rgo1, p. 107.
This species was based on a premaxilla and a fragment of
the maxilla, and these parts were figured as cited above.
The type now bears No. 2186 of the American Museum of
Natural History. Afterward a more complete specimen was
secured and described. This is now No. 1743 of this Museum
and is represented in Figure 42. The identification of this
with the type is doubtless correct. Only a part of a single
tooth of the type re-
mains, but this shows
the presence of the
ridges and _ furrows
which are so distinctly
seen in Cope’s second
specimen (Fig. 42, ¢.).
The skull of this
specimen is much
crushed and _ broken,
but nevertheless much
may be learned from
ita wlan eeneraliormat
‘ Fig. 42. Jchthyodectes multidentatus Cope. No. 1743.
resembles that of Gil- Skull x}; tooth x 2. ads., alisphenoid; 4. oc., basioc-
A cipital ; ef. 0., epiotic ; et#., ethmoid ; ex. oc., exoccipital ;
licus arcuatus (Cope), Jr., frontal : #x., maxilla; fa., parietal; fav., parasphe-
noid ; £7/., prefrontal ; fvo., prodtic ; A. sf., presphenoid;
although the bones are ptf., postfrontal; s. oc., supraoccipital ; sg., squamosal ;
¥ E vert., vertebra ; vom., vomer.
HOLINSON “thin As: “1m
that species, there is a strong upward flexure of the axis in
the basisphenoidal region. The snout is pointed, and there
is a high supraoccipital crest. The maxilla has been de-
scribed by Cope. It bears teeth, or spaces for them, to the
58 Bulletin American Museum of Natural History. [Vol. XIX,
number of about 50. Loomis states that there are about 44
teeth. The maxilla which he figures in outline (op. cit., p.
45) is slenderer than the one here figured and of a some-
what different form. Loomis also figures what he regards
as the mandible of this species.
In No. 1743 there is a deep excavation in the sides of the
skull just below the articulation of the hyomandibular. This
is seen also in the skull of Portheus, and is probably found in
the skulls of all members of the family. The supraoccipital
and the epiotic have been broken from the skull and replaced,
as shown. Just above the line of fracture is the very distinct
suture which passes between this bone and the squamosal.
Another suture starts well up on the front of the supraoccip-
ital crest, descends for a short distance, then turns back-
ward. The bone in front of and below it is undoubtedly the
unpaired parietal. Cope mistook the epiotic for the parietal.
The suture in front of and below the parietal is effaced by the
fracture. I have not been able to find a suture separating
the parietal from the epiotic, but such no doubt exists.
In a former paper (Zool. Bull., II, 1898, pp. 25-54) I have
announced such to be the position of the single parietal; but
its bounding sutures are not often easily seen. One speci-
men will reveal one suture, another specimen another suture.
In a specimen of Portheus, No. 2373 of this Museum, both
the upper and the lower sutures are distinct. Such a dis-
position of the parietals doubtless characterizes all members
of the family.
The palatine has been provided with a patch of small teeth.
The malleolus for union with the maxilla is low, broad, and
flat.
The interorbital septum of the species under consideration
appears to be occupied by a presphenoid; and above this
there were probably orbitosphenoids. The vomer is beset by
a patch of teeth.
As has been stated by Cope, the scapula and part of the
cleithrum are present. This author states that the width of
the cleithrum (‘‘clavicle”’) below the scapula is 40mm. This
shows that he mistook the upper for the lower end of the
4
1903. ] Hay, North American Cretaceous Fishes. 59
cleithrum; for the lower end is missing (Fig. 43). On the
inside of the cleithrum there is a precoracoid which ascends
from the coracoid three-fifths the distance to
me upper end of the cleitthrum, This pre-
coracoid, which is like that of Portheus, was
doubtless regarded by Cope as the coracoid.
There are two large convex surfaces for articu-
lation with the fin, the uppermost with the
large first ray, the lowermost with the first
baseost. On a level with the latter, but more
mesiad, are two pits, undoubtedly for the
reception of the next two baseosts. The
rope: interpretation Of these parts ts'made 75 Gea
: : : tus Cope. No. 1743.
easy by comparison with the shoulder girdle 4-7 cleithrum:
art. s., articular sur-
of a tarpon or salmon. face for fin ray.
Gillicus Hay.
The type of this genus is Cope’s Portheus arcuatus, later
called by him IJchthyodectes arcuatus. Dr. A. S. Woodward
(Cat. Foss. Fishes, IV, r90r, p. 101, pl. viii) has recently
described a second species, Gillicus serridens, from the Albian
epoch, Kent, England, under the name /chthyodectes serridens.
It differs in having the anterior mandibular teeth relatively
larger. The members of this genus are well characterized by
the falcate maxille, the reduced dentition, and the thin skull
bones.
PACHYRHIZODONTID.
Pachyrhizodus Agassiz.
Cope originally made this genus the type of the family
Pachyrhizodontide (Proc. Amer. Philos. Soc., XII, 1872, p.
343). Later he placed it in the family Stratodontide (Vert.
Cret. Form. West, 1875, p. 219). Loomis and Stewart regard
the relationships of the genus to be with the Salmonide.
Dr. A. S. Woodward in his latest volume places the genus
in the Elopide. To the present writer it seems best to retain
it and its related genera in a special family as Cope originally
did, until more is known regarding the anatomy.
60 Bulletin American Museum of Natural History. |Vol. X1X,
Pachyrhizodus caninus Cope.
Piate III, Fics. 1 AND 2.
Pachyrhizodus caninus Cope (E. D.), Proc. Amer. Philos. Soc., XII,
1872, p. 344; Rept. U. S. Geol. Surv. Mont. etc. 1872, p. 348;
Bull. U. S. Geol. and Geog. Surv. I, No. 2, 1874, p. 42; Vert.
Cret. Form. West, 1875, pp. 221, 276, pl. 1, figs. 1-4. — Crook
(A. J.), Paleontogr. XXXIX, 1892, p. 109. — Loomis (F. B.),
Paleontogr. XLVI, 1900, p. 262, pl. xxvii, figs. ro-12. — STEWART
(A.), Univ. Geol. Surv. Kansas, VI, 1900, p. 355, pl. lxx,"figs. 2-6.—
Woopwarp (A. §.), Cat. Foss. Fishes, IV, 1901, p. 44. — Hay
(O. P.), Bibliog. and Cat. Foss. Vert. N. A: r902, p. 387:
Pachyrhizodus latimentum Core (E. D.), Proc. Amer. Philos. Soc. XII,
1872, p. 346; Rept. U. S. Geol. Surv. Mont. etc. 1872, p. 348;
Bull. U. S. Geol. and Geog. Surv. I, No. 2, 1874, p. 42; Vert.
Cret. Form. West, 1875, pp. 223, 276, pl. 1, fig. 5; pl. li, figs. 1-7.
— Loomis (F. B.), Paleontogr., XLVI, 1900, p. 263, pl. xxv1,
figs. 7, 8. — Stewart (A.), Univ. Geol. Surv. Kansas, VI, 1900,
Pp. 357, pl. Ixviii; pl. lxx, figs. 9, 10. — Woopwarp (A. §.), Cat.
Foss. Fishes, IV, 1901, p. 42. — Hay (O. P.), Bibliog. and Cat.
Foss. Vert. N. A. 1902, p. 388.
Pachyrhizodus curvatus Loomis (F. B.), Paleontogr. XLVI, Igo0, p.
265, pl. xxv, figs. 6-8. — Woopwarp (A. S.), Cat. Foss. Fishes,
IV, 1901, p. 44. — Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A.
1902, Pp. 388.
The writer has ventured to unite the two species which
Prof. Cope has described under the names Pachyrhizodus
caninus and P. latimentum. The types of these are in this
Museum, that of P. caninus having the number 1881; that
of P. latimentum the number 1758. Besides these, there are
various jaws and other parts which were collected for Prof.
Cope by Messrs. Sternberg and Hill during the year 1877.
In attempting to apply to this material the characters as-
signed by Cope to his two species the writer has become con-
vinced that the differences are due partly to individual
variations and partly to distortions during fossilization. The
various collections indicate that the bones of this species were
soft and spongy, so that they easily suffered compression and
distortion. Jaw bones of the opposite sides of the same in-
dividual are sometimes so different that one is convinced
with difficulty of their identity. The groove which divides
1903. | fay, North American Cretaceous Fishes. 61
the symphyseal surface of the dentary of P. latimentum is
very distinct in specimens which on account of the small
height of the coronoid process would have to be assigned to
P. caninus. The height of the coronoid process will, with
little doubt, be found to vary in all degrees between the
measurements given by Cope for his two species.
The mandible of the species (Fig. 44) appears to have had
an external vertical surface and an inferior nearly horizontal
surface. In some specimens these are separated by a sharp
ridge; in others they are with difficulty distinguishable, a
condition probably due to distortion during fossilization.
Similarly, the maxilla has presented an external nearly flat
surface separated by a sharp border from a flat superior sur-
face. This surface, again, meets a flat palatal surface at a
sharp internal, or mesial, border. The section of the maxilla
e\ 3 Ad Ug ze 9 QQ DS ¥; ee ae ~~
GIO den a
Fig. 44. Pachyrhizodus caninus Cope. No. 1662. * 3. Mandible. azg., angular; arz.,
articular ; dex., dentary.
is, therefore, nearly triangular. In one specimen, however,
the maxilla of one side has the form described, while the other
is so distorted that its section is nearly parallelogrammic.
These modifications are mentioned in order to show the
necessity of guarding against giving too much value to varia-
tions in the forms of the bones of this genus.
Of the palatines and the pterygoids of this species the
writer has been able to learn little. Loomis (op. cit., pl.
XXvii, fig. 12) has figured what he regards as a palatine, but
it seems to be identical with a left maxilla in this Museum’s col-
lection. Besides, one would hardly expect to find a palatine
of a length so great that it would reach nearly to the quadrate.
In the National Museum at Washington there is a speci-
men of this species which I have been permitted to study.
62 Bulletin American Museum of Natural History. (Vol. XIX,
It appears to have been about six feet (1.83 m.) in length.
The skull measured about 275 mm. Unfortunately, the right
preopercular and some other bones are lying on the top of
the skull, so that the relations of the elements were not de-
termined. The preopercle had a length of at least 225 mm.;
and at the lower end, a part of which is missing, a width of
about 112 mm. A quadrate has a height of 85 mm. There
are 53 vertebre present, but they are considerably disturbed.
Of these apparently 27 belong to the caudal region. The
neural arches appear to have been slender and not high.
The tail is deeply forked. The lower lobe only is present.
Its length is about 375 mm., but a ray near the bottom
of the fork is only about 90 mm. long. There are about
Io rays in the lobe, not including 5 rudimentary rays on
the front edge. The first of the latter is a lunate bone, the
others are slenderer and straight or irregularly bent. The
terminal vertebre are not shorter than those farther for-
ward in the caudal peduncle and hence do not display the
crowded condition seen in Anogmius. There is a fan-shaped
hypural. The pectoral fin is
long and falcate. The anterior
ray measures close to 595 mm.
It has a width of 27 mm. at
the base and tapers gradually
to the tip. It is not divided
or segmented. Eleven rays
may be counted in this fin,
but probably a few are miss-
ing. The shortest one ob-
served measures 135mm. All
except the most anterior di-
vide distally into slender fila-
yy. ments. At the, base of tue
_ Fig. 45. Pachyrhizodus caninus Cope.
S. Nat. Mus. X 3. Shoulder girdle. arz. s., fin are seen two or three
articular surfaces for fin; cé., cleithrum ; cor.,
Fo eO ey eer Eee oar pct., pectoral baseosts. There appear to
be no remains of dorsal,
ventral, and anal fins. The shoulder girdle of the left
side is displayed from the inner surface (Fig. 45). It is
1903. | Hay, North American Cretaceous Fishes. 63
much like that of Zarpon, but the precoracoid is larger than
in the latter genus. On the other hand, it is smaller than in
Portheus. In the specimen under consideration it is about
125 mm. long. The shoulder girdle is here illustrated from
a rough sketch.
On Plate III, Fig. 1 is shown a view of the tail of this species
taken from No. 1900 of the American Museum. The tips of
the lobes are broken away so that the original dimensions
cannot be known, but the lobes are now respectively about
265 mm. and 335 mm.in length. There is a large terminal
fan-shaped bone, which supported the principal rays. On
each side there is a triradiate bone, apparently a modified
ray, lying on the terminal vertebral bodies. Ryder (Report
U.S. Fish. Com. for 1884, pl. vi, fig. 2) has figured a similarly
placed bone which he regards as growing out from a displaced
epural. In front of the lower lobe of the fin of Pachyrhizodus
here described is an excavation which probably has been
occupied by a lunate bone such as that referred to above as
being found in the specimen at Washington. Fig. 2 of Plate
III shows a nearly complete lobe. Its length is 435 mm.
In these fins the rays are few in number, large, and cross-
segmented. The fin resembles considerably that of Tarpon.
Its number is 1658.
Dr. Loomis has described a species which he calls P. curva-
tus. It is small, the tooth line of the dentary measuring only
50 mm. In this space are alveoli for 38 teeth. The maxilla
possessed alveoli for 41 teeth. The species appears to the
present writer to be a young individual of P. caninus. In
the type jaw of P. caninus are spaces for 38 or 40 teeth, and
in a maxilla I count at least 40 teeth. Professor Cope re-
garded this fish as probably a ground feeder, but the form
of the tail seems to indicate a swift, free-swimming, predaceous
animal.
No. 2041 of this Museum consists of 9 caudal vertebre
and apparently 15 rays of an unpaired fin. There can be
little, if any, doubt that the fin belongs with the section of the
vertebral column, but whether it is the anal or the dorsal
cannot be ascertained. The longest rays measure 147 mm.
64 - Bulletin American Museum of Natural History. (Vol. X1X,
in length, but the tips are broken off. From the anterior
they grow shorter and slenderer to the last. Probably
nearly the whole, if not the whole, of the fin is present. The
rays divide longitudinally into slender portions but show no
signs of segmentation. The nine vertebre have a length of
185 mm., and the fin has about the same length along its base.
Pachyrhizodus leptopsis Cope.
Pachyrhizodus leptopsis Cope (E. D.), Bull. U. S. Geol. and Geog.
Sutv. I, No. 2, 1894, p. 425 Vert: Cret: Porm, West) 1875-90.
225, 276, pl. li, figs. 8-8c. — Stewart (A.), Univ. Geol. Surv.
Kansas, VI, 1900, p. 354, pl. Ixx, fig. 1. — Woopwarp (A. S:),
Cat. Foss. Fishes, IV, 1901, p. 45.— Hay (O. P.), Bibliog. and
Cat. Fass Vert. N. A. 1902, p. 388.
Pachyrhizodus lepitopsis Loomis (F. B.), Paleontogr. XLVI, 1900, p.
264.
This species was based on a fragment of a right dentary
which presents the symphyseal surface. The specimen bears
the Museum’s number 1756. The species is characterized by
the large size of the bases of the teeth and the narrow symphy-
seal articulation. The bases on which the teeth rest are large,
fully as wide as long, and the empty spaces from which the
teeth have fallen are about circular. In P. caninus the teeth
are crowded, so that the tooth bases, measured across the
jaw, are wider than long, and the empty spaces are of greater
extent across the jaw than parallel with it.
Oricardinus sheareri Cope.
Pachyrhizodus sheareri Corr (E. D.), Proc. Amer. Philos. Soc. XII,
1872, p. 347; U. S. Geol. Surv. Wyoming; ete. 25726 presque
Bull. U. S. Geol. and Geog. Surv. I, No. 2, s874, p. 43; Vert.
Cret. Form. West, 1875, pp. 225, 276. — Woopwarb (A. S.), Cat.
Foss. Fishes, IV, 1901, p. 45.
Oricardinus shearert Cope (E. D.), Proc. Amer. Philos. Soc. XVII,
1877, pp. 177, 178. — Hay (O. P.), Bibliog. and Cat. Poss. Vert.
N. A. 1902, p. 388.
Pachyrhizodus sheari Loomis (F. B.), Paleontogr. XLVI, 1900, p. 264.
The type of this species is supposed to be a portion of the
left maxilla. A figure and section of the specimen is here-
1903. | Hay, North American Cretaceous Fishes. 65
with presented (Fig. 46). The anterior end of the fragment
is directed toward the right. The teeth are pleurodont, as
they are in Pachyrhizodus. They
have been crowded, as in Pachy-
rhizodus caninus, but none of the
crowns has been preserved. The
roots present at least one differ-
ence from the species of Pachy-
rluzodus,sofarasI have beenable pig. 46. Oricardinus sheareri Cope.
fosobserve. In the Watiem when “ori 7S - Pypen Manila and
the tooth separates from the root,
which is buried in the bone of the jaw and becomes anchylosed
with it, the ring-like edge of the root is very sharp. In 0.
shearert the remains of the root form a nearly flat ring around
the pulp cavity. This, looked at with a lens of high power,
shows radiating and concentric lines of dense bone. Nearly
all the teeth of the specimen appear to have been shed at
the time of its death; very few seem to have been broken
off afterward. The Museum number of the type is 1998.
Oricardinus tortus Cope.
Oricardinus tortus Cope (E. D.), Proc. Amer. Philos. Soc. XVII, 1877,
Pp- 177- — Woopwarbp (A. S.), Cat. Foss. Fishes, IV, 1901, p. 46.
— Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A. 1902, p. 388.
The type of this species is in the American Museum of
Natural History and has the catalogue number 2114. The
material consists of a part of the left mandible, including
the symphysis, and possibly about twenty vertebre. Figures
of the mandible are here presented showing it from the lingual
side (Fig. 47) and from the dental border (Fig. 48). In his
description, Cope deals much with the internal and external
ribs of this mandible, but it is difficult even with the speci-
men in hand to understand his meaning. The jaw has evi-
dently suffered some distortion and this has resulted. in
making some of the anterior teeth appear to lie on the outer
side of the jaw. The teeth are much like those of Pachyrhizo-
dus, but the symphysis is different and the jaw is narrower in
[ fanuary, 1903.) 6
66 Bulletin American Museum of Natural History. [Vol. X1X,
front, like that of Empo. Nevertheless, the narrowness may
be due to crushing, and cannot be relied on. The vertebre
accompanying the jaw may be said to be identical with those
Fig. 47. Oricardinus tortus Cope. No. 2114. Xx. Type.
Dentary. syz., symphysis.
Fig 49. ? Ovricar-
dinus tortus Cope.
No, 2114. X1. Two
caudal vertebra, lat-
Fig. 48. Same as Fig. 47. Shows tooth line. eral view.
of Empo in structure, and it is not improbable that they did
not belong to the individual that possessed the jaw. Two
caudal vertebre are figured here of natural size (Fig. 49).
Until more is known about the species it seems best to retain
it in the genus Oricardinus, of which it is the type.
ENCHODONTID.
Enchodus Agassiz.
Remains of fishes of this genus are very common in collec-
tions made in the Cretaceous of Kansas, and they likewise
occur in collections made in New Jersey. Fourteen species
have been described from these two States and another, E.
shumardi, from South Dakota. The greater part of the species
have been based on detached teeth. The most conspicuous
bone of the skull and the one most likely to be preserved is
the greatly swollen palatine, bearing a long fang. This bone
was regarded by Cope as the premaxilla, and the pterygoid,
which articulates with it behind, was supposed to be the
maxilla. The correct interpretation was afforded by Dr. A.
S. Woodward (Proc. Geologists’ Assoc., X, 1888, p. 315).
Dr. Loomis has more recently discussed the anatomy of the
genus, but in his restoration of the skull he has not repre-
sented the palatine as swollen nor drawn the boundary
between it and the ectopterygoid. He also describes the
wa
1903. | fay, North American Cretaceous Fishes. 67
palatine as a mass of osteodentine; but to the present writer
this mass, as also that composing the tritoral plates of Anog-
mius, appears to be merely very compact bone.
Dr. Loomis has probably given us the correct explanation
of the replacement of the palatine fangs. The new fang is
produced in front of the senescent one. Originally the latter
had stood on the very anterior extremity of the palatine
bone, but after the tooth had become affixed, the bone
prolonged itself in front of the base of the tooth and thus
provided a surface for the attachment of the next fang in
succession. When the new tooth has taken its position, its
predecessor, through absorption of its base, is loosened and
drops away, leaving a crescentic scar. Sometimes several of
these scars may be observed on the palatine. The new tooth
probably became firmly fixed before its predecessors fell
away; otherwise it would have been easily wrenched from
its moorings. It will probably also be found that there is
an alternation in the replacement of the fangs. Evidence of
this is found in the palatines of No. 2098. The right palatine
has a conspicuous process of bone extending forward over
the base of the fang. The left ah as has only the slightest
trace of such a process.
The large teeth on the anterior end of the pterygoid are
replaced also by the development of others in front of them;
and the scars resulting from the falling away of the old teeth
may be seen. On the other hand, the great fang on the
anterior end of the dentary is replaced by the growth of
another behind it, and the scars of former teeth lie in front
of the functional fang. Prof. Cope’s figure (Vert. Cret. Form.
West, pl. liv, fig. 3) shows the end of the right dentary from
without. The functional fang is broken off some distance
above its base. In front of it is an elevation of bone on
which stood the replaced fang. In the specimen, the surface
of the scar looks as if the old fang had only recently been
broken off. Cope’s figure 3a is unsatisfactory. Other spect-
mens of Enchodus confirm the conclusion that the new fang of
the dentary is produced behind the old one. Thus, while the
fang of the palatine is moving forward, that of the dentary is
68 Bulletin American Museum of Natural History. [Vol. XIX,
moving backward; but it is probable that growth of other
parts makes compensations, so that the two fangs are always
closely opposed to each other. Cope (op. cit., p. 301) has
described the mode of succession of the teeth of the dentary.
Enchodus ferox Leidy.
Enchodus ferox Letpy (J.), Proc. Acad. Nat. Sci. Phila. 1855, p. 397.
— Emmons (E.), Man. Geol. 2nd ed. 1860, p. 214, fig. 1824. —
Core (E. D.), Vert. Cret. Form. West, 1875, p. 277. — Loomis
(F. B.), Paleontogr. XLVI, 1900, p. 277. — Woopwarp (A. &.),
Cat Foss. Fishes, lV, ro0r, p: 204. — Eis. (O72) Bibliog and
Cat. Foss: Viert: NeAS 1902), p. aso.
Enchodus pressidens CopE (E. D.), Proc. Amer. Philos.-Soc. XI, 1869,
p- 241; Vert. Cret. Form. West, 1875, p. 277. — Loomis (F. B.),
Paleontogr. XLVI, 1900, p. 277. — Woopwarp (A. §&.), Cat.
Foss. Fishes, IV, 1901, p. 205. — Hay (O. P.), Bibliog. and Cat.
Foss. Vert. N. A. 1902, p. 389.
‘“Sphyrena” Morton (S. G.), Synop. Org. Rem. Cret. U. S. 1834, p.
a2 pl, sid, is on.
In the Cope Collection are two palatines belonging to the
genus Enchodus, both of which are labelled in Cope’s hand-
writing. One of these, No. 2251, is labelled “‘ Enchodus ferox’’;
the other, No. 2250, ‘“‘Enchodus pressidens, not typical.”’
Fig. 50 represents the specimen of E. ferox. A study of these
has led to the conclusion that the two species, E. ferox and
E. pressidens, are identical. The characters which are relied
on to distinguish FE. pressidens are the crescentic section of
the base of the palatine fang, the triangular
section of the middle of the tooth, the grooves
bounding the cutting edges, and the projec-
tion of the base of the tooth beyond the
anterior margin of the palatine.
As to the crescentic base, this results from
the pressure of the hinder side of the new
y oe pea anaes tooth against the base of the old tooth, and
ae ipolatine «and there is some reason for believing that the
concavity of the posterior side of the new
tooth diminishes somewhat after the old tooth has fallen
out. At any rate, such an explanation is suggested by a
1903. | Hay, North American Cretaceous Fishes. 69
difference found in the form of the bases of the two fangs of a
specimen of EF. petrosus. As regards the cross-section at the
middle of the tooth, I find no considerable difference. The
specimen labelled ‘“‘E. pressidens’’ agrees with the descrip-
tion of that species, but the “‘E. ferox’’ also has the inner
face considerably more convex than the outer, and the sec-
tions of these faces may be regarded as forming two sides
of a triangle, with the separating angle rounded off. The
grooves bounding the cutting edges of “FE. pressidens”’ are
hardly apparent, while there are indications of them in the
specimen called EF. ferox. Doubtless there were individual
variations in this character.
If Loomis’s explanation of the manner of replacement of
the palatine fangs is correct, as it quite certainly is, we can
see why in some cases the palatine bone projects beyond the
base of the fang, while in other cases the fang projects beyond
the bone. After the new tooth has taken its position in front
of the old tooth and at the very extremity of the palatine bone,
the latter proceeds to extend itself forward in order to provide
a seat for the base of the next tooth in succession. Thus we
sometimes get a palatine bone in one stage, sometimes in
another. In the “ferox”’ specimen the palatine had extended
considerably in front of the fang; in the “ pressidens”’ speci-
men the fang had only recently taken its place.
The specimen called E. pressidens is a little more than one-
half the size of that called E. ferox, the tooth being 32 mm.
long, that of E. ferox 51 mm. The palatine bone of the
‘‘pressidens’’ is also relatively slenderer than the other; but
all these differences are probably due to difference in age of
the animals.
E. ferox appears to differ from E. petrosus in three respects.
The inner face of the palatine fang is smooth, while in £.
petrosus it is coarsely striated. In E. ferox the cutting edges
of the palatine fang are minutely serrated; in EF. petrosus
they are smooth. In both species there is, on the outside of
the palatine, a broad shallow groove which runs from the
lower hinder portion of the bone upward and forward. In
E.. ferox this groove meets the upper edge of the bone over
70 Bulletin American Museum of Natural Flistory. [Vol. XIX,
the base of the fang; in E. petrosus, well behind its base.
The drawings of E. ferox furnished by both Morton and
Emmons of this species represent the teeth only and are very
unsatisfactory. I find no reason for questioning the correct-
ness of Cope’s identification of the tooth here figured as E.
jerox. E. ferox and E. pressidens Cope were both described
from the Cretaceous of New Jersey.
Enchodus petrosus Cope.
Enchodus petrosus Corr (E. D.), Bull. U. S. Geol. and Geog. Surv.
Terrs. I, No. 2, 1874, p. 44; Vert. Cret..Form. West, 1875, pp-
239, 278, pl. liv, figs. 4-7. — ? Loomis (F. B.), Paleontogr. XLVI,
1900, p. 278, pl. xxvii, figs. 13-15. — STEWART (A.), Univ. Geol.
Surv. Kansas, VI, 1900, pl. Ixx, fig. 11. — Woopwarp (A. $.),
Cat. Foss. Fishes, IV, 1901, p. 205. — Hay (O. P.), Bibliog. and
Cat. Foss. Vert. N. A. 1902, p. 380.
Tetheodus pephredo Core (E. D.), Bull. U.S. Geol. and Geog. Surv. I,
No. 2, 1874, p. 43; Vert. Cret. Form. West, 1875, pp. 237, 277)
pl. liv, figs. 1-3. — Woopwarp (A. 5.), Cat. Foss. Fishes, IV,
Igo1, p. 205. — Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A.
1902, p. 389.
Tetheodus pephero Loomis (F. B.), Paleontogr. XLVI, 1900, p. 278
(syn. of Enchodus petrosus).
The present writer agrees with Dr. Loomis in referring
Cope’s Tetheodus pephredo to Enchodus petrosus. The type of
this supposed species is in this Museum and has the number
1605. Cope’s description and figures are for the most part
COPrect:
Prof. Cope states that in Tetheodus pephredo there is no sur-
face for the attachment of a tooth and no scar or other trace
of the former existence of one. However, a close examina-
tion of one of the palatine masses shows that there are traces,
faint but undeniable, of at least four fangs which at different
times have occupied the lower border of the bone. More-
over, where we would expect to find a functional fang, the
surface is somewhat rough; while on the oral border of the
bone there is a ragged area which looks as if some of the bone
had been broken away. All this makes it appear probable
that in some conflict both fangs had been torn away at. their
1903. | Hay, North American Cretaceous Fishes. 71
bases, and that sufficient time had not elapsed for the scars
to be wholly healed. In front of and above the base of the
supposed former fang, on the anterior extremity of the pala-
tine, there is a process of bone which may be regarded as an
outgrowth preparatory to the attachment of a new fang. It
is rather thin from side to side, only about one half as thick
as would be required for the fang, but it is possible that its
thickness might become greater in due time. Whether or
not a new fang could be produced and effect an attachment
without the support and protection of a fang already in place
seems to be doubtful. It seems likely that every victim by
its strugglings would destroy any incipient connections that
had been formed.
Fig. 51 represents a damaged dentary bone of this species,
with two teeth nearly complete. The most anterior of these
is Shown two-thirds of the
natural size, in order to
display the rather strongly
developed ridges on the pos-
terior half of the tooth. On
the lingual face of the tooth =
these ridges are found well Fig. 51. Exchodus petrosus Cope. No. 2062.
forward, but here they are Den" Xi one tooth, x &.
rather short. Toward the hinder border éf this face they rise
well toward the tip of the tooth. About three millimeters
above the base of the tooth the ridges cease suddenly, and
the portion of the surface below them is provided with very
fine striations. The anterior fang is broken away, but its
outlines are restored from another specimen. The number
is 2062.
The surfaces by means of which the palatine bone comes
into contact with the bone which acted as its suspensory,
doubtless the prefrontal, deserve description. These articu-
latory surfaces are found at the hinder end of the palatine,
and those of Cope’s type are shown in Fig. 62; those of FE.
s@vus in Fig. 61. In the former we have two processes, one
below and directed upward and outward. ‘The posterior face
of this is smooth and forms one of the articulatory surfaces.
72 Bulletin American Museum of Natural History. (Vol. X1X,
Another process is considerably in front of the one just de-
scribed, and is directed upward. Its posterior face is flat
and smooth. The inner face of the lower and hindermost
process looks upward and inward, and 1s slightly convex and
smooth. It connects the two articulatory surfaces which are
directed backward and form a third surface. A thin per-
pendicular plate of bone has extended backward from the
inner border of the anterior articular surface near the base
of the posterior process, but it is now broken away. It is
seen in the figure of the corresponding parts of E. sevus. It
appears evident that the posterior process in E. petrosus has
been distorted, so that its upper surface is directed more out-
wards than in life. For the same reason, it is now lower
than originally. This is indicated by another specimen.
Through these smooth articulatory surfaces the palatine must
have had a very free movement on the prefrontal.
Portions of the palatine fangs of an Enchodus from the
Fox Hills Group of New Mexico are not distinguishable from
those of FE. petrosus. (Cope, Amer. Naturalist, XXI, 1887,
p- 566.)
Enchodus dolichus Cope.
Enchodus dolichus Corr (E. D.), Vert. Cret. Form. West, 1875, pp.
239, 278, 300, pl. liv, figs. 8, 8a; Proc. Amer. Philos. Soc. XXIII,
1885, p. 3. — Loomis (F. B.), Palzontogr. XLVI, 1900, p. 279,
pl. xxvii, figs. 16, 17. — STEWarRT (A.), Univ. Geol. Surv. Kansas,
VI, 1900, p. 377, pl. Ixx, fig. 12. — Woopwarp (A. S.), Cat. Foss.
Fishes, IV, 1901, p. 204. — Hay (O. P.), Bibliog. and Cat. Foss.
Vert. N. A. 1902, p. 389.
Of this species the type, a fragment of the palatopterygoid,
is in the Museum and bears the number 1820. There are like-
wise considerable portions of three skulls, including those
described by Cope on page 300 of his ‘Vertebrata of the
Cretaceous Formations of the West.’ One of these skulls,
Cope’s “No. 1,’’ No. 1837 of this Museum, is represented by
Fig. 52. It displays both palatines, the right much out of
its natural position, the left pushed backward about 25 mm.
Its great fang is crossed by the anterior end of the ectoptery-
goid. The long teeth of the latter bone are well shown.
1903. } Hay, North American Cretaceous Fishes. 73
The anterior end of the mandible is broken away. Above
the palatine is the right premaxilla with about 23 teeth and
spaces for others. Between the upper and the lower thirds
of the figure lie the occipital bones and the frontals. The
left articular runs forward beneath the left palatine. The
he a I
Cpt ——A]
fT
aS
Fig. 53. Enchodus dolichus
Fig. 52. Exchodus dolichus Cope. No. 1837. X 3. Coe No. 1685. X 4. den., den-
Disturbed skull. avg., angular; art., articular; den., tary; ec.pt., ectopterygoid; /*.,
dentary ; ec. f¢. ectopterygoid ; Tr frontal ; op.. oper- frontal ; mx., maxilla; pad, pala-
cular ; pal., palatine ; Awzx., premaxilla ; Gu., “quadrate, tine ; : pmx., premaxilla.
right quadrate is displaced so that its articular surface (above
qu.) looks upward.
Figure 53 represents No. 1865 of this Museum. This dis-
plays both palatines and their fangs; the anterior end of the
ectopterygoid and two teeth; the lower jaw, with its fang
and teeth of two sizes; both premaxille, with their small
teeth; and the anterior end of the frontal. Alongside of the
frontal is a fragment of a toothed bone which lies on a line
with the dental border of the premaxilla. It is probably a
portion of the maxilla.
The palatine fang was missing in the type. No. 1837
shows, as Cope has already stated, that the palatine fangs
are finely striated on the outer surface. On the hinder por-
74 Bulletin American Museum of Natural History. [Vol. XIX,
tion of the inner face of the left palatine and of two man-
dibular teeth we find a moderately coarse striation. Cope’s
specimen ‘‘ No. 2,’’ which is No. 1890 of this Museum, presents
both palatines, one with the fang complete. The ornamenta-
tion is as in No. 1837. The same is true of the fang of No.
1865, except that the striation, both on the inner and the
outer face, is somewhat coarser. The striation of the hinder
part of the inner face of No. 2385 is decidedly coarser than
that of any of the other specimens. All these teeth show
that E. dolichus differs from E. petrosus in having the outer
face with practically the same convexity as the inner. The
latter seems also to have attained a considerably larger size.
We must, however, keep in mind that there are likely to be
small specimens of E. petrosus.
Enchodus tetrzcus Cope.
Enchodus tetrecus Corr (E. D.), Vert. Cret. Form. West, 1875, p. 278.
— Woopwarp (A. S.), Cat. Foss. Fishes, IV, 1901, p. 205. —
Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A. 1902, p. 389.
Enchodus tetracus Loomis (F. B.), Paleontogr. XLVI, 1900, p. 277. —
STEWART (A.), Univ. Geol. Surv. Kansas, VI, 1900, p. 375.
This species is said to have been based on various teeth
from the Greensand of Delaware and New Jersey. One whole
tooth, which is labelled as the type, is in the
Museum and has been given the number
2248. There is another with
the distal end missing. Cope’s
description is sufficient, but it
is thought to be proper to pre-
sent here a drawing of the type
(Fig. 54). Figure 55 represents i
Fig. 54. Enchodus the paratype, an imperfect Fig. 55. Exchodus
tetrecus Cope. No. : 3 Z a NOs
ia Sie Type. tooth designated as No. 2240. ERE oe
Palatine fang and see ge i f d
crosssection «Lhe sharply defined ‘stratioosi. ne
of the very convex inner face
distinguish this species from E. ferox. The teeth are ap-
parently slenderer than those of E. petrosus, and the shallow
1903. | Hay, North American Cretaceous Fishes. 75
grooves running along the inner face, one close to and parallel
with each cutting edge, are somewhat more distinct; but
these characters are hardly satisfactory. On account of the
little that is known about E. tetrecus and on account of the
different geographical distribution, the two species may best
be regarded for the present as distinct.
Enchodus gladiolus Cope.
Cimolichthys gladiolus Corr (E. D.), Proc. Amer. Philos. Soc. XII,
HO 7/215 a Sat
Phasganodus ? gladiolus Corr (E. D.), Bull. U. S. Geol. and Geog.
Surv. Derrs.1, No. 2,,1872, p: 43; Vert. Cret. Form. West,. 1875
PP. 235; 277.
Enchodus gladiolus Corr (E. D.), Vert. Cret. Form. West, 1875, p.
301, pl. xlii, fig. 7, — Woopwarp (A. S.), Cat. Foss. Fishes, IV,
1901, p. 204. — Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A.
1902, p. 389.
The type of this species was a single detached tooth, and
this has not yet come to light in the examination of the Cope
collection. Cope states (Vert. Cret. Form. West, p. 301) that
he had a better specimen; but while he figured the type
tooth, he did not figure or describe the better example. In
the collection there is found a specimen which bears the label
in Cope’s handwriting ‘“‘ Enchodus ?gladiolus,” the interroga-
tion doubtless belonging, accord-
ing to Cope’s usage, with the
specific name. To these remains
have been given the number 1818.
The specimen presents the left
palatine, with its great tooth com-
plete; the left ectopterygoid, with Se Maier ree ek eee
several teeth; and the tip of the Ne. pee ie lence ;
left dentary with its fang. As
shown in Fig 56, the ectopterygoid has been turned so that
the teeth point in a direction opposite to that of the palatine
teeth. On another block and apparently belonging to the same
individual is shown the upper surface of the hinder half of the
skull. The palatine fangs differ from those of E. dolichus in
76 Bulletin American Museum of Natural History. [Vol. XIX,
having nearly the whole of both the inner and the outer face
coarsely striated. The ridges and intervening furrows are
easily seen with the unaided eye, while in the case of E.
dolichus this requires a close observation. The striation
subsides close to the anterior very thin edge. As in E.
dolichus, the two faces are equally convex. The pterygoid
and the mandibular teeth are similarly marked by coarse
ridges and grooves.
It is possible that this specimen is only an example of E.
dolichus with unusually coarse grooving of the teeth; but I
think that it is distinct.
Fig. 57 represents the upper surface of the skull. The
more anterior portions of the frontals have left their impres-
sion on the matrix but are not rep-
resented in the drawing. From each
epiotic region a grooved ridge runs
forward to the prefrontal region.
From this ridge, at the centre of
growth of the frontal bone, a less
prominent ridge runs outward and
backward to the middle of the squa-
Fig: 7. Bnckede: gations ™OSAl, “The epiotics ares promimens
Cope. No. 1818. 2. Hinder 3
part of skull. han epionc Thee and are connected by a sharp ridge,
Hensie 27> Postfrontal; #40 behind which the occiput drops off
steeply. The parietals are apparently
Separated by the small supraoccipital. The parietals seem to
form a narrow band along the ridge connecting the epiotics.
The sutures are very indistinct.
Enchodus sevus, sp. nov.
This species, which appears to be distinctly different from
any hitherto described, was collected near Elkader, Logan
County, Kansas, by Dr. W. D. Matthew, in 1897. The col-
lector regarded the beds as belonging to the Pierre formation;
but Dr. Williston, who is familiar with the locality, informs
me that the deposits belong to the Niobrara. The species
has been a large one, as is indicated by the following measure-
ments:
eS ee oS
1903. | Hay, North American Cretaceous Fishes. Te
Length of the lower jaw from the chin to the articular con-
GAS Bio otek id en a S Bye aratol CRUE GONE Re ae Ce 223 mm.
Meoht ol the jawiab tmercoronold; process. .-..-..-..4+«. 67 mm.
Breadth of the'skull at the postorbitals.- 2.5... a gees III mm.
Length of the palatine and prefrontal to the front of the fang 72mm.
Unfortunately the whole surface of the specimen has been
covered and in some places injured by a deposit of crystals
of gypsum, so that it is difficult to determine some structures.
Other structures and the general forms of the bones are dis-
tinct enough. The number of the specimen is 198.
The distinguishing character of this species is found in the
great palatine fang. In other American species of the genus
where the palatine fang is known the latter is compressed
laterally. In the present species the compression is nearly
antero-posterior. The species is nearest to E. petrosus, with
which it was at first identified.
The right palatine bone (Fig. 58) is in excellent condition,
barring the deposit of gypsum on its sur-
face. It has suffered little or no distortion
or compression. No part of the ectoptery-
goid adheres to it. The figure represents
the inner side of the tooth and shows the
position of the inner, or posterior, cutting
edge, The greater part of the fang is pre- Wig. «8, Exchodus so
served. The left palatine is distorted and 0°" Rete eae
has lost all but the base of the fang. sata Ppa
In E. petrosus a sharp cutting edge begins at the base of the
fang in front and runs downward to the tip (Fig: 59). This
is very distinct from the first, and it pursues its course near
the anterior midline of the tooth, separating an outer from an
inner face. The section shown in Fig. 59, 0, ought to have been
taken somewhat higher up on the tooth, in which case the
inner face (on the left) would have been somewhat more con-
vex, but it would still have differed much from Fig. 60,a. In
E. sevus (Fig. 60) there is a faint trace of a corresponding
edge and it may once have been stronger, but it lies much
nearer the inner side of the tooth. Another cutting edge
starts at the outer side of the base of the tooth; but, instead
78 Bulletin American Museum of Natural History. [Vol. XIX,
of soon getting near the middle of the hinder surface of the
tooth, as it does in E. petrosus, it forms the outer border of the
60
Enchodus
petrosus Cope. Oo
59
Fig. 59.
2062. 4. Left pal-
atine fang from in
front. The line, 64,
marks position of sec-
tion 6.
Fig. 60. LExchodus
sevus Hay. No. 198.
x4. Type. Right
palatine fang from in
front. The line aa
marks position of the
cross-section @.
tooth when this is looked at directly from
behind or front. These two edges divide
the surface of the distal end of the tooth
into two nearly equal faces, of which one,
somewhat smaller, is nearly anterior, but is
directed somewhat outward, while the larger
and somewhat more convex one is directed
backward and inward. About the middle
of the length of the tooth, where the section
(Fig. 60, a) is taken, the inner face is far
larger and more convex. The palatine bone,
from which Fig. 60 is taken, was only slightly
larger than that from which Fig. 59 was ob-
tained.
The palatine bone has about the size of
that of E. ferox, figured in this paper, and
the fang has probably had about the same
length. If we
measure the greatest diameter of this fang of E. ferox at a
point ro mm. below its base we find it to be ro mm., and
this diameter is the antero-posterior, while
the transverse diameter is not quite 8 mm.
At the same height the diameter of E. s@vus
is 10 mm. in the transverse axis of the cross-
section, and 9 mm. in the antero-posterior.
The same proportions and directions of the
axes are found in E. petrosus as in E. ferox.
The posterior end of the palatine (Fig. 61)
appears to differ somewhat from that of E.
petrosus (Fig. 62). It presents the same
smooth articular surfaces as are observed in
the latter species, but, as will be observed in
the figures, the lower process, a, of the bone
much thinner and higher than in E. petrosus.
Enchodus
No. 198.
Fig. 61.
se@vus Hay.
x 4. Type. Posterior
end of right palatine
bone. a, articular pro-
cess.
Fig. 62. Enchodus
petrosus Cope. No.
1608. X4. Type. Pos-
terior end of left pal-
atine bone.
in E. s@vus, is
This difference
may not be specific but due in some part to accidents of fos-
silization.
62, that of the left side.
Fig. 61 represents the bone of the right side; Fig.
1903.] Hay, North American Cretaceous Fishes. 79
Fig. 63 represents the lower jaw seen from the outside. It
is everywhere ornamented with radiating ridges and grooves.
In front are three deep notches, as in many species of the
Fig. 63. Exchodus sevus Hay. No.198. X 4%. Type. Lower jaw, outer
view. ang., angular; @art., articular; dex., dentary.
genus. Fig. 64 shows the lower jaw from the lingual side,
together with the ectopterygoid and quadrate. The two last-
mentioned bones are somewhat displaced. Attention is
called to the great raptorial tooth on the anterior end of the
Fig. 64. Exchodus sevus Hay. No. 108 X 4. Type. Lower jaw,
quadrate, and ectopterygoid ; inner view. amg., angular; art., articular; dex.,
dentary ; ec. #z., ectopterygoid ; gw., quadrate.
ectopterygoid. It is about 30 mm. long. The upper border
of the ectopterygoid is excavated in front and received the
lower border of the palatine.
-One preoperculum is present (Fig. 65). It appears to me_
to belong to the left side. It is very narrow above, but
broadens somewhat below. The exterior surface is convex
transversely and has ridges and grooves running lengthwise.
A conical process from its front border fits into a groove in
the hinder border of the quadrate. On the inner face of the
preoperculum there is a deep channel running nearly the
full length and opening forward. What I regard as the left
interoperculum is shown in Fig. 65, 7. op. A portion of its
80 Bulletin American Museum of Natural History, [Vol. X1X,
hinder border has been broken away. It is a rather heavy
bone and is ornamented with radiating ridges and grooves.
The anterior border has been furnished
with a row of about eight sharp processes.
It seems to have been overlapped by the
hinder border of the lower end of the pre-
operculum.
The upper surface of the head is present,
except the snout; but it is too much over-
lain with gypsum to permit accurate des-
cription. It resembles the skull of E.
faujast Agassiz, which is figured by Dr.
A. S. Woodward (Cat. Foss. Fishes, IV,
7a PA eters pl. xi, fig. 6). A deep and broad exca-
Hay. No. 198. x}. Type. vation runs along the middle of the head
Z. op., interoperculum; Z. of.,
preoperculum, F from the supraoccipital.
Stratodus oxypogon Cope.
Stratodus oxypogon Cope (E. D.), Proc. Amer. Philos. Soc. XVII, 1877,
p. 180. — Woopwarp (A. S.), Cat. Foss. Fishes, IV, 1go1, p, 189.
— Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A. 1902, p. 387.
The type of this species bears the number 2113. It con-
sists of the distal portion of the right dentary (Fig. 66, den.),
a portion of what Prof. Cope regarded as a maxilla (Fig. 66,
mx.) a piece of a palatine (Fig. 67), and eight vertebrae. Cope
2
CUCECT ERC) eee
LIED LLIP ZEAL
ae | poe A
Fig. 67. Stratodus
oxypogon Cope. _No.
2113. X 1. Type. Frag-
Fig. 66. Stratodus oxypogon Cope. No, 2113. X 1. ment of palatine, show-
Type. Jaws, outer view. dex., dentary ; x., maxilla. ing teeth.
has given sufficient description; but no figures have hitherto °*
been furnished of the specimens. The dentary and maxilla
are here shown as seen from the outer side, and the fragment
of palatine is represented from the toothed surface. Most of
the teeth are pressed against the surface in such a way as to
1903. | Hay, North American Cretaceous Fishes. SI
suggest that they were hinged; as Cope also inferred from the
form of the alveolar fosse. This author has described the
tips of the teeth as being simple; but in the case of some of
them I find the peculiar spade-shaped
apices which Cope has described in
the case of S. apicalis. Fig. 68 shows
two of the caudal vertebre. They
are much like those of Empo, but
lace ther lonpitudinalkrivs Of “that: Gio ce. “siefaus vaypocon
genus. Under the lens, however, a et ee eee opel
fine longitudinal striation is seen on
the sides of the centra. Dr. Woodward has placed this genus
in the Dercetide, but notwithstanding the union of the pari-
etals in the midline, it appears to the writer to belong to the
Enchodontide.
Empo Cope
This genus was erected by Cope in 1872 (Proc. Amer.
Philos. Soc., XII, p. 347). The type species is E. nepaholica,
later emended by Cope to nepeolica. The species belonging to
this genus have been referred by Loomis (Paleontogr., XLVI,
1900, p. 267) to Cimolichthys; and Dr. A. 8. Woodward has
more recently (Cat. Foss. Fishes, IV, p. 221) adopted the
same course. That the two genera are not identical the
present writer will not affirm; but the type species of Czmolich-
thys, C. levesiensis, is, as Dr. Woodward has said, not satis-
factorily definable, many parts of the skull not being known.
Likewise, we are in ignorance regarding various structures of
Empo. It is therefore not at all improbable that differences
of generic importance may yet be discovered in these types.
It is as well to be conservative in suppressing genera as in
establishing them. For these reasons the writer prefers to
retain Empo.
Empo nepaholica Cope.
IBF, Ih, Wes vib
Empo nepaholica Corr (E. D.), Proc. Amer. Philos. Soc. XII, 1872,
P- 347; Rep’t U. S. Geol: Surv. Mont. etc. 5th Ann. Rep’t, 1872,
p- 345. — Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A. 1902,
Pp- 387.
[ Fanuary, 1903.] 6
82 Bulletin American Museum of Natural History. (Vol. X1X,
Empo nepeolica Corr (E. D.), Bull. U. S. Geol. and Geog. Surv. I,
No. 2, 1874, p. 46; Vert. Cret. Form. West, 1875, pp..230; 279,
pl, oxlix, “te. 95. pl: J) fig, 85 ype. Uy ci ey pl eer
Stewart (A.), Univ. Geol. Surv. Kansas, VI, 1900, p. 332, pl. lix,
figs. 1-9; pl. lxi, figs. 2-5.
Cimolichthys nepeolica Loomis (F. B.), Paleontogr. XLVI, 1900, p. 271,
pl. xxvii, figs. 1-3. — Woopwarp (A. S.), Cat. Foss. Fishes, IV,
TOOL, ps 225, text figs, 8; 9.
Cimolichthys semianceps Corr (E. D.), Proc. Amer. Philos. Soc. XII,
1872, p. 351; Rept U.S. Geol. Surv. Mont.ete, 1872, pu 3205 —
Loomis (F. B.), Paleaontogr. XLVI, 1900, p. 273, pl. xxvii. figs.
4-6. — Woopwarb (A. S.), Cat. Foss. Fishes, IV, rgo1, p. 228.
Empo semianceps Cope (E. D.), Bull. U. S. Geol. and Geog. Surv. I,
No: 2; 1874, p: 46; Vert: Cret. Form. West; 1675, pp. 2so,0270-
pl. liii, figs. 1, 2, 6-9. — Stewart (A.), Univ. Geol. Surv. Kansas,
VI, 1900, p. 338, pl. 1xi, figs. 6-9. — Hay (O. P.), Bibliog. and
Catt, Hoss) Verta-NiAy 1902) p: 387.
Cimolichthys sulcatus Corr (E. D.), Proc. Amer. Philos. Soc. XII, 1872,
eno iGaie
Empo sulcata Corr (E. D.), Bull. U. S. Geol. and Geog. Surv. Terrs.
1, No. 2, =874, p. 46;
Empo contracta Cork (E. D.), Bull. U. S. Geol. and Geog. Surv. I, No.
2, 1874, p. 46; Vert. Cret. Form: West, 2675, pp. 232, 270, pre
lili, figs. 14-17. — STEWART (A.), Univ. Geol. Surv. Kansas, VI,
1900, p. 339. — Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A.
1902, p. 387.
Cimolichthys contracta Loomis (F. B.), Paleontogr. XLVI, 1900, p.
273, pl. xxvii, figs. 8, 9. — Woopwarb (A. §S.), Cat. Foss. Fishes,
IV, rgo1, p. 228.
Empo merrillait Cope (E. D.), Bull. U. S. Geol. and Geog. Surv. I, No.
2, 1874, p. 46; Vert. Cret. Form. West, 1675, ppw232s27 9700
liti, figs. 10-13. Hay (O. P.), Bibliog. and Cat. Foss. Vert.
N. A. 1902, p. 387.
Cimolichthys merrillii Loomis (F. B.), Paleontogr. XLVI, 1900, p.
272, pl. xxvii, fig. 7.— Woopwarp (A. S.), Cat. Foss. Fishes,
IV, 1901, p. 228.
The type specimen of Empo nepaholica is a fragment of the
hinder end of the palatine, which bears the bases of two large
teeth and parts of smaller teeth, or bases thereof. It is
figured by Cope as cited in the synonomy. The specimen is in
the American Museum, and has the number 1904. After-
1903. | flay, North American Cretaceous Fishes. 83
wards Cope identified as belonging to the same species much
more satisfactory materials. One lot of these, No. 1736 of
this Museum, was figured on plate lu, fig. 1, of his large work
of 1875. Another, No. 1735, furnished figures 3-5 of plate
liii of the same work. From these more complete materials
many additional characters of the species were determined.
In the paper in which EF. nepaholica was described, Cope pro-
posed also the new species Cimolichthys sulcatus, which he
afterwards made a synonym of F.nepaholica. The type of
this, No. 1882, is the left dentary, showing thirteen large
teeth. The distal end of this dentary afterwards furnished
figure 8 of plate 1 of the monograph of 1875.
The type of Cimolichthys semianceps consists of vertebre, a
palatine, portions of the dentaries with teeth, and some other
parts. The vomer mentioned by Cope has not been recog-
nized. The number of this type is 1989. Inthe monograph
just mentioned Cope figured other specimens which he had
identified as belonging to this species. One of these, now
No. 1740, furnished figures 1 and ra of plate liii; another,
No. 1741, furnished figure 2 of the same plate; while No.
1742 was represented by figures 6-9.
Empo merrillit was based on fragments now included under
the number 1737. There are present nearly the whole left
palatine, of which figure 11 of plate liii of Cope’s monograph
represents the anterior end; a considerable part of the right
palatine, one fragment of which is shown by figure 12; a
flat bone, represented by figure 10, and which is probably the
entopterygoid; and the vomer, represented by figure 13.
Empo contracta is likewise founded on fragments of jaws.
There are present the right palatine, except its anterior ex-
tremity; the anterior extremity of the left palatine; the
vomer; and the anterior portion of the left dentary. Most of
these parts are represented on plate liii of Cope’s work. The
Museum number of this lot is 1738.
It may be said that the figures which Cope has published
represent the originals adequately, and that nothing would be
gained by refiguring them. If the figures are in some cases
unsatisfactory the fault lies in the materials.
84 Bulletin American Museum of Natural History. [Vol. XIX,
What is the value of these species? £. contracta is said to
be characterized by the flatness of the “‘maxillary bone,” by
which is meant the palatine; but in the types the palatine
of E. merrillii is still flatter, especially at the anterior end.
To what extent the flatness of this bone in both species is
due to its original form and to what extent to distortion dur-
ing fossilization is hard to determine. There was probably
considerable individual variation among these fishes. As re-
gards the sizes of the teeth in different parts of the palatine,
the writer has found, in looking through the collection of
this Museum, so much variation that he regards the dis-
tinctions specified by Cope as of little or no value. Consider-
able differences are often to be observed on the opposite sides
of the head. Cope also found characters in the “‘tongue-
shaped pharyngeal bone,”’ which is really the vomer; but
after diligent comparison on the part of the present writer
the attempt to find specific characters in this part was
abandoned.
Prof. Cope has given extended and accurate descriptions
of E. nepaholica and E. semianceps; but when we seek for
the characters by which they may be distinguished from
each other, they are found to be rare and elusive. The
character which is most definitely stated is found in the
structure of the mandibular teeth. In E. nepaholica these
are said to possess no cutting edge on the posterior face.
Unfortunately the crowns of the teeth are nearly always
broken off, so that it is impossible to apply this test. The
types themselves cannot, at least now, be distinguished
in this respect. In another specimen I find a mandibular
tooth with a posterior edge, but there is no other indication that
it is not E. nepaholica. Both species possess such two-edged
teeth on the palatines. On the lower jaw of a very large
specimen in the Museum I find that the teeth are two-edged;
and observation makes it certain that if such teeth belong
only to E. semianceps this was not a smaller species than
E, nepaholica. Stewart says that the mandibular teeth of
specimens which he identified as E. nepaholica seemed to
have both anterior and posterior cutting edges.
1903. | Flay, North American Cretaceous Fishes. 85
Having made an earnest effort to apply to the specimens
in this Museum the characters presented by Cope as belonging
to his species and such other characters as could be found, the
writer has been compelled to abandon the attempt to retain
the species founded by Cope and has found it necessary to
reduce’ them’ all to”™ the’ one
having priority of description,
E. nepaholica.
Dr. Loomis has furnished an
excellent figure of the skull of
this species, seen from the side,
and other figures of the rear of
the skull. A view of a well
preserved skullis presented here
(Big. 69). The number ‘of the
Specimen is 2522. “The squa-
mosal region of the left side has
been damaged somewhat, and
the drawing of this part is com-
pleted from another skull, No.
1888. On the right side a ver-
tebra and some other elements
are lodged against the skull be-
hind the postorbital region. As
both Loomis and Woodward
have indicated, the very large
frontals (Fig. 69, jr.) extend Fig. 69. Emo nepaholica Cope. No.
xX 3. Upper view of skull. dex.,
: 2522.
close to the hinder end of the dentary ; e¢#., ethmoid; /~., frontal; #z2.,
maxilla; fa., parietal; fa/., palatine;
skull. However, there is. a pi., prenazilla: pt.0., pterotic ; At. or.,
narrow band of bone showing ee Sega maar oo Sets
behind each frontal on the upper
surface and extending from the supraoccipital outward to
the epiotic process. These bands are doubtless the parietals.
No suture is to be observed between the parietal and the
epiotic, but it may have been present. The supraoccipital
meets the frontals, thus separating the parietals. It sends
on each side outward and backward a process which passes
behind the inner end of the parietal. There is a deep fossa
den.
86 Bulletin American Museum of Natural History. {Vol. XIX,
intervening between the epiotic and pterotic processes. There
appear to be good reasons for drawing as we have, the lateral
boundaries of the frontals; but the suture between the squa-
mosal, sg., and the postorbital, pt. or., cannot be determined.
The frontals extend far forward, so that they occupy by far
the greatest portion of the upper surface of the skull. In
the median line in front is the ethmoid, eth. The exact
limits of this have not been determined. Posteriorly it ex-
pands and it appears to divide, sending a branch, na?., back-
ward close to the mesial border of each premaxilla; but it is
more probable that these lateral branches are distinct bones,
the nasals. The premaxilla, pmx., is a thin, elongated,
sculptured bone, having small teeth along the lower border,
except in front. It is ap-
plied closely to the palatine.
Its teeth appear to have
been directed forward. This
is shown in Figure 70, pmx.,
and the same appearance is
presented by so many speci-
Fig. 70. Empo nepaholica Cope. No, 17 2 ne
<4. Part of skull, lateral view. arz., eee mens that this position of
den., dentary ; ec. ft., ectopterygoid ; ex. fz., en-
topterygoid ; eth., ethmoid ; SEs frontal ; 72, pt., the teeth appears to be the
metapterygoid ; mx. . maxilla’; na ?, nasal? : pa.
parietal ; fyzx., premaxilla ;’ At. 0., postorbital : normal one.
SMX., supramaxilla.
The maxilla, Figs. 69, 70,
mx., is a long, compressed, toothless bone which forms the
posterior border of the mouth. Its anterior end overlaps for
a long distance the premaxilla, its extremity in both the speci-
mens figured here rising above the upper border of the pre-
maxilla. The same position is shown in the skull figured by
Loomis. Figure 71, No. 1969, shows the skull seen from
below. The parasphenoid is very broad in front. It has
possessed no teeth. The articular surface for the hyomandi-
bular is short. The vomer does not appear in this specimen.
It is the bone which Cope called and figured with doubt
a pharyngeal. Stewart calls it the ethmoid. It is possibly
consolidated with the ethmoid, but certainly both bones
are represented. It possesses a varying number, one to four,
of rows of teeth. Various specimens indicate that other
1903. | fay, North American Cretaceous Fishes. 87
bones within the mouth were furnished with small teeth,
but which they are has not been accurately determined. It
appears likely that the ectopterygoid
had minute teeth on its surface and some
larger ones near one border.
Ihave had the opportunity of study-
ing No. 4186 of the U. S. National
Museum. In this the cleithrum has its
external face presented. The first ray
of the pectoral is segmented transverse-
ly, and was not a spine. Its anterior
border is furnished with square notches
and teeth, as is seen in Cope’s work, pl.
lu, fig. 1. Onlifting the cleithrum from
its bed the base of the fin is seen more
distinctly. Besides the first ray about
seven others are present. The precora- _Fig.71. Zmfo nepakolica
ee F Cope. No. 1969. X 4. Skull
coid is present and is somewhat larger from below, 4.0c., basioccipi-
tal; ex.oc., exoccipital; /7.,
than in a salmon which has a jaw two- frontal: fa., parietal: pmx.,
thirds as long as that of the fish here Bia i
described. A portion of the coracoid is present and perhaps
some of the scapula. The presence of the precoracoid fixes
the position of the fish among the Isospondyli.
In No. 4719 of the U. S. National Museum there is a series
of 25 vertebre, together with ribs and the supports of the
pelvic fins. Of these, apparently 10 belong to the caudal
region, possibly only 9. Many ribs are present and several
of them in their natural relation with the vertebre. They
are strongly developed, broad, and with much broadened
heads. They resemble those of the common carp. They have
been attached to distinct parapophyses. In some cases the
latter have been freed by maceration and disturbance, and
falling out have left long and deep cavities along the sides
of the vertebrae. The hemal arches of the tail region are
apparently consolidated with their corresponding vertebral
centra.
The supports of the pelvic fins are preserved and a portion
of one fin (Fig. 72). It seems probable that the fin supports
88 Bulletin American Museum of Natural History. [Vol. XIX,
have not been removed from their natural position. If so
the base of the fin is placed between the 5th and 6th verte-
bre in front of the first caudal. The fin itself is overlaid
with fragments of ribs, so that the number of
its rays cannot be accurately determined, but
there were at least eight of them, the most
anterior one being apparently rudimentary.
At the upper ends of the 4th and 5th caudal
vertebre are two bones which resemble inter-
/ neurals with enlarged upper ends. They may
represent the supports of the dorsal fin. They
are rather weak and may be the more pos-
terior ones of the series. In the region of the
SIN
Fig. 72. Enipo ne-
paholica Cope. No.
Popioy, | AOR GSK niche
Mus. X 4. Supports
of ventral fins.
slender bones which may have been the sup-
ports of the anal fin.
No. 2032 of the American Museum of Natural History
furnishes most of the tail fin (Pl. I, Fig. 4). The principal
rays are large and coarsely segmented, but distally the rays
divide into extremely fine filaments. The lobes of this fin
were probably about 175 mm. in length.
The following fishes, as well as Spaniodon simus, described
on page 47 were, as we learn from Prof. Cope (Bull. U. S.
Geol. and Geog. Surv. Terrs., IV, 1878, p. 66), collected by
Dr. F. V. Hayden in the ‘‘ Niobrara Cretaceous of Dakota.”
No more accurate information has been afforded us regarding
the locality where these specimens were found; but on several
of the blocks of soft limestone, on which these fishes are pre-
served, some person has written in lead pencil the words
“Yankton, Neb.’’ From this label we may be quite sure that
the specimens were found in the region of Yankton, South
Dakota. We know likewise that the Niobrara deposits are
abundantly developed in that region.
These fishes are of great interest from the fact that they
belong to genera found in Upper Cretaceous deposits at Mount
Lebanon, in Syria, or to genera very closely related to those
of the latter region. It is greatly to be desired that further
search shall be made in the country about Yankton for more
three or four most anterior hemal arches are ~
1903. | fay, North American Cretaceous Fishes, 89
satisfactory specimens of those described by Cope, and for
other species which may be yet unknown. That fishes are
abundant in the soft limestone whence Dr. Hayden obtained
his specimens, is shown by Plate V, which reveals three types
on one side of a block, while a fourth type is found on the
other side, besides a part of a large undescribed fish.
The figures of the species furnished on the plates are of the
natural size and may be of some value in identifying other
specimens, but they are difficult subjects to illustrate.
DERCETID.
Triznaspis Cope.
This genus was established by Cope in the ‘Bulletin of
the U. S. Geological and Geographical Survey of the Terri-
tories,” Volume IV, 1878, page 67. The type is T. virgulatus
Cope. Dr. A. Smith Woodward unites the genus with Lepto-
trachelus, but the present writer is inclined, for the present,
to regard it as distinct on account of the backward position
of the ventral fins and the relatively short head. The type of
the genus Leptotrachelus, L. armatus, possesses a dorsal fin of
apparently about 12 rays, with the ventrals beneath the front
of this dorsal. In Tri@naspis the dorsal is still shorter, not
having beyond io rays, and the ventrals are placed a dis-
tance behind the origin of the dorsal equal to one-third of
the distance from the head to the dorsal.
Triznaspis virgulatus Cope.
PEATE VA etGe xe
Trienas pis virgulatus CoprE (E. D.), Bull. U. S. Geol. and Geog. Surv.
Terrs. IV, 1878, p. 67. — Hay (O. P.), Bibliog. and Cat. Foss.
Wert.sN- A. toe2, p. 307.
Leptotrachelus virgulatus Woopwarp (A. S.), Cat. Foss. Fishes, IV,
Igol, p. 187.
Of this species Professor Cope possessed, so far as can be
determined, only the specimen here figured (Pl. V, Fig. 1)
and a fragment of another. The present number of the type
90 Bulletin American Museum of Naturai History. {Vol. XIX,
is 2516. The describer concluded that the anal fin was absent
in the species, but the present writer finds no warrant in the
specimens for making any statements regarding this fin. The
length of the head, including the opercular apparatus, is con-
tained in the distance from the end of the operculum to the
beginning of the dorsal fin something over one and one-half
times. The head has not been prolonged into a beak such as
we find in the species of Leptotrachelus.
Leptotrachelus longipinnis Cope.
PuaTeE IV, Fic. 3, and PLaTE V, Fic. 4.
Leptotrachelus longipinnis CopE (E. D.), Bull. U. S. Geol. and Geog.
Surv. Terrs. IV, 1878, p. 68. — WiL.isToN (S. W.), Kansas Univ.
Quart., VIII, 1899, p. 115; Univ. Geol. Surv. Kansas, VI, 1900,
p- 382. — Woopwarbp (A. S.), Cat. Foss. Fishes, IV, 1901, p. 187.
—Hay (O. P.), Bibliog. and Cat. Foss. Vert. N. A. 1902, p. 397.
Of this species Cope possessed two specimens. Of these
his ‘‘No. 1”? bears the Museum’s catalogue number 2521 (PI.
V, Fig. 4) while his ““No. 2” has the number 2520 (PL. IV,
Fig. 3). The former displays indistinctly the head, with its
long beak, the neck consisting of much elongated vertebre,
the rays of the dorsal fin, and the ventral fins. Under a lens
traces of the pectoral fin may be found close to the head.
The other specimen shows more distinctly the dorsal and
ventral fins, and some ribs, triradiate scales, and longitudinally
directed hair-like bones in front of the dorsal. In No. 2521
a considerable part of the bones of the beak have been flaked
away, leaving only their impression on the matrix. The
letters su indicate approximately the end of this beak. PI.
V, Fig. 4 shows the vertebre of the neck forming a curve
at the left of the head. Above and at the left of this curve
is a blotch formed by the rays of another fish.
Besides the American species, six others may be recognized,
as follows: L. armatus Marck and L. sagittatus Marck, from
the Upper Cretaceous of Westphalia, L. triqueter Pictet, L.
gracilis Davis, L. hakelensis Pictet and Humbert, all from the
1903.| Hay, North American Cretaceous Fishes, gl
Upper Cretaceous of Mount Lebanon, and L. elongatus
(Agassiz), from the Senonian and Turonian of England.
It seems not unlikely that of the species of Leptotrachelus
those having a long dorsal fin, L. triqueter, L. longipinnts,
and L. gracilis, will have to be separated as a distinct
genus, while L. armatus and L. hakelensis will be retained in
Leptotrachelus.
MYCTOPHIDZA.
Myctophide JorLAN and EVERMANN, Fishes of North and Middle
America, 1896, p. 550.
Scopelide of most authors.
Leptosomus WVarck.
To this genus the present writer assigns the species from
the Niobrara of South Dakota, which Cope placed in the
genus Sardinius. The latter genus has the scales pectinated,
the pectoral fins with about 18 rays, the anal with about 20
rays, and about 45 vertebre. The American species agree
with Leptosomus in having, so far as can be determined,
cycloid scales, narrow pectorals, a short anal, and about 30
vertebre. Cope regarded Leptosomus as a synonym of Sar-
dinius, but Woodward properly separates it. The author last
named recognizes four species of this genus, L. guestphalicus
Marck and L. elongatus Marck, both from the Upper Cretaceous
of Westphalia, and L. macrourus Pictet and Humbert, and
L. minimus (Agassiz), both from the Upper Cretaceous of
Mount Lebanon.
Leptosomus nasutulus (Cope).
PLATE IV, FiGs. 4 AND 5.
Sardinius nasutulus Corr (E. D.), Bull. U. S. Geol. and Geog. Surv.
Terrs. IV, 1878, p. 70. — Woopwarp (A. S.), Cat. Foss. Fishes,
IV, 1901, p. 248. — Hay (O. P.), Bibliog. and Cat. Foss. Vert.
NAS G02;, p- 391.
Of this species there are in this Museum three specimens
as follows: The type No. 2512 (PI. IV, Fig. 4), No. 2513
(Pi IVe tie. 5), and No, o523.
92 Bulletin American Museum of Natural History. |Vol. XIX,
Most of Cope’s measurements are taken from the type, but
the distance from the snout to the base of the ventral fins is
taken from No. 2513. This was necessary because in the
type the ventrals have been washed forward to beneath the
lower jaw, as may be seen from the figure.
On the block containing No. 2523 are scratched in pencil
the words ‘‘ Yankton, Neb.”’ The other specimens are with-
out doubt from the same locality.
Leptosomus lineatus (Cope).
PLATE IV. Fic. 6 AND PuaTE V. Fie. 2.
Sardinius lineatus Cope (E. D.), Bull. U. S. Geol. and Geog. Surv.
Terrs. IV., 1878, p. 71. — Woopwarp (A. §S.), Cat. Foss. Fishes,
IV, 1901, p. 248. — Hay (O. P.), Bibliog. and Cat. Foss. Vert.
Nw As 1902, p. 30x.
This species was based on two specimens, No. 2538 (PI.
IV, Fig. 6) and No. 2511 (Pl. V, Fig. 2). The former specimen
displays the body as far back as the rear of the dorsal fin; the
latter specimen shows the hinder half far enough forward to
show the tips of the ventral fins; but neither specimen shows
both the dorsal and the ventrals. Hence, the exact relation
of these fins to each other can not be determined.
The block on which No. 2538 is preserved, and which also
bears Spaniodon simus, is marked ‘‘ Yankton, Neb.”’
Leptosomus percrassus (Cope).
PLATE V, PIG. 3;
Sardinius percrassus Core (E. D.), Bull. U. S. Geol. and Geog. Surv.
Terrs. IV, 1878, p. 72. — Woopwarp (A. S.), Cat. Foss. Fishes,
IV, 1901, p. 248. — Hay (O. P.), Bibliog. and Cat. Foss. Vert.
INGZAL 1 902.5p..20n.
This species is represented by a single specimen, No. 2510
(Pl. V, Fig. 3). It differs from the other species in having
a much deeper body. Although there is no record either on
the block bearing this species, or in Professor Cope’s descrip-
tion, there can be do doubt that it came from the same
1903. | flay, North American Cretaceous Fishes, 93
locality and deposit as the other species here recorded from
South Dakota.
Sardinius? imbellis sp. nov.
PLATE IV, Fic. g.
The block of soft limestone which bears the type of Cope’s
Sardintus nasutulus presents also parts of two other small
fishes which appear to be undescribed. One of these, No.
2550, lacks the head and the whole of the body above the
vertebral column, except a portion of the upper lobe of the
caudal fin. The other specimen, No. 2549, presents the body
from the front of the dorsal fin to the extremity of the caudal.
Perhaps it would be wise to refrain from describing these
Specimens, as Cope refrained. Nevertheless, they appear to
differ from any species yet described and to furnish characters
which will make it possible to recognize other specimens when
they shall be found. So far as can be determined from the
remains at hand, the species belongs to the Myctophide, and
it stands nearer to Sardinius cordiert than to any other related
form. From Sardinius it appears to differ in having fewer
vertebre, fewer rays in most of the fins, in the position of
the dorsal, and in the character of the scales. It appears
safer, however, to await the finding of additional and better
materials before proposing a new generic name. The longi-
tudinally divided specimen, No. 2550 (Pl. IV, Fig. 9), is taken
as the type.
From the caudal fin to the insertion of the pectoral 36
vertebre are counted. The whole number probably has been
about 40. Of these, 15 appear to have belonged to the caudal
region. The ribs are slender. The pectoral finis broad and
consists of 15 rays; but it is rather short and lacks somewhat
of reaching the ventrals. The latter fins are pressed down,
the one on the other, so that the exact number of rays cannot
be determined. Six may be counted. These fins are placed
nearer to the pectorals than to the anal. The dorsal is miss-
ing in the type. The anal has a long basis and consists of
I5 Or 16 rays. The caudal is deeply forked. No.evidences
94 Lulletin American Museum of Natural History. [Vol. XIX,
appear of any pectination of the scales. The body has been
rather deep.
The second specimen presents an anal fin of 16 rays. In
front of it is the dorsal, which appears to be supported by
12 interneurals. The front of the dorsal begins above the
tenth vertebra in front of the origin of the anal. Three or
four neural arches and two or three hzemal arches at the base
of the caudal are expanded somewhat. Besides the slender
neural arches and ribs, there are numerous fine intermuscular
bones. No part of the ventrals appears in this specimen.
Assuming both fishes to belong to the same species, it is evi-
dent that the dorsal fin is placed in the interval between the
ventral and the anal fins. The whole length of the type has
been close to 65 mm.
Niobrara Cretaceous, region of Yankton, South Dakota.
Rhinellus A gassiz.
Dr. A. .S. Woodward has properly, as it appears to the
writer, reduced Cope’s genus /chthyotringa to a synonym of
Rhinellus Agassiz. The scales along the lateral line of the
American species also may have been somewhat thickened,
and the dorsal fin has about the same number of rays as in
R. furcatus, the type of the genus.
Rhinellus tenuirostris (Cope).
PuiaTe IV, Fics. 7 AnD 8.
Ichthyotringa tenutrostris Corr (E. D.), Bull. U. S. Geol. and Geog.
Surv. Terrs. IV, 1878, p. 69. — Hay (O. P.), Bibliog. and Cat.
Poss. Vert. N..A.. 1902, p. 207-
Rhinellus tenutirostris Woopwarp (A. S.), Cat. Foss. Fishes, IV, rgor,
Pp. 260.
This species is based on two specimens, No. 2514, the type
(Pl, IV, Fig. 7), and No. 2515 (Pl. 1V, Pigss). Tue tomer
shows the head, with its long beak, and the body above the
vertebral column as far backward as the rear of the dorsal fin.
The latter, a small fish, displays the anterior portion of the
body from below. Both pectoral and both ventral fins are
presented.
1903 | Hay, North American Cretaceous Fishes, 95
The block on which this specimen is found is marked ‘‘ Yank-
ton, Neb.”’ Ofthe other valid species of this genus R. jurcatus
Agassiz is found in the Upper Cretaceous of Mount Lebanon
and Westphalia; R. ferox Davis and R. damoni Davis, in the
Upper Cretaceous of Mount Lebanon.
Explanation of the abbreviations employed in the figures
and on the plates to indicate the names of the bones and
other structures.
lant = anal fin, os. = orbitosphenoid,
als. = alisphenoid, pa. = parietal,
ang. = angular, pal. = palatine,
Zine Ss gail ihe wayyy par. = parasphenoid,
art. = articular, p- cor. = precoracoid,
art.s. = articular surface, pet. = pectoral fin,
bas. = baseosts, p.f. = pectoral fin,
b. br. = basibranchial, pmx. = premaxilla,
b. oc. = basioccipital, p. op. = preoperculum,
el. = cleithrum, pr.f. = prefrontal,
cor. = coracoid, pro. = prodtic,
den. = dentary, p. sp. = presphenoid,
det. = dorsal fin, pt.f. = postfrontal,
d.r. = dorsal fin rays, pt.o. = pterotic;
ec. pt. = ectopterygoid, pt. or. = postorbital,
en. pt. = entopterygoid, SC: = scapula,
Ep.0, | = epiotic, smx. = supramaxilla,
eth. = ethmoid, sn. = tip of snout,
ex. oc. = exoccipital, S. oc. = supraoccipital,
fr. = frontal, S.op. = suboperculum,
gl.h. = glossohyal, s.or. = suborbital,
ths 1a, = interhemals, sq. = squamosal,
i.n. = interneurals su. cl. = supracleithrum,
i. op. = interoperculum, vert. = vertebra,
m. pt. = metapterygoid, v.f. = ventral fin,
ra0b:e = maxilla, vom. = vomer,
na. = nasal, vom. t. = vomerine tooth,
op. = operculum, v.r. = ventral fin rays.
op. 0. = opisthotic,
ee eee, ae
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EXPLANATION OF PLATE I.
Fig. 1.—Protosphyrena aa (Cope). Page g. Pectoral f
x4. No. rgor. '
Fig. 2.—Protosphyrena tenuis Loomis. Page 15. Pectoral fin. x. i
No. 205. ral
Fig. 3.—Protosphyrena tenuis Loomis. Page 16. Distal end of Bee >
toral fin. X 4. No. 1620. :
Fig. 4.—Empo nepaholica Cope. Page 88. Part of caudal aoe x4
No. 2032.
wf
Butietin A. M. N. H.
Vou. XIX, Prats 1.
PROTOSPHYR&NA AND Empo.
C. L. Wright, Sc. & Imp New York
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EXPLANATION OF PLATE III.
Pachyrhizodus caninus Cope. Page 63.
Fig. 1.—Caudal fin and vertebre. X14. No. r1goo.
Fig. 2.—One lobe of caudal fin. X14. No. 1658.
Butietin A. M.N. H.
PACHYRHIZODUS.
C. L. Wright, Sc. & Imp.. New York.
VoL, XIX, Prats III.
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CreTaceous Fisnes
EXPLANATION OF PLATE IV.
Fig. 1.—Spaniodon simus Cope. Page 47. Nearly complete fish.
Type. X44. No. 2508. a.f., anal fin; d. f., dorsal fin; p. f., pectoral
fin; v. f., ventral fin.
Fig. 2.—Spaniodon simus Cope. Page 47. Anterior half of fish.
Paratype. X44. No. 2509. den., dentary; p.f., pectoral fin; pmx.,
premaxilla.
Fig. 3.—Leptotrachelus longipinnis Cope. Page go. Part of trunk.
Cotype. x1. No: e520. d.f., dorsal fin; v.7., ventral Gn.
Fig. 4.—Leptosomus nasutulus (Cope). Page g1. Nearly complete
fish. Type. +. No. e512...a, 7., anal tm? d.fdorsal ins wy.
ventral fin (displaced).
Fig. 5.—Leptosomus nasutulus (Cope). Page g1. Nearly complete
fish. <4. No. 2513. d.j., dorsal fia; ». j., pectoral fin; v.}., ven
tral fin.
Fig. 6.—Leptosomus lineatus (Cope). Page 92. Fish with tail miss-
ing. Cotype. X44}. No. 2538. d. f., dorsal fin; v. f., ventral fin.
Fig. 7—Rhinellus tenuirostris Cope. Page 94. Head and part of
trink. Cotype. XX}. No. 2514. d.j., dorsal fin.
Fig. 8—RKhinellus tenuirostris Cope. Page 94. Head and part of
trunk. Cotype. X 4. No. 2515. ».f., pectoral fin; v. f., ventral fin.
Fig. 9.—Sardinius ? imbellis Hay. Page 93. Lower half of fish.
Type. Xt} No. ess0. a. f., anal fm; p. 7. pectoral fin; wv. 7;
ventral fin.
BuLLetin A. M. N. H. VoL. XIX, PL. IV.
CRETACEOUS FISHES.
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EXPLANATION OF PLATE V.
Fig. 1.—Trienaspis virgulatus Cope. Page 89. Head and part of
trunk. Type. xX +. No, 2516. d.f., dorsal fin; p. f., pectoral fin;
v. f., ventral fin.
Fig. 2.—Leptosomus lineatus (Cope). Page g2. Hinder half of fish.
Cotype. Xi. No. 2511. a. f., anal fin; v. j., ventral fin.
Fig. 3.—Leptosomus percrassus (Cope). Page g2. Complete fish.
Type. xX}. No. 2510. a.jf., anal fin; d.f., dorsal fin; p. f., pectoral
fin; v. f., ventral fin.
Fig. 4.—Leptotrachelus longipinnis Cope. Page 90. Head, neck, and
part of trunk. Cotype. 4. No. 2551. d. j., dorsal fin; 9. 7.
pectoral fin; sv., snout; v. j., ventral fin.
Lying against the bend in the neck are some remains of another fish.
Bu._itetTin A. M.N H. Vor. XIX, Plate V.
CRETACEOUS FISIIES
C. L. Wright, Sc. & Imp., New York.
he Fauna of the Titanotherium B
at Pipestone Springs, Montana.
: 7 By W.D. Marruew.
AUTHOR’S EDITION, extracted from BULLETIN 2
Amevican Museum of Hatural History,
Vou. XIX, Arricte VI, pp. 197-226.
New York, May 9, 1903.
The tnickerbocker Press, Rew Wp
ork
Ae
Article VI.— THE FAUNA OF THE TITANOTHERIUM
BEDS AT PIPESTONE SPRINGS, MONTANA.
By W. D. MatrtHew.
The American Museum Expedition of 1902 in western
Montana had for object to make a further search in the Ter-
tiary deposits of that region, where Mr. Earl Douglas has
recently discovered many new and interesting fossil mam-
mals. Inthe White River formation near Pipestone Springs,
Mr. Douglas had found a very interesting micro-fauna, and
our collections at the same locality, which Professor Osborn
has kindly turned over to me for study and description,
enable us considerably to extend the list. 1 am indebted to
Mr. Douglas for the opportunity to examine the type speci-
mens of his various species, as well as for the information
concerning localities, etc., contained in the stratigraphic part
of his very excellent memoir recently published on the White
- River of Montana.
_ The majority of the species are small or minute forms, not
found in the Titanotherium Beds of South Dakota or Colo-
rado, where the scanty fauna is almost entirely of large ani-
mals, — Titanotheres, Elotheres, and Rhinoceroses. A few
small species have been described from Swift Current Creek,
o - Canada, based on very fragmentary materials. The Pipe-
___ stone Springs fauna is therefore of much interest, as it illus-
trates the direct precursors of the numerous small species of
the Oreodon Beds. In the species from the three successive
_ stages of the White River we have the most favorable oppor-
tunities for study of the details of evolutionary progress in a
given race that are presented among fossil vertebrata; for the
_ materials are abundant and complete, the succession is un-
- questionable, and the character of the beds, and hence the
a
local conditions of deposition, very uniform, so that we get
_ the same facies of the three faunas. It is doubtful how far,.
if at all, the Eocene deposits of the Rocky Mountain divide:
and foothills contain the same facies of their respective
197
a, , . > Fee ye “3 > a
art Wane epee, ee Eats
eee I ee
Ms
m3
.
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gf Udicys “ele ‘es
198 Bulletin American Museum of Natural History. [Vol. XIX,
faunas as do the Oligocene deposits of the plains. They con-
tain an important aquatic contingent, fish, crocodiles, and
water-turtles being comparatively abundant. In the White
River fauna .all these are absent, except in the sandstone
lenses, while a large element of it is apparently adapted to
open grassy plains; this is not found in the Eocene faunas.
But in the three zones of the White River a great part of
their respective faunas appears to be in direct and exact gen-
etic succession. We can therefore measure the amount and
direction of change during the Oligocene epoch in many series.
The amount of evolution as thus measured appears small,
but its direction somewhat constant. The species of the
Titanotherium Beds are all distinct from their successors in
the Oreodon Beds, but the difference is uniformly small.
Between the Oreodon and Leptauchenia faunas the difference
is often greater but less uniform, so far as present data go.
Some genera run through the three horizons (e. g., Cynodictis,
Paleolagus, Mesohippus, Cenopus, Leptomeryx). Others have
been found only in the two lower zones or in the two upper
zones, while many are as yet known from one horizon only.
Stratigraphy. — Mr. Douglas refers all the Tertiary at this
locality to one stage, correlating it with the Titanotherium
Beds of South Dakota. We find, however, a lithologic dis-.
tinction between the higher beds exposed north of the rail-
road, which resemble the Oreodon Beds of South Dakota,
Colorado, and elsewhere, and the lower beds exposed south
of the railroad, which resemble rather the Titanotherium
Beds of some parts of South Dakota. Likewise on Thomp-
son’s Creek, not far from the Pipestone locality, we were able
to distinguish between the Oreodon Beds exposed near the
head of a small northerly branch of the creek, and the Titan-
otherium Beds exposed on the main western branch. At
* Mr. Douglas has recently discovered fish remains in strata which he refers to the
White River epoch, so-called, in the Madison valley in Montana. But these strata are
quite different in character from the beds in which White River mammals are found,
apparently lacustrine or fluviatile in origin, and a very thorough search on his part
failed to reveal any mammals in them except a skull of the beaver Steneofiber. Ido not
understand that he considers them as of the same formation or origin as the mammal
beds, but merely as of equal age. The discovery of fish in them, therefore, does not at
all invalidate the fluviatile-eolian hypothesis of origin of the White River formation
maintained by Hatcher and myself. The same explanation probably applies to other
reported occurrences of fish in the White River.
1903.| Matthew, Fauna of Titanotherium Beds of Montana. 199
both localities the lithologic distinctions are confirmed by the
fossils found.
The Titanotherium Beds are soft, easily weathering, banded
clays, often sandy, crumbling to the usual weathered-clay
surface, varying in color from dark reddish brown through
buff to an almost greenish white. Cross-bedding is seen in
the sandier layers. They are not unlike the Titanotherium
Beds in South Dakota, but run to deeper and more contrasted
coloring. At Pipestone Springs they dip quite steeply toward
the north, lying up against the ancient crystalline rocks on
the south side of the creek. We found in them the following
fauna:
Marsuptalia.
Peratherium titanelix, sp. nov. Alhed to Peratheria of Middle Eo-
cene and Oligocene.
Insectivora.
Apternodus medievus, g. et sp.nov. Allied to ? Centracodon of Middle
Eocene.
Micropternodus borealis, ‘ *« Allied to ? Centetodon of Middle
Eocene.
Ictops acutidens Douglas More primitive than any Middle
‘“ thomsoni, sp. nov. Oligocene Leptictid.
Creodonta.
Pseudopterodon minutus (Douglas) Intermediate between the Oligo-
? Hyzenodon sp. cene Hyenodon and Middle
Eocene Sinopa.
Carnivora Fissipedia.
Bunelurus infelix, sp nov.
Cynodictis paterculus, sp. nov. Somewhat more primitive than
Middle Oligocene species.
Rodentia.
Ischyromys veterior, sp. nov. Somewhat more primitive than
Middle Oligocene species.
Cylindrodon fontis Douglas.
Sciurus vetustus, sp. nov.
Gymnoptychus minor (Douglas).
- minimus, sp. nov.
200 Bulletin American Museum of Natural History. |Vol. XIX,
Paleolagus temnodon Douglas. More primitive than Middle Oli-
+ brachyodon, sp. nov. gocene species.
Pertssodactyla.
Mesohippus westoni Cope. More primitive than Middle Oli-
gocene species.
Hyracodon sp.
? Ceenopus sp.
Titanotherium sp.
Artiodactyla.
Stibarus montanus sp. nov.
Bathygenys alpha Douglas.
Limnenetes sp. div.
Leptomeryx mammifer Cope.
; ? esulcatus Cope.
Leptotragulus profectus sp. nov. Advanced species of an Eocene
genus.
Lizards and Tortoises, sp. div.
Two species are reported by Mr. Douglas of which we
obtained no further evidence: Sciurus jefjersoni Douglas,
? Agriocherus maximus Douglas.
The Oreodon Beds are buff clays, somewhat harder than
the Titanotherium Beds, finer, not sandy, more calcareous,
and not unlike the Oreodon Beds of Dakota and Colorado.
They were very barren so far as we could discover, and the
only determinable fossils found at Pipestone Springs were:
Paleolagus haydeni, Eumys elegans, Mesohippus bairdit,
? Poébrotherium.
These are all characteristic species of the Oreodon Beds in
South Dakota, Colorado, etc.
The fauna from the Titanotherium Beds is a quite remark-
able one. Not a single species is identical with those of the
Oreodon horizon; all are either new, or have been described
by Douglas from the same locality, or by Cope from the same
horizon at Swift Current Creek. The majority of the species,
however, belong to genera of the Oreodon Beds, and these,
though fairly distinct, are not widely divergent from their
successors. We find that the Pipestone Beds are much
nearer to the Oreodon horizon than to the upper Uinta or
Diplacodon Beds. Fourteen genera are in common with the
1903.| Matthew, Fauna of Titanotherium Beds of Montana, 201
later horizon, while there is but one Uinta genus (Leptotrag-
ulus) and that represented by a rather divergent species.
This contrast is partly explained by the fact that the known
White River fauna is a very large one, while that from the
Uinta is comparatively small; partly also by considerations
of geographical distribution of the Oligocene mammals and
by different conditions of deposition in the Uinta and White
River beds. But, making allowance for all these, there seems
still to be a considerable gap between the Diplacodon and
Titanotherium faunas, while the latter is much closer to the
Oreodon fauna. It shows some marked differences, however:
(1) There are two new insectivore genera of the primitive
section (Zalambdodonta) of the order, which has hitherto
been practically unknown in a fossil state, unless the Eocene
species reported by Professor Marsh shall prove to belong to it.
(2) All the rodents are sciuromorphs! or lagomorphs.
Myomorpha, more abundant than sciuromorpha in the Oreo-
don Beds, have not yet appeared. They are unknown in the
Eocene, except Protoptychus, a form of doubtful affinities.
(3) The only Creodont from the Oreodon Beds is the highly
‘specialized Hyenodon. At Pipestone Creek we have a more
primitive type, intermediate between Hy@nodon and Sinopa.
At Swift Current Creek occurs Hemipsalodon (?=Pterodon),
also less specialized than Hyenodon. (Hyenodon itself occurs
also in the Titanotherium Beds.)
(4) Oreodon is not found, and two or three more primitive
genera (Bathygenys, Limnenetes, ? Agriocherus) take its place.
(5) Hypertragulus, common in the Middle and Upper Oligo-
cene, is not found, while Leptomeryx of the Lower and Middle
Oligocene is abundant and large.
(6) In place of Poébrotherium, the camel of the Oreodon
Beds, is found a pices yocont form, apparently the Eocene
genus Leptotragulus..
From the above facts we would infer that the Pipestone
Beds are at the base of the Oligocene, but above the Eocene,
accepting Osborn’s correlation of the White River formation
1 If Gymnoptychus be considered a sciuromorph, as it was by Professor Cope and is
by Dr. Hay
202 Bulletin American Museum of Natural History. |Vol. X1X,
with the Oligocene, and of the Uinta with the Upper Eocene.
They are probably of approximately the same age as the
White River beds of Swift Current Creek, Canada, with which
they have three species in common, probably a fourth (“‘ Pale-
olagus turgidus” from Swift Current Creek probably is P.
brachyodon),— a fair proportion out of so limited a fauna.
DESCRIPTIONS OF SPECIES.
MARSUPIALIA.
Peratherium titanelix, spec. nov.
Type, No. 9603, a lower law with p,; and m,_, and alveoli of the
remaining teeth except incisors.
This species is about the size of P. hunti of
the Oreodon Beds, and resembles it in the rather
short premolar region, the premolars and canine
small, close together without any diastemata.
The molars are similar to those of typical Pera-
theria from the Phosphorites, but the premolars
are very distinct, crowded, and reduced antero-
posteriorly, the cusps recurved instead cf sym-
Fig. 1. Peratherium metrical asin Peratheria from the Phosphorites,
titanelix. ‘Type speci- ; :
men, twice natural siz. and in P. fugax of Cope. The anterior part of
<i. outer, d*, inner view the jaw is rather short and deep and the canine
of teeth.
directed more upward than in P. fugax.
Measurements.
Petpet iis. stun a fa, shou. cose eres ee 7.3 mm.
entoth trp. aciaiese ve potk.c ok tts goes ate eee 4:2
Depth jaw wiidera: Sb Aosta eee 1.9
INSECTIVORA.
Apternodus medizvus, gen. et spec. nov.
Type, No. 9601, posterior half of a lower jaw with two complete
molars and the root of another.
Molars composed of high trigonid and minute basal talonid. Proto-
conid high, sharp, and triangular, paraconid and metaconid subordi-
nate. Dentition probably i; ¢, p,m; .
Talonid a small sharp cusp on m,;; On m, it is a minute postero-
internal basal cusplet. The third molar a little smaller than the
second; both are two-rooted, the anterior root wider transversely.
1903.| Matthew, Fauna of Titanotherium Beds of Montana. 203
The cusps are all high, sharp, trigonal in cross-section, the whole tooth
subtriangular with transverse and longitudinal diameters about equal
and vertical diameter consider-
ably exceeding either. The
heel is much smaller than
in Centetide, the protoconid
higher and larger in proportion
than in any other Insectivore
or Chiropter.
i es PY mt m2
m3
No. 9608, a lower jaw
without teeth, broken off
in front at about the same
point as the types, but
with the condyle complete,
appears to belong to the
same species. The condyle
iewviclelys cxpantled thams= | cyice aatueal ciee, the cuslines of onal jam
versely onvalevel with the (2a ce Co" oN Eee as
bases of the molars. The
angle in the type specimen is extended into a rather long
and stout flattened process with a sharp medial ridge on the
internal side. The coronoid is broken off in both specimens,
but was evidently high, stout, placed mostly external to the
toothrow, and directed upwards instead of backwards.
No. 9607, the anterior part of a lower jaw, with one pre-
molar preserved and the roots of other teeth, is provisionally
referred to this species. The premolar is either the third or
fourth; it is stout, two-rooted, composed of a round-conical
protocone and small postero-internal basal cusp. The an-
terior root is wider than the posterior. The premolar in
front of this was apparently similar but smaller, and was pre-
ceded by a small one-rooted premolar, and this by a larger
tooth, probably a canine. Behind the premolar are roots
of two teeth, one of which was probably m,, the other cer-
tainly a molar, judging from comparison of the corresponding
parts of the jaw in the three specimens.
The dentition is then probably c, p, m,, but may bec, p,m .
The weight of the anterior part of the jaw and doubtful
wie
204 Bulletin American Museum of Natural History. [Vol. XIX,
indications of a large alveolus lead to the suspicion that one
or more of the incisors was enlarged.
No. 9612, part of a lower jaw with the roots of the last two
molars, is also referred here.
This remarkable little jaw is quite unlike any described
species of Insectivore or Chiropter, except, perhaps, Marsh’s
Centracodon. So far as anything can be determined from his
brief description, the last molar of Centracodon is like the
second molar of Apternodus. Centracodon has four premolars.
Although the short, deep jaw would appear to ally it rather
with the Chiroptera, yet as the tooth without talonid is quite
unknown in this order, while it does characterize a section of
the Insectivora, I leave it provisionally in the latter group,
without attempting to assign it to any especial family. Its
nearest relatives are most probably the little Eocene Insec-
tivora from the Bridger Basin, described by Professor Marsh
in 1872. With the probable exception of these Eocene types
and of a single South American species, no fossil Insectivores
of the Zalambdodont division have hitherto been described,
although according to the Tritubercular theory this, as the
more primitive section, should have been more abundant
in ancient times.
Measurements.
ee eee irc Teen ere Ee heen yo Od oo Sod 4S 3.9 mm
Ma TONSIE it aie tye sank cee 6 See ee ee 2FO
PP RETALIS VS 2,3. le ot tee ae aec on ae cE or ee Oe 2.0
“height. om e205 ee hae eee eee 3.7
ME LONGI SS vido cid ktiaS Ae sp A eee 2.0
" WET ELNISI o35 <45/chctletie accep pena tear ese) ee ee 1.8
i Weigit. 5 oie. 20s eee ta hes eee ae ee 2.9
Depth of jaw. 4 cic ecaer ato oe ee ee Bas
Micropternodus borealis, gen. et spec. nov.
Type, No. 9602, a lower jaw with p,;-m, and alveoli of the anterior
teeth.
Dentition 3.133. Molars somewhat like those of Centetes in compo-
sition, with high trigonid and small, low talonid. Trigonid very wide
transversely with pr‘ considerably overtopping pat andme’ Talonid
with sharp posterior margin and low median ridge. Molars and
especially premolars, short, high,and recurved; p, sub-molariform, with
1903.] Matthew, Fauna of Titanotherium Beds of Montana, 205
small anterior and internal trigonid cusps and strong basal heel. P,'
much smaller and simpler, with small heel and no other accessory
cusps. P, is small and one-rooted, canine
small, incisors small, subequal. No dia-
stemata except a slight one behind pz,
Jaw rather deep in front. Second molar
slightly larger than the first, third much
smaller.
Like the preceding genus, this
must be placed among the Zalamb-
dodonta, with no very near relatives igs ai Mace abrer nate te rage
among living species, although it is _]P¢ specimen, twice natural size
not so strikingly different from mod-
ern types. Its nearest allies are also quite probably some of
the very inadequately described Insectivora from the Bridger
Basin, but neither it nor Apternodus can be considered as
possibly congeneric with any of the Bridger species, if Marsh’s
descriptions are correct.
Measurements.
Lower jaw, m, to incisive alveoli:..............-. 12.4 mm.
POETS POSEN Pe Tg eater oh swayed 0s nce aay aah wa 8.4
IO wer ma Glars Ut sacs ass cues nachsadal dies aus Sh conte 6.4
Lower molar, m, longitudinal, at base........... 1.8
. " MORCKANS VELSChts ee cunt einai I.9
i Mie 2 “Rei sht @v Crow s.ac sie cats ee ey
Ictops acutidens Doweglas.
A fragmentary skull and jaws, with some limb-bones of one
individual, and the upper and lower molars of another, con-
firm and extend the characters of this species as indicated by
Douglas.
The distinctions from previously described species are:
Dimensions fifteen per cent. less than any of the Leptictide
from the Oreodon Beds. First upper premolar one-rooted,
two-rooted in I. dakotensis and bullatus and in Leptictis hay-
deni. Supra-temporal crests widely separated anteriorly and
convergent posteriorly, instead of close together and parallel
as in all the later species. Upper molars and p* more
206 Bulletin American Museum of Natural History. (Vol. XIX,
constricted between the inner and outer cusps than in any de-
scribed Leptictid; cusps somewhat higher and last molar less
reduced than in any later species.
No. 9604, a fragmentary skull and jaw, with humerus,
radius, two phalanges, and a caudal vertebra, exhibits most of
the permanent dentition well preserved.
Upper jaw. — Incisors not known, Canine of moderate size, com-
pressed, somewhat ridged externally, with no indication of the in-
cipient heel seen in
Paleictops. P* one-
rooted, smaller than
canine. P? two-root-
ed, compressed, with
small posterior basal
cusp or heel. P3 three-
rooted, with strong,
well-separated de,
higher but less sep-
arated iv, and minute
antero-external basal
cusp; the protocone
much overtopping the
other cusps. P+ mo-
lariform, but ir not
yet as high as pr, de
equalling pv in height,
Fig. 4. Jctofs acutidens. Teeth, twice natural size. A,
, ‘ 1
crown view of upper teeth ; 4, external view of upper and smal] hy, and strong
lower teeth. No. y6o4, the lower molars from No. 9605. protostyle. First and
second molars with pa
and me of equal size, hy better developed than in p4, a small protostyle
on m™. M3 with reduced me and rudimentary fy. Atl molars and
p* wide transversely, with some constriction between inner and outer
cusps. M3 smaller than m* and m2, but not so much reduced as in the
later species.
Lower jaw. — Dental series continuous without diastemata. Three
small incisors. Canine small, incisiform, somewhat larger than in-
cisors. P, one-rooted, p, two-rooted, p, two-rooted, compressed,
with anterior and posterior cusps and small heel. The lower molars
are not preserved in this specimen. In No. 9605 m, and m, are pre-
served; the trigonid is high, composed of two equal well-separated
cusps, talonid much lower, bearing three posterior cusps, external, in-
ternal, and postero-median respectively, well separated from the
trigonid but not from each other. M, is a little longer than m, but
1903.] Matthew, Fauna of Titanotherium Beds of Montana. 207
much narrower, especially the talonid, in which the hypoconulid is
situated more behind the hypo- and entoconid instead of nearly be-
tween them.
The skull is wider between the eyes than those from the
Oreodon Beds, the postorbital constriction less pronounced.
The temporal crests begin on the posterior third of the frontal
bones, eleven millimeters apart, and converge rapidly on the
posterior half of the parietals.
The humerus is disproportionately smaller and more curved
than in J. dakotensis, the deltoid crest is not so wide nor does
it extend so far down. The radius is likewise strongly curved,
its distal end bearing two ill-separated subquadrate facets for
scaphoid and lunar, the scaphoid facet the wider of the two.
Two rather long and slender phalanges are preserved, one
somewhat compressed laterally. The caudal vertebra asso-
ciated is from the middle part of the series and indicates a
large, long tail.
This species is in some, but not in all respects intermediate
between Pale@ictops and the Leptictide of the Oreodon Beds.
The last molar is more reduced than in P. bicuspis, less than
in any later Leptictid. The first premolar is one-rooted, as
in P. bicuspis, while in the later species it is two-rooted. On
the other hand, the molars are more compressed and more
constricted medially, and the size is smaller than in either the
Wind River species or those from the Oreodon Beds; and the
temporal crests are further apart than in the later species,
while in the earlier one they are united into a sagittal crest.
Ictops thomsoni, spec. nov.
Two upper jaws, Type, No. 9606, Cotype,
No. 9606a, indicate a species closely allied
to Ictops acutidens, but distinguished by
smaller size, more compressed teeth, and
other characters of less importance. The
metacone on all the molars is decidedly sons” Gcawar ew OE
smaller than the paracone; in I. acutidens SPbshela *t- Type
they are nearly, and in other Leptictide
quite, equalin sizeon mt!-?. The protocone on p4~m3 is more
208 Bulletin American Museum of Natural History. [Vol. XIX,
compressed antero-posteriorly, and the constriction between
it and the outer cusps is more marked than in I. acutidens.
The hypocone is smaller on mt and m’, absent on m3 and
p+. The trittocone of p4 is smaller than in I. acutidens.
All these distinctions are exaggerations of the differences
between I. acutidens and the Leptictide of the Oreodon Beds,
but none of them ally it to Paleictops, in which the molars
and premolars (except p') are fully as complicated as in the
species of the Oreodon horizon.
The species is named in honor of Mr. Albert Thomson of the
American Museum Expedition of 1902, who discovered the
type specimens of this as well as those of four other new
species described in this article.
Comparative Measurements.
in Ness o Gs 2
Bs |ees| ba | ds | 22 | os
Fe | 2oc| sa | Se | 2a ae
3° . ZA ima Qn
Se | 6 Selec eee eon eee
i BA a ue} ate Or =
Ki a B =
Upper molar-premolar series........ 20.8 23.5 24.
: true molars and fourth p re-
ma molar. . Soro Te tree eel ees 10.4 THE 7) TAT 12.
m longitudinal. Ber sis (avrwuahe Jesshe yay 2.7 2.6 2.3 3-4 3-4 3:
i. Te TERE EDS 5000055 b 020006 3.0 4.0 4.0 Bas 4.0 4.7
LS rant thoyatexheibichhstMls Smog omoGUe Les 2.1 ir{6) 1.9 2.4
* >; ebransyerses.ee cen cress bese 3.6 Bas 3-2 320) 4.9
Lower molar-premolar series........ E. 20.0
true InOlarss ty ere eeoe E. 10.1
= soa Jhoyaverqubtabharlls on Gg odanouoc 2.7
NyeLTANSVEISee een 2.1
UE “< height “vase ote 3.7
1 Measurements from Leidy’s figure.
CREODONTA.
? Pseudopterodon minutus (Douglas).
Hyenodon minutus Douglas.
No. 9623, the upper jaw of a small Creodont of the family
Hyenodontide is referred provisionally to Schlosser’s genus,
and may. be identified specifically with the lower molar on
which Mr. Douglas based his species ‘“‘ Hyenodon”’ minutus.
This upper jaw, however, cannot be referred to Hyenodon, as
1903.| Matthew, Fauna of Titanotherium Beds of Montana. 209
-—
the first molar shows a sharp antero-internal angle and small
antero-internal cusp. It is not very close to Schlosser’s type,
and if, as Scott believes, Pseudopterodon is
founded on milk teeth of Hy@nodon, then 9" 4
our species represents an undescribed genus, iS N
which may well stand ancestral to Hyenodon, i
being directly intermediate between that , Hise 6. Penge.
genus and Sznopa. euch res
The animal was a little smaller than
Cynohyenodon cayluxt Filhol. P3 is two-rooted, slightly
compressed, set a little transversely in the jaw, moderately
high, with small anterior basal cusp and heel. fP4 is three-
rooted, the internal root well separated, median, supporting
a strong internal buttress to the protocone and a basal
cingulum, but no defined cusp. The antero-external cusp is
of moderate size, the postero-external developed into a short
cutting blade. Mt! is three-rooted, the inner root anterior
and well separated, bearing a wide buttress ridge and a small
internal cusp, which is worn off in the specimen. Only the
front part of the tooth is preserved. M? was larger than m!
and similar to it, judging from the character and position
of the two anterior alveoli which indicate it on the specimen.
? Hyenodont, indet.
An upper premolar of singular character which I am unable
to refer to any described species of Carnivore or Creodont. It
has two roots, the posterior one broadened inwardly so as to
support a median internal buttress tothe protocone. The pro-
tocone is very high, its height exceeding the antero-posterior
length of the tooth; somewhat compressed posteriorly, with
a well-distinguished posterior cusp and small posterior cin-
gular cusp. Cingulum obsolete except at anterior and pos-
terior ends of tooth. Appears to be the third or, perhaps,
second upper premolar of some Hyznodont, but not of Hye-
nodon or Pterodon.
CARNIVORA (FissIPEDIA).
Cynodictis paterculus, spec. nov.
Two lower jaws and parts of others, Nos. 9616, 9619, repre-
[Agril, 1903.] 14
210 Bulletin American Museum of Natural History. (Vol. XIX,
sent this species. Of these I take No. 9616 as type. Com-
pared with a quite large series of specimens, including the
types of C. gregarius and C. lippincottianus, these specimens
show certain constant differences, chiefly in the construction
of m,. This tooth is proportionately larger and longer, the
heel larger and wider, and the proto- and metaconids are
raised above the paraconid, instead of being nearly on a level
with it, as they usually are in C. gregarius. The shear of m,
is somewhat more transverse, and m, is a little less reduced.
The size is that of C. gregarius..
The above characters are slight distinctions indeed, but
their constancy in the very considerable series of specimens
compared makes them valid specifically. In Procynodictis
vulpiceps of the Uinta, the shear is more transverse, but m, is
smaller and its heel more reduced than in C. gregarius.
Measurements.
| Type, No. Cotype, No.
9616 961
eas Tee eee Aas Sei Sa eS 35 mm.
A Te ee ae,§ sr oe el woe lees oe sie) eebobedl, |)" a0
MM tonew< oS: ah Gris 9
Sie si SW ETS. 22.) ye: cae Meee ica een 1) ass 4.2
“height of pra Sees eee NEA [ru 87h
iM, Joaatte eos: ees ee ete See Lb es ae)
Seq RATAS Vin Meee Sat es he gh A Oe ee B55 2.3
iM. fonerk Sse ack peony at Oe 3
6 REARS co: pee ce eid sheer ie ee | 2.5
Bunelurus infelix, spec. nov.
No. 9620, part of a lower jaw with p,_m, and the stump of
m,, represent this species, which is with difficulty distin-
guishable from B. lagophagus. The protocone of m, is more
rounded, the shear a little more transverse than in Cope’s
species. The fourth premolar appears to be stockier and
longer than in the type of B. lagophagus, but it is not fully
formed in the jaw in that specimen, so the comparison is
questionable; the protocone is stout and round, with a small
1903.| Matthew, Fauna of Titanotherium Beds of Montana, 211
postero-external cusp, and external basal cingulum rising to
a small heel behind. The second molar is a minute crownless
stump.
Measurements.
2) aap eee eam eran 152.05 he, al acy a(o6)tse Ne hayie Fe oh ees 10.2 mm.
IE LORE eee eRe ete cB hel os citcfnie a nyo, 3 nhc ata Sele #i elie Bak
RSA BREN GENT a's. cy & Geala't ova Reb, 5 ROIS ORE CO ol eae 1.9
WILE Ou Sri ts Cabins 3 Lies ter oe eer en a oe a ae 5-9
Mai ATNS Ve mp eae Se de cs SLU PSs. cnet a usisahe a esis 2143
DepthwolgaweGemes hai ts ete. ci ik 2 hee cits aye wales 6
A larger Musteline is indicated by part of a jaw, No. 9621,
with p, and the roots of the front teeth init. It is about the
size of Cynodictis paterculus, but the premolars are higher,
shorter; the heel smaller, anterior basal cusp absent; and
other details join to show that it is a Musteline.
RODENTIA.
ISCHYROMYID&.
Ischyromys veterior, spec. nov.
The anterior part of a skull and some forty jaws or parts
of jaws, upper and lower, represent this species. It is con-
siderably smaller than a series of specimens from South
Dakota, which agree well with Leidy’s types of I. typus, and
the teeth are narrower throughout with higher cusps. It is
much closer to the variety or separate species from Colorado,
I. cristatus (Cope, 1872), which Cope has referred to I. typus
and described and figured in ‘Tertiary Vertebrata.’ From
this species I find a constant distinction in the last molar,
which in the Montana jaws has always a narrow heel with the
last crest imperfect internally, while in all the Colorado speci-
mens the heel is as wide as the rest of the tooth, and the third
(last) crest perfectly developed.
In the upper teeth a corresponding difference is to be seen
in the last molar, and also the valley between the anterior
and posterior inner cusps is well marked on all the teeth, dis-
tinct nearly to the base of the enamel, while in the specimens
212 Bulletin American Museum of Natural History. [Vol. XIX,
from Colorado and from South Dakota it is obsolete on p4+ and
on the molars does not extend so far down.
? CASTORIDA.
Cylindrodon fontis Douglas.
We have eight lower jaws and an upper jaw of this species,
some of which exhibit the tooth pattern, and enable us to
refer this curious little rodent provis-
ionally to the Beavers. The two lower
jaws on which the species was based
were of comparatively old individuals,
and the pattern had disappeared, so
Fig. 7. Cylindrodon fontis. that it was not possible for Mr. Douglas
Upper jaw, twice natural size. z ; : y
No. 9639. to determine its relationship.
The dentition is 14 c* pt m3 — four
cheek teeth in each jaw, as in Castoride, instead of five above
and four below as in Sciuride and Ischyromyide. The pattern
resembles that of Steneofiber more than any other related form,
consisting in the lower molars of a
deep and persistent external enamel
inflection, and three fossettes corre-
sponding in position to the internal | |
enamel inflections of Steneofiber and
Castor. Ofthese fossettes the median
is the most persistent; the median
and posterior are at first internal
enamel inflections, the posterior in-
flection becoming a closed fossette
at a very early stage of wear, while
the anterior fossette is closed from
the first.
It would appear from the history i
of those teeth that the enamel in- Rig, Bo) Gua
flections did not originate on the ne he eec ee eee eee Bae
sides of the tooth and become gradu- sie Cera SRE ee
ally deeper and more complicated as
the tooth became more hypsodont, the fossettes being a
secondary modification; but that the inflections and fossettes
1903.] Matthew, Fauna of Titanotherium Beds of Montana. 213
have both originated from the valleys and lateral notches of
a brachyodont molar, such as that of [schyronrys. Either of
these is easily derivable from a primitive tritubercular molar
such as those of the Tillodonta, but not easily from such teeth
as are displayed by Paramys and its allies, in which, as in
the squirrels, the teeth appear to be specialized and degen-
erate rather than primitive.
The upper jaw of Cylindrodon (No. 9639) is that of an old
individual, with but little indication of the pattern left on
the teeth. From what there is present it appears that the
pattern exhibited a strong median external inflection and an-
terior and posterior fossettes, and a trace remains on p4 of an
internal inflection. The pattern was probably like that of
Ischyromys, in having the external inflection of greater per-
sistence and depth than the internal. In other Castoride this
condition is reversed. The upper molars decrease in size from
before backward and are of rounded peg-like outline, fossettes
on all but mt. The incisor originates just above the roots of
p+ and m!, and is stout, not grooved, with a moderate dias-
tema between it and the grinding teeth. The antorbital fora-
men is small and the palate in front of it is narrow.
The depth of the jaw of Cylindrodon in its anterior portion
is a very marked character. IJschyromys comes nearest to it
in this respect.
SCIURID.
Sciurus (Prosciurus) vetustus, subg. et spec. nov.
Represented by an upper jaw, No. 9626, with complete un-
worn dentition.
The species is smaller
than S. relictus of the
Oreodon Beds, and at
least a third smallerthan
S. jefjersont Douglas of
the Pipestone Creek
Beds, or S. wortmant of
me ert Gurtince nannies Type
It is larger than S. ballo-
vianus of the John Day, the first molar (the only upper tooth
214 Bulletin American Museum of Natural History. {Vol. XIX,
preserved in the type of S. ballovianus) is considerably wider
transversely, its anterior cingulum much less developed, its
mesostyle or median external cusp (between the anterior and
posterior transverse ridges) more prominent. In none of the
other species is the upper dentition known, so that an exact
comparison is not possible.
Compared with modern Sciuride this species shows some
interesting points of difference. It is nearest to Sciurus, but
differs in several points of importance:
1. The cross crests on the molars are less complete, and are
partly broken up into separate cusps.
2. The third premolar is a much larger tooth and has a
small accessory posterior cusp.
3. The heel of m3 bears a short transverse crest and a strong
posterior marginal ridge. In Sciurus the posterior part of
m3 is a slightly concave flat basin.
4. The base of the zygoma is anterior to p3.
From Jamas and the other modern Sciuride it also differs
in the retention of the mesostyle, in addition to most or all
of the above-mentioned points. The ridges are not so high
as in Cynomys and Arctomys, and their patterns differ in
various other details. I have little doubt that with more
perfect material it will be necessary to place this and all the
other Oligocene Sciuri in a separate genus, nearest to Sciurus,
but retaining the above primitive features in the dentition,
and others of more importance in the skull. But with our
present knowledge the distinction in the last molar is the only
one which we can predicate of all the Oligocene species in
common, and this is only of subgeneric importance at best.
The last lower molar in S. relictus, ballovianus, and wortmant
exhibits a correspondingly ridged and unreduced heel to the
ridged and unreduced heel of the last upper molar in S. vetustus.
Upper dentition, p3sms_:....4...55. 3 eee 7.7 mm.
: molars mz -m3 4/5/0025. a. oe eee fiat
P3 transverseiditmg «haope ee ee 0.9
longitudinal. \.. te wee ee ee 0.8
P4 transverse )).30 a. 0. ae ae eee 22
“ longitudinal cv. wobhbc uh canoe (eee 2.0
M* ‘transverse. ij.5.045 ok Aeln. eie e Bar
1903.] MJatthew, Fauna of Titanotherium Beds of Montana, 215
Mar tranS VeESGae st.) tix cseremeeamerei elec arate tac ere ech Lave ahah 2.5 mm.
py LOnoNtit cima een erent a ae) Pegs seeogse acho ver site fe 1.8
IS icaaswrerSe. a a teeter he Wid case rer use! nanoeae 233
peso LOM SEITEN alee ey ge teas Eisus sid Sen a oneness 2.0
Width of palate, including molars............... 9.6
? HETEROMYID.
Gymnoptychus minor (Douglas).
Eumys minor Doucuas, Trans. Am. Phil. Soc. 1gor, 16.
The position of Gymnoptychus and Heliscomys has been
variously given by different authors; they are, in fact, rather
remote relatives of any modern type, and combine charac-
ters of Sciuride, Geomyide, and Heteromyide with others
peculiar to themselves or shared by the Ischyromyide. The
dental pattern strongly suggests that of /schyromys, but the
resemblance may be superficial; by simplification and hypso-
donty it might be
converted into a
Heteromyid pattern.
I place the genus in
this family on Scott’s
authority.
The present spe-
cies is intermediate
in size between G.
minutus and G. liolo-
phus, but nearer to ; ; , ;
mietcitter. The type races 24 oo es eee a ete
q teeth, No. 9630.
of G. liolophus re-
tains the milk dentition; a specimen of G. minor of corre-
sponding age shows a smaller and shorter d, and somewhat
narrower and smaller m,.
Seven lower jaws are referred to this species.
Measurements,
Type (Douglas). No. 9630.
Lower Centittiompy aiiin nti) ties clos) = 5-5 mm.
AIO LATS A, MMe etn afta yee are meee se 4.0
Pa Real SECS Gare ty hears hoch Get anet ste tusicnatel ay o¥e sea haggh ods a) ihe)
fF MorienacG iba. ao ab seen Ole cece Gna 1a I.4
IMEMGGAMSV.CESCs c-acrite skiers fee cacti ee oe E.7 1.7
pe mlOmontiGumalerret cieivsirct cy sis sere 6 ngs vigy'
Diastema behind imeisor. <1... =. «ss An
216 Bulletin American Museum of Natural History. [Vol. XIX,
Gymnoptychus minimus, spec. nov.
A smaller species than G. minutus Cope. Fourth lower
premolar larger in proportion
to the rest of the dentition,
its anterior and _ posterior
halves of equal width; while’
in G. minutus the anterior
half of p, is much narrower
than the rest of the tooth.
The first and second molars
Fig. 11. Gymnoptychus minimus, Type <
an fone times natural size. <A, in- are narrower than in G. MAN -
ternal, 41, crown view.
utus, and the whole molar
series is thus much more uniform in width.
Only one specimen of this tiny rodent was found by our
party.
Measurements.
Length of dentition, tip of incisorto m;......... 8.4mm.
nf “ molar-premolar ‘Series, py, 2.222 235
“ aingeigsnnolhesycrokasos wo hyo ag deSs ou Se 2.6
LEPORID.
Palezolagus temnodon Douglas.
This species is nearly related to P. haydeni of the Oreodon
Beds. We have for comparison a series of about seventy-
five jaws, upper or lower, but nothing more complete. Mr.
Douglas distinguished the species by the presence of an an-
tero-external groove on p?.1. On comparison of our series
with several hundreds of jaws of P. haydeni in the Amer-
ican Museum collections, we are able to add a number of
other changes less obvious on a single individual, owing to
the great variation that age brings about in the characters of
the teeth.
The size appears to be nearly constant, approximating that
of the smaller specimen described by Mr. Douglas, which, as
the first measured specimen, is the type of the species. The
molars, and especially the premolars, are less hypsodont than
+ Mr. Douglas says p%, but this must be a slip of the pen.
1903.| Matthew, Fauna of Titanotherium Beds of Montana. 217
in P. haydent. The internal enamel inflection on the upper
teeth is less deeply impressed and less persistent; it has dis-
appeared on all the older individuals, as it does in P. tur-
gidus, but at an earlier age; in P. haydeni only a few of the
very oldest animals have lost this inflection. The last upper
molar is larger than in P. haydeni. In the lower jaw p, has
a less persistent external enamel inflection, so that in old
individuals it becomes one-lobed, a character seen also in very
old individuals of P. turgidus, but which I have not seen in
any example of P. haydent.
Measurements.
iipper motar-premolar-senies: 402.) <r aace ee wears ra. Tam,
Lower - Fa uidhpiaih isfeiier'y tes 9 the emer tete BEE
Post-canine diastema in lower jaw............... 8
Palzolagus brachyodon, spec. nov.
Paleolagus ? turgidus, P.? triplex, Doucuas, Trans. Am. Phil. Soc.
r90x, Vol. XX, p..6. Not of Cope, except:
Paleolagus turgidus Cope, Geol. Sur. Canada. Contrib. to Can.
Palzeont. Vol. III (quarto), p. 5, pl. xiv, fig. 9. Not of previous pub-
lications.
This species is of the size of P. turgidus, and probably the
specimens referred by Mr. Douglas to that species and to P.
triplex really belong here. It is more brachyodont than tur-
gidus, and much more so than any other species of Palgola-
gus. P? is smaller and more conical, m3 appears to have been
larger, the internal enamel inflection less persistent. In the
lower jaw p, is shorter, more conical, and the inflection dis-
appears a little earlier than in P. turgidus.
Twelve specimens of more or less complete upper or lower
jaws represent this species in our collection.
Measurements.
Molar-premolar series, upper jaw (m3 estimated)... 16. mm.
a s et MIM LEER PARA J Nr a So. lia ak 3 Lei.
UF (aga hegre Us bia P25 ar SPC aS ERD, Bien RCP iene ge r55
See LLAMSWELSGy ey maya irre eet eie cite via ol obese eer stele o) ores 205
218 Bulletin American Museum of Natural History. [Vol. X1X,
It appears probable that this species and P. temnodon stand
in direct or almost direct genetic relationship to P. turgidus
and P. haydeni respectively. The occurrence of the species
of Pal@olagus is:
John Day Lepus ennisianus
; d Leptauchenia Beds P. agapetillus P. intermedius
White River Oreodon Beds P. hayvdent P. turgidus
Titanotherium Beds P. temnodon P. brachyodon
Oligocene
The evolution in Palg@olagus ran in parallel lines in the dif-
ferent species, some being more progressive in one character,
some in another, but none exhibiting either wide divergence
or retrogression. The characters in which progress is ob-
served, as I have remarked in a previous paper,? are:
r. Superposition of the Lepus tooth-pattern over the older
and simpler one inherited by Palgolagus. This pattern, show-
ing at the crown in the older species, bites continually deeper
into the tooth until it entirely replaces the older pattern
during the whole life of the animal.
2. Increase in length of teeth, molarization of anterior pre-
molars, and some reduction in size of m3.
3. Bending down of facial portion of skull on cranial por-
tion. This is associated with lengthening of neck and legs.
4. Increase in brain-capacity, in supra-orbital processes, etc.
5. Increase in size.
It will be observed that in the first and second characters,
our two species from the Titanotherium Beds are in all re-
spects more primitive than those of the Oreodon Beds. The
difference in size is trifling if any, the third and fourth charac-
ters cannot be observed in our specimens.
PERISSODACTYLA.
EQUIDA.
Mesohippus westoni Cope.
Parts of upper and lower jaws, fore and hind feet, and
many fragmentary jaws and teeth represent one or more
species certainly distinct from MW. bairdiz, which does not occur
* Bull. A. M. N. H., XVI, 1902, p. 306.
1903-] Matthew, Fauna of Titanotherium Beds of Montana. 219
in these beds. It is provisionally referred to Cope’s species,
known hitherto by an upper and two lower teeth from Swift
Current Creek, Canada. These specimens will be described
by Professor Osborn in a later paper.
HYRACODONTID2.
Hyracodon sp.
Two lower jaws and an upper molar belong to a species of
Hyracodon. I do not observe any important distinctions
from H. nebrascensis in the parts preserved; but in the ab-
sence of the diagnostic teeth (upper premolars) make no
specific reference.
ARTIODACTYLA.
LEPTOCHERID&.
Stibarus montanus, spec. nov.
No. 9668, a lower jaw containing the second, third, and
fourth premolars, and the first molar, enables
us to place the hitherto problematic genus
Stibarus in the Leptochceride.
Generic distinctions. — Molars like those of Lepto- Gy eee
cherus. Premolars much like those of Leptomeryx, je
but with lower and more rounded cusps.
=! : Bie ho aig = : Fig. 12. Stzbarus
Specific distinction. Third premolar with NO gitustlobus Cope,
posterior cingular cusp. The second premolar is type specimen, na-
. tural size, from the
long and laterally compressed, with three rounded Oreodon’ Beds of
2 : Northeastern Col-
ed ae in line, the an- orado, 4, external,
terior the smallest, 4’, crown view.
the median the high-
est. The third premolar has a similar
form and composition, but the median
and posterior cusps are somewhat
larger, and there is a cingulum around
the posterior end of the tooth. The
Fig. 13. Stébarus montanus, partof fourth premolar is wider but not much
lower jaw, type specimen, natural size. Over half as long as the third, it hasa
A, external, 41, crown view of teeth, ‘ ° F .
main cusp, protoconid, an anterior
cusp connected by a ridge with it, a postero-internal and postero-
external cusp, all worn off in the specimen and none marginal, and a
220 Bulletin American Museum of Natural History, (Vol. XIX,
posterior cingulum. The first molar has four chief cusps, the external
ones somewhat crescentic, the postero-internal smaller than the others,
and a posterior cingulum with small hypoconid. P, with diastemata
behind and in front of it.
No. 9670 shows the second and third molars of similar composition
to the first except that the hypoconulid in m, is equal to the other
cusps and posterior to them.
This species appears to be closely related to S. obtusilobus
Cope, but the posterior cingular cusp (heel-cusp) is lacking,
and p, has small diastemata before and behind it, while
there are none in S. obtusilobus. In size and other characters
it is identical as far as the type of Cope’s species permits
comparison.
Stibarus has been conjectured to be allied to the camels;
its actual position has, I think, never been suspected. It is,
in a way,a link between the Leptochceride and Leptomeryx,
and makes it more certain that the former is truly an artio-
dactyl family. ‘‘ Leptocherus’’ quadricuspis Hatcher is prob-
ably a species of Stibarus.
Measurements.
SORE oie eae ae 27.3 mm
Spa tha Bukiomomee ke ohed fe ee Ole width 2.1 mm.
oe BaDgticr vce s.6 eas eee 7.6 whey 27
pie ey en tas cen eg cles eri 4.9 ee)
SP PPCIN pnt cd eo, © aire eR ae 5-0 i) Avo
Depth of jawabelowaps seas. ose ole
i“ a Sa elias Sa Aone eee 10.
OREODONTID/&.
Bathygenys alpha Douglas.
We have three specimens referable to this genus and species:
parts of two upper jaws and one lower jaw. I identify these
with Mr. Douglas’s species in spite of wide distinctions in the
drawing of the teeth of his cotype specimen. These, if cor-
rectly drawn, could hardly be Oreodon teeth; they are fully
as narrow and trenchant as those of Leptomeryx. As, how-
ever, he compares the teeth to those of Merycocherus, which
are short and wide and crowded, I assume that the error is in
1903.| Matthew, Fauna of Titanotherium Beds of Montana. 221
the drawing, especially as the other drawings of the type and
cotype agree well with the description and with our specimens.
The best upper jaw shows the molars and fourth premolar.
The premolar is simpler than that of
Oreodon, lacking internal cingulum and
antero-external accessory ridge, and con-
sisting of external and internal crescent,
the former with slightly concave external
surface, the latter with a short postero-
internal cingulum. The molars are com- ees gee
posed of four crescents, no trace of the ae ee Mp:
paraconule remaining on m? or m3;
m! is a little worn, so that it is uncertain whether or not a
minute pl. existed. The anterior halves of mt and m? are
wider transversely than the posterior, the protoselene pro-
jecting further inward on the palate than does the hyposelene.
The posterior half of m3 is nearly as wide as the anterior half,
as in Agriocherus and the Uinta Oreodonts, instead of re-
duced in width as in Oreodon and the other Oreodontide.
The exterior surface of the inner crescents is slightly concave,
as in Agriocherus, instead of strongly concave as in the later
Oreodonts, or convex as in Protoreodon. The exterior surface
of the outer crescents is nearly flat.
The fourth lower premolar is oreodont in type, but differs
from Oreodon and resembles Promerycocherus, Merycocherus,
and Merychyus, in that the entoconid is a ridge extending
down and back from the deuteroconid, instead of a separate
cusp or distinct ridge. In Protoreodon the entoconid is rudi-
mentary. This tooth is a little narrower anteriorly than
p, in Oreodon, wider than in Protoreodon, Merychyus, or Pro-
merycocherus. The molars are proportioned nearly like those
of Oreodon, but are more brachyodont. They are shorter,
higher, and with more crescentic cusps than those of Pro-
toreodon.
Bathygenys is in most respects between Oreodon and Prot-
oreodon, but considerably nearer to the former. Some
characters point toward a nearer relationship with the
Merycocheerid group of Oreodonts than with Oreodom itself.
222 Bulletin American Museum of Natural History. (Vol. XIX,
More complete material is needed before its place can be de-
finitely determined.
Measurements.
Upper molars, m,25 9 <a. shaiasiee scree eS peta
‘« premolar p4, longitudinal........ 4. transverse 5.2 mm.
Lower TOGHM,PPjats 25 os 6 ae cen ee tie eS 16.
Upper molar m! longitudinal.......... BES a 6.8
a Sy A saale ia ee eee yee 5.8 ip Wag
ce ~ m3 ‘Dae eee 6.2 a ihe
? Limnenetes sp.
A number of lower jaws and parts of jaws may be provision-
ally referred to this genus, although the characters of the
teeth are nearer to those of Bathygenys than to Oreodon; while
Mr. Douglas describes the teeth of Limmnenetes as so like those
of Oreodon as not to need a separate description. The pre-
molars are narrower than those of Oreodon, the entoconid
ridge not separated from the deuteroconid on p,, and the
structure of p, is intermediate between that of O. culbertsont
and Merychyus elegans, but more brachyodont than either.
The lower molars are intermediate between those of Proto-
reodon, with conical internal cusps, and those of Oreodon
with fully crescentic internal cusps. Heel of m, narrow, as
in Protoreodon.
There is more than one species, and may be more than one
genus, among these specimens, and it is inadvisable to at-
tempt to place them very definitely at present.
HYPERTRAGULID&.
Leptomeryx ° esulcatus Cope.
The type of Cope’s species is an upper molar, probably m',
from the Titanotherium Beds of Swift Current Creek, Canada.
We have a large number of parts of lower jaws and separate
upper teeth, which belong to one or more species of Lepto-
meryx a little larger than L. evansi, but variable in size.
These are referred provisionally to L. esulcatus, because it is
within the limits of size, comes from the same geological hori-
1903.] Alatthew, Fauna of Titanotherium Beds of Montana. 223
zon, and agrees well enough with other first molars in our
series. The character by which Cope separated it from L.
evanst, the absence of
defining furrows to
the rib of the external
crescents, is exhibited
only on first molars,
and not on all “of
these; but two other Fig. 15. Leptomeryx esulcatus. Lower jaw, natural
characters of moreim- size, external view, composite, Nos. 9696, 9706, 9702.
portance are seen in our material, viz.:
1. The median internal cusp, a strong cusp in m3 of L.
evanst, and a smaller one on m? and m!, is quite small on m3,
and absent, or nearly so, on m? and m!.
2. In the third lower premolar the protoconid has two pos-
terior ridges, of which the internal one connects with the heel,
and the external one does not; while in L. evansz and other
species from the Oreodon and Leptauchenia Beds, the exter-
nal ridge connects with the heel, and the internal one does
not.
In the lower jaw I have observed no entirely constant dis-
tinctions, except in p,. The first premolar is small, and is
separated from p, by a diastema of about the same length as
that of L. evansi; the size of the jaws averages larger than
those of L. evanst from South Dakota, and all are larger than
L. evansi of Colorado. The height of crowns and proportion
of the teeth are about the same, and the premolar pattern,
allowing for individual variation, is identical, except as above
noted.
I have no doubt that better material will furnish more sat-
isfactory distinctions, but, except for the characters noted
above, | am unable to find any in the teeth.
Measurements.
LOWen Premolmsem soy wagers dey helt a Ske 2d oS a's eae 19 mm.
[Pron ermeemel ens. ! thy <3 t 0 eh A i er i 24
Upper molar, m3, antero-posterior............... 8.
ae
URATSVEESGS tie cards seat ane eles 8.5
224 Bulletin
American Museum of Natural History. |Vol. XIX,
Leptomeryx mammifer Cope.
A much larger species than the preceding, size about that
of Poébrotheri
um eximium. Distinguished from L. evansi by
the pattern of p,, which is like that of L. esulcatus, but with
Fig. 16. Leptomeryx mammifer. Lower jaw, natural size, external view. No.
9684, £3 and f, supplied from No. 9686.
the external
ridge of the protoconid more clearly separate
from the heel, and the postero-internal more clearly joined to
it. The first
canine nearly
lower incisor is large, second and third small,
as large as first incisor (larger than in L. evanst).
First premolar equally spaced between
c and p,; remaining premolars close
set. Molars an enlarged copy of those
Fig. 17. Leptomeryx mammi- of L. evansi. Parts of the feet of
Jer. Crown view of lower pre-
molars, natural size.
pa from No. 9689.
no important
No. 9687; Leptomeryces of appropriate size were
found at the locality; they show
distinctions either in fore or hind foot, from
L. evanst. The upper molars have a smaller median internal
cusp than those.of L. evansi. The size is nearly a third
larger, lineally.
Lower premolars: p.4. 2500) <n non ee eee 24. mm.
+ elitst prembplar, ‘pr... 6a «« acoisiek See ee 2
ppace between ‘@ Amd. ps. ...... sate See ee 17
Lower molars ays. SOP) ea eee 32
Complete lower dentition, estimated............. 78
Upper molar, longitudinal... ..-..'... s- 25a cee 9
“* EAIISWEESE. 224125 /5.006 poe ee II
CAMELID.
Leptotragulus profectus, spec. nov.
Parts of several lower jaws represent a species of Camelid
nearly as large as Poébrotherium wilsoni, but with brachyo-
1903.| Matthew, Fauna of Titanotherium Beds of Montana. 225
dont molars like Leptotragulus. No. 9681 (type) shows p,_m,;
No. 9682 (cotype) p,_, and the root of m,; No. 9683 a num-
ber of lower molars and milk molars. Nearly all these teeth
are unworn or very little worn.
The species is of about the same size as Leptomeryx mam-
mijer, but is distinguished by the camelid pattern of the pre-
Fig. 18. Lefptotragulus profectus. Type specimen, external view,
natural size; second and third molars supplied from another individual.
molars. The molars are very difficult to separate from those
of L. mammijer; they are a little wider and shorter-crowned,
with the crescents placed less obliquely, and m, has but a
vestigial postero-internal cusp. The fourth premolar has no
deuterocone, but two strong posterior crests from protocone
to heel sub-parallel, enclosing a narrow lenticular fossa. The
third premolar is similar, but more compressed; the second
has but one complete posterior ridge. Their pattern differs
from that of Poébrotherium chiefly in the completeness of the
inner posterior ridge, which in Poébrotherium does not reach
the heel on p, and p, and is entirely absent in py,
The cotype shows a moderate diastema, considerably
shorter than that of L. proavus, separating p, from the alve-
olus of a strong caniniform tooth.
The heel of the last molar in referred specimens is like that
of Leptomeryx or Leptotragulus, with postero-external cres-
cent and small antero-internal cusp. In Poébrotherium is a
posterior crest, not crescentic, and a small antero-internal cusp.
The species is about a fifth larger than L. proavus, with
which its premolar pattern corresponds quite closely, accord-
ing to Scott’s description. The molars are a little wider and
[April, 1903. ] 165
226 Bulletin American Museum of Natural History. |Vol. XIX,
a little more hypsodont, and the diastema both relatively and
absolutely less. It is throughout very suggestive of Poé-
brotherium, much
; more so thian
either Protylopus
(EF E
or Leptotragulus
Fig. 19. Leptotragulus profectus. Type specimen, crown proquus, in the
view of teeth, natural size; molars 2 and 3 supplied from
another individual. details and con-
formation of the molar and premolar cusps. It shows much
less resemblance to Hypertragulus. Unfortunately no upper
teeth can with entire certainty be referred to our species.
Those which are doubtfully referred have the Leptomeryx-
Poébrotherium pattern, with strong mesostyle and a rib on
the external face of the anterior external crescent, but none
on the posterior; they are less extended transversely than
those of Leptomeryx, much more than those of Poébrotherium.
The upper molars of Hypertragulus are very easily distin-
guished by the entire absence of mesostyle and equal develop-
ment of the external ribs on anterior and posterior crescents.
No upper teeth of this pattern were found in the Pipestone
beds. The upper molars of Leptotragulus proavus are not
known.
From the above facts I am inclined to believe that Lepto-
tragulus — this species at least —is more nearly related to
Poébrotherium than Professor Scott has supposed, and that
it has not much to do with Hypertragulus. It is probable in
either case that the caniniform tooth is the first premolar.
The species is really far nearer to Poébrotherium than is Pro-
tylopus petersoni; how much of the resemblance is due to
parallelism remains to be determined.
Mr. Gidley discovered last summer in the Oreodon Beds of
South Dakota, a brachyodont camel, which may be a direct
descendant of this species.
1 m2
Measurements.
LOWErpreniOlats, Pei, cree an pacer ee ee eee 25 mm.
Diastema in font Of ps thtteeaa <x cae ne Nee 10
Last lower molar {ak Wne GRR te Rect ee ee 15
vican
vu,
, extracted from BULLETIN.
>
atural BHisto1
eat
S
ae
Ni
S
os
Se
ene.
_ May 9, 1903.
Z
06
ageh
OL
By W. D. Matruew.
é
Nets Vani
VoL. XIX, ArticLe VII, PP. 227-229..
an Muiseum of A
1c
- AUTHOR’S EDITION
A Fol H. ”
Amer
The Knickerbocker Press, ew Work —
'
x
Article VII.—A FOSSIL HEDGEHOG FROM THE
AMERICAN OLIGOCENE.
By W. D. MaTrHew.
The Hedgehog family (Erinaceide) has hitherto been found
only in the Old World (Europe, Asia, and Africa). It includes
three living and five extinct genera, ranging from Lower Oli-
gocene to recent. It has been divided into two subfamilies,
one including the modern hedgehogs (Erinaceus) and one ex-
tinct genus (Pal@oérinaceus), the other with two modern
genera, Gymnura and Hylomys,— small rat-like East Indian
insectivores,— and four extinct genera, Necrogymnurus, Ga-
lerix, Lanthanotherium, and Tetracus. The subfamilies are
distinguished as follows:
: - ee -I.3. : :
Erinaceine. — Dentition 2" 2 s Palate imperfectly ossified.
Pelvis wide. Fur with spines.
Gymnurine. — Dentition : = ; - Palate completely ossified.
Pelvis very narrow. Fur without spines.
A related but more primitive family, the Leptictide, is
found in the American Eocene and Oligocene. The dentition
is unreduced (except that there are only two upper incisors),
the molars subtriangular and extended transversely, while in
Erinaceide they are subquadrate and not extended trans-
versely. The Leptictide might, however, without serious
straining of relationships, be included as a primitive sub-
family of Erinaceide, with which they agree well enough in
skeleton and in most skull characters. There are four de-
scribed genera, Paleictops from the Middle Eocene, Leptictis,
Ictops, and Mesodectes, from the Lower Oligocene of the
Western United States.
A true Hedgehog, of the Erinaceine subfamily, is repre-
sented by the front half of a skull from the Upper Oreodon
Beds of South Dakota, found by Dr. F. B. Loomis of the
American Museum Expedition of 1902. The dentition is that
of Erinaceus, but the teeth are less specialized, and in several
227
228 Bulletin American Museum of Natural History. [Vol. XIX,
respects resemble those of the Leptictide. The reduced pre-
molars and short facial portion of the skull exclude it from
the Gymnurine, but the teeth resemble quite nearly those of
Necrogymnurus,t and the palate does not show the defective
ossification of Erinaceus. The last molar is small and tritu-
bercular as in Hylomys, Necrogymnurus, and Galerix. In
Gymnura it is large and extended longitudinally; in Erinaceus
small and reduced to a transverse blade. The teeth are more
extended transversely than those of Erinaceus, and retain
considerable indications of the tritubercular form of molar
from which they no doubt originated. Their pattern, how-
ever, is definitely Erinaceid, with two equal outer and two
equal inner cusps, a smaller separate median cusp (meta-
conule), and an an-
tero-median ridge
from the antero-inter-
nal cusp (protocone)
to the antero-external
cingulum.
This hedgehog
forms a connecting
link between the Eri-
naceine and (symnu-
rine subfamilies, and
to some extent be-
tween Erinaceide and
Leptictide. Itseems
impossible to place it
in any of the described
genera, and it is there-
fore named:
Proterix loomisi, gen.
et sp. nov.
ig Es Proterix loomiisz. [ixes specimen, twice 3.1.3.3
natural size, A, anterior part of skull from above; A? it] OO I
palate from below. ; Dentition 1
enlarged. C! large, two-
rooted. P? small, one-rooted. P3 small, three-rooted with well de-
veloped deuterocone. P4 large, molariform, with small hypocone.
* Relying on Dr. Leche’s very careful figures and descriptions.
1903.] Matthew, Fossil Hedgehog from American Oligocene. 229
Mt and m? wider than long, quadrate with two external and two
internal cusps of about equal size and a small separate postero-
intermediate cusp (metaconule), the antero-internal cusp (protocone)
with a ridge running out towards the antero-external margin. M3
trihedral, small, not extended transversely, paraconid and metaconid
equal and well separated, no hypocone. Palate completely ossified,
its posterior margin as in Erinaceus. Skull bones arranged much as
in Erinaceus, a well defined sagittal crest; premaxille not reaching
frontal bones.
Measurements.
Maxillary dentition, c-m3 inclusive.............. 18.4 mm.
Transverse width of palate including molars...... 17.6
Depth of skull, junction of postorbital crests to
oy ear de Ais oal6 2 cok, pee ch Ae ee 16.8
Length of three true molars (antero-posterior).... 7.9
Antero-postemorn diameter Of mM... 0.6 fe. ese ew 2.4
Transverse + FO OO. EEO eR ae 4.8
Antero-posterior si PARTIR? seepciensiec eee omieests ee 2.9
Transverse ss RMP IO CIs. onto, shoe 3-9
Antero-posterior - BTINS eotaue.tcs.2 205 sy <peeonds 2.0
Transverse ts TN SIS eee eace eea eee gee 2.8
RANGE OF THE ERINACEID& AND LEPTICTID&.
Europe. Asia. Africa. North America.
Modern. Erinaceus. Erinaceus. Erinaceus.
Gymunura.
Hylomys.
WA |
Pleistocene. | Erinaceus.
Pliocene. Erinaceus.
Mies es ———_—_
Erinaceus.
Miocene. Galerix.
Lanthano-
therium.
Palzoérina- Proterix.
ceus.
: Leptictis
Oligocene. Tetracus. j )
2 Mesodectes.
Necrogym- Ictops.
nurus.
Eocene. | | Anisacodon
| ? Passalaco-
don.
| Palaictops.
AUTHOR'S EDITION, extracted from BULLETIN
OF THE ©
VoL, XIX, ARTICLE IX, pp. 387-394.
New York, Fuly 8, 1903.
The Rnicherbocker Press, “Rew Work
i
Oar pe, ee ER Pe Ree eT ar om,
Article IX,—ADAPTIVE SIGNIFICANCE OF THE
SHORTENING OF THE ELEPHANT’S SKULL.
By W. K. GRreGcory.
PLaTE XXIII anp 4 Text Ficures.
The following observations were made in connection with
Professor Osborn’s forthcoming memoir on American fossil
Proboscidea, for which were also prepared the figures here
used by his kind permission.
Weithofer* regards the changes in the elephant’s skull ob-
servable during individual growth as well as in phylogeny,
such as the compression and vertical heightening and deep-
ening of the skull, the wide separation of the inner and outer
tabule of the bones, and the cancellous condition of the diploé,
the forward shifting of the orbits from a point above the an-
terior grinders, etc., as primarily correlated with the prodi-
gious development of the tusks — weapons and crow-bars
whose effectiveness increased with and reciprocally hastened
the phyletic advance in body dimensions. Now the earliest
known proboscideans (Meritherium) possessed upper as well
as lower incisor tusks, and there are other grounds also for
inferring that the ancestors of Dinotherium probably pos-
sessed upper incisor tusks of small size, even smaller than
in the primitive Mastodon (Trilophodon) euhypodon; the pre-
sence of these may have initiated the shortening of the
head, but the final compression of the skull in Dinotherium
progressed notwithstanding the reduction and entire dis-
appearance of upper tusks, and this, together with the
underlying similarity of its skull to that of Mastodon and of
Elephas, shows thatssome other factor must also be repre-
sented in the extraordinary end results of the process of fore-
and-aft compression.
This factor seems to be the development of the proboscis.
This unique organ probably owes its existence partly to the
shortening reach of the head and neck which took place
1 Die Fossilen Proboscidier des Arnothales in Toskana. 4to, Wien, 1800.
[387]
388 Bulletin American Museum of Natural History. [Vol. XIX,
simultaneously with increasing stature and longer limbs,
partly to the necessity of reaching outside the lengthening
tusks, partly to the intrinsic advantages of a prehensile organ
of such manifold possibilities. On the one hand, we may
suppose, the enlargement of the trunk intensified the changes
due to the shifting and enlargement of the tusks, and on the
other hand inaugurated many of the peculiar conditions de-
scribed below.
The backward shifting of the weighty tusks and trunk,
compensating their adverse leverage, lessened the antero-
posterior space available for the grinding series, while the
work put upon the individual grinders increased with larger
bodies and longer lives. This may explain in part why the
simple grinders of Dinotherium, with their two, or at most
three, low, widely separated ridges, were constrained to
evolve into the wonderfully specialized and effective grinders
of the Mammoth, with ridges in great number, exceedingly
high, and closely appressed, and in which there is a so-called
horizontal succession in use, the unworn teeth being pushed
into place from behind. The weighty, rapidly heightening
molars, together with their immense, backward and upwardly
growing alveolar pouch, might thus be regarded as a third
factor in the evolution and individual growth history of the
skull. It is plain, however, that this factor was a minor one
and that the characteristic features of the skull were attained,
in great part, before the teeth had become so highly special-
ized; for the typical Mastodon americanus, with its great size,
and great tusk-and-trunk development, shows nearly the same
degree of backward and downward extension of the posterior
nares and hard palate as in Elephas, yet retains comparat-
ively primitive low-crowned grinders, and the considerable
space between the posterior border of the last molar and the
posterior lateral limits of the hard palate in this genus was
not nearly so much utilized for storing incoming grinders as
it is in Elephas.
Principally then, to the enlargement and backward shifting
of the trunk and tusks and the consequent fore-and-aft com-
pression of the whole skull, combined with the progressively
1903.] Gregory, The Shortening of the Elephant’s Skull. 389
widening separation of the inner and outer surfaces of the
bones for the sake of lightness and large area for the head
muscles, we may ascribe the following morphological con-
ditions, which attain the extreme of specialization in the
skull of the Mammoth. In the inferior view of the skull the
hard palate, contrary to what obtains in most Ungulates,
is tilted somewhat upward; the palatines become reduced
antero-posteriorly and shoved backward so as to diverge
widely posteriorly; the posterior nares, probably pari passu
with the anterior nares, have been pushed very far back?;
the enormous vertical pterygoid wing of the alisphenoid
wraps itself around and functionally replaces the hinder end
of the encroaching molar-tooth pouch; the foramen ovale of
the alisphenoid, which in the most primitive Ungulates is
anterior to the foramen lacerum medius, has been shifted
obliquely backward and outward, and becomes confluent
with it externally (Fig. 1)? ; the presphenoid, basisphenoid,
basioccipital thicken in the median plane and at diminishing
rates, the lower tabule of these bones growing downward to
a less and less extent as we pass backward, so that in the
adult the inferior surface of the basis cranii points sharply
downward, and forms, with the plane of the back of the
occiput, an angle greater than go°; the tympanic bulle, rela-
tively large and inflated in the young, flatten down and
become closely appressed to the skull, pointing obliquely
downward, forward, and inward, with the wider end toward
the transversely expanded occipital region (Fig. 1, 7y.). In
brief, the progressive brachycephaly of the skull has appar-
ently involved not so much a fore-and-aft shortening of the
individual elements as a readjustment and modification of
them, and secondly an expansion in the transverse vertical
planes.
The shortening and deepening of the temporal fosse, and
perhaps to some extent the above-mentioned expansion of
1 Among the Glyptodonts a similar backward and downward growth of the palate
and posterior nares seems to be correlated chiefly with the shortening of the skull, as
the anterior nares remain terminal.
2 This change must have taken place at a very early date in the history of the Pro-
boscidea, as it is already established in the primitive Mastodon (Trilophodon) productus
and also, if Kaup’s figures are here rightly interpreted, in Dinotherium.
390 Bulletin American Museum of Natural History. (Vol. XIX,
Fig.1. Inferior view of the skull of a young Asiatic elephant. About }:nat. size. From
a forthcoming memoir by Professor Osborn.
BONES. .
P. mx.—Premaxillary. Bo.—Basioccipital. : ;
M-x.— Maxillary. p. As.—Pterygoid wing of alisphenoid.
Mx. p~.—Maxillary pouch for molars. /t.—Pterygoid.
Ma.—Malar. Sg.—Squamosal.
Po.f.—Postorbital ridge of frontal. £x. 0.—Exoccipital.
pal. Mx,.—Palatine ledge of maxillary. pg.—Postglenoid ledge of squamosal.
/.—Palatine. p. ty.—Posttympanic ledge of squamosal,
Vo.—Vomer. which with Ze. forms a secondary external
Ps.—Presphenoid, auditory meatus.
#Bs.—Basisphenoid. Ty.—Tympanic bulla.
FORAMINA, ETC. :
zy. #.—Anterior process of tympanic. J. ¢. #.—Foramen lacerum medius.
tp. h.—Tympanohyal. /.0v.—Foramen ovale(confluent with /. 2. 7.)
euw.—Eustachian opening of tympanic, z. c. c.—Canal for internal carotid artery.
a. p. f.—Anterior palatine foramina (canals), J. st. mt.—Stylomastoid foramen,
Z. 0. f.—Infraorbital foramen. J. ¢. .—Foramen lacerum posterius.
p. x. —Posterior nares, c. f.—Notch, a vestige of condylar foramen
~. a. s.—Alisphenoid canal. (?) (confluent with /% Z. Z.).
TEETH.
7.7, —Tusk. p° (dm2).—Third premolar (or second deciduous molar of authors).
?* (dm 3).—Fourth premolar (or third deciduous molar of authors).
1903.] Gregory, The Shortening of the Elephant’s Skull. 391
the grinders and their alveolar pouch, have had a marked
effect on the relations of the bones and foramina of the
sphenoidal region: the obliquely placed external portion of
the orbito sphenoid (Fig. 2); has been squeezed into a long,
3 Wee ey,
M2 yn jh
4 whi J
a=
Fig. 2. Superior view of the same skull. From Osborn, Abbreviations as above ; also
La.—Lachrymal. A. ma.— Anterior nares. Na. s.—Narial sinus.
thin process; internally (Pl. xx111) the anterior edges of the
basisphenoid are directed outward and backward; both inter-
nally and externally the optic foramen, foramen lacerum an-
terius, and foramen rotundum, in the order named, are
392 Bulletin American Museum of Natural History. [Vol. XIX,
obliquely arranged on descending levels from within outward
and from in front backward, the whole region having been
thickened by the separation of the inner and outer tabule of
~¢ WN The WW !
So. Ly, JRkMeZ
Nisan ge 7/8) 2
Ze Lf 1, Z,
SS
= ————
——————
——
Fig. 3. Side view. From Osborn.
all the bones, and also sharing in the upward-and-backward
tilting of the nasal region and in the general fore-and-aft
Squeezing of the skull, the end result being that the foramina
have been pulled out into long tunnels running obliquely for-
1903.) Gregory, The Shortening of the Elephant’s Skull, 393
ward, outward, and downward; especially internally the
fore-and-aft extent of the alisphenoid proper is brief.
Internally the skull has shortened up, one might almost say
in bellows fashion, with the optic foramen on each side at the
apex of the internal transverse folding (Pl. xx111), the ridge
of the “‘lesser wing’’ of the human sphenoid. As the skull
has also expanded transversely, the general effect of the in-
ternal view of the skullis thus that of compression around the
\Sa
i in
Ws ty / Y)
Fig. 4. Orbito-sphenoidal region, left side. From Osborn. The view is obliquely from the
side and from below the malar bone (compare Fig. 3).
BONES.
Po. f.—Post-orbital ridge of frontal. Sq.—Squamosal.
Fry .—Frontal. p. As.—Pterygoid wing of alisphenoid.
O. s.—External process of orbitosphenoid. Mx. p.—Maxillary pouch for molars.
As.—Alisphenoid.
FORAMINA, ETC.
op. f.—Foramen opticum. J.r.and a. a. s.—Arcade leading to foramen
Jf. 2. a.—Foramen lacerum anterius. rotundum and anterior opening of the ali-
sphenoid canal.
center (represented by the basisphenoid) and increasing ex-
pansion toward the periphery — somewhat recalling the con-
ditions of the domelike human skull. Between the frontal and
temporo-sphenoidal fosse, which form a large trefoil as seen
from above, there is on each side, running obliquely outward,
forward, and upward, a prominent triangular buttress, the
transverse ridge mentioned above, to the formation of which
the orbitosphenoid, frontal, and parietal contribute. On each
3904 Bulletin American Museum of Natural History. |Vol. XIX.
side, at a point opposite the malars, where the skull is most
constricted laterally, these buttresses branch off externally
into symmetrically opposed arches in different planes, which
pass forward, backward, downward, or upward. The wedge-
shaped basisphenoid, situated between the inner ends of the
buttresses, is the veritable keystone of this converging system. |
The skull as a whole is thus highly adapted to resist the
severe strains put upon it. The occiput, both in ontogeny
and phylogeny, flattens out and rotates backward, spreading
both vertically and laterally, until at last it forms, as it were,
a great, functionally solid bed-plate, receiving the thrusts of
the opposite inverted arches into which the skull has been
resolved. Each pair of these symmetrically disposed arches,
which also connect with the system culminating in the basi-
sphenoid, reacts, of course, against some component of the
force transmitted either to or from the tusks, trunk, and
grinders, or when the forehead is used in pushing. The in-
numerable toughly constructed air cells of the diploé give
immense strength, lightness and especially resiliency. This
desideratum may also be the reason for the very loose articu-
lation of the malar with the zygomatic process of the squa-
mosal, which would also permit the facial portion of the skull
to bend back slightly, under pressure, toward the cranial
portion. :
“LNVHda1Tq OLLVISY ONONOX AO TINNS AO MALA AOLMALNT
“apFjes
‘quyxa'ay
BTN Lescepscouucnd climes Laut VAN ‘HN WW ‘YW Nizat1ng
gE HEN
from Mount Lebanon, Syria, ik
AUTHOR’S EDITION, extracted from BULLETIN _
OF aa
American Biuseum ot Ratural Bistovy, ss
VoL. X1X, Rearsie x pp. 395-452.
New York, Fuly 8, 1903.
be
vbocker Press, A
- The Tnicke
ew
Article X.—ON A COLLECTION OF UPPER CRETA-
CEOUS FISHES FROM MOUNT LEBANON, SYRIA,
WITH DESCRIPTIONS OF FOUR NEW GENERA
AND NINETEEN NEW SPECIES.:
Prares XXTV—XOXCX VII.
BwO iP) Ay,
In accordance with an arrangement entered into between
Professor H. F. Osborn, Curator of the Department of Verte-
brate Paleontology, and Professor R. P. Whitfield, Curator
of the Department of Geology, the writer has studied the col-
lection of fishes which was made in 1gor by Professor Alfred
Ely Dax in the Cretaceous deposits of Mount Lebanon, Syria.
This collection consists of several hundred specimens of fishes,
besides a considerable number of crustaceans and a few mol-
lusks. The result of the examination of the fishes has been
the discovery of a considerable number of undescribed species
and various additions to our knowledge of the structure of
species already described. A large part of the collection and
nearly all of the new species come from a village called Hajula.
This may be regarded as really a new locality, since I have
been able to find it mentioned only once in Dr. A. S. Wood-
ward’s Catalogue of the Fossil Fishes in the British Museum.
From a letter written by Professor Day to Professor Whit-
field we learn that Hajula and Hakel are each about twelve
miles nearly northeast of the seacoast town of Jebeil, the
ancient Biblus. Hajula is situated six miles south of Hakel;
and between the two villages there are two westwardly pro-
jecting spurs of Mount Lebanon and an intervening valley.
Professor Day estimates the elevation of these villages to be
1 The collection of fossil fishes, of which the new forms hereinafter described form a
part, was made by members of the staff of instructors of the Syrian Protestant Col-
lege at Beirut, Syria, during the year 1901, and donated to the American Museum of
Natural History by the Rev. L. Stuart Dodge. They now form a part of the Museum’s
exhibit in the Geological Hall as part of its chronological series, under the head ot the
Cretaceous formation, in the foreign series of fossil forms.
As Dr. O. P. Hay has recently been engaged in preparing a catalogue ot the fossil
fishes of the Museum collection and is familiar with the forms found at the Syrian lo-
calities, it has been considered advisable to place this collection in his hands for identi-
fication. While doing this he has found several interesting species not hitherto
known to science. These are described and illustrated in the following pages as a con-
tribution to the work of this department.— R. P. WHITFIELD, Curator oj the Department.
[395]
“te &
390 Bulletin American Museum of Natural History. [Vol. XIX,
between 2500 and 3000 feet. In both places there are clear
evidences of faulting by which the fish-bearing strata have
been let down into the midst of older strata. Those at Hakel
have been let down to the level of the hippurite limestone of
Lebanon, being above the Trigonia sandstone. Professor Day
thinks that the Hajula beds are an extension of those at
Hakel. This study of the fishes appears to show that the
horizon of the beds at Hajula is some what higher than that. of
the beds at Hakel. Professor Day says that one notable differ-
ence is found in the great abundance of crustaceans at Hajula.
According to Professor. Day, the fish-bearing strata at
Hakel are exposed over a space of half an acre or more on
the side of a valley, which slopes about 30° to the northeast,
the dip of the strata corresponding nearly with theslope. The
outcrop at Hajula is similar, but more broken up and irregu-
lar. At both localities the slabs of rock which lie on the sur-
face are more easily and perfectly split than are those which
are dug up from some depth.
Professor Day was not permitted to work at Sahel Alma.
Much credit is due to the Rev. D. Stuart Dodge, at whose
suggestion and expense this collection was made, and to Pro-
fessor Day, who has displayed great industry and good judg-
ment in gathering the materials.
At the close of this paper the writer will add a few words
on the age of the strata.
In the following pages most of the specimens are recorded
under two numbers, of which the second is enclosed within
parentheses. The first number is the one given to the speci-
men in the Department of Geology, while the number in
parentheses is the one which it bears in the general catalogue
of fishes. This catalogue belongs to the Department of Ver-
tebrate Paleontology.
Figure 2, Plate xxvi; figures 3 and 4, Plate saa eand
figures 4-6, Plate xxxvi1, have been reproduced from photo-
graphs made by Mr. A. E. Anderson. Figures 3 and 4, Plate
XXVI, are from drawings made by Mrs. L. M. Sterling. All
the other figures are from photographs made by Mr. Rudolph
Weber.
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 397
Class ELASMOBRANCHII.
LAMNID-.
Otodus sulcatus Geinziz.
PLATE X XVI, FIGURES 3 AND 4.
Otodus sulcatus Grinrrz (H. B.), Char. Schicht. u. Petrefakt. sachs.-
bohm. Kreidegeb., Nachtr., 1843, p. 5, pl. iv, fig. 2.
Lamna sulcata Woopwarp (A. S.), Cat. Foss. Fishes, 1, 1889, p. 398.
(Synonomy and literature.)
In the collection there is a single tooth of a shark, and this
I refer to the above species. The number of the tooth is
4508 (3867).
The height of the crown has been about 15 mm., but the
apex is broken off. The crown has resembled that of a speci-
men figured by Geinitz in 1875 (Paleontogr., XX, pt. 1, pl.
Ixv, fig. 4d). At the base of the crown, in front, the width
is 6.25 mm. At the base, the anterior face is somewhat con-
cave, but it soon becomes slightly convex. There are a few
short grooves at the base of the crown in front. The cutting
edges are very sharp. The posterior face is very convex, and
there are present numerous sharp coste of various lengths.
The lateral denticles are relatively large, close to 5 mm. high,
and are costate both in front and behind, although the an-
terior costz are feeble.
The root is relatively large, and its branches make a very
acute angle with each other. The outer borders descend
from the lateral denticles so as to be nearly parallel with each
other. In this respect the root resembles that of Sauvage’s
Odontaspis rochebrunet (Bull. Soc. géol. France [3], VIII, p.
437, pl. xiii, fig. 3). The tooth from Hakel differs from the
one just referred to in having a less slender crown and more
prominent lateral denticles. The length of one branch of the
root is ro mm.; of the other, 12 mm.
Collected at Hakel.
398 Bulletin American Museum of Natural History. [Vol. XIX,
PRISTIDA.
Sclerorhynchus Woodward.
PLATE XXIV, FIGURE 1.
Woopwarp (A. S.), Cat. Foss. Fishes, I, 1889, p. 76.
In the collection there is found the greater portion of the
trunk of a sawfish, No. 4502 (3686). To what species this
may belong cannot now be determined. The shagreen gran-
ules of this trunk are small, and are furnished with two or
three longitudinal ridges and intervening furrows. In two of
the species of this genus here described, S. sentus and S.
liram, no shagreen is satisfactorily observable behind the
rostrum, while in S. solomonis the shagreen appears to be
wholly smooth. Hence, while it is improbable that the trunk
belongs to the latter-mentioned species, it cannot be con-
nected with either of the others. The shagreen of the trunk
resembles that of Pristis perrottett.
The structure of the species of Sclerorhynchus here de-
scribed, especially the evidence that the gill-slits opened on
the lower side of the head, shows that the genus is to be re-
ferred to the Pristide.
To whatever species this trunk may belong, it gives us a
clear idea of the form of the body of the Upper Cretaceous
sawfishes. And this form is remarkably like that of Pristis.
As 1s to be observed from the figure (Ply xxiv Fie )2) pune
body is long and slender. The portion at hand begins ap-
parently near the base of the pectoral fin and ceases about
the root of the caudal fin. The length of the part is 280 mm.
It is not certain that any portion of the pectoral fin is seen.
The depth of the body anteriorly is 47 mm. Both the dorsal
fins and one ventral are presented. All are triangular in form
and approximately of the same length. The base of the first
dorsal is 36 mm.; the height, 2t mm. The apparent height
of the ventral finis 14 mm. At the base of each fin are seen
impressions of the cartilaginous supports of the fin. At the
base of the first dorsal there are at least 16 rays; at the base
of the second dorsal, a somewhat greater number; at the
base of the anal, at least a dozen.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 399
The vertebre have been well calcified. In front of the first
dorsal there are present 24 vertebre; from the front of the
first dorsal to the front of the second dorsal, 34; behind the
latter, 20. The diameter of those beneath the first dorsal fin
is 5 mm.
In the abdomen of the specimen just described, is seen the
skull and most of the vertebral column of a bony fish, prob-
ably Eurypholis boisstert (Pl. xxiv, Fig. 1).
It is interesting to observe how closely the sawfishes of the
Upper Cretaceous resemble those of our own day in most of
their characters, and yet how primitive is the condition of
their rostral teeth. We cannot doubt that our modern species
of Pristis have descended from forms closely like those found
at Mount Lebanon.
No species known to belong to Pristis has, I believe, yet
been found in Cretaceous strata. In my work, ‘Bibliography
and Catalogue of the Fossil Vertebrata of North America,’
page 316, I have credited Pristis curvidens Leidy to the Cre-
taceous of New Jersey; but on examining the matter more
closely, I have concluded that the deposits from which Leidy’s
Specimens were derived belong really to the Eocene.
Sclerorhynchus solomonis, sp. nov.
PLATE XXV.
The specimen on which this species is based, No. 4503
(3706), consists of the rostrum complete or nearly so, the
head somewhat damaged, and a faint impression of one pec-
toral fin. Itis the uppe: surface of the head which is directed
toward the observer. The plate will impart a sufficiently
clear idea of the form and proportions of the parts.
Dr. A. S. Woodward has described and figured the type of
the genus Sclerorhynchus, S. atavus (Cat. Foss. Fishes, I,
1889, p. 76, pl. it, fig. 1). The only part which this author
had at his command was a portion of the rostrum, with the
teeth along its borders. That the present species is distinct
from S. atavus is evident from various considerations.
The tip of the rostrum is slightly damaged, so that there is
A00 Bulletin American Museum of Natural History. |Vol. XIX,
some doubt regarding a small patch of scales seen there.
Measuring from the mouth to this patch of scales, which
appear to form the tip, the distance is 150 mm. The borders
of the rostrum have been straight or slightly convex. At
50 mm. in front of the mouth the width of the rostrum is
50 mm.; at a distance of roo mm. the width equals 35 mm.
Beyond this the rostrum appears to have narrowed somewhat
more rapidly. A glance at the rostrum described by Dr.
Woodward shows that it was of a different form, the lateral
borders being concave. In S. solomonis, at a point 45 mm. in
advance of the mouth, the head begins to expand rapidly, so
that at a line slightly in front of the mouth the width is 100
mm. In S. atavus the width in a corresponding position
could not have been more than 75 mm.
The preservation of the rostrum is due to its being com-
posed of a mosaic of minute hexagonal calcifications, such as
we find in the same cartilages of Pristis. If there was a
shagreen overlying these cartilages, it does not now show
itself. A stellate shagreen is present along the sides of the
head as far forward as where these join the rostrum. Over
the base of the fin rays the shagreen scales are polygonal, con-
vex, smooth, and enameled.
The rostral teeth of this species are quite different from
those of S. atavus, in size at least. The longest of those rep-
resented in Dr. Woodward’s figure are 7 mm. long, and there
are about 4 of them in a distance of romm. In S. solomonis
there are no teeth more than 3 mm. long, and there are 8 of
them in 1o mm. These teeth have a stellate base, as in S.
atavus, and they appear to have been directed somewhat
backward. For some distance beyond the base, for one third
or one half of its length, the tooth is terete; then the diameter
is suddenly increased, forming a sort of shoulder. The re-
mainder of the tooth is gently curved backward, slightly flat-
tened, and brought to an edge on the convex border. It is
apparently only the distal portion of the tooth which is
enameled. Toward the extremity of the rostrum the teeth
are somewhat smaller. Posteriorly, the teeth become very
small and are hardly to be distinguished from some of the
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 401
shagreen scales on the border. Some of these scales are also
stellate.
There are some indications of gill arches and of the base of
the pectoral fin, but they are not distinct enough for descrip-
tion. On the right side is seen the outline of the mouth, and
a few small teeth appear.
This specimen was collected at Hajula.
Named in honor of Solomon, king, philosopher, poet, and
naturalist. ‘‘And he spake of trees from the cedar that is in
Lebanon even unto the hyssop that springeth out of the wall;
he spake also of beasts and of fowl and of creeping things and
of fishes.”
Sclerorhynchus hiram, sp. nov.
PLATE XXVI, FicuRE 1.
The type of this species consists of the head, with probably
most of the rostrum, the pectoral fins, and the anterior por-
tion of the vertebral column. Its number is 4501 (3705), and
it was collected at Hajula.
The extremity of the rostrum is missing. The portion
present extends 103 mm. in front of the mouth. The form of
the rostrum corresponds quite accurately to that of S. atavus.
From the latter species S. hiram is to be distinguished prin-
cipally by the size and structure of the teeth of the rostrum.
Those of S. atavus reach a length, according to Dr. Wood-
ward’s figure (Cat. Foss. Fishes, I, 1889, pl. iii, fig. 1), of
7 mm.; and there are not more than 4 of them in a length of
tromm. InS. hiram they are not more than 3 mm. long, and
about 8 of them are found in a distance of 10 mm.
In S. atavus each rostral tooth is said to comprise ‘‘a high,
round base, crimped, and having a somewhat stellate ap-
pearance when viewed from beneath; upon this is fixed a
backwardly directed crown, compressed to an anterior and
posterior edge.’”’ In S. hiram each enameled crown is fixed on
a high, round base, but this base is not crimped, but perfectly
smooth, and no evidences have been observed of any stellate
appearance. Each crown is curved, but, in addition to this,
[ Fune, 1903.| 26
402 Bulletin American Museum of Natural History. [Vol. XIX,
the crown as a whole is directed backward so as to make an
angle of 45° with the axis of the pedicel produced. As in S.
atavus, the crown is flattened to anterior and posterior edges.
Asin S. atavus, again, the teeth appear to graduate posteriorly
into the dermal scales; but this occurs at a greater distance
in front of the mouth than in the species from Sahel Alma.
At so mm. in front of the mouth the rostrum is 46 mm.
wide; at a distance of 100 mm., 35 mm. wide; at the mouth,
75mm. wide. The rostral cartilages have been well developed
and have left their impress on the matrix. These and all the
other cartilages present a mosaic of hexagonal calcifications.
From the point where these cartilages begin to narrow, in
front of the mouth, a band of stellate shagreen runs backward
along the sides of the head. In a few other places the sha-
green appears to have been circular in form and smooth. The
mouth resembles that of the living species of Pristis, and it has
a width of 50mm. The teeth are small, and some of them
appear to have been furnished with one or more sharp ridges,
one of which was doubtless the cutting edge.
The position of the gill arches is seen in the figure. On the
outer borders of each are seen the rays which supported the
gill septa. There can be no doubt that the gill slits opened
out on the lower side of the body, and that hence the genus
belongs to the Pristide.
This species is dedicated to Hiram, king of Tyre, the friend
of Solomon, who furnished for the latter cedar-trees and fir-
trees from the forests of Lebanon.
Sclerorhynchus sentus, sp. nov.
PLATE XXVII, Ficure tr.
The only portion of this fish that is known is a part of the
rostrum. This fragment, numbered 4504 (3864), has a total
length of 153 mm. The distal end of this rostrum is missing.
The proximal end of the specimen is supposed to have come
close to the mouth. From the distal end, where the width is
29 mm., the specimen expands gradually until, at a distance
of 80 mm. from the distal end, the width has become 45 mm.
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 403
From this point backward the width again diminishes until
at the proximal end the width is about 35 mm. It might be
supposed that immediately behind the widest part of the
rostral cartilages the shagreened skin would part from the carti-
lages and pass outward and backward to the sides of the
head. There is, however, no indication of such expansion of
the head, and a few teeth are found at a little distance behind
this broadest part of the rostrum.
The rostral teeth resemble most those of S. solomonts, but
they are still smaller, not exceeding 2.5 mm. in length, and
numbering 10 in ro mm. The base does not appear to be
distinctly stellate. From S. /iram the teeth differ in that
the enameled blade does not form any considerable angle with
the pedicel.
The form of the rostrum is quite different from that of S.
solomonts, in that it does not contract so rapidly toward the
distal end, and it has evidently been longer in proportion to
its width. The rostrum has been composed of small, hexag-
onal, smooth calcifications. Most of these are now removed
from the fossil, only their imprints remaining. A peculiar
feature, one not found in either S. hiram or S. solomonts, is
the presence of two rows of denticles throughout about the
anterior two-thirds of the fragment, one row on each side of
the midline. These denticles appear to have had a height of
about a millimeter. Only their bases are seen, the remainder
being buried in the matrix. The bases are stellate on their
hinder borders, but not in front. The denticles are placed
about 3 mm. apart. Whether they were on the upper or the
lower side of the rostrum I am unable to determine. These
doubtless belonged to the shagreen, and they appear to be all
of the dermal structures, except the teeth, that now remain.
It will, perhaps, be profitable to note certain differences in
the forms of the rostra of the three species described in this
paper. In S. hiram the rostral cartilages have a width, where
widest, of 44 mm., and the mouth is placed 55 mm. behind
this. In S. solomonis the greatest width is 55 mm., and the
mouth is only 40 mm. behind this widest portion. In S. sen-
tus the rostrum has a maximum width of 45 mm., and the
404 Bulletin American Museum of Natural History. |Vol. XIX,
mouth must have been at least 75 mm. behind this. As will
be observed from the figures, the rostrum of S. solomonts tapers
toward the distal end more rapidly than that of either of
the two other species.
Locality, Hajula.
RHINOBATIDA.
Rhinobatus eretes, sp. nov.
PLATE XXIV, FIGURE 2.
The type and only known specimen of this species was col-
lected at Hajula. It has the number 4500 (3715). As may
be seen from the figure, Pl. xxiv, Fig. 2, the specimen is quite
incomplete, neither the extremity of rostrum nor the tail
being present. The species has probably resembled R. tenu1-
rostris Davis in having had a much-prolonged snout, but this
is uncertain. There appear, however, to be characters which
are sufficient to distinguish the present form from the Sahel
Alma species just mentioned.
In general, the type has about the same size as the type of
R. tenutrostris. The distance from the pectoral arch to the
mouth is the same in the two; the distance between the inner
borders of the hinder lobes of the pectoral fins is a little greater
in R. eretes. The differences noted between the two species
are the following:
The concavity of the sides of the head opposite the gill
arches is considerably greater in R. eretes than in R. tenutros-
iris. In the latter the pectoral fin extends forward to a point
somewhat in front of the mouth; in R. eretes it lacks about 25
mm. of reaching a point opposite the mouth. In the type of
R. tenutrostris the pectoral measures, fore and aft, 180 mm.;
in R. eretes they measure 103 mm. It is thus seen that the
latter species has a much shorter fin in proportion to the size
of the animal.
It is evident that the ventral fins are also relatively smaller
than in RX. tenuirostris. In the latter the length of this fin
along the body is 75 mm.; in R. eretes the hinder extremity
of this fin is broken away, but the whole length could not have
been more than 60 mm. The anterior border of the ven-
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 405
tral of R. eretes is about 30 mm. and the breadth 20 mm., as
compared with 50 mm. and 25 mm. respectively in R. tenu-
rostrts.
We cannot be certain what was the form of the apex of the
pectoral fin of the species here described, since, after taking
into account the remnarits of the right and the left fins,
there yet remains about 30 mm. of the border unrepresented,
and this includes the apex.
The mouth of R. tenutrostris appears to have been smaller
than that of R. eretes, the former being said to measure about
33 mm. from side to side, while the latter measures 40 mm.
In the mouth of R. eretes there are seen several rows of teeth.
They each measure a millimeter in length parallel with the
jaw, and each has a thin cutting edge, in the middle of which
is a conical point.
The vertebre have a diameter of about 7 mm. Davis
states that those of R. tenutrostris have a diameter of 0.1 inch,
but this is obviously an error. The length of those of KR.
eretes is 2.6 mm.; and, according to this, there would be about
ts of them between the pectoral and the pelvic girdles. Davis
states that those of R. tenutrostris have a length of .15 inch,
and that there are 14 of them between the two girdles.
Over the greater portion of the body of the specimen here
described the shagreen has been removed. There is, however,
a band of stellate scales along the margin of the rostrum; and
further toward the midline, apparently on the upper side of
the rostrum, there are three or four rows of similar scales.
Along the greater part of the border of the front lobe of the
pectoral fin the shagreen has become smooth, and each scale
nearly circular in form. Probably this represents the general
character of the shagreen.
RAJIDA.
Raja whitfieldi, sp. nov.
PrATE SCoevVi Ent,
This new species is represented in the collection by three
specimens, No. 4505a (3707), No. 45056 (3708), and No.
406 Bulletin American Museum of Natural History. [Vol. X1X,
4505c (3709), all from Hajula. Of these the first-named is
taken as the type, inasmuch as it presents a greater portion
of the body than either of the others does. Even in this, the
tail is missing and a part of the right side is gone.
The disk is broad and rounded. The snout is slightly
drawn out, but its tip is rounded. The greatest width across
the pectorals is 156 mm. The distance from the snout to the
hinder border of the pectoral girdle equals 82 mm.; from the
snout to the pelvic girdle, 117 mm. There appear to be 15
vertebre between the pectoral and pelvic girdles.
The disk is everywhere covered with a very fine shagreen,
and no asperities are anywhere visible. On the upper surface
of the snout there are a few enlarged scales, each nearly 2 mm.
across. On the upper surface, on each side of the midline
and over the pectoral girdle, is a patch of scales, some of
which are about .5 mm. in diameter.
On each side of the head, where the scales have been broken
away so as to expose the mouth, may be seen a few small
teeth. The impressions of the gill arches are faintly seen;
likewise those of the eyes and the nasal cavities.
The other specimens add little or nothing to our knowledge
of the species.
Three other species of Raja have been described from
Mount Lebanon. Raja expansa (Davis), from Hakel, has a
very broad disk, and the pectoral fins are acute at their outer
angles. This species was regarded by Davis as belonging to
Rhinobatus (Trans. Roy. Dublin Soc., (2), III, 1887, p. 486,
pl. xviii). Raja primarmata A. S. Woodward (Cat. Foss.
Fishes, I, 1889, p. 85, pl. iv, figs. 1-3), from Sahel Alma, also
has the outer angles of the pectorals acute. Raja minor Davis
(op. ctt., p. 493, pl. xxi, fig. 2), from Sahel Alma, is either a
very small species or the young of a species otherwise un-
known. The pectorals are rounded. There is little or no
shagreen on the disk. The disk is very broad.
This species is named in honor of Professor R. P. Whit-
field, Curator of the Department of Geology of this Museum,
who is the author of many important memoirs on paleon-
tology, among them one entitled ‘Observations on some
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 407
Cretaceous Fossils from the Beyrut District, Syria, in the Col-
lection of the American Museum of Natural History, with
Descriptions of some New Species’ (Bull. Amer. Mus. Nat.
Hist., III, 1891, pp. 381-441, pls. iva—x).
Class PISCES.
BELONORHYNCHID?
Stenoprotome, gen. nov.
The writer finds it difficult to determine with any certainty
either the relationships or the characters of this genus. The
following characters are given provisionally :
Body furnished with large tuberculated bony scutes. Ver-
-tebral centra not developed. Head elongated, the snout slen-
der, obtuse at the apex. Teeth of moderate size, conical.
Opercular? bone furnished with a long, curved spine. Type,
Stenoprotome hamata sp. nov. Derivation of name, orevos,
narrow, and zpotouy, the face.
Stenoprotome hamata, sp. nov.
PLatE XXVI, FicurE 2; PuateE XXVII, FIGURE 2.
The specimen which forms the basis of the following de-
scription was obtained at Hakel, and the number is 4509
(3863). The head is the part most satisfactorily preserved,
and figures are here presented of both the counterparts. The
most striking feature of the fish is the possession of two long,
curved spines, one on each side of the head. Each of these
ends in a sharp point; and just proximad of the point is a
sharp barb. The distal end of the spine resembles closely the
point of a fishhook. The writer has not been able to deter-
mine conclusively what bone supports this spine. On looking
at Coccodus it is suggested to one that the spine is homo-
logous with the lateral spine of that genus, but further con-
sideration makes it evident that the present form has no
relationships with the pycnodonts; and the close attachment
of the spine to the side of the head and its evident great ex-
tension forward indicates that it is rather the opercular bone.
408 Bulletin American Museum of Natural Frstory. [Vol. XIX,
The length of the head, from the apex of the snout to a line
joining the hinder borders of the lateral spines, is 35 mm.
The apex of the snout is rounded and only 3.5 mm. wide.
The skull lies with the upper surface toward the viewer. The
bones are so closely united that their limits cannot be dis-
tinguished. The surface is almost everywhere covered with
tubercles, sometimes scattered, but usually arranged in rows
more or less regular. A few of those on the upper surface of
the snout form short spines.
Along the borders of the snout, for about 20 mm., there are
seen, at intervals of two or three millimeters, what appear to
be teeth, but which are possibly only enlarged marginal
tubercles. Between the larger ones are others of smaller size.
Some medium-sized ones are found at the apex of the snout.
At a considerable distance behind the head is seen a stout
bone bearing four or five teeth larger than those of the ros-
trum, the largest about 1.5 mm.in length. This bone seems
to the writer to be a portion of the lower jaw, which has
been displaced.
As may be seen from Pl. xxv, Fig. 2, orb., there are,
between the bases of the spines, two rings of bone. These
appear to be the sclerotic rings and to indicate the position
of the eyes; but, incomprehensibly enough, these have been
overlain by some bones of the upper surface of the skull.
The appearances of the fossil are not consistent with the sup-
position that the lower surface of the head is presented. There
is probably some distortion here.
Behind the bases of the spines the fossil contracts for about
to mm. Whether this region belongs to the skull or not is
not easily decided. There appear to be five bones here, an
elongated median one, extending the full length of the area,
and two others on each side. Of the latter, the hindermost
sends an arm forward along the outside of the more anterior
one. ‘These bones can be distinguished on the specimens only
by close inspection.
Behind the area mentioned, at a, Pl. xxvu, Fig. 2, there
is a bony mass whose surface is covered with ridges which
converge to a point at one side. These ridges may repre-
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 409
sent either the sculpture of a bony scute or a number of rays
of a dorsal fin. On one side of this, b, is a bony scute whose
axis runs obliquely to the axis of the head; while at c there
is another scute whose greater axis is transverse to that of
the head. Still farther away, at d, is found the supposed
lower jaw. Around this are some remains apparently of fin
rays. Finally, at e, is seen a very large bony plate, 33 mm.
long and 21 mm. wide. One end is narrowed and rounded
off. Near this plate also there are seen scattered fin rays.
All the plates are tuberculated. .
Nowhere are there any certain evidences of vertebral centra
orribs. There seem to be some evidences of neural or hemal
arches.
PYCNODONTID:.
Coccodus lindstreemi Davis.
PLaTE XXIX, FiGuRE 1.
Coccodus lindstremz Davis (J. W.), Jour. Geol. Soc., XLVI, 1890,
p. 565, pl. xxii—Woopwarp (A. S.), Cat. Foss. Fishes, III, 1895, p.
268.
Of this not well-known species there are in the collection
three specimens. One of these, No. 4517a (3698), with its
counterpart, presents the head. The second, No. 4517)
(3699), much damaged, shows a part of the head, with the
occipital spine, and a portion of the abdominal region. The
third, No. 4517¢ (3793), also presents the head and a com-
plete occipital spine. The first-mentioned specimen is here
neured (Pl xxix, Pig: rj) On the hinder border of the
spine there are 14 denticles. The region below the orbit has
been covered with bony plates, whose surface was ornamented
with more numerous and smaller tubercles than the other
portions of the head. Behind the occipital spine is seen a
series of fin rays, 10 in number. They are slender, and pre-
sent distinct evidences of segmentation, but none of longi-
tudinal division. They extend downward beneath the bony
covering of the region and toward the neural spines, or be-
tween them. No. 4517c (3793) shows the presence of the
410: Bulletin American Museum of Natural History. |Vol. XIX,
same rays, and here they seem to be longitudinally divided.
They appear to form a feeble anterior dorsal fin.
Through a fracturing of the snout some of the teeth, ap-
parently those of the splenial bone, are exposed. The rows
cannot be counted, but the teeth themselves are very much
smaller than those of Coccodus armatus. At the tip of the
lower jaw is seen a small, conical, pointed tooth.
The specimens are from Hakel.
Coccodus insignis, sp. nov.
PLATE XXIX, FIGURES 2-5.
Of this species there are in the collection several specimens,
all from Hajula. The following six are especially to be
mentioned: Nos. 4516a (3666), 4516b (3700), 4516¢ (3701),
4516d (3702), 4516¢ (3703), and 45167 (3794). Of these, No.
No. 45166 (3700) and No. 4516d (3702) are to be regarded as
the types. The former consists of a somewhat damaged fish
which has been flattened from above downward, and may be
taken as showing the form of the fish when seen from above.
Only the tip of the tail fin is missing. The total length is
80mm. On each side is seen a broad, hooked spine, a part of
the shoulder girdle. The head is pointed in front, and it and
the anterior body region expand backward to the ends of the
spines mentioned, so as to be wedge-shaped. These lateral
spines appear to be much broader than the corresponding
ones of C. armatus, as figured by Davis (Trans. Roy. Dublin
Soc., III, pl. xxx, fig. 1) and Woodward (Cat. Foss. Fishes,
III, p. 267). The anterior, or outer, border of each is finely
denticulated. The upper and lower surfaces are ornamented
with fine ridges, which start from the base and converge to
the point. Anteriorly these ridges become tuberculated. In
front and behind, the base of each spine passes into anterior
and posterior processes of the pectoral arch. Lying in the
curve of the hinder border of the pectoral spine and on the
matrix from which a portion of the spine has been removed
are seen abundant remains of pectoral fin rays; but these
have been much disturbed. The rays of the dorsal are so
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 411
much disturbed that their number cannot be determined;
but they are not numerous. Somewhat in front of a perpen-
dicular from the origin of the dorsal fin are seen the ventral
fins and their supports. The latter are 6 mm. long and rather
slender. So far as can be determined, there are only 5 fin
rays in each, and the outermost of these is short and claw-
like. The divided rays are also segmented. The anal fin is
disturbed, and the caudal is missing. Another specimen, No.
45167 (3794) (Pl. xxix, Fig. 2 v. f.), appears to have an
additional divided ray in the ventral fin.
In No. 45166 (3700) two or three rows of teeth can be seen.
Nothing more can be said of them than that they resemble
those of C. armatus. There are indications given by the
neural and hemal arches that there were 15 or more vertebral
segments.
The block bearing this specimen has been broken along the
length of the fish in such a manner as to expose the occipital
spine, which was buried in the matrix. This is represented
by Fig. 3 on Pl. xxix. It is readily seen to be different from
that of either C. lindstremz or C. armatus, being broad antero-
posteriorly and relatively short. The posterior edge is finely
denticulated; the anterior edge is nearly smooth. The lat-
eral surfaces are ornamented with fine ridges, which rise from
the base and either terminate in the borders or ascend to the
apex. Just behind this spine there is another process of bone
which may be either another spine or a ridge passing across
the rear of the skull. Its hinder border overhangs the an-
terior vertebre.
The co-type, No. 4516d (3702) (Pl. xxx, Fig. 4), is a small
fish having a length of 97 mm. from the snout to the end of
the caudal fin. The specimen is especially interesting because
it presents the shoulder girdle from below. It is difficult to
determine what sutures exist in this region. The lower ends
of the right and left halves of the girdle join in the midline,
and here the bones are 8 mm. wide, fore and aft. The suture
between them is very distinct for a part of the distance across
the bridge, but it then becomes indistinct. Whether or not
these bones are separated by suture from the bases of the
AIl2 Bulletin American Museum of Natural History. [Vol. XIX,
lateral spines I have been unable to determine with any cer-
tainty; but they are possibly distinct bones.
Dorsal and anal fins are present, but their rays are dis-
turbed.
What are probably vomerine teeth are presented. There
are three rows on one side of the midline and one row on the
other, and there were not less than 5 rows. Those of one of
the rows farther from the midline are compressed laterally
and each forms a longitudinal cutting edge. The other teeth
have a part of their triturating surface mammillated and the
borders finely crenulated.
' No. 4516 (3703) is a fish which has been spread out either
by crushing or by inflation by gases during decomposition.
The space occupied by the notochord is enlarged so that the
bases of the neural arches and those of the ribs are from 6 to
II mm. apart. This specimen shows that both the neural
arches and the hemal arches had their proximal ends ex-
panded against the notochord. The expansions of the neural
arches join and form a continuous covering for the upper side
of the notochord. The ends of the hemal arches probably did
not come into contact with each other or with the neuralarches.
Beneath the dorsal rays I count 1o interneural supports.
There appear to have been 8 anal rays. There are traces of «
both pectoral and ventral fins. In one gill chamber are seen
the impressions of four series of gill filaments.
No. 4516a (3666) shows the fish as seen from the side, and
thus gives us an idea of the elevation of the head and body.
The total length is r20 mm. The height of the body at the
pectoral spine is 30 mm. Only a faint impression of the
occipital spine remains. The rays of the dorsal fin cannot
be counted. The lower rays of the caudal are the longest and
the fin ends rather bluntly. The anal appears to comprise
8 rays. The ventrals are distinctly displayed. There is a
continuous line of bones along the upper side of the noto-
chordal region, the bases of the neural arches. Eleven neu-
ral arches are counted from the middle of the back to the
base of the caudal fin. Vomerine and splenial teeth are seen,
but the number of rows cannot be determined.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria, 413
No. 4516¢ (3701) lacks the hinder portion of the body (Pl.
XXIX, Fig. 5). The fish presents the body as seen from above,
but damaged somewhat, the roof of the skull being gone, as
well as the occipital spine. The head is 40 mm. long, from
the snout to a transverse septum formed apparently by the
shoulder girdle. Splenial teeth are present, three rows on
each splenial. Six teeth are found in each row, and there
were probably three or four more. The ones in front are
small. The inferior transverse portion of the shoulder girdle
is seen, on the left side, passing across beneath remains of
probably gill arches. Faint indications of pectoral fin rays
are seen on one side.
No. 45167 (3794) appears to deserve description and illus-
tration (Pl. xxix, Fig. 2). The inferior surface of the body
lies toward the viewer, and the apex of the occipital spine has
been found by excavating on the opposite side of the block.
From the base of one pectoral spine to the other, a bar of
bone crosses the body. It presents a fractured edge toward
the abdominal surface, and has, in all probability, been a
ridge, or plate, of bone which extended upward from the in-
ferior transverse portion of the shoulder girdle.
Behind the right (left in the figure) pectoral spine are seen
the neural arches with their conjoined expanded proximal
ends. Each arch is seen to have a wing-like expansion in
front of the spine. Crossing the upper ends of the hinder-
most spines are seen some interneural supports of the rays of
the dorsal fin. Immediately behind the bar of bone, passing
from one pectoral spine to the other, are seen some confused
ribs. Then come the supports of the ventral fins and the fins
themselves, and immediately behind and above these, the
expanded ends of the hemal arches. Behind each pectoral
spine are seen the remains of pectoral fin rays. Twelve of
these may be counted behind the right spine. The proximal
ends of these rays are to be seen on the matrix mesiad of the
base of the spine. This proves that the pectoral fins were
inserted above the spines.
In front of the transverse bar of bone mentioned above
is seen the parasphenoid. In front it appears to receive
414 Bulletin American Museum of Natural History. [Vol. XIX,
between two prongs the hinder end of the vomer. On the
latter there appear to have been four rows of teeth, but many
of these teeth are broken away.
The counterpart of specimen No. 4516c (3701) presents the
parasphenoid bone. Posteriorly this seems to join a basi-
sphenoid. On the lower surface of the latter, in the midline,
there is a short, pointed, downwardly directed process.
All the specimens were collected at Hajula.
ELOPID.
Holcolepis attenuatus (Davis).
Clupea attenuata Davis (J. W.) Trans., Roy. Dublin Soc. (2), III,
1887, p. 580, pl. xxxiii, fig. 4.
Osmeroides attenuatus Woopwarp (A. 8.), Ann. and Mag. Nat. Hist.
(7) II, 1898, p. 409; Cat. Foss. Fishes, IV, rgo1, p. 19.
No. 4526 (3781), from Hakel, is a specimen which the writer
is unable to distinguish from Davis’s Clupea attenuata, a
species hitherto known only from Sahel Alma. In the present
specimen the total length is 95 mm.; to the base of the
caudal fin,77 mm. The head, including the opercular appara-
tus, is about 23 mm., but the extremity of the snout is miss-
ing. The depth is only 15 mm. ‘There are a few more than
50 vertebre, of which not more than 20 belong to the caudal
region. ‘There are 20 interneurals supporting the dorsal fin.
Davis reports the presence of only ro rays in the dorsal, but
this is doubtless anerror. This fin, in our specimen, is equally
distant from the occiput and the base of the caudal fin. The
ventrals are inserted below the front of the dorsal. The anal
is disturbed, but there are 7 supporting interhemals present.
It is entirely behind the dorsal. The bones of the head are
smooth. Nothing can be determined regarding the size or
the form of the scales.
There are various reasons for not identifying this specimen
as H. sardinioides (Pictet). It is entirely too slender to be
AH. lewist (Davis). The dorsal and ventral fins of the latter
are farther backward, and there are said to be 35 vertebre in
the abdominal region behind the operculum.
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 415
Dr. A. S. Woodward has called our attention to the fact
that the name Holcolepis antedates Osmeroides as a name for
the species of this genus (Cat. Foss. Fishes, IV, p. 11).
ICHTHYODECTIDA.
Eubiodectes, gen. nov.
Teeth in sockets? Vertebre with lateral grooves; the
centra pierced by the notochord. Some of the anterior pec-
toral rays expanded distally, and longitudinally divided.
Anal fin elongated, falcate in front. Type, Cluirocentrites
libanicus Pictet and Humbert. Derivation of name, evBuos, .
well-living; and 8yxrys, a biter.
Eubiodectes libanicus, (Pict. and Hums.).
PLATE XXX, FIGURE I.
Chirocentrites libanicus PicteT and HumBert, Nouv. Rech. Poiss.
Foss. Mt. Liban, 1866, p. 88, pl. xiii—Davis (J. W.), Trans. Roy.
Dublin Soc. (2), III, 1887, p. 585.
Ichthyodectes libanicus Woopwarp (A. S8.), Cat. Foss. Fishes, IV,
IQOI, Pp. I05.
This species has been referred by Dr. Woodward to the
genus Ichthyodectes. However, a comparison with J. anaides,
the type of Ichthyodectes, will convince one that the Syrian
species belongs elsewhere. In J. anazdes the anterior pectoral
ray is very stout, but it does not expand distally; and I know
of no evidence that its distal end was split into fine filaments.
It formed rather a sort of spine, like that of Portheus. The
succeeding rays were considerably smaller than the anterior
one and were distally divided, as is usually the case with
such rays. In I. anaides the notochord seems not to have
passed continuously through the centrum of the vertebra;
but in the Syrian species under consideration there was free
communication between the concave ends of the centrum.
In this Syrian species the anal fin is very long; we have no
proof that it was so in the American species of [chthyodectes.
For these reasons I place the Chirocentrites libanicus of Pictet
and Humbert in a distinct genus. I have not seen the teeth,
416 Bulletin American Museum of Natural History. [Vol. X1X,
nor have I found any statement that they are lodged in
sockets. If they are not in sockets, this will furnish another
character to distinguish the species from [chthyodectes.
In the collection there are several portions of this fish, but
no complete specimen. None of these furnish the teeth. One
example, No. 4506a (3681), shows the fish from apparently
just behind the pectoral girdle to the base of the caudal fin.
This length is 400 mm. In this distance there are about 54
vertebre, but these have been disturbed anteriorly. The
depth at the origin of the anal fin is r20 mm. The anal fin
begins 186 mm. in front of the base of the caudal. There are
apparently about 8 strong rays in front; behind these the
rays are weak and, in the fossil, form an obscure fringe along
the body. There are altogether 34 interhzemal fin supports.
The whole length of the anal base is 137 mm.
So far as indicated by the specimen, the dorsal fin begins
Imo mm. in front of the base of the caudal fin and opposite
the beginning of the hinder third of the anal. Only ro rays
can be counted, but there may have been others posteriorly.
Large interneurals are present in front of the fin, but there
seem to have been no rays attached tothem. The neural and
hemal spines are long, slender, and sigmoid in form. The
vertebre both of the abdominal and caudal regions are some-
what higher than long, about 7 mm. long and 9 mm. high.
The side of the centrum is excavated by a pair of rather deep
pits or grooves. Where the centra happen to be split longi-
tudinally they show that the notochordal perforation is a
millimeter or more in diameter.
About 95 mm. in front of the dorsal fin, and therefore not
far behind the head, there is, on the back, a rounded mass of
bone. This seems to be connected with the distal ends of
some rather stout interneurals. This mass is about 27 mm.
long, 10 mm. high, and 5 mm. thick from side to side.
Most of the scales are removed from the fossil. They
appear to have been about 15 mm. high, and their exposed
portion about 8 mm. fore and aft.
Another specimen, from Hajula, No. 4506b (3646), con-
firms the above description, except that it does not include
1903.]| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 417
the region of the bony mass behind the head. There are
apparently 13 dorsal rays. A considerable portion of the
body is covered with scales. No. 4506c¢ (3799), from Hakel,
exposes on the matrix an impression of the caudal fin. It has
been deeply forked. The larger rays have been obliquely
segmented; the median ones split up into smaller filaments.
No. 4506d (3866) is part of a fish which had a depth of at
least 125 mm. It presents the ventral fins; but no other fins
are present to afford us a means of locating the position of
the ventrals. These fins appear to have been about 30 mm.
long.
No. 4506¢ (3823), from Hakel, is about as large as No.
4506a (3681). It seems to present some traces of the ven-
trals at a point 75 mm. in front of the anal fin. There are
also some faint indications of the bony mass on the back, to
which reference has been made above.
No. 3899, of the Cope Collection, is probably from Hakel.
The specimen resembles much the one figured by Pictet and
Humbert, but not so much of the body is present. One pec-
toral fin is well exposed, mostly as an impression on the ma-
trix (Pl. xxx, Fig. 1). The longest ray, the most anterior,
has a length of 65 mm., and its distal end is slightly more
than 7 mm. wide. The next ray, somewhat shorter, is 9 mm.
wide distally. The other rays are successively shorter and
narrower. All the rays, for a great part of their distal por-
tions, are longitudinally divided. There appear to be some-
what more than ro rays. No teeth are seen in this specimen.
CTENOTHRISSIDA.
Ctenothrissa signifer, sp. nov.
PLATE XXXI, FIGURES I AND 2.
Two fishes which were collected at Hajula belong evidently
to an undescribed species of Ctenothrissa. The number of
the type is 4521a (3651). With its counterpart, it furnished
all parts of the animal. A second specimen is numbered
45210 (3813).
[ Fane, 1903.] 27
418 Bulletin American Museum of Natural History. (Vol. X1X,
The total length of the type is 80 mm.; to the base of the
caudal fin, 55 mm. The body is deeper than it is in C. vexil-
lifer (Pictet), the greatest depth, 30 mm., being contained in
the length to the base of the caudal somewhat less than two
times; and in the distance from the pectoral arch to the base
of the caudal, one and one fourth times. The length of the
head, 21 mm., is less than the greatest depth of the body.
The height of the dorsal fin is fully 30 mm., and when de-
pressed its distal end would reach beyond the base of the
caudal fin. It contains 20 rays, including the anterior un-
divided ones. The anal fin has a height of 13 mm., but the
rays are broken and zigzagged, so that it is evident that
originally the fin was still higher. There are present in it
13 Or 14 rays.
The body is slightly turned, so that both ventral fins come
into view. These sweep backward along the lower border of
the body and cross all the anal fin-rays. The pectoral fins
are only dimly seen.
The maxilla is provided with short conical teeth. This
bone and the supramaxilla resemble those of C. radians, as
figured by Dr. A. S. Woodward (Cat. Foss. Fishes, IV, pl. x,
figs. 2, 3). The cheeks and the opercular bones are covered
with scales.
No. 4521b (3813) shows only the posterior half of the body.
The dorsal fin is depressed and overlaps the base of the caudal
fin. The anal rays pass somewhat behind the last caudal
vertebra. Some of the rays of the ventrals extend beyond
the origin of the anal.
This species differs from C. vexillijer in having a shorter
and deeper body, much higher dorsal and anal fins, and lon-
ger ventral fins. _
No. 4522 (3802) is a small fish from Hakel which I identify
as C. vextllifer. The length to the base of the caudal is 35
mm.and the depth is 14mm. The fish is therefore consider-
ably slenderer than the specimens of C. signifer. The pecto-
ral fin in this Hakel specimen also is long, as in those from
Hajula; but the dorsal is not more elevated than usual.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 419
DERCETIDi.
Leptotrachelus serpentinus, sp. nov.
PrATE XXX Frcure x.
This species is represented in the collection by two speci-
mens, No. 4511a (3683), which was collected at Hajula, and
No. 4511) (3739), which was obtained at Hakel. The first
mentioned specimen is regarded as the type. It lacks a large
part of the caudal region and the whole of the head, except
a part of the opercular apparatus. The length of the speci-
men in its present condition is 260 mm., and the total length
in life could hardly have been less than 300 mm., and was
probably more. The distance from the opercular region to
the ventral fins is 134 mm. The diameter (probably the
horizontal) at the ventral fins is contained in the distance
from the operculum to these fins thirteen times. In L.
triqueter the diameter at the ventral fins, as shown by Pictet
and Humbert’s figure (Nouv. Rech., pl. xiv, fig. 1), is con-
tained in the part of the body in front of the ventral fins about
seven times. We have in both of these cases apparently the
breadth of the body and not its height. L. serpentinus, there-
fore, appears to have been a much slenderer fish than L. tr7-
queter. This slenderness is shown also by the bodies of the
vertebrze, which are more than three times as long as the
diameters of their articular ends. In L. triqueter the bodies
are said to be twice as long as deep.
There are 31 vertebre between the operculum and the ven-
tral fins, about the same number as in L. triqueter. The ver-
tebral centra are much constricted. Each vertebra of the
abdominal region sends out on each side two long processes,
which diverge from the middle of the centrum. The broader
one is directed outward and forward. Near its end there is
articulated to it the head of a long, slenderrib. The posterior
and narrower process is directed outward and backward. Its
distal end approaches very closely the rib-bearing process of
the next vertebra behind. In the region of the ventral fins
these processes are nearly 5 mm. long. These lateral pro-
cesses are found on about 12 of the vertebre behind the ven-
420 Bulletin American Museum of Natural History. |Vol. XIX,
tral fins. The posterior process appears to be the one first
reduced, but both soon disappear. From the figures and de-
scriptions of L. triqueter we must conclude that there is only
a single process on each side of each centrum.
There are present about 6 or 7 rays of one pectoral fin, and
these are about 15 mm. long. A broad bone having a pos-
teriorly directed process lies just behind the operculum. This
may belong to the pectoral arch. The dorsal fin is missing,
unless it is represented by two or three rays which lie above
the ventral fins.
One of the ventral fins is present and appears to contain 6
rays. Its supporting bone is 7 mm. long and 2.5 mm. wide
posteriorly, and is pointed in front. No remains of the anal
fin are seen.
As may be seen from the figure, on the upper side and be-
hind the ventral fin, there is a row of triradiate dermal scutes
running along one side of the body. A similar row is found
on the other side, but the bone is broken away and only the
imprints of the scutes are left on the matrix. These scutes
continue for some distance in front of the ventral fins. They
appear to have a longer anterior branch than do those of L.
triqueter. There are also numerous fine intermuscular bones
throughout the length of the fish.
The condition of the vertebree and dermal scutes occupying
some distance behind the head is not easy to determine, on
account of the presence there of the bodies of three small
fishes. It is possible that these had been swallowed, but
their presence there is more probably accidental.
The specimen collected at Hakel is a fragment 98 mm.
long. It is from the portion of the body behind the ventral
fins. It presents no new features.
_ From Leptotrachelus gracilis Davis (Trans. Roy. Dublin
Soc., III, 1887, p. 623, pl. xxxviii, fig. 3), this species differs
in having a much less slender anterior abdominal region and
probably a longer post-pelvic region. Davis’s species likewise
had quite different lateral vertebral processes.
Collected at Hajula and Hakel.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria, 421
ENCHODONTID.
Enchodus marchesettii ? (Kramberger).
PLATE XXX, FIGURES 2 AND 3.
Eurygnathus marchesettii KRAMBERGER (D. G.), Djela Jugoslav.
mulcad. CVI, 1695, p. 34, pl. vi, eZ.
In the collection are two specimens of an Enchodus which
seems to be distinct from FE. longidens; and, since there are
no species of fishes known to be common to Sahel Alma,
where the latter species is found, and Hakel, where E. mar-
chesettt was found, it appears to be best for the present to
retain the two species as distinct. It is proper to state that
I have had no specimens of FE. longidens for direct compari-
son, and I have not been able to see Kramberger’s description
and figure of his species. The present identification is there-
fore wholly provisional.
The two specimens are numbered respectively 4507a (3779)
and 4507b (3859). The former (Pl. xxx, Fig. 2) presents the
head, except a portion of the lower jaw, and the body to the
rear of the anal fin, except a portion of the back. No. 4507)
(3859) consists of the body and tail from the beginning of the
dorsal fin. The two specimens have been almost identical in
size, and they supplement each other quite completely.
The head of No. 4507a (3779) is 55 mm. long to the hinder
border of the operculum. The lower jaw has been broken
away just below the tooth-line, leaving the teeth, but render-
ing it impossible to determine the depth of the jaw. The
teeth have been slender, some of them quite long; and they
are furnished with a few sharp grooves, especially distally.
The premaxilla has been of moderate size, and furnished with
a long, now missing, fang. The skull and opercular bones
appear to have been ornamented as in EF. longidens.
In front of the dorsal fin are three dermal scutes. These
are of an elongated oval form, with pointed ends. The an-
terior and largest is 8 mm. long and a little more than 3 mm.
wide. From the centre of each, ridges radiate to the circum-
ference. Below each scute there is seen a plate of bone which
422 Bulletin American Museum of Natural History. (Vol. XIX,
appears to be an expansion of an interneural bone. This has
probably formed a support for the scute. The first inter-
neural support of the dorsal fin is similarly expanded. Along
the position of the lateral line is seen a succession of small
thin scutes, each not more than 2mm.long. These are to be
observed as far backward as the rear of the dorsal fin, where
they lie close to the upper border of the vertebre. It is
probable that they continue to the base of the caudal. I
have observed no traces of the hooked dermal scutes that are
described as occurring at the base of the tail of some species.
The pectoral fin has had a length of at least 25 mm., and it
must have had somewhat more than 15 rays. The ventrals
arise about 30 mm. behind the pectorals and slightly behind
the origin of the dorsal fin. Its rays are somewhat damaged
in both specimens, but they were at least 17 mm. long. The
number of dorsal fin rays cannot be accurately determined,
since some are wanting in both specimens; but there were
probably 16, as in E. longidens. The anal fin was supported
by 19 rays, so that the fin does not appear to have differed
from that of E. longidens. The caudal fin is deeply forked,
and some of the external rays are rather strongly developed.
In the specimen showing the head there are 24 vertebre in
front of the origin of the dorsal fin, and 15 behind this point;
therefore 39 altogether, possibly 40. Dr. A. S. Woodward, in
his description of Enchodus longidens (Cat. Foss. Fishes, IV,
Pp. 199), states that there are 24 vertebre in the caudal region
of the latter species. In our specimens from Hakel, the
twenty-fourth vertebra from the base of the caudal fin will
be on a line joining the base of the ventral fins with the an-
terior portion of the dorsal. Not more than 19 or 20 verte-
bre can be fairly assigned to the caudal region. This may
furnish us with one difference between E. longidens and E.
marchesetttt. Another will probably be found in the different
forms of the dorsal scutes, those of E. longidens being broadly
oval, those of our Hakel specimens elongated oval.
Besides the specimens described above, there are in the
collection three others which probably belong to the same
species. One of these, No. 4527a (3735), is from Hakel; the
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 423
others, No. 45276 (3840) and No. 4527¢ (3832), are from Ha-
jula. The first-mentioned presents the head and the body to
behind the ventral fins; the two others, only the heads. In
the first, the lower jaw has a width of 11 mm. and a length of
45 mm. The teeth behind the anterior fangs appear to be
compressed to edges and to be striated and grooved. The
fangs are slender. No dorsal scutes are present. In No.
4527¢ (3832), the jaw is 12 mm. wide and apparently 55 mm.
long. The teeth, upper and lower, are mostly damaged, but
they are rather coarsely striated; and many of them, even
the large fangs, are furnished with edges.
No. 45276 (3840) and its counterpart furnish the head of a
large individual. The length of the head to the hinder bor-
der of the operculum is 120 mm. The length of the lower
jaw is 80 mm.; its depth is 21 mm. The head has been
crushed from above. Both tumid palatines are shown, each
with a long slender fang. The fang is nearly smooth on the
outer side, but coarsely striated on the inner side. Whether
cutting edges are present is not certain. The lower fang is
likewise striated on the inner surface. This belongs possibly
to a different species, but this cannot now be demonstrated.
Specimens from both Hakel and Hajula.
MYCTOPHID.
Osmeroides Agassiz.
Osmeroides Acassiz (L.), Poiss. Foss., V, pt. ii, 1844, p. 103.—
Pictet (F. J.), Poiss. Foss. Mt. Liban, 1850, p. 27.
Sardinioides Marck (W. v.), Zeitschr. deutsch. geol. Gesellsch., X,
1858, p. 245.—Woopwarb (A. S.), Cat. Foss. Fishes, IV, rgor, p. 236;
Foss. Fishes English Chalk, rgo02, p. 32.
Dermoptychius Marcx (W. v.), Paleontogr., XV, 1868, p. 287.
The employment of the name Sardinioides for this genus is
an evident violation of the law of priority; a law recognized
by all naturalists, but obeyed with reluctance by many when
their prepossessions are attacked; and the only means now
provided for deciding between rival systematic names. The
type of the genus is O. monasteri Agassiz.
424 Bulletin American Museum of Natural History. [Vol. XIX,
The species which are satisfactorily determined as belong-
ing to the genus are the following: Osmerotdes monastert
Agassiz, O. crassicaudus (Marck), O. megapterus Pictet, O.
pusillus (A. S. Woodward), O. woodward1, nom. nov. (= Sar-
dinioides attenuatus A. S. Woodward), O. macrophthalmus
(Marck), and the two new species below described, O. ponti-
vagus and O. ornatus.
Three other species of doubtful value, according to Dr. A.
S. Woodward, have been described by von der Marck, O.
macropterygius, O. minutus, and O. tenuicaudus.
Osmeroides pontivagus, sp. nov.
PLateE XXXIII, Figures 1-4.
This species is represented by at least 15 individuals. The
size is small, and the fish is closely related to O. puszilus,
which has been described as Sardinioides pusillus by Dr. A.S.
Woodward from Sahel Alma (Cat. Foss. Fishes, IV, p. 240,
pl. xvi, figs. 2, 3). O. pontivagus differs from the latter in
several important respects. Dr. Woodward states that in O.
pusillus the serrations of the scales are deep and conspicuous.
In O. pontivagus, on the contrary, they are very obscure, and
it is only in favorable situations on the matrix, and with the aid
of a good lens, that they can be seen at all. There are also
more rays in the dorsal fin than there are in the species from
Sahel Alma, 12 or 13, instead of 10 or 11. There are quite
certainly 9 rays in the anal fin. The ventrals are inserted
somewhat behind the origin of the dorsal fin, and they show
the presence of 7 rays. Several specimens present at least 13
rays in the pectoral.
The mouth of this species is large, the articulation of the
lower jaw being placed well behind the orbit. No. 4524)
(3846) has the mouth widely opened and shows the slender
and toothed premaxilla as forming the whole of the upper
border of the mouth (Pl. xxxiu, Fig.1). There is a patch of
teeth on the palatopterygoid arch. The maxilla is expanded
at the distalend. There were probably 8 branchiostegal rays
on each side. There are about 30 vertebra, not more.
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 425
No. 4524a (3845) (Pl. xxxi1, Fig. 2), is taken as the type
of this species. It has a length of 60 mm.; to the base of
the caudal, of 47 mm. The head is 16 mm. long, and the
depth is 20 mm.
Among the specimens which I am compelled to refer to this
species there is much variation in the depth of the body. On
Pl. xxxin, Figs. 1, 2, are represented two specimens which
have very deep bodies. Fig. 3 of the same plate shows an-
other specimen, No. 4524¢ (3855), whose body has less depth.
In this, the depth of the body, 15 mm., equals the length of
the head... The distance from the snout to the base of the
caudal finis 50mm. No structural differences are to be seen.
Still slenderer specimens occur, as No. 4524¢ (3852), which is
37 mm. long from the snout to the base of the caudal fin,
while the depth is only 1o mm. Between the extremes there
are all gradations in relative depth.
In some of the slenderer specimens there is a tendency tow-
ard a deepening of the fins. In No. 45247 (3841) (Pl. xxxi11,
Fig. 4), about 50 mm. long to the base of the caudal and 15
mm. deep, the rays of the dorsal fin extend backward two
thirds of the distance from the fin to the base of the caudal.
The anal is rather deep, while the pectoral rays fully reach
the base of the ventrals, and the rays of the latter lack but
little of attaining the front of the anal. In No. 4524¢ (3842)
from Hakel, the dorsal and anal fin-rays extend backward to
the base of the caudal.
In some, but not all, of the slender individuals, the scales
appear to be quite thick; but this condition may be due to
-some peculiarity of preservation.
The slender specimens described here resemble somewhat
O. woodwardt (= Sardintoides attenuatus), described by Dr. A.
S. Woodward, from Hakel (Cat. Foss. Fishes, IV, p. 241, pl.
xii, fig. 5). However, the latter is a more elongated fish,
having the length of the head contained in the distance be-
tween the pectoral arch and the base of the caudal fin three
times. It also has the scales conspicuously serrated, and only
Io or 11 rays in the dorsal fin.
All of the specimens, except two, are from Hajula.
426 Bulletin American Museum of Natural History. [Vol. XIX,
Osmeroides ornatus, sp. nov.
PLATE XXXIII, FicurReE 5.
No. 4518 (3870), from Hakel, appears to belong to an un-
described species of Osmeroides. Only a single specimen has
been found in the collection. The total length is 48 mm.;
to the base of the caudal 36 mm. The length of the head,
including the opercular apparatus, equals 14 mm. The
greatest depth is 12.5 mm. It will be seen, therefore, that
the depth is less than the length of the head, and is contained
in the distance from the pectoral arch to the base of the
caudal fin less than twice. There are 25 vertebre behind the
operculum; hence not more than 30 altogether. There are
II or 12 in the caudal region. The dorsal fin arises 7.5 mm.
behind the occiput. The number of its rays is uncertain, but
there are probably not more than 1o. The ventrals are in-
serted below the anterior half of the dorsal. The rays of the
anal cannot be counted, but the fin is short. The caudal is
deeply forked. The pectoral fins are delicate and inserted
well above the ventral border.
The scales are thick and deeply serrated, as may be seen
on the matrix, and even in some places where the scales over-
lie one another. The operculum is conspicuously ornamented
with coarse ridges and rows of tubercles, which radiate from
the articulation of the bone with the hyomandibular. There
seem to be similar ridges on the other opercular bones and
apparently on the cheeks.
The mouth appears to have been relatively small, the ar-
ticulation of the lower jaw being advanced to a perpendicular
line from the front of the orbit. The lower jaw is only 5 mm.
long. The orbit is rather large.
This species differs from O. megapterus, a Sahel Alma
species, in having fewer vertebre, 30 or fewer instead of 4o.
From O. woodwardi Hay (= S. attenuatus A. S. Woodward),
from Hakel, it differs in being less elongated and in having
coarsely serrated scales. It appears to resemble most O.
pusillus, described as Sardinioides pusillus by Dr. Woodward,
from Sahel Alma, but the latter is a more robust species, with
1903.] ay, Cretaceous Fishes from Mount Lebanon, Syria. 427
evidently a larger mouth. Nothing in the description indi-
cates that its opercular bones are ornamented like those of
O. ornatus. |
Collected at Hakel.
Acrognathus Agassiz.
Acrognathus Acassiz (L.), Poiss. Foss., V, pt. ii, 1844, p. 108.—
Woopwarp (A. §S.), Cat. Foss. Fishes, IV, r901, p. 243; Foss. Fishes
English Chalk, 1902, p. 36.
The type of this genus is A. boops Agassiz. The type speci-
men of this species is practically the only known example.
It is refigured in Dr. Woodward’s publication of 1902, cited
above. Unfortunately this author has not figured his A.
libanicus, from the Cretaceous of Sahel Alma. The following
species is believed to be congeneric with the one described
from Sahel Alma, but it is doubtful if either belongs to the
genus Acrognathus.
Acrognathus dodgei, sp. nov.
PLATE XXXIV, FIGURE 3.
The type of this species is No. 4520a (3673). It has a total
length of 88 mm.; to the base of the caudal fin68 mm. The
head and the opercular region together measure 20 mm. The
depth is 21 mm. The rays of the dorsal fin are somewhat
confused, but there appear to be 12 interneural supports.
The pectoral fin is wanting in the specimen. The ventrals
are slightly in front of the origin of the dorsal, but this is
probably due to slight displacement. The rays are stout.
The anal is somewhat damaged, but it was evidently short.
The vertebre number 32 or 33. The scales are thick.
The premaxilla is furnished with small teeth. The man-
dible is high posteriorly. Its length is ro mm. The articu-
lation is brought forward to beneath the eye.
At the base of the caudal fin, both above and below, are
about half a dozen reduced rays, and these are preceded by
a very short, but stout, modified ray, which appears to have
428 Bulletin American Museum of Natural History. (Vol. X1X,
been pointed at both ends. It resembles a similar ray seen
in the same situation in Microcelia day.
Another example, No. 4520b (3696), shows the ventrals
placed slightly farther backward than in the type. The pec-
toral fin has 10 rays. Other specimens, referred to the same
species, present the pectoral fin as placed considerably above
the ventral border.
It is probable that normally the ventral fins of this species
are inserted opposite the middle of the dorsal fin.
This species differs from A. boops Agassiz and A. libanticus
A. S. Woodward in having a shorter head and a smaller orbit.
All the specimens, about 10 in number, are from Hajula, ex-
cept No. 4520¢ (3836), No. 4520d (3767), No. 4520¢ (3730),
which are from Hakel.
Named in honor of Rev. D. Stuart Dodge, the donor of the
collection here described.
Nematonotus longispinus (Davis).
Pirate XXXIV, Ficure 2; PratE XXXV.
Pseudoberyx longispina Davis (J. W.), Trans. Roy. Dublin Soc.
(2), 31887; p-<522, pl. xv, figs 2:
Nematonotus botte (in part) Woopwarp (A. §.), Cat. Foss. Fishes,
IWS BOYS 7)s AGE
Among the fishes collected by Professor Day at Hajula there
is a considerable number which appear to me to be different
from Nematonotus botte, and which I have no means of dis-
tinguishing from the species described by Davis, as above
cited. The latter has been identified by Dr. A. S. Woodward
as N. botte; and his procedure may prove to be correct. If
so, the form here described will require a new specific name.
These Hajula specimens differ from N. botte especially in
the great length of one of the anterior dorsal fin-rays. It also
appears to attain a much greater size than does the species
from Hakel. Six specimens are especially to be mentioned,
as follows: Nos. 4510a-7 (3655, 3663, 3664, 3678, 3679, 3723).
No. 4510a@ (3655) has a total length of 95 mm. The pec-
torals and ventrals appear to be larger than they are in speci-
1903.] May, Cretaceous Fishes from Mount Lebanon, Syria, 429
mens of N. botte. The pectoral seems to have had 13 rays.
Of these the third is the largest, its breadth being 1 mm, and
it is broken off at a distance of 16 mm. from its base. The
ventral fin appears to have comprised 8 rays. These are
broken off at a distance of ro mm. from the base, but they
have evidently extended much farther. The dorsal probably
possessed 12 rays; there are 12 interneurals. Of the rays,
the third is greatly elongated, extending nearly to the tip of
the caudal fin. Inthe analthere are 8 orgrays. The caudal
is deeply forked. There appear to be 28 vertebre. The
scales are large. I count 4 above the lateral line.
No. 4510) (3663) has the caudal region and the caudal fin
wanting. The total length must have been close to 170 mm.
The head is 39 mm., the depth of the body, 47 mm. The
elongated dorsal ray is only 50 mm. long, but had doubtless
in life been longer. The pectoral rays are 24 mm. long, but
may originally have been longer. The stomach and intes-
tines are represented by their phosphatic contents, the former
having formed an S-like loop.
No. 4510¢ (3664) lacks the body in front of the dorsal fin
and most of the remainder of the body below the vertebral
column. This individual has had a total length of about
200 mm., a giant when compared with the described speci-
mens of N. bottg. From the fork of the caudal fin to the
front of the dorsalis 107 mm. The third dorsal ray has no
remarkable length, but it appears to have had the extremity
broken off.
No. 4510d (3678) (Pl. xxxiv, Fig. 2), lacks most of the
caudal fin. The specimen measures 70 mm. to the base of
the caudal. The third dorsal ray has a length of 56 mm.,
and would extend nearly to the extremity of the caudal fin.
The head is 24 mm. long; the body 37 mm. deep. The long-
est pectoral ray is nearly as long as the head.
No. 4510¢€ (3679) is a nearly complete fish, but it has suf-
fered some weathering. The total length is ro2 mm.; to the
base of the caudal fin 76 mm. The longest dorsal fin-ray
equals 62 mm. No. 45107 (3723) (Pl. xxxv), has a total
length of 170 mm. The elongated dorsal ray runs to the
430 Bulletin American Museum of Natural History. [Vol. XIX,
edge of the block bearing the fish; the part remaining meas-
ures 77 mm., and would have overlapped somewhat the
caudal rays.
Other specimens of the genus found at Hajula are referred
to N. botte. Some of these are quite certainly such, but
others may belong to N. longispinus; being either quite
young individuals, or having had the elongated dorsal ray
broken off. If the form here called N. longispinus is really
N. botte, it appears strange that more individuals have not
been found at Hakel, Davis’s specimen being the only one
found there possessing a greatly elongated dorsal.
Microceelia dayi, sp. nov.
PLATE XXXII, FIcuRE 3; PLATE XXXIV, Picure x.
There is in the collection a considerable number of speci-
mens of what must be regarded as a distinct species of the
genus Microcelia. Of these, No. 4525a (3692) is taken as
the type (Pl. xxxiv, Fig. 1). The total length of this-is 98
mm.; to the base of the caudal fin 75 mm. All portions of
the body are represented, although in places the scales are
flaked off, leaving only their imprint on the fine matrix. The
length of the head, including the opercular apparatus equals
23 mm. The depth of the body, greatest between the pec-
toral and the ventral fins, is 28 mm.; but other specimens
seem to indicate that this is not natural. From the shoulder
girdle to the base of the caudal fin is 53 mm. There are 42
vertebre behind the pectoral arch, of which 24 belong to the
caudal region. Altogether there are 49 vertebree. Each ver-
tebral centrum is strengthened by about 5 longitudinal ridges
on each side. The jaws are tightly closed, so that the teeth
cannot be observed. The gape is evidently large and‘con-
siderably oblique, the length of the lower jaw being 15 mm.
The orbit is large; the diameter of the eye, as shown by the
sclerotic bones, is 7mm. There are 25 or 26 rays in the dorsal
fin, including 5 short, undivided ones in front. I count 22
supporting interneurals. In the anal fin there appear to be
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 431
18 or 19 rays, but there are 20 interhemals present. The
middle of the back, from the occiput to the dorsal fin, is occu-
pied by a line of crimpings, such as have been mentioned by
Dr. A. S. Woodward in the case of M. libanicus. These have
the appearance of narrow folds or ridges, which are directed
upward and backward, each ending in a point. What the
relation of these crimpings is to the median row of scales has
not been determined.
The pectoral fins are missing from this specimen. The
ventrals are inserted somewhat in front of the origin of the
dorsal, but they may have been moved slightly forward
through distortion.
Another specimen, No. 4525) (3677), is a slenderer fish than
the type, but this elongation may not be wholly natural.
The head has a length of 19 mm., while the depth is only 16
mm. The length of the head is contained in the total length
of the fish 4 times, asin the type. The fin rays agree closely
with those of the type. The ventrals are placed beneath the
origin of the dorsal. Fifteen rays are counted in the pectorals
and 13 in the ventrals. In front of the interneural bones
which support the dorsal fin rays there are at least 12 others
which have no rays. These have the upper ends expanded
and turned somewhat forward. These are seen also in the
type of the species. Both specimens show also an enlarged,
free, and spine-like ray in the midline above and below and
just in front of the first caudal rays. It was wholly, or almost
wholly, buried in the flesh. There are on each side at least
9 branchiostegal rays.
No. 4525¢ (3798) is a third specimen of this species. It is
nearly complete, and it has a length of 84 mm.; to the base
of the caudal of 65mm. The head is 21 mm. long; the depth
is 19 mm. There are 49 vertebre. The origin of the ven-
trals is beneath that of the dorsal fin. There are apparently
26 rays in the dorsal, and these are supported by 23 inter-
neurals. The anal fin is supported by 20 interhemals. The
crimped scales in front of the dorsal are seen.
No. 4525d (3803) has a length of r10 mm. No. 4525¢
(3816) shows unusually well the crimpings in front of the
432 Bulletin American Museum of Natural History. (Vol. XIX,
dorsal (Pl. xxx1, Fig. 3, cr. sc.). These appear to consist of
squarish masses whose broad sides are applied to the similar
masses in front and behind, while the narrower sides look
outward and upward. There seem to be about three or four
_of such masses for each median scale.
All the specimens are from Hajula. The species is named
in honor of Professor Alfred Ely Day, of the Protestant
Syrian College at Beirut, who made the collection which is
described in this paper.
Rhinellus delicatus, sp. nov.
The type of this species is a small and probably young fish.
Its number is 4530 (3661). The total length is only 32 mm.;
from the snout to the base of the caudal fin is 26 mm. It is
also quite slender, the greatest depth being only 2.5 mm.
The head is injured, so that little except the upper and lower
jaws remains. The whole head seems to have been 7 mm.
long. The jaws are slender and are 4 mm. long, but it is not
certain that the extremities are not broken off. No teeth are
to be seen. Pectoral, ventral, dorsal, anal, and caudal fins
are all present. The pectorals have a length of 4mm. The
ventrals are inserted 4.8 mm. behind the base of the pectorals.
The number of the rays cannot be determined. The origin of
the dorsal fin is about 2 mm. behind that of the ventrals. The
rays are delicate, and those that can be counted are 8 in
number, but these probably do not represent the whole num-
ber originally present. The anal fin is placed halfway be-
tween the dorsal and the caudal fins. The number of its rays
cannot be determined. The caudal fin is deeply forked.
There appear to be 43 vertebre. These, especially the
more anterior ones, are somewhat longer than deep. No ribs
are seen. Scales are present, but their boundaries are indis-
tinguishable.
This fish differs from others of the genus in having the
origin of the dorsal fin much nearer the occiput than to the
base of the caudal fin.
This species comes to us from Hajula.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 433
GONORHYNCHIDA.
Charitosomus hakelensis (Davis).
Spaniodon hakelensis Davis (J. W.), Trans. Roy. Dublin Soc., III,
1987; DP. 501, pl. xxxiv, fig. 4.
Charitosomus hakelensis Woopwarp (A. S.), Ann. and Mag. Nat.
Hist. (7), II, 1898, p. 412; Cat. Foss. Fishes, IV, rgor, p. 274.
Some specimens of this species appear to have the ventral
fins situated immediately below the origin of the dorsal. No.
4523@ (3746) is from Hakel. It is much elongated and very
slender, but this form may be due to distortion. The an-
terior portion of the head is missing. The ventrals are placed
just below the origin of the dorsal. No. 4523b (3817) has
apparently 44 vertebre, and the ventrals are situated slightly
behind the front of the dorsal.
In the Cope collection there is a small lot of fishes from
Mount Lebanon. There is no record of the locality, but all
the species are those found at Hakel. Among these are two
specimens of this species. No. 3895 is a nearly complete fish,
only the anal fin and a portion of the caudal being gone. The
length to the base of the caudal is 82 mm. The elements of
the snout are injured, so that not much information regard-
ing them is to be obtained. Dr. A. S. Woodward’s conclu-
sions regarding the small size of the mouth are confirmed.
No teeth are observed. In this specimen also the ventral
fins are below the front of the dorsal. There are 44 verte-
bre. Where the scales are present they form an incrustation
whose elements cannot be distinguished. Where they are
flaked away from the matrix but little impression is left.
There are, however, faint lines which seem to indicate that
the scales were of an elongated diamond-shaped form, ending
behind ina sharp point. Nothing is seen to suggest the spiny
scales of Gonorhynchus.
The other specimen, No. 3894, is without the head and the
caudal fin. The length from the pectoral arch to the base of
the caudal has been more than roo mm. The ventral fins
are placed below the middle of the dorsal. Each ventral fin
has plainly 8 rays. There are at least 11 rays in the pectoral
[ Fune, 1903.] 28
434 Bulletin American Museum of Natural History. [|Vol. X1X,
fin. Where the incrustation of scales remains there is seen
a number of very distinct narrow, longitudinal, whitish lines
along the body. These probably indicate rows of scales. I[
count about ro of these lines below the vertebral column, and
apparently 5 or 6 above it. Possibly this species was longi-
tudinally striped. No additional information regarding the
scales is to be obtained.
ANGUILLID.
Urenchelys A. S. Woodward.
Urenchelys Woopwarp (A. S.), Ann. Mag. Nat. Hist. (7), V, 1900,
p. 322; Cat. Foss: Fishes, IV, rooxr, p. 337; Poss) Fishes; Buslichy
Chalk, 1902, p. 30.
Urenchelys germanus, sp. nov.
PLatE XXXVI, FicuRE 1; PLatE XXXVII, FIGURE 7.
There are several specimens of eels from Hajula which are
referred to this supposed new species. None of these are
complete fishes, more or less of the caudal region being miss-
ing in allof them. No. 4515a (3654) is regarded as the type
(Pl. xxxvi, Fig. 1). Of this individual, perhaps about 25
mm. of the extremity of the tail is gone. The remainder of the
animal is well preserved. Anteriorly the head and trunk are
viewed from above; from just in front of the anal fin back-
ward the fish lies on its side. This species is closely related to
U. avus, described by Dr. A. S. Woodward from the Upper
Cretaceous of Sahel Alma (Cat. Foss. Fishes, IV, p. 337, pl.
Xviii, figs. 1, 2), and there are few characters by means of
which it may be distinguished. _However, it appears that the
part of the trunk which lies in front of the anal fin has a
greater number of vertebre, and it is longer in proportion to
the length of the head than in U. avus. The latter is stated
to have about 35 vertebre in front of the anal, whereas U.
germanus possesses Over 40. This portion of the trunk is also
about 5 mm. longer than that of U. avus relatively to the
length of the head.
For these reasons I regard the Hajula specimens as form-
ing a distinct species.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 435
There is in none of the individuals any trace whatever of
the ventral fins, although the fishes must have been enclosed
in the matrix without any disturbance of their parts. This
is indicated by a brown stain on the matrix, which shows that
the outline of the body is unbroken in the region where the
ventrals would have been located. It is further evident that
they are not specimens of Anguillavus quadripinnis, since the
head of the latter is longer and more pointed; furthermore,
in the specimens referred to Urenchelys there are no indica-
tions of plates in the region of the lateral line.
The head of the type of U. germanus has a length of 24 mm.;
the portion of the trunk between the head and the anal
fin measures 46 mm., and the part of the caudal region re-
maining equals 74 mm. Between the occiput and the anal
fin I count 43 vertebre. The neural arches of this region are
broken away. Slender ribs project on each side. Behind
the origin of the anal fin are 47 vertebre, making go in all.
In case the caudal region was more than twice as long as the
portion of the trunk in front of the anal, as is the case with
U. avus, there must have been considerably more than 100
vertebre in the vertebral column.
The upper surface of the skull closely resembles that of
Anguilla. Vomer, ethmoid, united frontals, the parietals, and
probably the supraoccipital and epiotics can be located, al-
though the sutures cannot always be seen. The opercular
apparatus is well developed, showing apparently all the ele-
ments found in Anguilla, and possessing the peculiarities of
the latter genus. The lower jaw is two thirds the length of
-the cranium. The palatopterygoid bar seems to have the
structure seen in Angutllavus. Many teeth are seen on both
the maxilla and dentary.
Both pectoral fins are well displayed, and each had at least
18 rays. As already stated, there are no evidences of ventral
fins. The dorsal fin is seen to come as far forward as the 13th
vertebra from the occiput. Just behind the head there are
8 vertebre in 10 mm.; in the anterior portion of the cau-
dal region there are only 6 in this space. From this and
other specimens it is seen that the notochord passed uninter-
436 Bulletin American Museum of Natural History. [Vol. XIX,
ruptedly through the vertebral centra. The neural spines of
the caudal region are moderately stout and are smaller at
their bases than in the middle of their length. The hemal
spines are slender and tapering, and have long, delicate flesh-
bones attached to them.
No. 4515) (3762) appears to have only 40 or 41 vertebre in
front of the anal fin. In No. 4515c¢ (3695) we have what
seem to be the most anterior rays of the dorsal fin, and these
are placed over the 12th vertebra behind the occiput. An-
other specimen confirms this view.
No. 4515d@ (3790), from Hakel, probably belongs to this
species, since it is too large to belong to U. hakelensis. This
presents the vertebral column seen from above. In the case
of some of the vertebre the neural arches have slipped to one
side of their centra, so that the upper surfaces of the latter
are exposed. The right and left portions of each arch are dis-
tinct, never having been co-ossified (Pl. xxxvit, Fig. 7).
No. 4515¢ (3860) presents the head and the trunk to a
short distance behind the origin of the anal fin. Here also
there are 43 vertebre in front of the anal. The head is flat-
tened from above, and its hinder portion is broken away.
Numerous short blunt teeth are seen on the dentary and the
maxilla. There are also traces of vomerine teeth. Eleven
branchiostegal rays are counted on one side. They extend
far backward and the distal ends of some are curved up-
ward. Both pectoral fins are displayed, and each seems to
have possessed at least 20 rays. There are no traces of ven-
tral fins, although the conditions seem favorable for their pre-
servation had they existed.
ANGUILLAVIDA, fam. nov.
Apodes with well-developed cleithrum, pectoral arch, pec-
toral and ventral fins, and a distinct caudal fin. Dorsal and
anal fins extended. Palatopterygoid arch developed. Scales
rudimentary or absent; in some cases a row of enlarged plates
on each side, probably on the lateral lines. Ribs present.
One genus, Anguillavus.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 437
Anguillavus, gen. nov.
The characters of this genus are included in those of the
family. The type of the genus is Anguillavus quadripinmis.
Anguillavus quadripinnis, sp. nov.
PLaTE XXXVI, FIGURES 2 AND 3.
Of this species there is recognized in the collection only a
single specimen, No. 4512 (3796), collected at Hajula. This
specimen lacks the whole of the caudal region. It is lying
on the block with the dorsal surface directed upward. The
following dimensions are noted: Length of the head to the
occiput, 25 mm.; to the posterior border of the opercular ap-
paratus, 33 mm. Length from the snout to the thirty-eighth
vertebra, 90 mm.; height of the body, 14 mm.
The suspensory apparatus of the jaws is extended laterally.
Hyomandibular and quadrate are present, and from the dis-
tal end of the latter the mandible runs forward and projects
somewhat beyond the snout, as in Angutlla. Parallel with
the premaxillz on each side are the maxille. Starting from
the quadrate there runs forward on the left side a very dis-
tinct palatopterygoid arch. The pterygoid portion termi-
nates 7mm. behind the snout. For a great part of its length
this portion has a width of only one millimeter, and the width
is occupied by two distinct bones. Of these, the inner is re-
garded as the entopterygoid, the outer, as the ectopterygoid.
The anterior ends of these bones lie on the upper surface of
the parasphenoid, as do also the corresponding bones of the
other side; but this position is probably due to some shifting
during decomposition. Immediately in front of these ptery-
goids there is, on each side, a small, scale-like bone which may
be the prefrontal. Outside of these, underlapping the anterior
ends of the pterygoids, and extending forward toward the
snout, are two delicate bones, one on each side, and these I
regard as the palatines. The appearance of these bones in-
dicates that they were already much reduced. There are
almost certainly no teeth on the arch thus constituted. A
bone resembling a supramaxilla lies above the maxilla.
438 Bulletin American Museum of Natural History. [Vol. XIX,
Behind and mesiad of the palatopterygoid arch are seen
the anterior ends of the stout hyoids. Slender branchioste-
gals appear to be attached to these near their anterior ends.
Some of the upper branchiostegals are widened out at their
posterior ends and curled upward, but not so conspicuously
so as in Anguilla. The opercular bones appear to have re-
sembled those of Anguilla, but mostly only impressions of
these on the matrix remain.
The bones of the roof of the skull have been considerably
injured, so that the exact limits of the elements cannot be
determined.
Fragments of the cleithrum are present. The pectoral fins
are well preserved, and the rays, apparently 16 in number,
had a length of nearly 9 mm. The number of the baseosts
cannot be determined. ‘There are distinct evidences of the
scapula and the coracoid.
Both ventral fins are present, attached to their supporting
bones (Pl. xxxv1, Figs, 2, 3,v.7.). The latter are only slightly
more than 2 mm. long. The extremities of the fin rays are
broken off. There appear to have been about 8 rays in each
fin. These fins are placed 65 mm. behind the occiput, and
were doubtless only a short distance in front of the anal fin.
Owing to the position of the fish, the dorsal fin is not displayed.
There are 31 vertebre preserved, the last of which lies over
the ventral fins. The centra are constricted, and 5.5 of them
occupy the space of romm. The ribs are slender and rather
long, some of them s.5 mm.
There are evidences of the presence of two rows of bony
plates, one on each side of the body. These probably occupy
the position of the lateral line. On the right side these are
shown as impressions on the matrix, and, beginning just be-
hind the cleithrum, continue as far as the specimen is pre-
served. On the left side, they may be followed throughout
the greater part of the abdominal region, and in several places
are represented by thin bony scales. These plates, or scales,
are relatively large, six of them being found in ro mm. No
other scales or plates are seen on the body.
Collected at Hajula.
1903.] May, Cretaceous Fishes from Mount Lebanon, Syria. 439
Anguillavus bathshebe, sp. nov.
Prate XXXVII, Ficure tr.
The type of this species is No. 4513a (3704), while No.
4513) (3685) is regarded as the paratype. The type is from
Hakel, the paratype is from Hajula. No. 4513¢ (3868) is a
second specimen from Hajula.
The type is a nearly complete fish, wanting only the tip of
the snout and the extremities of the rays of the caudal fin
(Pl. xxxvu, Fig. 1). The paratype presents the hinder half
or more of the body. The total length of the type is 168 mm.
Two millimeters may be added for the missing portion of the
snout. The whole head would then measure, to the occiput,
12 mm.; to the cleithrum, 19 mm. The dorsal fin has evi-
dently had its origin 13 mm. behind the occiput, since there
the first rays are seen, and in front of these a stain on the
matrix plainly reveals the outline of the body. The dorsal
fin continues backward to near the caudal fin, but is plainly
distinct from the latter. The caudal is fan-shaped, rounded
at the extremity, and is supported by 5 or 6 hypural bones.
These characters are confirmed by the paratype. The anal
begins 59 mm. behind the occiput and continues to near the
caudal fin. The ventral fins have their origin 50 mm. behind
the occiput. In the type only one of these fins is preserved,
and an injury to the matrix has removed both the supports of
these fins. Another flaking away of the rock occurs just be-
hind the ventral; but these rays cannot belong to a detached
portion of the anal, since they have the structure of ventral
rays, and not that of anal rays. Moreover, the course of the
hinder portion of the intestine is shown by its fossilized con-
tents, and this passes above the rays in question. In the
paratype both ventrals are in their normal position and
attached to their supports. Each of these fins is about 5 mm.
long and is composed of 8 rays.
There are apparently 44 vertebre in front of the origin of
the anal fin and 57 behind its origin. The neural arches are
long and low, and each is overlapped in front by the preced-
ing arch. The centra are long, constricted in the middle
440 Bulletin American Museum of Natural History. [Vol. XIX,
of the length, and they are permitted the notochord to pass
continuously through them. In the abdominal region there
appear to be no neural spines. These are first seen as low
and rather broad plates just over the origin of the anal fin.
They soon become considerably higher and quite slender, but
expanding somewhat toward their distal ends. The hemal
arches are well developed, the spines of many of the anterior
ones being expanded at their distal extremities. Only the
merest traces of a few ribs are seen in the specimens.
The rays of the dorsal and anal fins are slender, and there
are, on an average, two and one third interneurals and inter-
hemals for each vertebra.
The cleithrum is preserved in its natural position. It is
sigmoid in form, and is pointed at both the upper and the
lower end. The substance of the pectoral fins is wanting, but
there is present what appears to be the imprint of one on the
matrix.
So far as preserved, the head resembles that of Anguzlla.
The hinder portion of the lower jaw is present, articulated to
the quadrate. It is not certain that any part of the palato-
pterygoid arch is seen. A strongly developed hyoid arch is
present, and to each side are attached at least 15 branchios-
tegal rays. These are long and slender, some of the upper-
most ones being broadened at their hinder ends and turned
upward. The opercular bone is probably represented by a
flake of bone.
Both the specimens appear to have been enveloped in a
fine matrix without any part having been disturbed. The
lower outline of the body is distinctly marked by a brown
stain. This stain and minute patches of similar stain on the
sides of the abdominal region may indicate the existence of
scales, but of these there is no other evidence. There are no
traces of enlarged scales along the sides of the body, such as
are found in A. quadripinnis.
This eel resembles rather closely Davis’s Urenchelys hakel-
ensis, and at first it was regarded as such. A closer study
has, however, made it certain that the two are distinct species.
In U. hakelensis the dorsal fin probably came forward to the
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 441
occiput. According to the published figures of U. hakelensis,
the vertebral centra are higher than long. This is confirmed
by the examination of three small specimens in this collection.
In A. bathshebe, on the contrary, the centra are longer than
high, longer, in fact, than the height of the centrum and the
neural arch taken together. The figures of U. hakelensis
show that the portion of the body in front of the anal fin
measures 38 mm., the portion behind this fin to the base of
the caudal, 53 mm. In A. bathshebe the two portions of the
body are practically equal. Of course, the presence of ven-
tral fins in A. bathshebe and their absence in U. hakelensis
constitutes the most important difference between the two
species; but the other differential characters given may en-
able us to distinguish specimens when the region of the ven-
tral fins is injured or absent.
Named in honor of Bathsheba, who attained the distinc-
tion of being a wife of one great king and poet and the mother
of another great king and poet.
ENCHELIIDA, fam. nov.
Apodes destitute of cleithrum, of all paired fins, and, so far
as known, of all median fins. Opercular apparatus greatly
reduced. Vertebral centra apparently diplospondylous. No
scales.
Enchelion, gen. nov.
Characters included in those of the family. Type E. mon-
tium. Derivation, eyxéAcor, a little eel.
Enchelion montium, sp. nov.
Pirate XXXVII, FicurRES 2-6.
The types of this species are No. 4514@ (3765) and No.
4514b (3766). These specimens are both from Hakel.
No. 4514@ (376s) (Pl. xxxvu, Fig. 2) presents a vertebral
column extending from the extremity of the tail forward to
an unknown distance behind the head. The length of the
442 Bulletin American Museum of Natural History. |Vol. X1X,
part of the column present is 143 mm. No. 4514b (3766)
(Pl. xxxvit, Figs. 3, 4) presents the head and 45 mm. of the
vertebral column. It has belonged to a somewhat smaller
specimen than the other. The striking feature of the verte-
bral axis of this form is that all the vertebre, from the head
to the tail, have been represented each by two rings or
“bodies,” apparently just such a condition as we find in the
middle of the caudal region of Amza among living fishes and
in that of Eurycormus of the Jurassic. That this condition
exists in the species before us is shown by the fact that
throughout the series only alternate vertebral centra possess
neural spines, while in the caudal region only those centra
bear hzmal spines which have neural spines.
In view of my results in the study of the vertebral column
of Amia (Field Columbian Mus. Pubs., Zodl. Ser., I, p. 37,
1895), I hold that a vertebral ring possessing a neural arch
and one without such an arch together constitute the equiva-
lent of such a vertebra as we find in ordinary fishes; and that
the archless ring belongs, not with the ring situated imme-
diately in front of it, but with the one just behind it. In the
species before us the rings, or “‘bodies,’’ which have no neural
spines appear to possess neural arches, and thus seem to differ
from the corresponding elements in Ama. However, in the
tail of Amza the rings which have no neural arches have their
upper halves formed from ossifications which develop on the
upper surface of the notochord and on each side of the mye-
lon; and there appears to be no reason why these ossifica-
tions should not sometimes grow upward and form an arch
over the myelon. Such an arch would not, however, be the
equivalent of the arch which develops the spine, since this
arch is formed, in Ama at least, by a pair of bones distinct
from the centrum on which it rests.
For reasons detailed in the paper referred to, I shall call
the ‘centrum’ which is devoid of either neural spine or
hemal arch an epihypocentrum; that provided with either or
both arches, a pleurohemacentrum. The former corresponds
with what Dr. Zittel and others call the hypocentrum; the
latter with what is called a pleurocentrum.
1903.] Hay, Cretaceous Fishes from Mount Lebanon, Syria. 443
In the species before us I find 17 vertebral rings in a length
of ro mm. in the larger specimen, and 23 in the same length
of the smaller one. The pleurohemacentra are usually some-
what shorter than their companion rings. Both have their
neural arches low and extending backward so as to overlap the
arch just behind it. The epihypocentra present a short back-
wardly directed spine and a shorter process which extends for-
ward somewhat over the archinfront. The pleurohemacentra
possess neither process. Both kinds of centra send downward
a rather broad, short process, that of the epihypocentrum
being apparently somewhat longer. No ribs are found ar-
ticulating with any of these processes. Both the pleurohe-
macentra and the epihypocentra are constricted about the
middle. I have not been able to determine whether or not
they permitted the passage of the notochord.
When we come to examine the caudal region we find the
pleurohemacentra not especially different from those of the
abdominal region (Pl. xxxvu, Fig. 6). The inferior processes
are, however, somewhat smaller. On the other hand, the epi-
hypocentra are furnished with well-developed hzemal arches.
@iese cam be seen to continue to near the tip of the taut
Altogether there are close to 275 vertebral rings preserved
in the larger specimen. Of these about 170 belong to the
caudal portion of the body. This would indicate that in the
tail there are 85 complete vertebre.
No traces are seen anywhere of either fin rays or of inter-
neural or interhemal bones. All were probably absent. The
extremity of the tail is shown only by a faint impression on
the matrix, and there are no traces of a caudal fin. Like-
wise, there are no indications anywhere of scales.
The head of this species appears to be remarkably small.
Its length in the smaller specimen, measured to the occiput,
is only 4.5 mm.; but, including the branchiostegal rays, 9.5
mm. Its heightis1.;mm. The snout appears to have been
conical. In the jaws are small teeth in more than a single
row. The branchiostegal rays are slender, much longer than
the skull, and the hinder ends are somewhat curved upward.
No opercular bones or pectoral arch is present.
444 Bulletin American Museum of Natural History. (Vol. XIX,
BERYCIDA.
Pycnosterinx levispinosus, sp. nov.
PLATE XXXVI, FIGURE 4.
This small species was found at Hajula. The number of
the type and only specimen is 4528 (3671).
This has been a very short-bodied, elevated, and compressed
fish. It is possible that distortion has somewhat diminished
its length, but of this there is no evidence. The length from
the snout to the base of the caudal fin is 29 mm., while the
greatest depth is 30 mm. The length of the head to the bor-
der of the operculum is 12.5 mm. The body has been cov-
ered with scales of moderate size. Whether or not these were
ctenoid is yet uncertain. There is some indication that they
were feebly ctenoid. The scales have also covered the bases
of the dorsal and anal fins.
The dorsal fin has been composed of 7 or 8 spines and ap-
parently 16 soft rays. Seven spines are present, but it seems
probable that the most anterior one has been eroded away.
The spines are stout and entirely smooth. The tip of the
hindermost has apparently been segmented, but its size
makes it proper to count it with the spines. Through erosion
of portions of the bone it is seen that the spines and the sup-
porting interneurals have been hollow, their lumen being now
filled with calcite. It is also revealed that the notochord
passed continuously through the vertebral centra.
The anal fin has had 3 or 4 spines and 10 to 12 soft rays.
It seems probable that the most anterior one has been weath-
ered away. These spines are stout and smooth.
The caudal fin has been forked. The pectoral fin is present
in its natural position, but the rays have been damaged. The
ventrals arise below the pectoral. The spine has been stout,
and' long enough to reach to the first soft rays of the anal fin.
It is now represented mostly by its imprint in the matrix.
From the other species of the genus found at Mount Leb-
anon, except P. dubtus, this species differs in the greater
number of dorsal spines and smaller number of soft rays.
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 445
From P. dubtus it differs in having fewer soft rays in the
dorsal and anal fins, and in having these smooth, instead of
ribbed.
The possibility that this fish is specifically identical with
the one here described as Azpichtys formosus has not escaped
the attention of the writer. Were the dorsal spines pres-
ent in the latter species, this question could be settled
more satisfactorily. It is regarded as belonging to Atpichtys
rather than to Pycnosterinx, because the scales are smooth-
edged, and there appear to be thickened scales along the
border of the abdomen. Aside from the generic differences,
Pycnosterinx levispinosus has almost certainly been a deeper-
bodied fish. If we measure the distance from the anterior
anal spine to the anterior soft rays of the dorsal in the two
fishes, we find that in P. levispinosus this measurement is
equal to the distance from the base of the caudal fin to the
snout; while in the case of Azpichtys formosus the dimension
will be equal only to the distance from the caudal base to the
front of the opercular apparatus. Again, the ventral spine of
P. levispinosus extends backward to the first soft ray of the
anal fin, while in A. formosus it reaches only to the first spine.
Lastly, there do not seem to have been any thickened scales
on the abdomen of P. levispinosus.
CARANGID.
Aipichtys formosus, sp. nov.
PLATE XXXII, FIGURES 2 AND 3.
Of this supposed new species there is in the collection only
a single specimen, No. 4519 (3831). This is incomplete, the
head and the anterior portion of the body being broken away
from about the articulation of the lower jaw to the middle of
the dorsal fin. As a consequence, various characters are un-
determined, and we cannot be wholly certain regarding the
generic position of the fish.
No dorsal spines remain in the specimen. Of the articu-
lated rays there are 13 present; possibly, but not certainly,
all that were possessed by the fish. The anal fin comprises
446 Bulletin American Museum of Natural History. [Vol. XIX,
4 short stout, spines and 12 articulated rays. The longest anal
spine is only 7 mm. long, and all the spines are entirely smooth.
The ray which I recognize as the first articulated one is very
stout and resembles one of the spines, but it is divided and
articulated at the distal end.
The base of the left pectoral fin is present. The pelvic fin
has its origin directly below the pectoral. The first ray is
long and thick, the length being 13 mm. and the extremity
reaching the anal fin. There are at least 5, and possibly 6
or 7, articulated rays; but the number cannot be definitely
determined. The caudal fin is forked, but the extremities
of the rays are not exposed. Seven branchiostegal rays are
counted. There are 14 caudal vertebre.
The scales are thin, and their hinder borders have a per-
fectly smooth edge. They are rather large, there being about
8 longitudinal rows above the vertebral column and 12 below
it. The greatest height of the body equals 38 mm.; the dis-
tance from the pectoral arch to the base of the caudal is about
28 mm.
This fish has been assigned provisionally to the genus
Atpichtys, but it possibly belongs to Acrogaster, of the Bery-
cide. Nothing is known regarding the structure of the head,
and the number of the pelvic soft rays is uncertain. The
rather large scales suggest Acrogaster; while, on the other
hand, there appears to be a series of thickened scales along
the lower edge of the abdomen, as in Atpichtys.
From all the described species of Azpichtys this differs in
having a smaller number of anal rays.
From Acrogaster heckelit this fish differs in having a greater
number of anal spines and a smaller number of articulated
rays.
Collected at Hajula.
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 447
List of the Fishes found at the Three Fish-bearing Localities of
Mount Lebanon.
Sahel Alma. Hakel. Hajula.
HEXANCHID.
Heptranchias? gracilis.| |
SCYLLIIDA.
Scylliorhinus elonga-
tus.
Scylliorhinus curti-
rostris.
Scylliorhinus tumidens.
Mesiteia sahel-alme.
LAMNID#.
Scapanorhynchus
lewisi1. |
Scapanorhynchus
elongatus.
Otodus latus. Otodus sulcatus.
SQUALID.
Squalus latidens
S. ? primevus.
SQUATINIDA.
Squatina crassidens. | |
PRISTIDA.
Sclerorhynchus atavus. Sclerorhynchus
solomonis. ;
Sclerorhynchus hiram.
re sentus.
RHINOBATID#.
Rhinobatus tenuiros- | Rhinobatus maronita. | Rhinobatus eretes.
tris.
Rhinobatus inter-
medius. |
Rhinobatus latus.
RAJIDZ.
Raja primarmata. ; ;
‘““ minor. Raja expansa. Raja whitfieldi.
DASYATID2.
Cyclobatis oligodac-
Cyclobatis oligodac-
tylus.
tylus.
Cyclobatis major.
448 Bulletin American Museum of Natural History. (Vol. X1X,
Sahel Alma.
Holcolepis gracilis.
os attenuatus.
blondeli.
elongatus.
latus. .
Thrissopteroides tenui-
ceps.
Thrissopteroides pul-
cher.
Istieus lebanonensis.
Spaniodon
oe
Histiothrissa crassi-
pinna.
Hakel.
BELONORHYNCHID#&?
| Stenoprotome hamata.|
PYCNODONTID2.
Coccodus armatus.
* lindstroemi.
Xenopholis carinatus.
Palzobalistum goedeli.
MAcCROSEMIIDA.
syriacus.
Petalopteryx
iq dorsalis.
OLIGOPLEURID.
| Spathiurus dorsalis.
ELoPID&.
Holcolepis attenuatus?
om sardinioides.
lewisi.
ae
ICHTHYODECTID.
CTENOTHRISSID®.
Hajula.
Coccodus insignis.
Holcolepis lewisi.
| Eubiodectes libanicus.| Eubiodectes libanicus.
Ctenothrissa vexillifer.| Ctenothrissa signifer.
Ovalis. -
CLUPEID2.
Pseudoberyx syriacus.
— botte.
grandis.
Scombroclupea
macrophthalma.
Scombroclupea
gaudryi.
Diplomystus
brevissimus.
Diplomystus birdi.
“e
Pseudoberyx syriacus.
Scombroclupea
macrophthalma.
Scombroclupea
gaudryi.
Diplomystus
brevissimus.
1903.| Hay, Cretaceous Fishes from Mount Lebanon, Syria. 449
Sahel Alma
Enchelurus syriacus.
Pronotacanthus sahel-|
alme.
Leptotrachelus
triqueter.
Leptotrachelus gracilis.
Enchodus longidens.
i major.
Pantopholis dorsalis.
Osmeroides megap-
terus.
Osmeroides pusillus.
Acrognathus libanicus.
Leptosomus macrurus.
Opistopteryx gracilis.
Rhinellus furcatus.
S ferox.
damoni.
oe
Charitosomus major.
oh lineolatus.
libanicus.
Chirothrix
ne lewisi.
Urenchelys avus.
[ Fune, 1903.]
Hakel
HALOSAURID&
NoOTACANTHID&.
DERCETID&.
Leptotrachelus
serpentinus.
Leptotrachelus
hakelensis.
ENCHODONTID&.
Enchodus marche-
settil.
Eurypholis boissieri.
Halec microlepis.
Prionolepis cata-
phractus.
Prionolepis laniatus.
MycTopHID&.
Osmeroides wood-
wardi.
Osmeroides ponti-
vagus.
Osmeroides ornatus. -
Acrognathus dodgei.
Leptosomus minimus.
Nematonotus bottz.
. longispinus.
Microccelia libanica.
GONORHYNCHID&.
Charitosomus
hakelensis.
CHIROTHRICID.
Telepholis? tenuis.
Exoccetoides minor.
ANGUILLID&.
| Urenchelys hakelen-
si
Ss.
Hajula
Leptotrachelus
serpentinus.
Enchodus marche-
settii?
Eurypholis boissieri.
Halec microlepis.
Prionolepis cata-
phractus.
Osmeroides ponti-
vagus.
Acrognathus dodgei.
Leptosomus minimus.
Nematonotus botte.
: longispinus.
Microccelia day.
Rhinellus delicatus.
lee eaaee minor.
| Urenchelys germanus,
29
450 Bulletin American Museum of Natural History. [Vol. XIX,
Sahel Alma. Hakel. Hajula.
ANGUILLAVID2.
Anguillavus bath- Anguillavus quadri-
shebe. pinnis.
Anguillavus bath-
shebe.
ENCHELIIDA.
| Enchelion montium.
BERYCIDZ.
Acrogaster heckel1.
= davisi.
Pycnosterinx russeg- Pycnosterinx levi-
geri. spinosus.
Pycnosterinx discoides.
gracilis.
dubius.
elongatus.
latus.
Hoplopteryx syriacus. | Hoplopteryx lewisi. Hoplopteryx lewisi.
stachei.
oblongus.
Dinopteryx spinosus.
ce
cc
(a3
ce
STROMATEID.
Orosoma sahel-alme.
a intermedium.
pulchellum.
ce
CARANGID.
| Aipichtys velifer. Aipichtys formosus.
minor. minor.
An examination of the lists given above shows that the
beds at Sahel Alma contain an assemblage of species which is
quite different from that found at either of the other localities;
furthermore, that the fauna of Hajula closely resembles that
of Hakel. From Sahel Alma there have been collected 62
species; from Hakel, 50 species; and from Hajula, 34 species.
Of those obtained at Sahel Alma it is not certain that a single
species has been collected at either of the other localities.
It is possible that Holcolepis attenuatus, of Sahel Alma, oc-
curs also at Hakel, and that Enchodus longidens is common
to all three localities; but this is not yet proved. On the
other hand, out of the 34 species found at Hajula, 20 are
1903.] May, Cretaceous Fishes from Mount Lebanon, Syria. 451
common to this place and Hakel. This indicates that the
fish-bearing beds of these two localities are on nearly the
same geological level. In case the deposits are not synchro-
nous, can we reach any conclusion regarding their relative ages?
We observe that the Hajula fauna is related to that of
Sahel Alma by the presence of three genera which are not
found at Hakel, viz., Sclerorhynchus, Rhinellus, and Pycno-
sterinx. Furthermore, that pycnodonts, especially abundant
during the Jurassic, are well represented at Hakel, but by
only a single species at Hajula; that, of Pseudoberyx, there
are three species at Hakel, and only one, so far as we know,
at Hajula. While too great reliance must not be placed on
these observations, they appear to indicate that the beds at
Hakel are the lowest; that these are followed at no great dis-
tance by those at Hajula; and that those at Sahel Alma are
considerably above the latter. That the latter beds are
higher in the series than those at Hakel has been the conclu-
sion of most geologists. This is further confirmed by the
occurrence of Macrosemiide, one oligopleurid, and many
pycnodonts at Hakel; while at Sahel Alma there are few or
none of these, but many Berycide, altogether 12 species.
The large number of sharks found at Sahel Alma, mostly be-
longing to living genera, may be regarded as indicating a
more recent time than that during which the beds of the
other localities were deposited.
Explanation of the Abbreviations employed on the Plates.
‘a. f. anal fin. d.r. dorsal fin ray.
ant. pr. anterior transverse pro- d.sc. dorsal scutes.
cess (Leptotrachelus). ep. hy. epihypocentrum.
br. branchiostegals. h. a. hemal arch.
br. ar. branchial arches. hy. hyoid.
c.f. caudal fin. hym. hyomandibular.
el. cleithrum. int. intestine.
eran. cranium. i. op. interoperculum.
er. sc. crimped scales(Microcelia). lat. sc. lateral scutes.
den. dentary. mes. pt. mesopterygoid.
derm. sc. dermal scutes. met. pt. metapterygoid.
adi dorsal fin. mo. mouth.
452 Bulletin American Museum of Natural History. (Vol. X1X,
mx. maxilla. pro. p. propterygium.
na.cp. nasal capsule. p. sp. pectoral spine.
nar. mares. pt. pterygoid.
n. sp. neural spines. qu. quadrate.
oc. occiput. rost. rostrum.
op. operculum. rost. t. rostral teeth.
orb. orbit. shag. shagreen.
par. parasphenoid. Ss. op. suboperculum.
pelv. gird. pelvic girdle. spl. t. splenial teeth.
pl. h. pleurohzemocentrum. v. a. neural arches.
ae
Pp. op. preoperculum. v.c. vertebral centra.
post. pr. posterior transverse v..f. ventral fin.
process. v. f. s. ventral fin support.
Mounr LEBANON CRETACEOUS FISHES
ie \
EXPLANATION OF PLATE XXIV.
Fic. 1.—Sclerorhynchus sp. undet. Page 398. Trunk from near pec-
toral fin to near root of caudal fin. #2. No. 4502 (3686).
d. j’, anterior dorsal fin; d. 7.”, posterior dorsal fin; v. f.,
ventral fin. Anteriorly is seen the skeleton of a swallowed
fish.
Fic. 2—Rhinobatus eretes Hay. Page 404. Part of head and part of
trunk. Type. X22. No. 4500 (3715). Or. ar., branchial
arches; mes. p., mesopterygium; met. p., metapterygium;
p. }., pectoral fin; pelv. gird., pelvic girdle; pro. p., pro-
pterygium; shag., shagreen; v./f., ventral fin; vert., verte-
bre.
NOH.
BuLietin A. M,
——ESEO
SeIqss
i Be o%
/
Pe oe.
5 Se
paid ajed
i
i
| as
i
FISHES
LEBANON CRETACEOUS
MouNT
r LEBANON CRETACEOUS Fisnns
2a sl J a
ss a
= — i ale A) ee ee ee ey
as a wy 4
ij al ee + a 7 ras nl
=a ed)
4 Coe hp
= ‘ %
Sclerorhynchus solomonis Hay. ie ie ede, an
cranium and pectoral fin. Type. X}. No. 4503 (37 6).
cranium; mo., mouth; pro. p., portion of propter gium
shagreen. On the left side of the rostrum are
rostral teeth. ae
.
ay
awe
BULLETIN A, M. N. H. Whojyes 2A 12 os Mina D< NOE
Mount LEBANON CRETACEOUS FISHES
a +e ‘ 2
~ an)
Wamalitg r (itn ie | Aas ia ais ’ /
Ae wey
Fay
aa
Lane
wt +
2
a ae
we
ae
©
BUILens A: M,N A,
mates il2
to t1sq bas bsoH .yop ogsT iy
(E088) Gozp .0
Mount LEBANON Caeracrous FIsuKs
EXPLANATION OF PLATE XXVI.
Fic. 1.—Sclerorhynchus hiram Hay. Page 401. Head, part of ros-
trum, and pectoral fins. Type. X#%. No. 4501 (3705).
br. ar., branchial arches; mo., mouth; mes. p., meso-
pterygium; met. p.,metapterygium; nar.,nares; p./f., pec-
toral fin; pro. p., propterygium; rost., rostrum; rost. t.,
rostral teeth; shag., shagreen.
Fic. 2.—Stenoprotome hamata Hay. Page 407. Head and part of
trunk. Type. X+. No. 4509 (3863).
Fic. 3.—Otodus sulcatus Geinitz. Page 397. Tooth wanting the tip.
Front view. X+. No. 4508 (3867).
Fic. 4—Side view of same tooth.
BULLETIN A: MW. N HH. VO. ek PLATE osoNow le
Mount LEBANON CRETACEOUS FISHES
5 a
M
; ~Botrerin A,
f
oa
wn
2
Ss)
=
]
~~
4%
zz
<
o
3)
|
=)
me
pm
=
“
EXPLANATION OF PLATE XXVII.
Fic. 1.—Sclerorhynchus sentus Hay. Page 402. Portion of the ros-
trum. Type. X+. No. 4504 (3864). The rostral teeth
are shown on the right side of the anterior end.
Fic. 2—Stenoprotome hamata Hay. Page 407. Head and several
dermal scutes. Type. X+. No. 4509 (3863). 4a, 0, c, @,
dermal scutes; d, supposed lower jaw.
SEX, Prare, Xvi
VoL.
INS Jal,
M,
BuLiETIN A,
" bey, OF
ho SL A a
LEBANON CRETACEOUS FISHES
MouNtT
Beurierin A. M. N. H,
Mount LEBANON CRETACEOUS FISHES
7 \ ;
a a ,
° 4 7 m 4 Ah ae - A
> -
a a ae a 4
= - -
as
’
By
oe EXPLANATION OF PLATE XXVIII.
Raja whitfieldi Hay. Page 405. Head and trunk. Type.
; 4505b (3708). mo., mouth; na. cp., nasal capsule;
- fin; rost., rostrum; v. f., BES fin; vert., vertebra.
a i
Burretin A. M. N. H. Vor. XIX, Plate XXVIII.
Mount LEBANON CRETACEOUS FISHES
2 F
i
;.
|
4 P «
g FO “wo
Sed .etvsd 45
te
ot enitnew det .grp, bed ee argent aS
stom ,.qz jmit Isetob Xb -(poye) \Orep .o“% 4X
we , “i Men voit Istotosq ..{ -4 ;bromedqesisq ...wy ;smiqe
=? droga nit [srt cov ee
Tees ‘atte Istiqioo to wsiV OIE. 43st .vsH
G2 20 jtyqiono ..20 .(ooys) ddtz, .o%~
, ae “it Isztob
po8T .ysH zsrgtens ewhomol0—.¢ .017
(10ve) al oe 1X
Mount LEBANON CRETACEOUS Pieri
laine
Fic.
EXPLANATION OF PLATE XXIX.
1.—Coccodus lindstremi Davis. Page 409. Head. Xt. No.
4517@ (3698). d. f’., supposed anterior dorsal fin; oc. sp.,
occipital spine; orb., orbit.
. 2.—Coccodus insignis Hay. Page 413. Fish wanting the tail.
X+. No. 4516f (3794). d. f., dorsal fin; . sp., neural
spine; par., parasphenoid; p. 7., pectoral fin; v. 7., ven-
tral fin; v. 7. s., ventral fin support.
3-—Coccodus insignis Hay. Page 410. View of occipital spine.
Cotype. X}. No. 45266 (3700). 0€., eGcipub; 06. SPp.,
occipital spine.
Fic. 4.—Coccodus insignis Hay. Page 411. View of fish from below.
Fic.
Cotype. Xt. No. 4516d (3702). a. f., anal fin; d. 7.,
dorsal fin.
5.-—Coccodus insignis Hay. Page 413. Head and part of trunk.
X}. No. 4516¢ (3701). p.f., pectoral fin; p. sp., pectoral
spine; spl. t., splenial teeth.
BurLerin A. M. N. H. Vor
. XIX, Plate XXIX.
Mount LEBANON CRETACEOUS FISHES
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EXPLANATION OF PLATE XXX.
Fic. 1.—Eubiodectes ibanicus (Pict. and Humb.). Page 417. Pec-
toral fin. XX}. No. 3899, Dept. Vert. Pal.
Fic. 2.—Enchodus marchesetttt 2 (Kramb.). Page 421. Head and
trunk. xX}. No. 4507a (3770). Go7., anal in; 4.7,
dorsal fin; d. sc., dorsal scutes; Jat. sc., lateral scutes; p. j.,
pectoral fin; v. f., ventral fin.
Fic. 3.—Enchodus marchesettii ?(Kramb.). Page421. Trunkand tail.
X+. No. 4507b (3859). a. f., anal fin; v. f., ventral fin.
Burrerin A. M. N. H. MOr Exo Pram sxexuxe:
Mount’? LEBANON CRETACEOUS FISHES
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EXPLANATION OF PLATE XXXI.
Fic. 1.-—Ctenothrissa signijer Hay. Page 417. Completefish. Type.
X1. No. 4521a (3651). a. f., anal fin; br., branchios-
tegals; den., dentary; op., operculum; v. 7., ventral fins.
Fic. 2.—Counterpart of same fish as above. pp. f., pectoral fin; mx.,
maxilla.
Fic. 3.—Microcelia dayi Hay. Page 431. Head and part of trunk.
Type. Xt. No. 4s25e (3836). a. fs) anal Gams crease.
crimped scales; den., dentary; d.f., dorsal fin; zmt., intes-
tine; op., operculum; »p. 7., pectoral fin; v. f., ventral fin.
LX. Pras XX
VoL,
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BuLitetin A, M,
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CRETAC
LEBANON
Mount
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EXPLANATION OF PLATE XXXII.
Fic. 1.—Leptotrachelus serpentinus Hay. Page 419. Trunk. Type.
x+. No. 4511a@ (3683). ant. pr., anterior transverse pro-
cesses of vertebrae; derm. sc., dermal scutes; op., oper-
culum; p. 7., pectoral fin; post. pr., posterior processes of
vertebree; v. f., ventral fin.
Fics. 2 and 3.—Azpichtys formosus Hay. Page 445. Fish and its
counterpart, without head. Type. X+. No. 4519 (3831).
a. f., anal fin; br., branchiostegals; op., operculum; p. }.,
pectoral fin; v. 7., ventral fin.
BuLretin A. M. N. H. Vor. XIX: PrarE XXXII.
SITIES RIS PTT OS
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EXPLANATION OF PLATE XXXIII.
Fic. 1.—Osmeroides pontivagus Hay. Page 424. Complete fish. X+4.
No. 45240 (3846). a. f., anal fin; br., branchiostegals;
den., dentary; d. f., dorsal fin; hy., hyoid; mx., maxilla;
p. f., pectoral fin; pmx., premaxilla; v. f., ventral fin.
Fic. 2.—Osmerotdes pontivagus Hay. Page 425. Complete fish. Type.
X+. No. 4524a (3845). a.f.,analfin; den., dentary; d. f.,
dorsal fin; p. 7., pectoral fin; v. f., ventral fin.
Fic. 3.—Osmeroides pontivagus Hay. Page 425. Completefish. xX.
No. 4524¢ (3855). a. f., anal fin; den.,,dentary; d.*7.,
dorsal fin; p. 7., pectoral fin; v. 7., ventral fin.
Fic. 4.—Osmeroides pontivagus Hay. Page 425. Completefish. x}.
No. 4524f (3841). a. 7., anal fin; br., branchiostegals;
den., dentary; d. j., dorsal fin; p..f., pectoral fin; v. 7.,
ventral fin.
Fic, 5.—Osmerotdes ornatus Hay. Page 426. Type. X+. No. 4518
(3870). a. f., anal fin; d. 7.,dorsal fin: v7. ventral fin.
Buiretin A. M. N. H. Vor. XIX. Pratre XXXIII.
Mount LEBANON CRETACEOUS FISHES
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EXPLANATION OF PLATE XXXIV.
Fic. 1.—Microcelia dayi Hay. Page 430. Complete fish. Type. XH.
No. 4525a@ (3692). a. f., anal fin; den., dentary; d. }.,
dorsal fin; p. f., pectoral fin; v. 7., ventral fin.
Fic. 2.—Nematonotus longispinus (Davis). Page 429. Fish wanting
tail. X+. No. 4510d (3678). a.f., anal fin; den., dentary;
d. f., dorsal fin; orb., orbit; p. f7., pectoral fin; v. f., ventral
fin.
Fic. 3.—Acrognathus dodgei Hay. Page 427. Complete fish. Type.
X+. No. 4520a (3673). a. f., anal fin; br., branchios-
tegals; den., dentary; d. f., dorsal fin; mx., maxilla; orb.,
orbit; pmx., premaxilla; v. f., ventral fin.
3ULLETIN A, M. N. H. VoL, XIX, Pratre XXXIV.
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S35 Fa
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EXPLANATION OF PLATE XXXV.
Nematonotus longispinus (Davis). Page 429. Complete fish. 4.
No. 45107 (3723). a.f., anal fin; den., dentary; d.r., long dorsal
ray; op., operculum; p./7., pectoral fin; v. 7., ventral fin.
BuLreTin A. M. N. H. VoL. XIX, PLATE XXXV-_
Mount LEBANON CRETACEOUS FISHES
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Mount LEBANON CRETACEOUS FISHES
EXPLANATION OF PLATE XXXVI.
Fic. 1.—Urenchelys germanus Hay. Page 434. Nearly complete fish.
Type. X+. No. 4515a (3654). a. f., origin of anal fin;
cl., cleithrum; den., dentary; d. f., origin of dorsal fin; hy.,
hyoid; hym., hyomandibular; 7. op., interoperculum; mx.,
maxilla; op., operculum; pal., palatine; p. f., pectoral fin;
p. op., preoperculum; pt., pterygoid; qu., quadrate; s. op.,
suboperculum.
Fic. 2.—Anguillavus quadripinnis Hay. Page 437. ‘Anterior half of
fish. Type. X+. No. 4512 (3796). 06r., branchiostegals;
den., dentary; hy., hyoid; int., intestine; lat. sc., lateral
scutes; mx., maxilla; op., operculum; ». f., pectoral fin;
pt., pterygoid; qu., quadrate; v. 7., ventral fins.
Fic. 3.— Ventral fin of the same specimen as Fig. 2. Page 437.
Enlarged 7 diameters. v. f., ventral fins; v. 7. s., ventral
fin supports.
Fic. 4.—Pycnosterinx levispinosus Hay. Page 444. Complete fish.
Type. Xi. No. 4528 (36772).
oe
Butietin A, M. N, H. Vou. XIX, Prats XXXVI-
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Mount LEBANON CRETACEOUS FISHES
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EXPLANATION OF PLATE XXXVII.
Fic. 1.—Anguillavus bathshebe Hay. Page 439. Complete fish. Type
Xt. No. 4513a@ (3704). a. f., anal fin, origin and end of;
br., branchiostegals; c. 7., caudal fin; cl., cleithrum; den..,
dentary; d. f., dorsal fin, origin of; hy., hyoid; zn#., in-
testine; p. f., pectoral fin; v. 7., ventral fin.
Fic. 2.,—Enchelion montium Hay. Page 441. Fish wanting head.
Cotype. X+. No. 4514a@ (3765).
Fic. 3.—Enchelion montium Hay. Page 442. Head and anterior part
of trunk. +. No. 4514b (3766).
Fic. 4.—Part of same specimen as Fig. 3. Page 442. Enlarged 4
diameters. or., branchiostegals; den., dentary.
Fic. 5.—Part of same specimen as Figs. 3 and 4. Page 442. En-
larged 4 diameters, showing alternation of epihypocentra
and pleuro-hemacentra.
Fic. 6.—Part of same individual as Fig. 2. Page 443. Vertebre 88
mm. behind anterior end, enlarged 9 diameters. ep. hy.,
epihypocentra; ha., hemal arches; n. sp., neural spines;
pl. h., pleuro-hemacentra.
Fic. 7.—Urenchelys germanus Hay. Page 436. Portion of vertebral
column. 3. No.4515d (3790). vw. ¢., vertebral centra;
v. a., displaced neural arches.
Vor. XIX, Prate XXXVII.
BuLtetin A. M. N. H.
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,
Species of Fossil E:dentate from
Santa Cruz Formation of Patagonia. —
~ By Barnum Brown,
Vou.. XTX, ARTICLE X¥,-pp.- 453-457.
New York, Fuly 8, 1903.
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Article XI.— A NEW SPECIES OF FOSSIL EDENTATE
e FROM THE SANTA CRUZ FORMATION OF
i PATAGONIA.
i. By Barnum Brown.
In the autumn of 1898 the American Museum of Natural
_ History made arrangements with the Princeton University
Museum for the writer to accompany the third expedition to
Patagonia, under the leadership of Mr. J. B. Hatcher. The
expenses of this trip were largely defrayed by Professor Henry
Fairfield Osborn.
A large collection, comprising nine nearly complete skele-
tons and nearly a hundred skulls with skeletal material, was
F secured by the writer from exposures on the Rio Gallegos, on
Bs the seashore south of Rio Coy, and from the bluffs along the
coast south of Rio Santa Cruz. This material represents most
A of the families that lived in such great numbers during the
; Santa Cruz period, and contains several new species.
By previous arrangement with Professor Scott and Mr.
“_ Hatcher, the portion of this material belonging to species
eS already known will be described in the Princeton memoirs,
Es while the new species will be described in the American Mu-
= seum Bulletin.
Ra Pai ci ae
@ Eucinepeltus complicatus, nov. sp.
a _ This species is founded on an adult skull with cephalic
r -_. shield, No. 9248 of the collection of the American Museum of
Natural History. The type was found in the talus of cliffs on
Rio Gallegos, near Mr. Felton’s residence.
Comparison with the type of the genus, Eucinepeltus pete-
satus, has been possible through the kindness of Professor W. B.
Scott, the advance sheets of whose memoir I have examined.
It differs from the type of the genus in the following charac-
ters:
Eucinepeltus petesatus. Eucinepeltus complicatus.
Pattern of teeth: rst to 3rd non-lobate. 1st to 3rd-lobate.
Cephalic shield: 9 plates, not all pitted. 11 plates, all pitted.
[453]
454 Bulletin American Museum of Natural History. [Vol. XIX,
The cephalic shield is composed of eleven plates in four
rows, arranged in the following order from the anterior to the
posterior end: 2 in the first, 3 in the second, 4 in the third,
and 2 in the fourth rows. Each plate has a central pit which
is of pronounced character in the two median plates of the
third row. These pits have a circular shape, with raised
margin, very rugose sides, and a small cone at the bottom of
the pit.
The sutural borders present a prominent ridge, very rugose,
with deep paired holes on either side of ridge, excepting the
sutures separating the four posterior median plates which are
well defined, but do not show raised edge or holes.
The outline of the shield is not as circular as in E. petesatus,
and the border is more distinctly emarginated at the junction
of the first and the second rows. ‘The plates in the first row,
also the median plates of the second row, in the present
species are smaller than in EF. petesatus, while the four pos-
terior median plates are relatively larger.
The teeth differ from those of FE. petesatus, especially in the
anterior part of the jaw. The first molar is rather large, of
elliptical contour, though the grinding surface is broken away,
set obliquely to the dental series so that the anterior teeth of
the two rows approach each other more closely than any of the
following teeth. M? is larger, less elliptical, and obscurely
trilobate, only one internal groove being prominent, with
faint indication of posterior internal and external grooves.
M? is much larger and more distinctly trilobate internally.
M* is distinctly trilobate, the lobes separated by deep sulci;
the anterior lobe in each tooth showing a groove on the an-
terior face near the external border. M®58§ are of the same
pattern as M?.
Most of the sutures are obliterated, so that little can be said
of the cranial bones or of their proportions. The cranium is
broad and depressed. The forehead is flattened and very
wide, ending in a distinct postorbital process. The rostrum
is very broad at the base, narrowing rapidly to the narial
opening. The muzzle is heart-shaped. The zygomatic arch
extends out widely from the skull and is very deep, with
hitaas
1903.] Brown, A New Fossil Edentate from Patagonia.
455
—inf. zyg. proc.
Fig. 1. Eucinepeltus complicatus. Type skull and casque. (No. 9248.)
Ge.
456 Bulletin American Museum of Natural History. (Vol. XIX,
prominent postorbital process. The descending process is
triangular at the base. The parietals are very rugose, pitted
with large foramina, gently convex transversely and longi-
tudinally. The sagittal crest is prominent. The lambdoidal
Fig.2. Eucinepeltus complicatus. Type casque. (No. 9248.) X 3.
crest extends out over the occipital plane in a heavy ridge,
curving in to join the sagittal crest. The occiput is about
half as high as wide; deeply incised for foramen magnum.
The bony palate is perforated by numerous large foramina,
1903.] Brown, A New Fossil Edentate from Patagonia. 457
with a large foramen opposite the posterior of M4. This fora-
men leads into a deep canal, extending forward nearly to the
premaxillary. The palate is straighter than in Propaleohoplo-
phorus australis, and forms with the superior line of the skull
a more acute angle than in the last named genus.
Measurements.
Cephalic shield: -length in median line.................2.- 130 mm.
_ ee RECEP OR WICC is. ans oe welde. oheuics a hada tees 133
Teeth
WICRPR IC CHODI os 2 -y.'- | 8 mm. Wid Gis: c. scams ee ent
M? *, 18 oS aS 9 Le ce iat ene
M$ SS ene A ean ie 12 + 6.6
M* ea coccre AY ae ra 13 Ce CR Oa Sut 7
M® ems ers St 15 7 7
M¢ "Fi keener ae 15 pin alee Oat ee 7.8
M’ ease tae eae 14.4 pT ae Re 7.8
M® oy tS eee nee 14 (ae Meate ea cranes 7.8
Piepeedenual sere Memoth sows uiG)sbieued chive dale ees e slotows Iog mm.
SUS Til, Tees aw eraete Wei 210 0 a a a ee eae ae 176
paemlenci of median basal line. .....2.- 1) 662. os ne be 156
4 ve occ. condyle to anterior end of premaxillary... 175
Cranium, length to anterior rim of orbit.................. 125
ms width behind zygomatic arches................. 102
Skull, maximum width over zygomatic arches.............. 135
Ar CLO METHICAILY 02.5 ioe). cc wd bleh oo bea tien cele ne 48
i UGE RPP et rrel 2% las 8 Aechapslereyehv vaca teyaioe em Me 89
Ayeomavextrenme length insides... 0... i ese c eee eteeane 77
Bee MAME VeEDICAIMGIATMeTCL : ..5 .ccls ot). vo << a, 4 ete 28
pw wlemoha GL descending process... 0.0... 0. + abe ween 45
“breadth a eg oS Ae ark a eens aan bee cee 20.5
PAlates leriot deat me GAT MAT Cas, 6 ee Siva p veh ecole ate oo + wiete oirie 117
e VOGT USVI ERMA Mere hy MeL ne cn ena b stata cities ale avaand wleos 14
a Eee a oe aang ol n> ccm ists og Bx ala pied ons 25
eee SceI ESTELLE AMIN ified ap Si ois svn doo wm ayes soars slate wel Bin a 43
© PCR EOS AS Es aoc) 4 ale aie aya ir yon vw a allt Wess sot 63
G MEAT ATIDCTIOL CtlG i.e cirlsle s Glee ole susie oye tans - 41
Cranium, width at postorbital constriction... ............. 49
IRKe®., IKeTaNSt bu} 5 Bere conto & co ROTC ELE eee epee or ic TOR 53
Forehead, width across postorbital process................ gl
AUTHOR'S EDITION, extracted from BULLETIN
ae Amevican Museum of Hatural History, —
Sato aE: : OF THE
3 a Vou. XIX, ARTICLE XII, pp. 459-464.
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Article XII.—_ORNITHOLESTES HERMANNI, A NEW
COMPSOGNATHOID DINOSAUR FROM THE
UPPER JURASSIC.
By Henry FarrFIELD OsBorn.
The type skeleton (Amer. Mus. Coll. No. 619) of this re-
markable animal was discovered at Bone Cabin Quarry, near
Medicine Bow, Wyoming, by the American Museum Expedi-:
tion of t900. It was removed and transported to the
Museum with the greatest care, and worked out, restored, and
mounted under the direction of the head preparator, Mr.
Adam Hermann, in recognition of whose many services .to
vertebrate paleontology the species is named.
The material embraces: the skull; 45 vertebre, including
3 cervicals, 11 dorsals, a complete sacrum, 27 caudals;
the complete pelvic girdle; representative portions of both
fore and hind limbs,—all belonging to one individual; our
knowledge of the manus is chiefly derived from another
specimen (Amer. Mus. Coll. No. 587).
PRINCIPAL CHARACTERS.
The entire length of the skull and vertebral column as re-
stored is 2.22 m. (7 ft. 34 in.); the height at the pelvis is
yom. (2:2: in.):
The vertebral formula, except in the sacrum, is still unde-
termined.
The most distinctive feature is the narrowing of the manus
and the great elongation (.172 m.) of the metapodials and
phalanges of the second digit, suggesting the rapid grasping
power of agile and delicate prey. This feature, combined
with the prehensile character of the somewhat enlarged an-
terior teeth, the extreme lightness of the skeleton, the cur-
sorial structure of the hind limbs, the balancing power of the
tail, suggest the hypothesis that the animal may have been
adapted to the pursuit of the Jurassic birds; in allusion to
this supposed habit the genus may be named Ornitholestes, or
‘bird robber,’ as suggested by Dr. Theodore Gill.
459
460 Bulletin American Museum of Natural History. [Vol. XIX,
A possible objection to this hypothesis is that the teeth,
while distinctively prehensile, are not so serrate or trenchant
as in Celurus. They are, however, quite as sharp as in the
varanoid and other lizards which are known to capture and
feed upon small birds.
The premaxillary contains 4 teeth, the most anterior of
which is the largest tooth in the upper jaw. The maxillary
retains to teeth, of which the fifth is the largest. In the
dentary are 12 teeth. In both jaws the teeth occupy a
rather short space, a little more than one third the entire
length of the skull, and gradually decrease in size posteriorly.
The premaxillary teeth are slightly worn on the posterior
surface. There are two antorbital openings, a smaller within
the maxillary, and a larger bounded posteriorly by the co-
alesced lachrymal and jugal. The orbits are very large,
bounded posteriorly by the united postorbito-frontal, which
connects by a slender bar with the squamosal. The depres-
sion of the quadrate extends the latero-temporal fenestra
vertically. The jaw is relatively long and slender, with ses-
sile coronoid process; the sutures have not been determined.
The cervicals are gently opisthoccelous, the dorsals are amphi-
ccelous, the posterior face being slightly more concave than
the anterior; the caudals are gently amphiccelous. The
neural arch only of the supposed fifth cervical is preserved.
The supposed tenth and eleventh cervicals are moderately
elongate, slightly opisthoccelous, with separate attachments
for the capitulum on the anterior portion of the centrum, and
for the tuberculum on the broad diapophysial expansion of
the neural arch; the zygapophyses are large and the neuro-
central suture is faintly indicated. In the supposed second
dorsal or thirteenth presacral the capitulum is still borne on
the centrum; behind the capitulum is a pit (paraccele), a feature
also observed in the supposed fifth dorsal; the diapophysis is
narrow; the head of the rib gradually rises to the junction
between the centrum and neural arch, as in other dinosaurs.
The four sacrals are firmly coalesced. The sacral ribs are
still suturally distinct and attached chiefly at the sides of the
centra, although the third sacral rib partially overlaps the
The unshaded areas indicate the portions restored.
1
9.
Am. Mus. No, 619.
Type.
Fig. 1.—Ornitholestes hermannti.
461
462 Bulletin American Museum of Natural History. |Vol. XIX,
posterior portion of the second centrum. The forked chev-
rons apparently begin at the posterior side of the third cau-
dal, the chevrons being intervertebral; the hemapophysial
canal is apparently closed in the first chevron. The firmly
coalesced caudal ribs of the anterior caudals are broad and
backwardly directed. With the supposed thirteenth caudal
there begins a gradual elongation of the zygapophyses, which
reaches a great development between the sixteenth and
twenty-fourth, the prezygapophyses being greatly elongated
and partly encircling the somewhat smaller postzygapophyses
of the preceding vertebre. At the same
time the chevrons become depressed, bifur-
cate in front, with a deep posterior keel.
The pelvic girdle is distinguished by the
deep symphysial union of the pubes, the
considerably more slender ischia, which are
in contact distally and proximally exhibit
two hooked processes; the ilium has a very
broad pubic and narrow ischial peduncle,
the pre- and post-acetabular portions of
the depressed crest are subequal. A very
distinctive feature of the postacetabular
crest is the hollowing out and inferior
expansion into a broad concave plate.
In the fore limb there was some question
as to the determination of the ulna and
radius. The humerus is longer than these
elements, measuring .127 m. The few
phalanges preserved enable us to associate
with this animal a relatively complete
manus belonging to another individual, in
which the striking elongation of the second
heey coy (pers digit, the. conipatative sleademlesssOumne
Nicaea. «third and the atrophy Gh tke oumheean
be clearly made out. The palmar view of
the manus somewhat suggests that of the two-toed sloth.
The terminal phalanges are elongate, recurved, and laterally
compressed, with a distinct lateral claw-groove. The some-
a
1903. | Osborn, A New Compsognathoid Dinosaur. 463
what crushed femur (.207 m.) is much longer than the
humerus and somewhat longer
than the tibia (.159 m.). The
pes has the typical tridactyl
arrangement, the phalanges be-
ing more rounded and less de-
cidedly curved than those of the
manus; the median metatarsal
measures .I17 m.
GENERIC AND SPECIFIC
CHARACTERS.
Skull with two antorbital
openings; four premaxillary and
ten maxillary teeth, non-serrate;
Ewememedentary teeth. Four ager Mus Noacty, XE
coalesced sacrals. Mid- and
posterior-caudal vertebre with greatly elongate zygapophyses;
manus narrow with greatly elongate digits, second digit of
manus enlarged, fourth digit vestigial, fifth digit wanting.
Fig. 3.—Ornitholestes hermannt.
Left manus.
AFFINITIES.
The affinities of the animal are evidently with the light-
limbed, slender-jawed group of Theropoda, for which the
subordinal name Compsognatha Huxley will probably be
found applicable, as distinguished from the large Megalo-
sauria. The exceptional rod-like elongation of the pre- and
' postzygapophyses in the mid- and posterior caudals strongly
suggests affinity to Ornithomtmus of the Upper Cretaceous.
Ornttholestes, however, is a much less specialized form, lack-
ing the peculiar compression of metatarsal III which charac-
terizes the Cretaceous genus. It is distinguished from the
contemporary Celurus by the non-serration of the teeth, by
the relatively short cervical vertebre, by the less extreme
hollowness of all the vertebre. From the contemporary
Hallopus it is distinguished by the less elongate character of
the metatarsals. Comparison with smaller foreign Wealden
464 Bulletin American Museum of Natural History. [Vol. XIX.
dinosaurs also serves to show its distinctness. The sacrum
differs from that of Aristosuehus in the possession of four
primary sacral ribs. The elongation of the second digit of
the manus resembles that in Archeopteryx, but was evidently
for a different purpose.
.
A New Three-Toed Horse.
By J. W. GIDLeEy.
AUTHOR'S EDITION, extracted from BULLETIN
OF THE
Le
Amevican Museum of Aatural
History,
, fuly. 24, 1903.
New York
VoL. XIX, ARTICLE XIII, pp. 465-476.
sie
nw
ickerbocker Press,
The Rn
Article XII.—A NEW THREE-TOED HORSE.
By J. W. GipLEy.
The Expedition of 1902, sent out by Prof. Henry F. Osborn
to explore the Miocene exposures in South Dakota, had the
good fortune to obtain, besides other material, a complete
skeleton of a large three-toed horse (Amer. Mus. Coll. No.
9815), associated with incomplete skeletons of five other in-
dividuals, undoubtedly of the same species.
This splendid specimen, which represents an undescribed
genus and species, was discovered by Mr. H. F. Wells, a mem-
ber of the party, in the upper Miocene deposits on Little
White River, near Rosebud Agency, South Dakota.
The characters presented, especially in the teeth, if inter-
preted according to former authors, would undoubtedly
place the present species in the genus Hipparion. However,
as indicated by a careful study of this new material, and of
the abundant material of other Miocene horses in the Ameri-
can Museum collection, together with a comparison with
specimens and descriptions of the European forms, it seems
probable that the genus Hipparion is limited in distribution
entirely to the Old World, and that the American species for-
merly referred to this genus should be placed in a group dis-
tinct from Hipparion.
Before describing the new skeleton, therefore, the writer
wishes to point out the chief characters which distinguish the
~ Old World from the New World forms.
The characters common to both groups are as follows:
(z) Column of protocone of the upper molariform teeth en-
tirely surrounded by cement; (2) the lower molars of the
milk dentition possess a median external basal tubercle; (3)
each foot possesses three complete toes, the lateral ones being
much reduced.
The characters which especially distinguish the true Hip-
parion are: (1) Protocone cylindric or subcylindric through-
out the greater part of its length. (2) Enamel borders of the
[yuly, 1903.) 465 30
466 Bulletin American Museum of Natural History. [Vol. X1X,
fossettes of the upper molariform teeth very elaborately folded.
(3) The middle portion of the external walls of the meta- and
paracones is flat or slightly convex. (4) The external median
tubercle in the lower milk molars is relatively high and circu-
lar in cross-section.
The American group differs from Hipparion in the follow-
ing characters: (1) The protocone is relatively larger and
elliptical in cross-section, or with the outer wall flat to con-
cave. (2) The enamel foldings are in general comparatively
simple. (3) The external walls of the meta- and paracones
are concave. (4) The external median tubercle of the lower
milk molars is elliptical in cross-section and less prominent
than in Hipparion. (5) The limbs and feet, so far as known,
indicate a comparatively more slender construction of the
long bones and especially longer proportions of the meta-
podials. There is also apparently a relatively greater reduc-
tion of the lateral digits in the American genus.
These characters, as stated above, have the more signifi-
cance from the fact that nearly all the American species are
Miocene, while those of the Old World are of Pliocene age.
It will be seen that in the development of the protocone and
the ectoloph in the upper teeth, and the proportions of the
feet and limbs, the American species, though coming from an
older formation, are more progressive than the Pliocene
species of Europe; while in some other respects, especially
the complicated foldings of enamel in the upper teeth, the
Old World species are more progressive. The reasons for
separating these two groups are further strengthened by the
fact that there are apparently no species common to both
hemispheres.
From the foregoing it seems apparent that a new term is nec-
essary for the American species hitherto referred to Hipparion,
and they may be distinguished by the name Neohipparion.
The following description is based on the complete skeleton
above referred to and is named in honor of Mr. William C.
Whitney ,whose generosity made possible the expedition which
secured this very valuable acquisition to the present knowl-
edge of American fossil horses.
1903. | Gidley, A New Three-Toed Horse. 467
Neohipparion whitneyi, gen. et sp. nov.
Generic characters. — Protocone free, except at base, as in Hzp-
parton. Protocone comparatively large and much expanded antero-
posteriorly. Enamel foldings simple. The median external basal
column present in the lower milk molars as in Hipparion, but much
shorter and more expanded anteroposteriorly. Lateral digits much
reduced.
S pectfic characters.—Size about equal to Neohipparion occidentale, but
enamel foldings much more simple, even more simple than in N. affine.
N. whtineyi further differs from N. affine in the much stronger develop-
ment of the styles of the ectoloph. Protocone relatively large and
very much elongated in cross-section anteroposteriorly. Outer wall
of the protocone flat and slightly folded inward, as is usual in Equus
caballus. Metapodials very long and slender. Lateral digits greatly
reduced, their terminal phalanges not extending to the distal end of
the first phalanx of the median digit.
Although the collected material representing the Miocene
horses of America is very abundant, it is, for the most part,
so fragmentary and the different parts of the skeleton so un-
certainly associated that most of the species are known only
from the teeth. Hence the present specimen, though not in
the line of ancestry of any of the living horses, may serve as
a standard for comparison of equal value with Mesohippus
batrdit, so fully described. by Scott,' and may be described in
detail as follows:
I. THe DENTITION.
Dental formula I.3, Ct, P.4, M.%. In proportion to the
other parts of the skeleton the teeth are very large compared
with those of Equus caballus. The molars and premolars of
both jaws are heavily cemented.
The Upper aw. — The tooth-crowns, though strongly hyp-
sodont, are of moderate length. Incisors much shorter than
in Equus. P' is placed well back, extending but little for-
ward of the anterior lobe of p? to which it is closely appressed
on the inner side. This position brings it in opposition with
the anterior lobe of p, of the lower jaw.
The external styles of the molars and premolars are as
1 Journal of Morphology, Vol. V. 1891, pp. 301-342.
468 Bulletin American Museum of Natural History. |Vol. XIX,
strongly developed as in Equus caballus and here is no trace
of the external median ribs of the meta- and paracones.
These last two characters are apparently common to all the
species of Neohtpparion and may be of generic importance.
Though the teeth in the present specimen are worn just to
the stage when they present the most complicated pattern of
enamel folding, they are very simple in this respect.
The Lower Faw. — The incisors, like those of the upper jaw,
are only moderately long-crowned, and are all fully cupped.
The external pair is smaller than the others. The first pre-
molar (p,) is entirely wanting. The molars and premolars
show an advanced stage of progression in the greatly flattened
external walls of the para- and.hypoconids. The antero-
external enamel fold of the protoconid, except in p»,4s
strongly developed. The lower border of the jaw is very
much curved.
The Milk Dentition. — There is no trace of p' in the milk
series of one of the specimens, but another associated speci-
men possesses this tooth reduced to a mere vestige. In the
upper molars the protocones are strongly developed and free
as in the adult. They are elongated in cross-section, though
not to the degree shown in the permanent series. The lower
molars possess a little tubercle arising from the cingulum be-
tween the para- and hypoconids. This conule is much
shorter than in the Hipparion of Europe, but broader antero-
posteriorly, being elliptical in cross-section. The outer walls
of the para- and hypoconules are flattened, but in less degree
than in the permanent series.
Ik THe: SkuLe:
There are many primitive characters observable in the
skull, the most prominent of which are the vertical thickness,
general shortness, and consequently the relatively large space
occupied by the molar-premolar series. The orbit is placed
well forward, its anterior border being above the posterior
half of p?. The anterior projection of the masseter ridge
extends forward to the middle of m'. The position of the
infraorbital foramen is between p* and p*. The facial pit is
1903. | Gidley, A New Three-Toed Horse.
409
broad and its borders are not clearly defined. The anterior
palatal foramina are small and do not extend back of the
canines. The anterior border of the posterior narial notch is
opposite the middle of m’. The vomer overlaps the anterior
end of the basisphenoid.
Measurements of Teeth.
; Anteroposterior.
|B ifeWanke) Kerns We) 9h oe rein Cente tere er eat 9.5 mm.
“cc “ec 2 a “ic
De ite e Speveree rss, talent 29.5
oe
BP tse vers arcpeis Gueee apc steeoraal 2 25 ie
- Dinter Onis Cees fer eet 25 ot
. gr ELISA estes Cones Phin Bie Wake Me 22 os
i PTT ee Sr sods aera ewer ae 24 be
te a3 m?
Motalklencth ofiSeries: ce an cess <= ee ee ee
Width across external incisors
Anteroposterior diameter of protocone, p*
“ yy a 7" Pes eee Meas
$ a : as Dae eee ae
a - : > TL ho Wee
a iy oy TI Arn ts eee
zs iS Se = LOCLMOe RY OSE oiage
Anteroposterior.
Diametersiol Peewee] slot eee oer 24 mm.
. YAO) Faure. © oe ersiees ciara Ons ox 25
a Ce Wits Got are Shy Berit a yy EE 25 :
i Bi 8 nde PAS OA ae ROTO 24 ci
ee poapionlPnte Cas rene ann ERE ete SS
2 Sn 1 See cn ey owt caesar One error 22 .
Milk dentition (Upper)
Anteroposterior.
DiameterstObiGip > satel a eiaeiee re 31.5 mm.
at isl oh gana ope Pee eee ee ergo 26 -
‘i P Ghoce queda so ogabonmanene 20 +
Skull Measurements.
Motal lenoth on Skotos sce foe goa enh ere
as 7 So RPRGUCIUG’S pti Stents padtn cere ns sem «aa narnia
Width of palate between first molars.............--
eS an a at flarrowest poimt...... 2. 25 wel eee
Length of diastema between canine and external incisor
By = e external incisor and p!...
Greatest width of posterior nares........-.-.-++++++:
VWiliiva Hiliconr Gurq oll ates pu GAS miRRepe eee Dice cue rarinatres O96 43 14 ol
Condylessits. sae oa ses oe Gone Dee reams
Transverse.
7 mm.
23.5 ae
25 oe
e. “c
25-5
2 ae
23 ce
152 mm
oe
55
8 mm
ac
9-5
TOs5
ae
9
9-5
Os5 | une
Transverse.
1r.5 mm.
13 “
T2 oe
Io oe
9-5
Wi ue
Transverse.
21.5 mm.
ee “
205 F
44
20 oe
20 x
67.50
28 -
128 ‘a
ae
54
470 Bulletin American Museum of Natural History. [Vol. X1X,
Ill. THE VERTEBRAL COLUMN.
The vertebral column of the type specimen was found com-
pletely articulated, even to the tip of the tail, hence the for-
mula can be given without a possibility of error. It is as
follows: Cervicals, 7; dorsals, 18; lumbars, 6; sacrals, 6;
eanudalso-7.-= 16) a:
The cervicals are long and slender and are horse-like in
general appearance. They resemble much more those .of
Equus than those of Mesohtppus, but show some intermediate
characters.
The atlas differs from Equus in the following characters:
The exterior pair of anterior foramina observed in the atlas
of the horse are not bridged over in Neohtpparton, but are
represented by open notches at the anterior borders of the
transverse processes. The median ventral tubercle, for the
attachment of the longus colli muscle, is very strongly de-
veloped.
The axts is comparatively longer and of less vertical thick-
ness than in either Mesohippus or Equus. Compared with
[quus the odontoid process is not so deeply spout-like, and is
proportionately narrower. The anterior vertebral foramina
are comparatively large, opening directly into the side of the
neural canal, and are not directed forward asin Kquus. In
Mesohtppus the foramen seems not to be inclosed, but is open
anteriorly. The spine is divided posteriorly into two diverg-
ing ridges which, extending backward and downward, merge
into the posterior zygapophyses on either side. In Mesohip-
pus the spine is not divided posteriorly, but extends back-
ward, ending in a strong high tubercle.
The 3rd, 4th, and 5th cervicals, except for their greater com-
parative length and more delicately formed processes, differ
but little from those of the modern horse.
The ventral surface of the 6th cervical is flat, turning down-
ward laterally into the wing-like transverse processes, which
are more strongly developed than in Equus. There is only a
trace of the median keel, so well developed in both Mesohip-
pus and Equus.
1903. | Gidley, A New Three- Toed Horse. 471
The spine of the 7i/ cervical is comparatively higher than
in Aquus, but more reduced than in Mesohtppus.
The spines of the anterior dorsals are long and slope back-
ward at a greater angle than in Equus.
The spines of the /umbar vertebre are comparatively high
and narrow. The transverse processes of the third lumbar
from the sacrum are comparatively long and do not articulate
with those of the second lumbar from the sacrum.
The rubs are remarkably like those of the zebra in form,
except the more posterior ones, which are proportionately
longer and have a greater curvature.
The sternum, which is well preserved in the type specimen,
is very characteristic. It is composed of six bony segments, as
is usual in the horse, but the ventral keel, so highly developed
in the living horses, is entirely wanting, except in the two
anterior segments, and it is only weakly developed in these.
The ventral surfaces of the third, fourth, and fifth sections are
flat and are widest transversely. The xiphisternum is the
largest of the series. The anterior portion of the ventral sur-
face is broad, concave, becoming narrower posteriorly where
the edges turn upward, making the posterior third of the
ventral surface convex.
The cartilaginous ribs are composed mainly of spongy bone
as in the horse, hence are preserved in the type specimen.
IV. THE Fore Limes.
The scapula is distinguished from Equus by: (1) relatively
narrow prespinous fossa; (2) the narrowness of the neck; (3)
the prominent vertical ridge or thickening of the subscapular
area beneath the postscapular border.
The humerus is widely distinguished from that of Equus by:
(x) the depth and narrowness at the proximal and distal ex-
tremities; (2) the shallowness of the bicipital groove; (3) the
absence of the groove in the lesser tuberosity; (4) the sharp
definition of the grooves and convexities of the ulno-radial
trochlea.
The radius is proportionately long and slender and, except
472 Bulletin American Museum of Natural History. [Vol, XIX,
for modifications of the shaft, is much lke that of Equus.
The shaft of the radius, in its median portion, is concave
behind, forming a sharp angle with the posterior border of
the inner face.
The shaft of the w/na is continuous, but very much reduced,
and firmly coalesced with the radius.
The carpus as a whole is more rounded in contour, and the
transverse diameter is proportionately less than in Equus.
The articulation of the scaphoid and magnum is peculiar.
On the distal face of the scaphoid the facets for the articula-
tion of the trapezoid and magnum are in form and position
practically the same as in Aquus, except that the magnum
facet extends further backward and curving downward ends in
a conical tooth-like process, which in the flexed position of
the carpus fits into a corresponding depression in the mag-
num. In Lquus the scaphoid and magnum do not touch each
other when the foot is fully flexed.
The convex portions of the radial facets of the scaphoid and
lunar occupy a relatively larger part of their proximal sur-
faces thanin Lkquus. The cuneiform is proportionately small-
er than in either Mesohippus or Equus. The pisiform is
short, thin, and broad, resembling that of Hquus in propor-
tions. The trapezium is very small and rudimentary, and
articulates principally with the trapezoid, though there are
two other small facets which indicate that the trapezium
articulates slightly with the scaphoid proximally and with the
rudimentary metacarpal I distally. The unciform is relatively
high and narrow, and projects below the distal face of the
Magnum.
The metacarpus consists of one principal and two much
reduced lateral members, metacarpals III, Il, and IV, and
two rudimentary bones representing metacarpals I and V.
These rudimentary metacarpals are about equal in size and
are reduced to mere nodules of bone, which articulate prin-
cipally with the second and fourth metacarpals respectively.
Metacarpals II and IV are nearly as much reduced in size as
the splint bones in Hquus. Their proximal ends and the
greater part of their shafts are placed well behind metacarpal
1903. | Gidley, A New Three- Toed Horse. 473
III. Their shafts taper to very slender proportions, but
expand again distally, forming articular ends which support
small lateral toes. Compared with Mesohippus and Equus,
metacarpal III is very long and slender. The distal end is
keeled entirely around, but is not so strongly developed on
the distal surface as in Equus. The lateral toes are much
shorter than the median toe, their extreme points reaching
only to about four fifths the length of the first phalanx of the
latter. The terminal phalanx of the median toe, compared
with that of Kquus, is proportionately longer and more com-
pressed laterally in front. The palmar surface is heart-shaped
in outline, with the apex deeply cleft. The proximal end
is moderately high. The articular facet is placed at about
the same angle as in Aquus. Two processes extend outward
and backward, one on either side of the articular face. These
processes are thin, with rounded edges, and each is perfor-
ated by an arterial foramen.
Veen Ein p) ices
The femur is slender and the shaft is relatively long. The
second and third trochanters are placed relatively nearer the
proximal end of the shaft than in either Mesohtppus or Equus.
The groove for the ligamentum patella is comparatively
longer and narrower than in Equus. The tbia exceeds the
femur in length, but in other respects is like that of Hquus.
The fibula is as much reduced as in Equus. The remnant of
the distal end is entirely fused with the distal end of the tibia.
The tarsus is relatively deeper anteroposteriorly than in
Equus, but is essentially the same in other proportions. The
shallow, irregular pits are beginning to form on the broad,
flat facets of the astragalus, navicular, cuneiforms, and the
proximal end of metatarsal III, but are, for the most part,
only indicated by slight depressions and roughened patches
on the bone surface. In Equus these pits are deeper and
their boundaries are well defined, though varying greatly in
size and form in different individuals.
The metatarsals are very long and slender, metatarsal III
equaling the femur in length. The lateral digits are reduced
474 Bulletin American Museum of Natural History. |Vol. X1X,
in about the same proportion as those of the fore-foot. The
phalanges are much like those of the fore-foot, except that
the terminal phalanx of the third digit. is somewhat smaller
and comparatively narrower than that of the fore-foot.
VI. PELVIs.
The pelvis shows a marked stage of advancement. It dif-
fers in no essential way from that of Equus, the proportions
throughout being about the same as in Equus caballus.
As the comparisons in the foregoing description have been
mainly with Mesohippus batrdit and Equus caballus, it may
prove instructive to give here a comparative table of measure-
ments taken from skeletons representing the three genera,
and tables showing comparative proportions. In the latter
the Virginia deer is included.
COMPARATIVE SKELETAL PROPORTIONS.
DORSO-LUMBAR SERIES TAKEN AS STANDARD.
Mesohippus batrdit.
Length of skull and neck .So
a mebaclk
a ‘ fore limb
+ oy Inwhayel
Neohtpparion whitneyt.
Length of skull and neck 95
sii ~ back it |
* “ fore limb 1.16 |
laine ima) i
Equus caballus.
Length of skull and neck 1.03 l
a Se pacik i |
“ fore limb 1.30
Tania! erat |
Virginia Deer.
Length of skull and neck 1.
~ back ity
‘ fore limb 1.46 |
7 layirayel “8 Tuan 12a gk : r
ce
ce
1903. |
Gidley, A New Three-Toed Horse.
475
COMPARATIVE PROPORTIONS OF LIMBS.
Length of fore limb
ay “humerus
“radius
sé
a3
Length of fore limb
3 “humerus
“radius
ce
ce
Length of fore limb
9 “humerus
“ radius
a3
ce
Length of fore limb
t “humerus
“radius
“ metacarpals
“e
“<
Length of hind limb
iF “femur
“ tibia
“ metatarsal I11
a3
“ec
Length of hind limb
s “ femur
“ tibia
“metatarsal 111
ae
Length of hind limb
5 “ femur
“tibia
‘“ metatarsal 111
ce
ce
Length of hind limb
ie “ femur
Es eri ohitee)
“‘ metatarsals
ce
ce
‘““ metacarpalir
“ metacarpal 111
“metacarpal 111
Mesohippus bairdii.
Fore 1. limb.
Neohipparion whitneyt.
19
saul
21
|
Equus caballus.
Mesohippus bairdit.
Hind
rT. linailo:
|
|
cltt7/ |
Neohipparion whitneyt.
Te
476 Bulletin American Museum of Natural History. [Vol. XIX.
COMPARATIVE SKELETAL MEASUREMENTS.
S
ees
a, 3...
os of
a8 | 23
mm mm
Skeleton: Total length, premaxillaries LOviSChiia sy, 82 ces Wee Sete ah ea eer
‘ hereht. spimevobesd idorsals- eet eee tenders AGG? i ercketaens
breadth across iliave es. ee wae eae See 135 290
Axial Skeleton: Skull, incisors to occipital condyles............ 175 378
“ transverse, zygomatic arches........... 80 130
Vertebral column, 30 presacral vertebre...... 645 | 1280
4 GELVICals, abOvals s.lecise eter 195 446
midcervical centrum....... 30 80
DS 1GOrsalS ahoball: csemeen at 280 605?
4th dorsal} centrum: o4.05 4: 20 32
i height of spine.... 53 135
PAikescalloehesu nour ae odo gey ean 170 240°
2d lumbar centrum........ 23 40
6Sacrals:“totalosee sees: 100 170
Catidals: ‘totale.§ sac era ? 325
Ribs, sth, length, outer measurement Bets ah ried 175 285
Si eo Ce Seog ike aan merge rire 190 365%
= 13th be Ke Cn ae oe ae 135 350
OC rerthi * "Fi es hevaatoitlone Yates aya ae 252
Appendicular Skeleton: Hind limb, total length............... 506 950
Femur, total length’... 2 Soe cere 170 255
circumference of gearts. aorta eaten By 100
Tibia, total length. . Sete 170 200
‘“” circumference of shaft. norte 50 90
Pes, os calcis to Dig. III, Ph. 3. Pes FRE 190 400
transverse, tarsals...... Aadaetens 20 30
““ anteroposterior, tarsals. . bee Bisa 22 34
Metatarsal III, total length. . 08 255
e circumference ‘of shaft... 32 70
Fore limb, total length. . eon cater LOS EN SEOMO
Scapula, length “Aree, Bieta lige ty Neat 130 250
greatest width........ bistesnete 80 128
‘Euimerius) totalilengthy aneurin eeee IIS 190
; circumference of site st Musee 47 85
Radius, total length. . Re. ahete 122 245
circumference of shaft. 38 80
Wine, totalblengt ile sewer ieee terete rene rene 157 300
Manus, TunarttolDissii, eb aee eee 130 355
transverse, carpals. . eats ove 19 40
“anteroposterior, carpals gee oes 15 30
Metacarpal III, total length.:......... 80 215
1 Thirty-one presacral vertebra.
4 Seventh rib.
2 Highteen dorsals.
Equus
caballus.
3 Six lumbars.
5 Five sacrals.
“AUTHOR'S EDITION, extracted from BULLETIN aes
OF THE
ee Fe
oe re ee
Article XIV.— ON TWO SPECIES OF PLATYGONUS
FROM THE PLIOCENE OF TEXAS.
By J. W. GIpLEy.
Among the very interesting fossils obtained by the Ameri-
can Museum expedition of r90r from the Blanco beds, are
two specimens of peccaries. Both are referable to the genus
Platygonus, but represent two distinct species. One is prob-
ably referable to P. bicalcaratus Cope, but the other is clearly
distinct from any described species. Both specimens consist
of upper teeth unassociated with those of the lower series.
Platygonus bicalaratus Cope.’
This species was founded on a posterior portion of a molar,
m, of the lower series. Unfortunately there were no charac-
teristic portions of the upper teeth associated with the type,
hence the upper dentition of P. bicalcaratus is not known.
As shown by Cope’s figures, the crests of the lower teeth are
BIN
Fig. 1.—Platygonus bicalcaratus ? Cope. Right upper molar series. Amer. Mus. No.
10701, Xf.
very high, and the conids forming them are subequal.
These characters, together with the agreement of size, have
led the writer to refer provisionally the present specimen
1Geol. Surv. Texas, 4th Ann. Rep., 1892 (1893), pp. 68-70, Pl. Di fl lea oF aise
477
478 Bulletin American Museum of Natural History. [Vol. AIX,
(No. royor Coll. Am. Museum, Fig. 1) to Cope’s species,
P. bicalcaratus, from the same beds. Although the teeth
present the generic characters of Platygonus they differ strik-
ingly from any species hitherto described in which the upper
teeth are known.
The chief distinguishing characters are as follows: (1) The
posterior and anterior crests of the molars are high and com-
pletely divided by the cross valley. (2) The cones forming —
the crests are comparatively wide apart at their summits; thus
when they become a little worn the upper molars of this
species present very much the appearance of the lower molars
of the tapir. (3) The posterior heel in m* is entirely wanting.
Measurements.
Anteroposterior. Transverse.
Diameters, eee gta ein AANSOT ni) santa,
~3 “
i‘ De aks oaks os ete 12 K 13 *
De sys ol SSkv ne ete eee 12 T4
i. 14 MS ROSES ie nen ot nSnchc Oe: «1h 36 1G 15 i
‘ 2 oe oe
TALC Wek ed ack ee eee aE eee 19.5 17.5 “
19.0 PRM Ene Aon) HAS 38 osha. 20 10
Platygonus texanus, sp. nov.
The second species, hitherto undescribed, is represented by
the ve a palate (No. 10702, Am. Mus. Coll., Figs. 2 and 5),
iy “iN
|
Fig. 2.—Platygonus texanus. Type. Amer. Mus. No. 10702. X t.
1903. | Gidley, Species of Platygonus from Texas. 479
containing the complete upper molar-premolar series, parts
of the alveoli of the canines and incisors, and a portion of
the upper anterior part of the skull.
Though coming from a later horizon,' the somewhat primi-
tive quadritubercular character of the molars suggests in this
species a close relationship to P. rex Marsh.?, However, com-
paring it with a cast of Marsh’s type, which the writer has at
hand, the following differences are very clearly shown: (1)
The four principal cusps of the molars are proportionally
shorter. (2) The cusps are subequal in the anterior and pos-
NIN
Fig. 3.—Platygonus vetus. Leidy. Amer. Mus, No. 2724. X tf.
terior pairs, and are comparatively wide apart at their sum-
mits. In P. rex the cusps of the outer side of the molars
are perceptibly higher than those of the inner side. (3) The
posterior heel of m* is much more strongly developed. It is
very weakly represented in P. rex. (4) The size of the molars
indicates a species larger than P. rex.
Compared with P. vetus Leidy (Figs. 3 and 4)the chief dis-
tinctions are as follows: (1) The cones of both the molars and
premolars are proportionally lower and, in the molars, more
1 The type locality of Platygonus rex Marsh is eastern Oregon. Marsh gives ‘the
horizon as Pliocene, but it is almost certainly Upper Miocene.
2Am. Jour. Sci. (3) xlviii, 1804, p. 273-
480 Bulletin American Museum of Natural History. [Vol. XIX,
simple. (2) The molars are proportionally wider trans-
versely, especially at the summits of the cones. (3) The
posterior heel of m* is wider transversely, and is more dis-
Fig. 4. —Platygonus vetus Leidy. Amer. Mus. No. 2724. X }.
Fig. 5.—Platygonus texanus. Type. Amer. Mus. No. 10702. X 3.
tinctly separated by a cross valley from the posterior crest of
the tooth. (4) The canine is proportionally much thicker
anteroposteriorly than in P. vetus, as indicated by the
alveolus. (5) The backward extension of the palatines is
much greater than in any species in which this character is
known, the anterior border of the palatal notch being situ-
ated nearly three times farther back of the posterior molars
than in 2. 72,
1903. | Gidley, Species of Platygonus from Texas. A481
Measurements of Type.
Anteroposterior. Transverse.
Diameters pgs ware tlaiote Avo an oe: Ir.5 mm. ro.5 mm.
ro Drenthe tes. ene eheged tet ates STAC et 12
rs | Oia seat, miigeese isha shame enor 14 “ 14 s
i. Ua tA ates hte ateteannd tac ote ely 555 aun
- TNS SPs cle ay AS eM ae ae ace 20 iS 20 2
+ TMS Mase Ma rae eee ee Tae 2G hy Altai 6h
Motaltlenothiomseniessatesat te eee Ior mm.
Position of palatal notch back of posterior molars... .. 48 mm.
Measurements of two upper molars of Platygonus rex Marsh.
Anteroposterior. Transverse.
Diameterse mae a ease eee Gena 19 mm. 16 mm.
te AS Pei cre een ces eco as ae ie if) eee
(These measurements are taken from Marsh's figures and confirmed
by the measurements of the cast.)
[¥uly, 1903-] 31
“dont, from the he Plewstocene
por Texas.
_ AUTHOR'S EDITION, extracted from BULLETIN
OF THE
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Vou. XIX, ARTICLE XVII, pp. 491-494. ~
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Article XVII. GLYPTOTHERIUM TEXANUM, A NEW
GLYPTODONT, FROM THE LOWER
PLEISTOCENE OF TEXAS.
By Henry FairFIELD OSBORN.
Pirate XLIII.
The existence of Glyptodonts in the United States was
made known by Cope ' in 1888 in his description of Glyptodon
petaliferus, based on a single carapace scute or plate from
the Lower Pleistocene Equus Beds of Texas; and by Leidy ?
in 1889 in his Glyptodon septentrionalis, based on carapace
and caudal scutes from Florida; Leidy also described 3 and
figured from Florida plates of the G. petalijerus type. The
American Museum Expedition sent out by the writer, with the
aid of the William C. Whitney Fund, under the leadership of
Mr. J. W. Gidley, was fortunate in securing the nearly complete
carapace, pelvis, sacrum, caudals, and complete tail armature
of an individual in fine preservation. It proves to represent
a new genus and species, combining characters of several of
the South American forms of the Pleistocene and Miocene
periods. The specimen was skillfully brought in by Mr. Gidley
and prepared and mounted under the direction of Mr. Adam
Hermann. The writer is especially indebted to Prof. W. B.
Scott for pointing out several of the features in which
Glyptotherium differs from the known South American genera.
Also to Mr. W. K. Gregory for the preparation of the manu-
script for the press.
The specific distinctions given below are derived by com-
parison of (1) the specimens referred by Leidy to G. petalijerus
and of (2) Cope’s description, which runs as follows: “*
with the circumferential areas of the rosette but little orate
than the central one. The former are regularly pentagonal,
oe Teepe ore ao saree 1880, Se ane Description of Vertebrate Remains
Ss the Peace Creek of Florida,’ Trans. Wagner Free Inst. of Phila., Vol. II, 1889.
‘9 tbid., pl. iv, fig. 9; pl. vi, fig. z.
[491]
492 Bulletin American Museum of Natural History. [Vol. XIX,
the latter regularly hexagonal, and they are separated by well-
defined grooves.’’ Cope’s type specimen is not available for
comparison; it may prove that Cope’s description and Leidy’s
reference are both misleading.
Glyptotherium texanum, gen. et sp. nov.
The carapuce measures 1450 mm. (4 ft. g in.) along the
dorsal curvature anteroposteriorly, and 1920 mm. (6 ft. 4 in.)
from side to side; the tail armature is 620 mm. (about 2 ft.
1 in.) long, having a circumference proximally of 705 mm.
The larger plates in the dorsal region measure 60 mm. in the
longest diameter, which is oblique to the axial line of the
carapace. The marginal plates may be estimated at 84, or
42 as.counted on the more perfectly preserved left side.
Eight of the posterolateral marginal plates are pointed or pro-
jecting; all the others have an even, gently convex, border.
The central plates are quadrate in the lower portion of the
shield, irregularly pentagonal and hexagonal in the upper,
all being characterized by a large central circular area sur-
rounded by from 7 to 12 smaller peripheral areas, all sepa-
rated by grooves. From 35 to 36 transverse rows of these
plates may be counted along the top of the carapace, and 34
at the side, practically the same number asin Glyptodon clavipes
Owen.! The six anterolateral rows are firmly united, but the
seventh to fifteenth are separated by deep grooves and ad-
mitted of some freedom of motion; these plates overlapped,
the borders being bevelled. A similar freedom is observed in
Panocthus.
The tatl armature is composed of eighteen circles of plates,
of which the anterior fifteen are arranged in paired rings and
the posterior three are coalesced into a single triple piece or
terminal cone. There are thus eight of the movable rings, of
which the seven anterior are composed of two rows of simple
flattened plates, and the eighth of a single row of elongate
_* Cf. Lydekker, R., Contributions to a Knowledge of the Fossil Vertebrata of Argen-
tina, Pt. II., pll.i-v. Paleontologia Argentina, III. La Plata, 1894.
1903. ] Osborn, A New Glyptodont from Texas. 493
plates with three pieces intercalated anteriorly. Behind these
eight distinct rings is the terminal cone, composed of three
rings of plates; of this codssified portion the anterior ring con-
tains eight pieces with two anterolateral pieces; the mid-ring
is composed of six pieces, and the terminal ring is irregularly
composed of four pieces. Thus the tail comprises eight com-
plete rings and a terminal cone, the same number as in
Glyptodon clavipes as described by Lydekker. The rings
gradually increase in length from 60 mm. anteroposterior
measurement, and decrease in diameter from 230 to 183 mm.
The terminal cone is laterally compressed, measuring 75 mm.
transversely and 132 anteroposteriorly. The posterior bor-
ders of the ring plates are gently scalloped, not pointed as in
Glyptodon clavipes.
Within the carapace and caudal rings were found one
sacrocaudal and thirteen free caudal vertebre, with an imper-
fectly developed fourteenth, and seven chevrons—all per-
fectly preserved. Of these vertebre the posterior ten, as
appears from measurement and from the deflected transverse
processes, were fitted within the tail-sheath, there being thus
a vertebra for each ring, while the anterior three articulated
with the peculiar sacrocaudal vertebre, in which the greatly
elongated transverse processes or ribs extend outward to
codssify with the posterior plates of the ischia. The first free
caudal has a transverse diameter of 302 mm., and distinct
lateral articulations as facets for the posterior borders of the
last sacrocaudal and of the ischium; the neural lamine are
elevated, the pre- and postzygapophyses are elevated and
vertically placed; the neural spine is low; caudals 2 and 3 were
also well within the carapace, with transversely extended
spines; in caudals 4-11 the transverse processes are deflected,
downwardly and forwardly directed; the neural arches,
zygapophyses, and spines diminish in distinctness. Caudals 12
~13 lack all processes. A single chevron of the narrow type,
similar to the most anterior chevron in G. clavipes, was found
with the specimen; it measures 130 mm. vertically. Six
stout chevrons with shallow, obtusely forked inferior pro-
cesses, anteroposteriorly expanded distally, are placed beneath
494 Bulletin American Museum of Natural History. [Vol. XIX,
caudals 5-11. These chevrons have an entirely different a
from the deep, narrow chevrons in G. clavtpes,
Both ossa innominata, as well as the entire dorso- and
caudo-sacral complex, are preserved.
GENERIC AND SPECIFIC CHARACTERS.
The hexagonal osseous plates of the carapace of this species
resemble those figured by Leidy (op. czt., pl. iv, fig. 9; pl. vi, fig.
1) as G. petalijerus, but differ specifically in: (1) the relatively
large size of the central area; (2) the smaller size and irregular
form of the peripheral areas; (3) the shallowness of the cir-
cular and-radiating grooves.
As regards generic distinction, this animal is very primitive
and simple in its tail structure, which strongly suggests. that
of Propaleohoplophorus and other Santa Cruz (Miocene) types,
but it appears to show some degeneration in the sculpturing
of the carapace plates, in which the central and marginal
areas are not so sharply defined as in G. petaliferus, Panoc-
thus, and other types. The imbricating lateral plates suggest
those of Panocthus. The shallow caudal chevrons are rather
like those of Sclerocalyptus (Hoplophorus) than the deep
chevrons of Glyptodon. The general shape of the carapace is
also rather like that of the Sclerocalyptus type than that of
Glyptodon.
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Article XXII.— A NEW GENUS OF GROUND SLOTH
FROM THE PLEISTOCENE OF NEBRASKA.
; By Barnum Brown.
Puates L anp LI.
Paramylodon, gen. nov.
This genus is founded on a nearly perfect skull and lower
jaw in the American Museum collections (No. 2780), with
associated skeletal material including five cervical vertebre,
tibia, fibula, calcaneum, astragalus, lunar, middle digit of
manus, and ribs, found by the Expedition of 1897 near Hay
Spring, Nebraska. Professor Henry F. Osborn has placed
this material in the writer’s hands for description.
The following characters distinguish it from allied genera:
Skull elongate; muzzle inflated; dentition ¢; first upper molar the
largest of the series; last lower molar trilobate; first lower molar
without opposing tooth.
Paramylodon seems to have been less specialized than
Mylodon, retaining features of the older, more primitive
sloths. From the long nasals it seems improbable that it
had a proboscis, while the greatly inflated muzzle, and the large
movable premaxille, indicate a large prehensile lip. The re-
duction of the twelfth nerve ' shows a less specialized tongue
than in Mylodon. The rounded condyles, with the greater
part of the articular area on the ventral surface, and the
aspect of the foramen magnum, opening obliquely to the long
axis of the skull instead of backward, show that the head was
carried more at right angles to the vertebral column than in
Mylodon. The long calcaneum with posterior end resting
flat on the ground, and the astragalar facet looking forward,
indicate a primitive foot more flexible at the ankle than in
the contemporaneous Mylodon.
The sum of these characters points toa difference in feeding
habits and indicates that Paramylodon was a grazer.
1 As indicated by the small condylar foramen in the skull.
[569]
570 Bulletin American Museum of Natural History. [Vol. XIX,
SyYNopsIsS OF ALLIED GENERA.
Mylodon.* Grypotherium. Paramylodon.
Number of teeth in skull.... 5 4 4
Number of teeth in lower jaw 4 4 4
Character of last molar of
LOWER JAW ace sue ClaucuerHsteke bilobed bilobed trilobed.
Character of first molar of
LOWED AW crevice esse g vein elliptical, cylindrical, elliptical,
opposes m2 opposes m+ long, not
opposing
any tooth.
Character onuiizzler. meer slightly contracted, inflated,
contracted, with osseous thin and
rugose. interseptum. smooth.
1840.
1840.
1890.
GENERIC CHRONOLOGY.
Glossotherium Owen, Foss. Mam. Voy. Beagle, p. 57. No
species named. Type, the back of a skull from S. America.
Mylodon Owen, tbid., p. 86. Type, M. harlant, based on a
jaw found in Kentucky, which had been previously referred
by Harlan to his Megalonyx laqueatus. Referred species, M.
darwint, based on jaws, etc., from S. America.
Mylodon robustus OwEN. Skull and skeleton. Glossotherium
here made a synonym of Mylodon.
Lestodon Gervais. Type, L. armatus, which (fide Lydekker) is
not generically distinct from Mvylodon.
Grypotherium REINHARDT. Founded on skulls of Mvylodon
darwint.
Pseudolestodon GrErRvats and AMEGHINO. Type, Mylodon
gracilis = Lestodon myloides.
LYDEKKER, in his Catalogue of the Fossil Mammalia in the
British Museum, correctly states that M. harlani is the type
of the genus, and divides Mylodon into two groups: one
represented by M. darwini, the other including M. harlanz,
along with WM. robustus, M. littsomi, M. armatus, and M.
gracilis. In the first, ‘the first tooth in each jaw is ex-
tremely minute and wears horizontally.” His reason for
placing M. harlani with the second group is, perhaps, that a
jaw shorter in front than the length of the dentition is indi-
cated in that species, as is the case in M. robustus, etc., while
in M. darwint the predentary portion of the jaw is much
elongated.
Neomylodon AMEGHINO. Type, N. ltstat.
1 M. robustus.
1903. | Brown, A New Genus of Ground Sloth. 571
1900. Roru and SmirH Woopwarp identify Neomylodon with Grypo-
therium, which they hold to be generically distinct from M.
robustus, etc. (presumably from Lydekker’s second group as
a whole, and therefore by inference from the typical Mylo-
don).
1900. AMEGHINO objects to the use of Grypotherium on the ground
that it was founded on the type species of Mylodon, which
he erroneously supposes to be M. darwinzt.
The type of M. harlant should be in the Columbia University
Museum, but has been lost or mislaid. A cast of it, in the
Museum of Williams College, has been loaned to the writer
through the kindness of Professor Cleland. Comparison of
this cast with a series of skulls of the subsequently described
genera and species shows that M. harlani is not congeneric
with G. darwint and is nearest, on the whole, to M. robustus
This classic species, therefore, is properly referred to Mylodon
and the characters of the genus may be derived from it.
Grypotherium appears to be a valid genus, but its name may,
perhaps, be antedated by Glossotherium. Lestodon is a valid
genus; Pseudolestodon is not separable from Mylodon.
Paramylodon nebrascensis, sp. nov.
SKULL.
Basal View. — The upper dentition comprises eight teeth
set in two diverging rows at such an angle that the space
separating the two anterior is twice that separating the two
posterior teeth.
The first molar is the largest and simplest of the series, hav-
ing the form of an elongated ellipse, with the longer diameter
twice that of the shorter. It is greatly curved antero-posteri-
orly, the convex surface in front; the sides are straight. The
longest diameter of the tooth is parallel to the dental series,
and the outer side forms a line with the posterior teeth. The
crown is worn deepest in the middle.
The second molar is bilobed; the anterior lobe stands at
right angles and the posterior lobe oblique to the series. The
tooth is triangular, with the angles rounded. The anterior
side is gently convex; the inner side is marked by a deep
572 Bulletin American Museum of Natural History. |Vol. X1X,
sulcus, a little in front of the middle; the outer face is marked
by a corresponding sulcus opposite but not quite so deep.
These sulci, on opposite sides, form the constriction that
divides the tooth into two lobes. Its greatest length is
oblique to the line of the series.
The third molar is bilobed but somewhat modified from the
preceding. The anterior side is gently convex with a faint
indication of a groove in the middle. The inner side is short-
est with a deep sulcus near its middle, while the outer side is
longest with a less defined sulcus. The outer anterior angle
is convex, as in the preceding tooth, but much broader. The
greatest diameter is oblique to the series.
The fourth molar is shaped like the figure 8; the anterior
lobe is larger and slightly oblique to the dental series. The an-
terior side is convex; the outer and the inner sides are nearly
parallel, and near the middle of each is a sulcus that forms the
constriction dividing the tooth into two lobes; the posterior
side is convex but not as wide as the anterior side. The
greatest diameter is parallel to the series. The plane of the
alveolar outlet slopes from without, inwards and downwards;
the space separating the alveoli of the first two is about twice
that separating the succeeding teeth.
The maxillary part of the palate is of a triangular form with
the base turned forward; gently convex from the palatines
forward to a line connecting the anterior borders of the first
molars, from which point it bends downward to the end of the
maxillaries. The surface is pitted with several rows of small
deep foramina parallel to the alveoli, and the anterior palatine
foramina are situated on either side of the median premaxil-
lary notch, passing forward, in two shallow grooves on each
side, to small notches on the inner anterior border. The
median maxillary suture presents a raised ridge, on either side
of which is a shallow groove extending from the antepenulti-
mate molar back to the palatines. The anterior border of
the maxillaries presents two rounded thickened surfaces,
separated by a deep median notch, for the articulation of the
premaxillaries, which must have been large and movable,
indicating a large prehensile lip. Directly in front of the
1903. ] Brown, A New Genus of Ground Sloth. 573
first molar, marking the boundary of the premaxillary sur-
face, is a small deep notch, back of which there is a small
expanded area marking the widest part of the maxillaries.
The palatine is concave and smoother than the maxillary
surface; the median sutural line is not prominent. Anteri-
orly the palatines join the maxillaries in an irregular suture,
anterior to the alveoli of the last molars; laterally it unites
with the maxillaries just internal to the alveoli. The pos-
terior palatine foramina, situated on the outer side near the
origin of the descending pterygoid processes, are the only
openings in the palatines. Posteriorly they expand to unite
with the orbitosphenoids and are separated-by a wide enter-
ing notch.
The pterygoids will be described from the outer aspect.
Internally they are convex, with a rugose posterior border
forming the lateral boundaries of the large posterior narial
opening.
The sphenoidal bones are not suturally defined from one
another: that part forming the roof of the nasal depression
between the pterygoids is broadly arched from side to side,
and wider than in Mylodon; the surface is smooth with two
longitudinal channels parallel to the base of the pterygoids.
Posteriorly the narial opening is bounded by two rough sub-
elliptical tuberosities.
The basiocctpital is a broad concave plate, the posterior
edge of which is deeply incised, forming the lower boundary
of the foramen magnum; the palate is pierced by a small
vascular foramen immediately in front of the condyles. The
anterior condylar foramina are large, but not more than half
the diameter of those in M. robustus. External to the con-
dylar foramen the basioccipital continues in a strong rough
tuberosity forming the posterior inner boundary of the articu-
lar depression for the stylohyal, and also the posterior bound-
ary of the jugular foramen. The sides of the basisphenoid
descend rapidly in front of the jugular foramen, ending in the
large tuberosities before mentioned as bounding the posterior
narial opening.
The condyles are rounder and shorter than in Mylodon,
574 Bulletin American Museum of Natural History. (Vol. X1X,
without the encroaching ridge which bounds the articular sur-
face in that genus. The convex surface of each condyle is
inclined a little outwards; the inner edges bounding the fora-
men magnum converge from the upper to the lower boundary.
The condyles stand away from the skull more than in Mylodon,
and the greater articular area is on the lower surface. The
foramen magnum looks downwards as well as backwards.
The tympanic is lost, leaving exposed the petrous bone
wedged in between the sphenoid, exoccipital, and squamoso-
temporal bones. The inner portion is a subcompressed,
conical protuberance uniting with the basisphenoid in a
straight line, forming the outer and posterior margin of the
carotid canal. In Mylodon the basisphenoid is distinctly
emarginated for its reception. On the outside and posterior
to the conical process the petroso-temporal sends down a
rugged process forming the anterior boundary of the jugular
foramen and the depressions for the stylohyal. In front it
bounds the posterior part of the tympanic cavity. The bony
canal of the Eustachian tube expands where it communicates
with the narial aperture, separating the pterygoids from the
sphenoid protuberances.
Side View.— As in Mylodon the side view of the skull
presents the form of an elongated parallelogram. The occipi-
tal plane inclines forward as it rises to the upper surface of
the skull. The top of the cranium presents a nearly straight
line slightly depressed at the posterior end of the nasals. The
muzzle ends in a slightly curved line; nasals and maxillaries
about the same length above, with protruding palate. Ven-
trally the basicranial outline is interrupted by the greatly
expanded pterygoid.
The supraocctpital element forms nearly the whole of the
posterior region of the skull, joining the parietals in a trans-
verse lambdoidal suture, which forms the crest but does not
encroach upon the coronal surface of the skull. From the
condyles the broad occipital plate rises upward and forward,
first at an angle of about 75° to two-thirds its height, and then
forward to the lambdoidal suture at an angle of 60°. It is
divided into two equal areas by a prominent occipital crest.
1903. | Brown, A New Genus of Ground Sloth. 575
On each side of this crest there is a shorter parallel ridge ex-
tending from above to about the middle of the skull. Im-
mediately above the left condyle there is a small foramen.
A deep groove separates the condyle from the occipital plate.
Wedged in between the occipital and temporal regions is
the prominent mastoid, about half as wide as it is long, bounded
above by a continuation of the suture that separates the ex-
occipital from the supraoccipital. The outer margin is raised
into a prominent ridge forming the lower posterior boundary
of the temporal fossa. Immediately inside this ridge there is
a deep channel running upward, ending in the mastoid fora-
men. The entire posterior surface of the mastoid, between
the raised outer border and the exoccipitals, is a depressed
rugose area. The inner part of the lower end of the mastoid
is a cup-like depression for the articulation of the stylohyal,
which is not nearly so deep or so extensive as in Mylodon.
The temporal fossa is uniformly smooth with an extensive
depression just above the mastoid, and a large protuberance
above this area, as in Mylodon.
The zygomatic process of the temporal is a stout trihedral
bone, the upper edge of which is nearly straight; the external
surface is nearly flat and gently convex. The under surface
is broad and flat for the articulation of the lower jaw, without
any distinct glenoid cavity, thus allowing a great forward and
backward movement of the jaw. The anterior end of the
zygoma terminates in an obtuse point.
The malar undoubtedly articulated loosely with the maxil-
lary and the zygoma, but unfortunately is missing on both
sides.
The most prominent bone of the skull is the pterygoid, which
extends downward and outward in a broad plate ending in a
very wide convex border, more prominent than in any allied
genus. Posteriorly it descends from the sphenoid at an angle
of twenty degrees, forming the gently rounded distal end,
then sweeps upward, in a quarter arc of a circle. The external
surface is rugose with a prominent ridge parallel to the
zygoma dividing it into two depressions. In the posterior
part of the upper one is the large foramen ovale, which opens
576 Bulletin American Museum of Natural History, [Vol. XIX,
forward and upward in a small canal at the upper border of
the pterygoid, partly defined by the overhanging projection of
the temporal. The posterior upper border is marked by deep
channels and ridges.
The frontal is the largest bone of the skull, forming the
middle half of the upper surface of the cranium, articulating
in front with the nasal, maxillary, and lachrymal, below with
the lachrymal, maxillary, and orbitosphenoid, and behind with
the parietal and squamosal. Externally it presents a smooth
surface of irregular curves expanded in front at the postorbital
process, extending downward and inward to unite with the
maxillary below the lachrymal, with a deep overhanging fold
exterior to the optic foramen, as in Glyptodon. Above the
optic foramen it descends outward slightly to unite with the
orbitosphenoid.
The orbitos phenoid is not suturally defined but represents a
depressed area, bounded on the outside by an overhanging
wall of the frontal containing the optic foramen and foramen
rotundum separated by a thin wall of bone.
The lachrymal presents an irregular outline, the superior
border of which rises anteriorly above the malar in a rugose
surface, in the centre of which is the lachrymal foramen. The
posterior portion is much thinner and extends back in a
truncated point to a line connecting the postorbital process
of the frontal and the posterior margin of the last molar.
Below the lachrymal foramen there is a deep rounded pit, ©
with a raised cone in the center, for articulation with the
malar.
That part of the maxillary presented in side view is of a
quadrate form, convex on the outer surface, and extends
nearly to the anterior end of the nasals, with which it forms
a vertical cross-section. Posteriorly it unites with the orbito-
sphenoid at the beginning of the canal that leads into the
optic foramen; above, with the lachrymal, frontal, and nasals.
The malar process is a stout projection set obliquely to the
maxillary, forming the anterior buttress of the malar and
enclosing the infraorbital foramen. Across the middle of the
maxillary, beginning above the infraorbital foramen in a
1903. | Brown, A New Genus of Ground Sloth. 577
curved line from the lachrymal foramen to the middle of the
muzzle, there is a line of foramina of decreasing size.
Top View. — The length of the skull from the condyles to
the upper anterior border of the maxillary is about the same
as in the type of M. robustus. In the mastoid region it is
slightly narrower, while across the anterior end of the zygoma
at its widest point, behind the postorbital process and at the
muzzle, it is much narrower. The upper surface is uniformly
smooth; gently convex transversely; nearly flat in the parie-
tal region, and slightly depressed from the postorbital process
forward, but much less than in Mylodon. The face is much
narrower than in M. robustus and the nasals are more highly
arched; in the type, the nasals are crushed down into the
narial opening. At the anterior end the nasals are slightly
deflected, terminating in an outer rounded edge, the median
halves presenting wide notches with two central, extended
points and articular upper faces, to which were probably
attached an incipient osseous interseptum. No well defined
ridge separates the top of the skull from the temporal fossa,
this surface merging into the sloping sides of the skull with-
out any marked separation. The posterior boundary is
formed by the thickened deltoid ridge, behind which are seen
the sloping occipital region and the condyles.
LOWER JAw.
Dentition. — The first molar is smallest in diameter and
simplest of the series: it is ellipsoid in cross-section and tapers
to a point, the internal and posterior borders of which are worn
down, leaving a crescentic anterior border. This tooth pro-
jects far above the crowns of the succeeding teeth (canine-
like) and has no opposing tooth in the upper jaw.
The second molar presents an irregular trapezoidal outline
with rounded angles, the posterior one most produced. The
external side is convex, while the internal side is strongly
concave, the concavity marked by a deep sulcus in the middle.
The anterior and the posterior sides are concave, the latter
more strongly pronounced. The sulci of the internal and the
[ October, 1903.] 37
578 Bulletin American Museum of Natural History. [Vol. X1X,
posterior sides tend to divide the tooth into two lobes. The
posterior lobe is worn obliquely. The greatest length is
oblique to the dental series.
The crown of the third molar is broken off, but the root
gives the shape and dimensions of this tooth accurately. The
external side is slightly convex, nearly straight. The inter-
nal side is slightly concave with indication of a groove in
the middle. The anterior and the posterior sides are con-
cave, with a faint groove in the middle of each. The greatest
length is oblique to the dental series.
The fourth molar is the largest and most complicated of the
series. It is composed of three lobes: a large anterior lobe,
oblique to the dental series; a posterior lobe, more than half
the size of the anterior lobe, transverse to the series, and a
much smaller, less defined middle lobe, connecting the two
and parallel to the series. The anterior lobe is convex on the
internal and the external sides, concave on the anterior side,
with a faint double groove in the middle, and concave on the
posterior border, drawn out to form the connecting middle
lobe, which unites with it on the internal half of the posterior
border. The external and the internal sides of the posterior
lobe are convex, and the posterior side is convex, with a
slight median groove; the anterior side is convex, uniting
with the middle lobe on the outer half. The middle lobe is
well defined on the outer face by two deep sulci, and less
marked on the internal side by two broad shallower sulci.
The anterior and posterior borders are drawn out to form the
connecting isthmuses, the latter being twice the width of the
former. The posterior and half of the middle lobes of this
tooth opposed the last molar of the upper series, and the
crown is nearly flat. The anterior lobe opposed the second
molar of the upper series, and the crown is worn in a cres-
centic surface.
Ramus, etc. — The ramus of the lower jaw is long and robust
but contracts rapidly to the symphysial region. It is thick
and massive in the dental portion and expanded in a thin
vertical plate posteriorly. The external surface is concave —
rounded both vertically and horizontally. The internal sur-
1903. | Brown, A New Genus of Ground Sloth. 579
face may be divided into three regions: the anterior, which,
from the second molar to the symphysis, is convex vertically
and concave horizontally; the middle, from the second to the
last molars, which is nearly flat; and the posterior, back of
the last molar, which is deeply concave. The lower border is
rounded and thickened below the dental series, thinning out
anteriorly and posteriorly. The upper or alveolar border is
wide and rugose, and on the outside of the jaw there is a
shallow depression extending the entire length below the
alveoli.
The anterior expanded end presents an obliquely sloping
symphysis, nearly half as wide as deep, extending anteriorly
in a deep rounded border. From a front view this anterior
border has the outline of a keel-like projection extending for-
ward in the middle of the jaw. Below this keel is a rounded
mammilloid process, much less prominent than in Mylodon.
The portion of the jaw in front of the first molar is longer
than in VM. robustus and the width of the expanded portion is
much less. The dental region is not nearly so deep as in
Mylodon.
The posterior part of the jaw is expanded into a thin deep
plate, divided into the coronoid, condyloid, and angular pro-
cesses. The point of the angular process is broken off; the
lower margin is convex and the inner side presents a deep
concavity bounded below anteriorly by the inferior inflected
border of the process and a ridge that extends forward and
upward to near the alveolus of the last molar; below, posteri-
orly, the inner border is roughened by processes; above, it is
bounded by the rounded border of the dental canal. The
outer surface of the angle is convex and rugose.
The coronoid process is broken at the point but seems to
have been nearly straight, slightly convex on the anterior
border.
The condyle is long and narrow, set obliquely to the vertical
plane of the coronoid. The greater part of the condyle over-
. hangs the supporting plate of bone, only about one fourth
extending on the outer side of the ramus. It rises from a
triangular base, the outer angle of which begins below the
580 Bulletin American Museum of Natural History, (Vol. XIX,
notch separating the coronoid from the condyle, and passes
backward and upward in a sharp ridge, forming the outer
articular border of the condyle. The articular surface is
slightly rounded.
The dental canal begins in a deep elliptical depression at a
point below the origin of the condyle and passes forward to
the entry of the large dental foramen. It continues forward
in two divisions: the smaller and shorter, opening on the
outer side of the jaw, opposite the posterior border of the
last molar; the larger, passing forward and dividing into three
branch canals that open on the outer side of the jaw where it
bends forward to form the symphysis. These foramina are
about ro mm. apart, and the middle one is the largest. The
smaller ones are about one fourth the diameter of the larger
one. Near the depression that gives rise to the dental canal
is a narrow deep channel that passes downward and forward
under the upper part of the ridge bounding the concavity of
the angle where it divides and comes to the surface in two
foramina 20 mm. apart.
TIBIA.
The tibia is a short, massive bone with a flattened shaft.
The proximal end is greatly expanded laterally and in width is
about three fourths of the entire length of the tibia. In
general appearance and proportions it agrees with the type
of M. robustus with the following exceptions. The external
condylar facet is distinctly pyriform. In M. robustus it is
circular. The fibular facet is elliptical and slightly concave;
oblique to the condylar surface and sloping from without in-
ward and downward. Anteriorly it nearly meets the superior
surface. Posteriorly they are separated by a convex facet
for the articulation of a sesamoid. The distal articular sur-
face is divided as in M. robustus, but the external semi-elliptical
compartment is convex, while in M. robustus it is flat. The
external malleolus is not as prominent as in M. robustus.
FIBULA.
The fibula is a subprismatic bone enlarged at both ends.
The outer surface of the shaft is convex; the inner surface
1903. | Brown, A New Genus of Ground Sloth. 581
flattened, slightly convex and rugose, the posterior border thin
and rounded. The upper end of the inner surface is excavated
with a rugose border. The proximal end is truncated ob-
liquely, presenting two facets: the outer and smaller for the
articulation of a sesamoid; the inner (a very long surface
convex in its narrowest dimension) for articulation with the
tibia. About one half of the proximal end of the fibula on
the outer side presents a rugose border and excavated surface
for the attachment of the peroneus longus. The tibial facet
is prolonged backward in a hook-like process. The distal end
is expanded in an irregular quadrilateral form on the inner
surface and a convex, very rugose, pitted obtuse point. The
inner surface presents a deeply excavated concavity and two
articular facets; the lower part of the concavity is occupied
by the flat, oblique, tibial facet, which continues uninter-
ruptedly into the vertical plane of the astragalar facet. The
posterior distal third of the malleolus is rugose. In M.
robustus the tibial and astragalar facets are separated by a
transverse concavity.
ASTRAGALUS.
The astragalus, aside from being much larger than that of
M. robustus, agrees with it in form and proportion with the
following exceptions: The navicular facet is deeply excavated
in the middle, with convex edges continuing uninterruptedly
into the cuboidal facet. In M. robustus it is flat on its upper
half and convex on the lower. The calcaneal facet is elongate
and triangular to a greater extent than in M. robustus, with
‘scarcely a perceptible constriction in the middle.
CALCANEUM.
The base of the calcaneum forms a nearly straight plane,
concave on its posterior triangular surface, and of greater
length proportionally than in M. robustus, with a broadly ex-
panded posterior end. The astragalar facet occupies a more
anterior position than in M. robustus, while the large tendi-
nous grooves on the outer side of the bone are similar, but
the tuberosities marking their boundaries are more prominent.
In other respects it agrees with M. robustus.
582 Bulletin American Museum of Natural History. [Vol. XIX,
LUNAR.
The transverse surface of the radial facet is convex in this
form and the posterior extension of the same surface is
broadly expanded, while in M. robustus it is nearly flat
transversely and contracted on the posterior surface.
Digits oF MANus.
The three phalanges of the middle digit are preserved, and
differ considerably from those of M. robustus. The claw
process of the ungual phalanx is somewhat deeper than wide,
of an ellipsoid outline. The under surface is not separated
from the sides by sharp ridges toward the distal end, as in
Mylodon, and it is only toward the base that these ridges are
shown. The middle phalanx resembles that of Mylodon,
though the proximal facet is much deeper. The proximal
phalanx is very much shorter than in M. robustus; the
outer surface is twice the width of the inner surface and is
expanded on the lower outer side. The proximal articular
facet presents a deep concave channel for the articulation
of the trochlea of the metacarpal; on the inner side of this
channel, continuous with its inner edge, is a narrow, con-
cave articular surface. The outer surface of the trochlear
depression presents a sharp ridge. On the ventral surface
there is a sesamoid facet on each side of the trochlear depres-
sion, the outer being the larger. The distal articular facet
is a wide shallow channel, concave from side to side and con-
vex vertically. Below the distal articular surface there is
a rugose depression one fourth the height of the articular
surface.
MEASUREMENTS.
Skull.
mm
Letieth of ms: 4/3 om abs 4 oa ee eae eee 20
Wraidtha. 8" Une emai te) gehen, oe ee 15
Rene ths m2. olen eh sees Ae se 26
Wridth "5 Scat aces inh Sevan ¢ eis oe 23
Lengthy “*~ a's 055 e< ciee) naeecnhe SE one re ee 23
Width 8D hagoe es og ae One ae 25
Length ** mits. 222 Satan oe ee oe 30
Width te. Sea NY en ee eee 17
1903. | Brown, A New Genus of Ground Sloth. 583
mm.
Lenpth of dental sere: 1... fcc .s-e artas eon ek ee 133
“““skull from condyles to end of maxillary..... 470
Depth of occiput in vertical line to condyles......... 136
Width across muzzle at widest part....... 028
Width at narrowest part back of apace Beeec ee. 89
Width of condyles. vino Vein cnet oat cen oee eee 128
Mandible
ere Glas Gi is eee heehee exe wa aoe 3 harass HE St eee 20
RVC CLG LU Meroe eet rac: otic ort wei seen eee ect tee eam 14
Length ‘ a eaday ee srScedales yor ate Wale? ancl Setens gains Wee sent PP Ee 24
NAAMG IE Stee Se oon ek en ea are Serer Ree eA. 22
LASS Tea) RIN 23 RR ae aOR a are RR en BEEN a ae edhe ne a0
VPC Lie We pane Streit iwstens ots 5 Sd nt binant Be Sea ee ty |
eee GRNge = SIME ye ltteer a aN ORG 2s See ke siamese eee 56.4
IVVit lita: ee owes torcyicn ae ce teal Rint te tah ee ee 24.4
Length of dental series, alveolar measurement....... 180
Length of jaw from condyle to extreme point........ fait
GET St GO Las yrtia Gy NSLS can ci isles ine wellag & Maeases moh eee ae ae 120
Waidthrofantentoniend: i. «cl:cee se ane 6 of pot cereale 445
Tibia
ID Grat qe ai eee Cees oe ee Ie near 5G 3 oat 275
Promitdaliend across widest path. ease crceis o/-teies ein eget
Distal os - * Paice Hose. one ee ESO
Fibula.
UE SEV SGI a oetar ove serch ars cot fs oye sr aca Os geete Ne Deepaieeeios 280
Wadth of«prosmalrend rt. c) 0 ntens «acral acters ature 105
Wadthrok distal ends sved Ge cra lara ah ale ee 70
Astragalus
Greatesté breddbiiiy « siachte cto So hies steams aelere i tae rae 150
OS SWAN eo AOn rE OD nS Cor At och 127
Calcaneum
METAS Gili meas dete tect and Aten ie Ae nee Ele, cs ye wa Os eee etoege 240
Wad Ghsacross 1b aseicei sei crsicrseere soccer aatone eae Repotaeeg= 130
Height at posterior articulation of astragalus......... 123
Third Digit of Manus.
Ungual phalanx, oon af Digncdh Sa ois oh tate aie eanas Be ceeeens 180
Second PT OP ae entrarecra rte cola OOD po 7c 63
4 rs Eins KA aeiple sa ownor on bd toc oma OS 47
Proximal “ METIS EEN, osnoce <rmcd Av ew coual 218 hee) ne 33
ss oy WHIGTOSACKOSS MilG@ Gli. cys) eters ei sivienels 46
PARAMYILODON NEBRASCENSIS
Cype skull x i
Lhe Frresh-Water Tertiary of Northwest-
ern Lexas. American Museum
Eixpeditions of 1899-1901.
sy]. W. GIDLey.
AUTHOR’S EDITION, extracted from BULLETIN
OF THE
American PMluseum of Aatural History,
VoL. XIX, ARTICLE XXVI, pp. 617-635.
New York, November 21, 1903.
*divog auaco1y peord Ay Surmoys
“NONV() AUNNHATAY Wadd)
‘JIT S1v1g ‘XIX “10, ‘HON ‘WY Nizating
Article XXVI.— THE FRESH-WATER TERTIARY OF
NORTHWESTERN TEXAS. AMERICAN MUSEUM
EXPEDITIONS OF 1899-1901.
By J. W. GIDLey.
Pirates LII-LVIII.
In the spring of 1899 Professor Henry F. Osborn sent an
expedition, under the leadership of the writer, to the Llano
Estacado or Staked Plains in northwestern Texas, for the
purpose of exploring the Miocene, Pliocene, and Pleistocene
beds previously reported from that region by Professor E. D.
Cope and Mr. W. F. Cummins, of the Texas Geological Survey.
This expedition met with such success that a second and third
expedition were sent to this region in the successive summers
of 1900 and 1901. ‘The results of the work of the three years
are comprised in this report.
The following brief itinerary of the three expeditions seems
necessary to a clearer understanding of the region explored
and the relations of the different localities visited.
EXPEDITION OF 1899.
Clarendon, the county-seat of Donley County, a little town
on the Fort Worth and Denver City Railway, was the chief
base of outfitting and supplies, and the initial starting-point
of the three expeditions.
The writer, with Mr. Alban Stewart as assistant and Mr.
Alfred Brown as cook and teamster, left Clarendon July 1,
1899. , Going north about ten miles the first camp was estab-
lished on Barton Creek, where a thorough exploration of the
surrounding country was made.
As reported by Cope and Cummins, the beds occupying the
tops of the divides in this vicinity are true Miocene and,
though of not great vertical thickness nor extensively exposed,
are very rich in fossil remains. Several good fossils were
found at this locality, the most important being a partial
skeleton, in a splendid state of preservation, of AZastodon
[617]
618 Bulletin American Museum of Natural History. [Vol. XIX,
productus (No. 10582, American Museum Collection) including
the skull and lower jaws.
Leaving this locality the party returned to Clarendon, then
going west about twenty-five miles to the head of Mulberry
Cafion turned southeast about three miles and established a
camp at the top of the bluffs a little to the west of the mouth
of the cafion. It was at this locality that Cope and Cummins
reported a new geological horizon which Cummins called the
‘Goodnight Beds’ and which Cope considered the same as the
Paloduro beds, placing them, in time, between the Loup Fork
(Miocene) and Blanco beds which are true Pliocene.
The party remained several days in this locality and the
writer explored both sides of the cafion for its entire length,
failing to find any evidences of a break in the stratigraphic
continuity of the deposits on opposite sides of the cafion, as
reported by Cummins.
From Mulberry Canon the party traveled southeast, crossing
the South Fork of Red River near the mouth of Mulberry
Creek, continuing south to a point nearly due east of Silverton,
the county-seat of Swisher County, turned west and ascended
the steep and rugged escarpment to the top of the Staked
Plains. Continuing west, passing the head of Rock Creek, a
third camp was established at the head of Tule Cafion.
The deposits here and extending east to and beyond Rock
Creek on the south side of the cafion, and for some distance
along the north side of the cafion as well, are of Pleistocene
formation. At the head of Tule Cafion was found a specimen
of Elephas tmperator (No. 10598, American Museum Collec-
tion), consisting of a complete fore limb, part of the fore foot,
the lower jaw and upper teeth, besides a few vertebre and
ribs.
Returning to the head of Rock Creek, seven miles to the
east of the Tule Cafion camp, the party established a fourth
camp. At this locality was made the splendid find of fossil
horse skeletons (Equus scotti) already described by the writer.*
This practically ended the season’s work, the party return-
ing to Clarendon by the way of Canyon City at the head of
* Bull. Am. Mus. Nat Hist., Vol. XIII, pp. 114-116; Vol XIV, pp I34-137.
1903.] Gurdley, Fresh-water Tertiary of Northwestern Texas. 019
Paloduro Cafion, then to Amarillo and back along the line of
the railroad.
EXPEDITION OF 1900.
The writer was accompanied on the second expedition by
Mr. Hans W. Zinsser, of Columbia University, who proved
both a valuable assistant and agreeable companion.
A little preliminary work was done this year at the old
locality north of Clarendon, but the main object of this ex-
pedition was to explore the eastern escarpment of the Staked
Plains south from Silverton, and especially to examine the
Blanco beds at Mount Blanco. Accordingly the party, leav-
ing Clarendon July 26, 1900, started southwest, taking nearly
the route followed the previous year as far as the main divide
between Mulberry Creek and South Fork of Red River, then
crossing the South Fork of Red River, several miles west of
the crossing point of the previous year, ascended the steep
escarpment to the top of the Staked Plains at a point nearly
north of Silverton. Going east about fifteen miles to the
point at which the party ascended the previous year, we
turned south along the top of the escarpment, examining the
bluffs for a distance of forty or fifty miles, then taking a
southwest course across the Plains went direct to Mount
Blanco.
The exposures at Mount Blanco are true Pliocene, and,
though of small extent, are very rich in fossil remains. The
specimens of most importance found at this locality were a
nearly complete skull and lower jaws of Dzbelodon marificus
(No. 10622, American Museum Collection), and, the following
year, a partial skeleton of a Glyptodont consisting of a nearly
complete carapace, tail pieces, pelvis, sacrum, lumbar and
caudal vertebre. This specimen has since been described by
Professor Henry F. Osborn,’ under the name Glyptothertum
texanum, gen. et Sp. nov.
On finishing the work at this place the party went directly
north to the head of Rock Creek, where a second excavation
was made in the bank or quarry, from which the horse skele-
1 Bull. Am. Mus. Nat. Hist., Vol. XIX, 1903, PP- 491-404.
620 Bulletin American Museum of Natural History. [Vol. XIX,
tons were taken the previous year, resulting in the finding of
additional material of Equus scotts (Nos. 10628, 10629 and
10630, American Museum Collection).
Finishing up the season’s work here the party again returned
to Clarendon, practically by the same route followed the pre-
vious year.
EXPEDITION OF Igot.
The third expedition to the Staked Plains was made possible
by the kind generosity of Mr. William C. Whitney, who do-
nated a sum to the American Museum to be used, under the
direction of Professor Henry F. Osborn, for the collecting and
placing on exhibition of fossil horse material to illustrate the
evolution of the horse in America. The object, therefore, of
this last expedition was to make a more extended exploration
of the Miocene and Pliocene exposures with the hope of
obtaining some new material of the three-toed horses of these
periods.
Accompanied by Mr. William Kendal as assistant and Mr.
James Morton as cook and teamster, the writer left Clarendon
July 1, going first to the old locality between Barton Creek
and Salt Fork of Red River. Wishing to extend the explora-
tions further to the north and east the party continued on
north about five miles to the head of Petrified Cafion, camping
there a few days, then turning east traveled down Whitefish
Creek about twenty-five miles to Skillet Creek, making another
camp at this place. At the head of Petrified Cafion were
found the skulls and parts of skeletons of twelve three-toed
horses. Unfortunately, however, the bones were badly
crushed and broken and covered with a hard limestone con-
cretion which is very difficult to remove without injury to the
fossil.
On the divide east of Skillet Creek the writer obtained three
important specimens, a skull and lower jaws of a Mastodon
(No. 10673, American Museum Collection) of the WM. productus
type, both fore feet complete of a second individual (No.
10672, American Museum Collection) of the same species, and
a skull with a few skeleton bones of the big dog Dinocyon
1903.|] Gidley, Fresh-water Tertiary of Northwestern Texas, 621
gidleyt (No. 10671, American Museum Collection) subsequently
described by Dr. W. D. Matthew.
The creeks have cut deeply into the underlying strata of the
Triassic in this locality, and erosion has entirely obliterated the
Miocene deposits from many of the divides.
The deposits that remain in this vicinity differ greatly in
character from the exposures along Barton Creek and to the
west. They represent much more the appearance of the
underlying red beds of the Triassic, from which the materials
composing them are apparently derived.
Finishing the work at this locality the party made a second
visit to the Blanco beds at Mount Blanco. Except for a few
days of fruitless search in the vicinity of the mouth of Tule
Cafion, the remainder of the time was occupied in a second
careful examination of the deposits at this place and in ex-
ploring Blanco Cafion to its mouth. The most important
specimens found on this second trip to Mount Blanco were the
Glyptodon specimen referred to above and a new species of
Platygonus, since described by the writer? (Platygonus texanus
No. 10702, American Museum Collection).
Returning to Clarendon, practically by the route taken the
year before in going from Clarendon to Mount Blanco, the
three years’ work in northwestern Texas was at an end.
GEOLOGICAL NOTES.
Professor W. F. Cummins, in the very complete and interest-
ing reports of his explorations in northwestern Texas,3 has so
fully, and, for the most part, accurately described the general
geological character and stratigraphy of the Staked Plains,
that an attempt at any such extensive and detailed description
here would be an unnecessary repetition of much of Cummins’s
work. Cope‘ has given detailed descriptions of the fossils,
taken from various localities, apparently verifying Cummins’s
determination of the various beds. It is, therefore, the inten-
1 Bull. Am. Mus. Nat. Hist., Vol. XVI, 1902, pp. r29- 136.
2 Bull. Am. Mus. Nat. Hist., Vol. abe 1903, P- 478
® Geol. Surv. Texas, 3rd Ann. Rep., 1891 (1802), pp. 129-200; 4th Ann. Rep. 1892
8 1790-238.
a be eee Texas, 4th Ann. Rep., 1892 (1893), pp- 11-87. Proc. Acad. Nat. Sci.
Phila., Vol. XLV, 1804, pp. 63-68.
622 Bulletin American Museum of Natural History. (Vol. X1X,
tion of the writer to discuss, in some detail, only the more
important localities with the purpose of correcting some
obvious errors made by Cummins, both in the distribution
and correlation of these beds, and to present briefly some of
the writer’s observations and conclusions which are at variance
with those of both Cope and Cummins. These conclusions are
briefly as follows:
(1) The fossil-bearing formations are fluviatile, not lacus-
trine, in origin.
(2) The Blanco has a limited distribution.
(3) The Goodnight (Paloduro) Beds are not a valid divi-
sion.
(4) The principal deposits forming the Staked Plains are of
the Miocene epoch but older than the Loup Fork stage.
(5) The fossil-bearing beds in the locality north of Clarendon
and at Mulberry Cafion correspond in age with the Loup Fork
formation, as shown by the fossils.
PLEISTOCENE.
Rock Creek Beds. = Sheridan (Equus) Beds.
The Sheridan or Equus division of the Pleistocene beds, as
reported by Cummins, are extensively exposed, especially
along the south side of Tule Cafion and at the head and on
both sides of Rock Creek, a small tributary emptying into
Tule Cafion from the south.
Cummins’s section of Tule Cafion, taken near the head, is as
follows:
G: Pine -white sand. 0. cite ee eee ae 2icuteet
5. Coarse sand, with) pebbles: et-is) sanea = ae eeerenen 20
4). Blush’ clay cf ia’. vie aie ate ete ae eee eee ee oe ene
3. ‘Coatse Samndl.2 A015 ih See eae gee eee BOs
a. JRed dish ielaiy:: 20 81's iat ate eee eee ae Conny
re Triassic sandstone’)! cme cae ae, eee mee ey
The upper four strata of this section (Nos. 3-6) belong to
the Pleistocene age. But the section is representative of these
beds only in a general way. Other sections taken at different
2 Local name proposed by the writer.
“agg02s snnby Jo Suoja[ays [eVIed xXIs udyRI oJAaM YOrYM wWosy Arrenb ayy uvas st punoisai10j s[pprui aya uy
‘SVXEL ‘'OO UAHSIMS ‘MAAUD MOOY AO AVA]T] LV ‘SHynAsOaxXyY ANAOOLSIATY
‘TIT @2v1d *X1X “10A ‘HN WY Nizating
1903.] Gidley, Fresh-water Tertiary of Northwestern Texas. 62 3
points show great variations in details of character and in the
relative thickness of the different strata. No. 1 belongs to the
Triassic, which apparently underlies the whole of the Staked
Plains. No. 2 of the section, designated by Cummins as
“Reddish clay,” is probably of Miocene age, as will be shown
later. Unfortunately no characteristic fossil remains have
been found in this stratum to fix definitely its geological
position. However, it is older than the Sheridan beds, as are
also the strata which bound them on the north and south.
The Sheridan beds are unconformable with those of this older
formation and apparently mark the course of an ancient
stream which, after scooping out a channel or narrow valley in
the older formation, refilled it again in Pleistocene times.
Fig. 1, page 625, is a diagrammatic cross-section across Tule
Cafion near its head.
There is nothing in the character of the deposits to indicate
beds of lake formation. On the contrary, the distribution of
the beds, which are nowhere very wide but extend several
miles east to the edge of the Plains, indicates, rather, an allu-
vial origin. The sharp cross-bedding- of sand, gravel, and clay,
which the writer observed at certain points in the formation,
and the peculiar distribution of the coarser gravels, all indicate
the depositions of a river or smaller stream rather than those
of a lake. A further indication of an alluvial derivation of
these beds is that the fauna represented consists wholly of
land forms, and some of the bones show weather checking.
The wind, carrying large quantities of fine sand and dust from
the surrounding plains, may also have played a very important
part in forming these deposits.
Following is a list of species from the beds at Tule Cafion,
as given by Cope!:
TESTUDINATA. DIPLARTHRA.
Testudo hexagonata Cope, Equus excelsus Leidy,
‘“ laticaudata Cope. ‘* semtplicatus Cope,
EDENTATA. ‘* tau Owen,
Mylodon ? sodalis Cope. ‘“ major Dekay,3
PROBOSCIDIA. Holomeniscus sulcatus Cope,
Elephas primigenius Blum.? - macrocephalus Cope.
1 Rep. Geol. Surv. Texas, 1892 (1893), p- 87.
2 Elephas primigenius, probably E. wmperator.
3 Equus maior = E. complicatus.
624 Bulletin American Museum of Natural History. (Vol. XIX,
The following additional species were taken from these beds
by the American Museum party:
Elephas imperator, Platygonus sp. ind.
Equus scottt, Large carnivore, gen. et sp. ind.
PLIOCENE.
Blanco Beds.
The Blanco beds have been correctly referred to the Pliocene
age, as is shown by the peculiar fauna they contain; Cummins
has, however, given them a much wider distribution than is
evident from a close study of the formations in that region.
He says of these beds’: “This formation constitutes the
eastern scarp of the Staked Plains from the Double Mountain
Fork of the Brazos River on the south to Paloduro Canyon on
the north”
Cope also reported exposures of these beds northeast of
Miami, Roberts County, Texas, more than one hundred and
fifty miles to the north of Mount Blanco. His identification
was founded on some fossil horse teeth which he referred to
Equus cumminsit Cope. .This species, as has been shown by
the writer in a former paper,” is not referable to the genus
Equus, and is indistinguishable at present from some species
of the Loup Fork Protohtppus, hence the correctness of Cope’s
identification of Blanco beds at Miami is, at least, doubtful.
Cummins was probably led to error by including in his
section of the Blanco beds strata properly belonging to beds
of an older age.
Cummins’s sections of the Blanco beds, taken at two differ-
ent localities, are as follows3:
At Mount Blanco.
12% WO... comedic Oe oe ee eee 8 feet
ri. (lard limestones as. oe ee ee eee 2 ie
ro: Sandstone. oo 2... 20 28 eee. Oe eee oie aay
g.. Stalactitic limestone yo4). 405. see ee A ee
8; Caleareouis sandstone ae 5250/2 eee eee ae
7: White Sandy clay: :.¢:5i6 15 eee 3Ome
6. White. diatemaceousvearth 4 eee Aa
§; Packsantd on os /eeh ss tos tie eee ne ee eae 20)
* Geol. Surv. Texas, 4th Ann. Rep., 1892 (1893), p. 201.
* Bull. Am. Mus. Nat. Hist., Vol. XIV, pp. 126, 127.
* Geol. Surv. Texas, 4th Ann. Rep., 1892 (1893), Pp. 200, 201.
‘oour[g JUNO JO samy Nos ayim-jpey auc)
‘sadagq OONVWTg
“AIT S31V1d “XIX “TOA ‘HON (IN ‘YV Nizaqing
.
ay
9
&
a 3 a
‘
.
2 a
=
190 3-| -Gidley, Fresh-water Tertiary of Northwestern Texas. ©25
Ao NVihttex diatomaceous carthees see a eee eees 8 feet
Se Greent SAnGynClany-,s sa 312 3, fot ts aay ee aon ke ae BX) 0
Bee EEE Cla eames a o'srayadtusevcieae le aL ep se es ay
Tt REO CIS H CLAN tm a stel wod eisai gee sme ee eee ed oreee Oma
Three miles north of the old town of Docum, in Dickens
County:
bese Wi Ge camCyGlany:, ciira,cic . <.steice seers arctan 6 feet
Am Witte Giatomaceous Garth. cme ci. otedtcts ertiltale Bye
Ben DUT LEY Glavg ct ye rate. os! «lays. s/apess arene eater kerma ae at
PEW erciatomaceOus Carblic tc. cise eee delle ee Aaa
Tepe dis hysamchyp Clay arse <1. svcus eeus «ees ereun ie oneveuerertes 1S)
These sections are valuable only in giving the approximate
thickness and general character of the deposits. No. 1, of
both sections, is wrongly included in the Blanco series, as are
also Nos. 9, 10 and 11 of the section taken near Mount Blanco.
These strata belong probably to the Miocene age and are
apparently continuous with the beds they so much resemble
at Tule Cafion and other portions of the Staked Plains.
A complete section of this older formation is represented
by Cummins’s section taken “‘one fourth of a mile northwest
of H. C. Smith’s ranch and one mile north of Mount Blanco”’:
Sel Vineay ect Cin Shh eo man hs Cet PR Ree LORE ear epee ein ho 2 feet
5. SiAMeenine WhoWENHONS 5 6 cep Has aawoDos onto bec LO
t. Red clay (same as No. 1 of previous section) 1... 30 “
The diagrammatic section, Fig. 2, page 626, taken by the
writer across Blanco Cafion and through the Blanco beds,
shows both of Cummins’s sections taken near Mount Blanco
and illustrates their relations to each other.
It is thus seen that the Blanco beds, at Mount Blanco, like
the Rock Creek beds, apparently occupy a comparatively
narrow valley or basin formed for their deposition by ancient
erosion of the older beds. Like the Rock Creek beds also they
extend a long distance in one direction, being traceable south-
eastward for fifteen or twenty miles to the edge of the Plains.
Though the deposits differ in character from those of the Rock
Creek beds and the fauna indicates an earlier age, here, as at
Tule Canofi, there is a total absence of any proof of a lake
origin for these beds and many evidences of river or stream
deposition.
I Cummins refers to the section taken near Mount Blanco and given above.
*a]TUI I 0} ‘ur € noe a]eds TeIUOZTIOY
+] OST 0} “ULI yNoge aTeIS TeoNIIDA “eIeIIS SuTAyIapun pu spaq oourjg jo uoyrsod Surmoys ‘oouvyg UNO Je uoUKD OoULTg ssotOe UOIOSS ONeMUILISLIGE “Z ‘BI
. pe aK is = | ©) QU (\ & oH ,
pe iy nec IN (MENS CI Mi TW WS
Oe ha ny Varn -\ | A oe
SS Seg NONV) OONV14 —_————————— waadD HSIALWO
Saad OONV19
[626]
‘apr 1 03 “ur € ynoqe afeos [ejuozt40y *3y OSr 0}
“UTT INoge a]voS IIe A “ees SutA[iepun pue spaq (ueprlisys) 42219 yoy Jo uontsod Surmoys ‘pray sy sesu ‘uoweD apn, ssor9¥ Uodes O4euUWeIseIqG “I “SI
ee eee
RST eat
Pasa
Ge /
TEEN
Sdad NVGIUAHS
1903.] Gidley, Fresh-water Tertiary of Northwestern Texas. 627
The occasional beds of diatomaceous earth are easily ac-
counted for by supposing that there were in this ancient
valley occasional ponds filled with clear water, enduring for
various periods of time, partially or totally isolated from the
stream that ran through the valley, such as exist at the present
time in the West, especially in the Sand-hills country of
northern Nebraska and southern South Dakota. The diato-
maceous deposits are for the most part quite impure and con-
tain great quantities of remains of rushes and pond grasses,
indicating that these ponds were never of any great depth and
probably occasionally received an overflow from the stream
in times of freshet.
Following is the list of species, as reported and determined
by Cope from the Blanco beds‘ and also as reported and de-
termined by the American Museum Expeditions:
Cope.
TESTUDINATA.
Testudo turgida Cope,
“ pertenuis Cope.
AVES.
Creccoides osbornit Schuf.
EDENTATA.
Megalonyx leptostoma Cope.
CARNIVORA.
Canimartes cumminsu Cope,
Borophagus diversidens Cope,
Felis hillanus Cope.
PROBOSCIDIA.
Tetrabelodon shepardiu Leidy,
Dibelodon humboldti Cuvier,
: tropicus Cope,
precursor Cope.
“cc
DIPLARTHRA.
Equus simplicidens Cope,?
cummins Cope,?
minutus Cope,?
Platygonus bicalcaratus Cope,
Pliauchenia spatula Cope.
Total number of species, 16.
American Museum,
TESTUDINATA.
Testudo campester.
EDENTATA.
Glyptotherium texanum,
Megalonyx sp.,
Mylodon sp.
CARNIVORA.
Amphicyon (?Borophagus).
PROBOSCIDIA.
Dibelodon mirificus,
ns tropicus.
PERISSODACTYLA.
Neohipparion sp.,
Pliohippus simplicidens.
ARTIODACTYLA.
Platygonus bicalcaratus,
= texanus,
Pliauchenia spatula,
a sp.
1 Rep. Geol. Surv. Texas, 1803, D-
2 Equus simplicidens = Pliohtpp
pp. 123-128 and p. r4o.,
Te ay Cae ;
us simplicidens; Equus cumminsu = Protohippus
sp.?, and Equus minutus = Protohippus sp.? See Bull. Am. Mus. Nat. Hist., Vol. XIV
628 Bulletin American Museum of Natural History. [Vol. XIX,
MIOCENE.
(?) Goodnight (Paloduro) Beds.
In 1893 Cummins proposed a new horizon, the Goodnight
beds, placing it, in time, between the Loup Fork and Blanco
divisions. In thus placing these beds he says: ‘I do this on
both stratigraphic and palzontologic grounds. In making a
stratigraphic section of the country, I found that the Good-
night fossil beds were above the conglomerate bed of the Loup
Fork section made at Clarendon, the Clarendon beds being
near the base of the Tertiary and below the conglomerate,
while the beds at Goodnight were nearly at the top of the
plains.”’
Following are Cummins’s sections taken-at the typical lo-
cality on opposite sides of Mulberry Cafion near its mouth:
North Side.
1. White sandy clay, concretionary.............. 60 feet
2. (Sandy: Clays Nive s. ok te tk ee OL eee ACwas
3; Reddish ‘sandy clay otc. she eee 25.‘
4, Conglomerate, eross-pedded=aseenioeee rer eniaae 20) ws
5. Inedyclay,.to the base anya. .10 eee eae Sons
225 feet
South Side, about one mile from the preceding section.
t. Concretionary yellow limestone............... 4 feet
2. Yellowish clay, with small concretions......... 12 Nar
3. ‘Coneretionary limestone. 1.0) 4.4: 6. 2. Pe Phat ua is
4. Yellowish sandy clay.. ae AO
5. Hard stalactite lameeeee reais ite conenes
dal fracture sir soar ¢ eon eee Ue ee yeah
6. Yellowish sandy clay, with small concretions, less
concretionaryat base. eee steer Sle) i
7. eright red Clayjjccase se. ee te ee eee eee AOumen
&. Solt dark redisandstone..0 0 set ooo Ons
0: Soft limestone sc. cetera ae ee eee yeas?
ro» Red clay, with white spotsacs: 7a eee ee Ome
226 feet
A careful study of this region has compelled the writer to
disagree with Mr. Cummins both as to the correctness of his
“uOUB ayy jo sapts ajtsoddo uo eyes jo Ayinutuod puv uortsod [ejuozuOY Surmoys
“NONV)) AUYAATOIN
ONTL EHO AE SEBS Saloy\ ‘HN ‘WW ‘Y Nizating
te. ;
b ‘—s A
i ! is Peay 3 1
iv . (er i ee ie, - 7 iy
: s
i LY 7 V ¥ fo
rien -
z
f
f -
j
‘
I
+
L if
i i
i
,
.
?
‘
4
a
f
.
)
?
i
iy ck
i
? 4
1903.] Gidley, Fresh-water Tertiary of Northwestern Texas. 629
observations and in his interpretation of the strata in this
locality. The writer could find no warrant for making any
separation of the beds at Mulberry Cafion, either on strati-
graphic or paleontologic grounds.
Briefly, Cummins’s stratigraphic grounds for separating the
Goodnight beds from the Loup Fork division are as follows:
(1) That the Loup Fork beds in the vicinity of Clarendon were
overlaid by the heavy cross-bedded conglomerate layer which
underlies the upper series of strata at Mulberry Cafion. (2)
That there is a marked difference in his two sections taken on
opposite sides of Mulberry Cation. He says of these sections:
“Tt will be apparent upon examination of these two sections
that there is a marked difference between them. The heavy
bed of conglomerate on the north side of the canyon, No. 4 of
the section, does not occur on the south side, nor was there
any gravel on that side to show that the conglomerate bed had
ever been there.”’
This statement is incorrect, for the writer found an abun-
dance of gravel on both sides of the canon and had no trouble
in tracing the conglomerate layer (No. 4 of Cummins’s section)
across to the south side of the cafion where it is exposed in two
localities, showing a maximum thickness of at least fifteen feet.
This bed of conglomerate and sand is nowhere of great width,
hence it does not appear in every section on either side of the
cafion. The writer also found this coarse conglomerate bed
appearing again in the Clarendon locality, resting directly on
the eroded surface of the Triassic and underlaying the Miocene
beds, but not overlaying them, as reported by Cummins. There
are scattered patches of loose gravel partially covering the
Miocene deposits in the vicinity, but it is superficially distrib-
uted and nowhere can be said to be in its original bed of
deposition. It was probably this frequent occurrence of loose
gravel which led Cummins to believe that the conglomerate
bed had overlaid the Miocene in this locality.
Cummins’s two sections taken at Mulberry Cafion are mis-
leading, for no two sections, even though taken on the same
side of the cafion, agree in detail. There is, therefore, no such
real difference in the two sides of the cafion as these sections
630 Bulletin American Museum of Natural History. [Vol. XIX,
apparently show. The main divisions of either section are
traceable around the head of the cafion and are continuous
with corresponding strata of similar character on the opposite
side.
From the foregoing facts it seems evident that Cummins’s
separation of the Goodnight beds on stratigraphic grounds is
scarcely admissible.
Cummins’s palzontologic grounds for the separation of these
beds is based on Cope’s determination of the small collection
of fragmentary fossils taken from the vicinity of Mulberry
Cafion on the south side.
Cope’s determination of these fossils is as follows ?:
A phelops sp., Hippidium inter polatum Cope,
Protohippus lenticularis Cope, gy ? spectans Cope,
Protohippus sp., Equus eurystylus Cope,
- ? perditis Leidy, ‘““ ? simplicidens Cope.
In referring to this list Cope? stated that two of these genera
are characteristic of the Loup Fork beds, but are not known
to extend higher; that Equus, on the contrary, had never been
found in the Loup Fork formation; and that Hippidium was of
uncertain horizon. Of the species Cope said ‘‘the three which
are identifiable are new to science.”
As shown by the writer in a former paper 3 E. simplicidens
Cope and E. eurystylus Cope can not be referred to the genus
Equus, but on the contrary are distinctly Miocene in character
and only referable to Miocene genera. Hippidium is probably
an exclusively South American genus, and the species referred
to it by Cope are probably referable to the genus Protohippus.
The three species from the Mulberry Cafion locality which Cope
considered identifiable are as follows: Protohippus lenticu-
laris Cope, Hippidium interpolatum Cope, and Equus eury-
stylus Cope.
The first of these species is, perhaps, authentic, but it is
referable to the genus Neohitpparion 4 and not to Protohippus.
*Geol. Surv. Texas, 4th Ann. Rep., 1892 (1893), p. 45.
?Geol. Surv. Texas, 4th Ann. Rep., 1809 (1892), p. 45-
*Bull. Am. Mus. Nat. Hist., Vol XIV, 1901, pp. 123-126.
*The genus Hipparion was founded on an European type, though many American
species have since been referred to it. The writer, however, has separated generically
the American group from the Old World Hipparion, giving to the American group the
name Neohipparion. (Bull. Am. Mus. Nat. Hist., Vol. XIX, p. 467)
~
MULBERRY CANON
ANAIOIN SDISSVIEL
Fig. 3. Diagrammatic section across Mulberry Cafion near its mouth, showing position of Panhandle (Lower or Middle Miocene) beds and the underlying strata.
Vertical scale about x in. to 250 ft,; horizontal scale about 2 in. to r mile.
[631]
—
SS Se
MIOCENE
Nil
a
i
FCS YES EATEN
GGELIEZLEELLOVEEGLOLELA EA:
CLYY EEL ELELEELZ:
Se A ne ee
eee a
,
ote
LOUP FORK
Wirz
al
\\l
|
a
Fig. 4. Diagrammatic section of the Clarendon locality, showing position of Clarendon (Loup Fork) beds and Panhandle (Middle or Lower Miocene) beds,
extending west to the top of the Staked Plains. Vertical scale about 1 in. to 300 ft.; horizontal scale about t in, to x mile.
632 Bulletin American Museum of Natural History. (Vol. XIX,
The species does not indicate a later phase than the Upper
Miocene, for the writer found, in the Clarendon locality,
several specimens, including a skull with complete dentition,
which are indistinguishable from Protohtppus lenticularis Cope.
The second species, as already mentioned, is indistinguishable
from some species of Protohippus. The third species of this
group, Hipparion eurystylus Cope, was founded on lower teeth
and is not distinguishable from specimens found in the Clar-
endon locality. It is quite possible that Cope’s type of this
species represents the lower dentition of Neohtpparion lenti-
cularis, with which it corresponds in size.
Thus it will be seen that the palzontological evidence at
hand not only fails to prove a new horizon for the so-called
Goodnight beds, but, on the contrary, seems to prove con-
clusively that they are identical in age with the beds in the
vicinity of Clarendon, which Cope recognized as Loup Fork
(Upper Miocene) deposits.
There is little doubt, then, that there is no break at Mulberry
Cafion, either in strata or fauna. Hence the Goodnight beds,
as a new horizon, should be abandoned.
Upper MIOCENE.
Clarendon Beds: ‘‘Loup Fork” Stage.
The deposits at the locality north of Clarendon belong un-
doubtedly to the Miocene epoch. The fauna indicates a close
relationship with the Loup Fork formation.
This locality is east of the Staked Plains proper, but con-
nected with them through long, low divides. The fossil-bear-
ing strata do not, however, follow these divides back to the
Plains, and is it impossible to say, owing to so much of the
country in that direction being covered by recent deposits
which are now more or less grass-covered, whether they ex-
tended any great distance to the westward or not. The
* Local name proposed by the writer. The name Loup Fork was first proposed as
a formation name, and its subsequent extended use has given rise to so much con-
fusion that it seems better not to employ it in the sense of a time division, but to limit
its use to the formation occurring in the Loup River, Niobrara, and White River val-
leys for which it was originally used. The Clarendon beds are of approximately the
same age as the Loup Fork beds, as judged by the known fauna, but cannot be re-
garded asa part of the same terrane, and they differ considerably in structure and
composition.
“SasioY paoj-aa14) Jo SUOJI[AHxS [enied PU S[[NYAS [kIaAVS UdHv}] BIOM YoOryM Woy spag uopuare[>) Ul UOlRARDXa Surmoys
‘NONVD GalMdINtag 40 avay
IAT SLV1d “XTX “109A ‘HN ‘W ‘WV Nigating
1903.] Gidley, Fresh-water Tertiary of Northwestern Texas. 033
exposures, however, show an extensive distribution to the
northeast.
Here again, as at Tule Cafion and Mount Blanco, the pecu-
liar formation of the deposits indicates, though in a somewhat
different manner, an‘ alluvial origin. Though distributed over
a wider area in every direction there are running through these
beds several narrow channels of sandy clay. The main body of
the beds consists for the most part of cross-bedded sands and
sandstones intermixing more or less and cross-bedding with
the clays. These channels all take a direction nearly east and
west, or approximately the same as that of the streams drain-
ing the country at the present time. Some of them are trace-
able for long distances. It is in these peculiar beds of sandy
clays that all the fossils of this region occur.
Cummins’s section of the Clarendon locality is as follows:
TMM VV A La EtG tae Seat liven CL ety ea vant st sii.) eu syalonege serieireicls pss eouteveteia hewn 20 feet
2. Sandy clay, with many rounded siliceous pebbles of differ-
CMMGESI ZECCA erey MINS, co ierae es ai, Ghat e wseuarins 4c, = ha lod tiakepemae ete ons 20
RNG UE ONSEN SADT A nates awe Fon no 5600 a erat) s (oS ors «sei @ joc oko ale tare een tate Titey
Peep htnchii ithe Guytiid GexG atts «oc cnc ice 20s yal sna: aval els stlel ea ald ween tales ACM ie
5. Yellow sandy clay, with the sand more or less predominat-
ing in places. In places the sand is hardened, while in
others the clay is more or less concretionary.. 250. ~
6. Alternating beds of bluish clay and white 2x Gl (ust
IBtGhGl eS Mcod oat eae oe Cer Ceucbo iG Aeron tent ub Dorion co.cc ZOU
400 feet
Cummins has here placed the Loup Fork formation at the
very bottom of this section of 400 feet of deposits. A careful
study of this region, however, does not warrant such a dis-
position of this stratum. In reality this stratum (No. 6)
belongs properly at the top of the above section, and the
explanation is simple. Nowhere is bed No. 6 overlaid by
any of the upper strata of the section; hence to obtain this
section Cummins probably included the beds west to the top
of the plains, and because No. 6 was at a lower level concluded
that it ran under the beds to the west. This, however, is
erroneous. The writer found several places where this fossil-
bearing stratum lies unconformably on the eroded surface of
634 Bulletin American Museum of Natural History. [Vol. XIX,
beds resembling the lower portion of the beds to the west which
Cummins identified as probably Goodnight beds. It seems
certain, therefore, that the beds to the west are older than
the fossil-bearing strata under discussion. This apparent in-
consistency in level is due to the heavy erosion of the older
beds before the Upper Miocene deposits were laid down.
The following species have been reported from the Claren-
don locality:
REPORTED AND IDENTIFIED BY
COPE.
A phelops fossiger Cope,
Protohippus perditis Leidy,
‘ parvulus Marsh,
nov.),
Protohippus mirabilis Leidy?,
m placidus Leidy,
Hippotherium 2 affine Leidy,
£ occidentale Leidy,
Procamelus robustus Leidy,
“ gracilis Leidy,
nov.),
Blastomeryx gemmifer Cope,
Tetrabelodon serridens ? Cope.
fossulatus Cope (sp.
leptognathus Cope (sp.
OBTAINED AND IDENTIFIED BY
THE AMERICAN MuSEUM
EXPEDITIONS.
Mastodon productus,
“ sp.
Dinocyon gidleyt,
is meandrinus,
Macherodus sp.,
Teleoceras sp.,
Hipparion 3 lenticularis,
Me occidentalis,
Sp.,
Protohippus perditis ?
« Sp.”
Pliohippus ? sp. nov.
Procamelus sp.,
Pliauchenia sp.
“e
MIDDLE OR LOWER MIOCENE.
Panhandle Beds.4
As to the age of these older beds it is difficult to say, owing
to the present lack of palzontologic evidence, just where they
should be placed.
Lower or Middle Miocene.
It is probable, however, that they are
Some lower teeth of Merycocherus and a lower tooth of
Procamelus taken from these beds by the writer prove them
to be not older than the Lower Miocene.
Though differing somewhat in detail in the different locali-
ties, these older beds in the vicinity of Clarendon, at Mul-
berry Cafion, at Mount Blanco, and at Tule Cafion and Rock
*Protohippus mirabilis = Merychippus mirabilis.
2 Hippotherium = Neohipparion.
3 Hipparion = Neohtpparion
* Local name proposed by the writer.
Buiretin A. M. N. H. Vou, XIX, Plate LVI
| OKLAHOMA
AREA OF ee RATION
don prodiqctys"@) DDilno
i
|
i S, BLANCO
sles adon@ <3) ina sed mirificus
Sy
34 \
NEL PASO A) ae 8
‘ %
“Cos 9 .
Pe ure
. S
AUSTIN
Vox
\ es :
\ es “2, 37
. . ai GALVE STON
% Elephas imperator® VICT OBIAZ
aes
J 7 io
MEXICO & y
SS
=
ey. (
y
N
Scale: + = 100 Miles “4. a
MAP oF TEXAS.
Showing area of Explorations.
BucceTin A. M. N. H. VoL. XIX, Pirate LVIII.
OOTY
3
s = S25, bits < Ein: (1) Dinocyon gidl
3s * Pals -0.4.04 procdetus és
Re CI
=
oy
=
PleistocenesRecen
Pliocene ( Blanco)
Miocene (Loup Fork)
Miocene
M. 1a |
Triassic
l=lacustrine, r=fluviatile, e=eolian. Explorations of the
American Museum
1899-1901, under
sakae LUE Osa Expedition of 1899. J.W. Gidley.
ce Ay ls tea oy » 1900.
ca ee see » » 1901.
10 20 30 40 50 60 70 80 90
0
Scale: = —
——= 100 Miles.
Pleistocene = Rock Creek Beds.
Pliocene = Blanco Beds.
Miocene (Loup Fork) = Clarendon Beds.
Miocene = Panhandle Beds.
1903.] Gidley, Fresh-water Tertiary of Northwestern Texas. 635
Creek are evidently of the same age. They closely resemble
each other in a general way, and the formations are traceable
from one locality to another around the irregular escarpment
of the Plains. Numerous wells dotting the Staked Plains
show everywhere the existence of these beds. They are of
nearly uniform thickness, and form practically the whole area
of the Staked Plains.
SUMMARY.
Following is a summary of the conclusions reached by the
writer from this study of the formations of northwestern
Texas:
_(1) There has been no great disturbance or change of level
in the region of the Staked Plains since the close of the
Triassic, hence the strata of the Triassic which underlie this
whole region are for the most part nearly horizontal, and the
country at the beginning of the Miocene was comparatively
level.
(2) The Panhandle (Lower or Middle Miocene) beds were
comparatively evenly distributed over the vast area now
occupied by the Staked Plains and in addition extended
westward to the Rocky Mountains in New Mexico, and spread
out to the eastward over a much greater territory than they
now occupy. These deposits seem to be, at least partially,
lacustrine in origin.
(3) All the formations of the Staked Plains that are of more
recent date than the Lower or Middle Miocene are represented
by comparatively small areas, and are fluviatile, or eolian and
fluviatile, in origin. These later depositions are represented
by the Clarendon beds in the vicinity of Clarendon, the
Blanco Beds at Mount Blanco, and the Rock Creek beds at
Tule Cafion and Rock Creek.
Skull of Triceratops Serratus.
By RicHarp Swann Lu tt, Pu.D.
AUTHOR’S EDITION, extracted fom BULLETIN
OF THE
American Blusewm of Hatural History,
VoL. XIX, ARTICLE XXX, pp. 685-695.
New York, December 24, 190}.
b:
; The tknickerbocker Press, Rew
hd
Article XXX.— SKULL OF TRICERATOPS SERRATUS.
By RicHARD Swann LULL, Ph.D.
1eWGN IID
The American Museum Expedition of 1902, under Mr. Bar-
num Brown and the writer, which was sent by Professor
Osborn into the Laramie formation of Montana had the good
fortune to secure, among other material, a fine specimen of
Triceratops serratus Marsh. The exact locality in which the
specimen was found was in the wall of Hell Creek Cafion,
some twenty-five miles from the Missouri River, and one
hundred and thirty-five miles northwest of Miles City,
Montana. The unconsolidated sand matrix has been entirely
removed from the skull, thus affording an exceptional oppor-
tunity for the study especially of the remarkably preserved
palate.
Through the courtesy of Professor Charles E. Beecher the
writer was permitted to study the type skulls of Triceratops
prorsus and of T. serratus which are preserved in the Peabody
Museum at Yale University. This confirmed the opinion
already formed that the American Museum specimen is refer-
able to the latter species. The agreement between the speci-
mens is close, the main points of difference being the inferior
size of the type specimen, which is evidently that of a younger
animal, and that the median ridge of the parietal crest or frill
is not so prominent in the American Museum specimen; nor
are the bony projections along the ridge quite so conspicuous as
in the type; but in general proportions, the form and arch of
the frill, the shape of the orbit and other points mentioned by
Marsh in his specific definition the resemblance is very close.
Triceratops serratus Marsh.
Marsu, O. C., 1890, Amer. Jour. Sci. (3) XX XIX, p. 81.
Marsh, O. C., 1890, Amer. Jour. Sci. (3) XX XIX, p. 425, pl. v, fig. 2;
pl. vi, figs. 1-6.
Marsu, O. C., 1896, Sixteenth Annual Report U. S. Geol. Survey, p.
208; pl. Ix, fig. 3; pl. 1xi, figs. 7, 9, ro.
[685]
686 Bulletin American Museum of Natural History. [ Vol. XIX,
Materials. — The skull lacks only the distal portions of the
postfrontal horn cores, the nasals and their horn core, and a
portion of the premaxillary bones. The rostral bone was
found displaced but a short distance to the rear on the right
side of the muzzle, while on the other side lay the left mandi-
ble in perfect condition. The codssified right angular and
articular, together with portions of both splenials, were found
beneath the skull. One badly preserved humerus, half of
another, a radius, five metacarpals, three phalanges, a fibula,
and fragments of a scapula complete the list. The specimen
is No. 970 of the American Museum fossil reptilian collection.
THE SKULL.
The condition of the sutures, the fact that the rostral bone
had not ankylosed with the premaxillaries, and that the so-
called epoccipital bones, the lozenge-shaped ossicles around
the margin of the frill, were not sufficiently codssified with the
latter to prevent the loss of some of them, give evidence that
in spite of the enormous size of the animal it had not yet
reached maturity. The maxillary teeth have dropped out of
position with one exception, a tooth which lay deep in the
alveolar channel of the right side. Other teeth, found loose
in the matrix, were clearly of the upper series and are shown
in position in the photograph (Plate LIX). The rostral bone is
highly rugose, due to the impressions of blood-vessels over its
surface showing it to have been closely sheathed in horn. The
forward border is a full, gentle curve, while the inferior margin
is straight and nearly horizontal when the bone is in position,
as in most Testudinata. This, together with the form of the
predentary bone, which curves upward towards the tip, would
seem to indicate a cutting beak very turtle-like in aspect, as
one would be led to expect from somewhat similar feeding
habits, rather than the trenchant, downwardly curved, rap-
torial beak usually given to the restored Triceratops. The
fact that in Chelydra, where the upper beak is hooked, the bone
which supports it is of similar form, may be taken as corrobora-
tive evidence.
BuLvetin A. M. N. H. VoL. XIX, Prate LIX.
TRICERATOPS SERRATUS JZarsh.
Palatal aspect of skull, with frill. »%< ~;. (Extreme length about 6 ft. 4 ins.).
1903. | Lull, Skull of Triceratops Serratus. 687
° nd LL GL a
Palatal Aspect.
The premaxillaries are only in part preserved. The max-
allaries are heavy bones uniting anteriorly in the median line,
in front of the narial fenestre, by a pronounced dentate
suture. Anteriorly the premaxillaries overlap them above
and posteriorly they bifurcate, one limb, the dorso-lateral, run-
ning obliquely outward and backward to join the jugal, while
the postero-ventral limb unites posteriorly with the ptery-
goids. The maxillaries thus form the anterior and about two
thirds of the lateral walls of the large narial fenestre. In the
palatal face lies the alveolar channel, sculptured transversely
into a number of shallow grooves, incomplete sockets in which
the teeth were formed, forty in the right channel and forty-two
in the left. The dental channel, is 482.8 mm. in length with
an average width of 30 mm., which is somewhat less than its
original measurement, owing to crushing. Other measure-
ments may be found in the table on page 694. As in the
mandible, a row of dental foramina runs along the inner face of
the maxilla, one foramen being opposite each alveolar groove,
through which passed the blood-vessels needful for the rapid
forming of teeth in the dental magazine. The external face
of the bone also bears two such foramina.
The vomer, or ‘prevomer’ as determined by Broom,} is a
slender rod-like bone bridging fore and aft the space of the
narial fenestra. Anteriorly it is dilated into a flattened
rhombic expansion articulating by a squamous suture with
the united maxillary bones. Passing backward there appears
a median ventral keel giving the bone in its narrowest part, _
about the middle, a triangular section. Further to the rear
the lateral edges bend downward to the level of the median
keel and then rise again to their former level, where they give
rise to thin plate-like expansions which are embraced at their
posterior end by the pterygoid bones. Dorsally viewed the
vomer is seen to become trough-like, the depression being
about the width of the shaft of the bone and running the
1 Broom, R., Proc. Linn. Soc. N. S. W., 1902, pt. 4, PP- 545-560.
Triceratops serratus Marsh, Palatal aspect of skull. x. A, rostral bone, War., narial vacuity,
2.pt.s., interpterygoid space, D.g., dental groove, ? Ews., Eustachian groove (?).
[688]
1903. | Lull, Skull of Triceratops Serratus. 689
length of the expanded posterior portion. There is no trace
of paired elements in the vomer.
The palatines bound laterally the posterior part of the
narial fenestre, and are somewhat triangular in shape, with
the base of the triangle meeting the maxillaries in a squamous
articulation somewhat overlapping the dental foramina.
Posteriorly they are bounded by the pterygoids, and the
anterior portion runs upward over the jaw until it ends in a
large vacuity on the dorsal side. This vacuity is further
bounded anteriorly and externally by the maxillary, and
posteriorly by the pterygoid, and it lies above a point one
third of the distance from the posterior end of the dental
channel.
The pierygoids are large and irregular with peculiar channels,
probably the eustachian canals, running obliquely from the
articulation with the posterior end of the maxillaries to the
median line; these channels are formed by thin, overarching
ridges of bone which in their mid-length almost meet. The
pterygoids form the postero-lateral margins of the narial
fenestre in the rear of the palatine bones and embrace the
hinder.end of the vomer. Anteriorly they are bounded by the
palatine and maxillary bones and possibly by the ectoptery-
goids, though the last-mentioned cannot be located in this
specimen'; the ectopterygoids are not suturally separated
from the pterygoids themselves. Posteriorly the pterygoids are
met by the basisphenoid and in thé median line they nearly
embrace the parasphenoid, or ‘ vomer’ of Broom. Laterally
they are broad and thin plate-like expansions which pass out-
‘ward and backward to meet opposing processes of the quad-
rate, though the precise limit of the pterygo-quadrate suture
is not everywhere distinct.
The quadrates are well developed and firmly fixed in place
by the pterygoids within and the quadratojugals without.
They also pass backward and upward, forming, with the
quadratojugals, the lower boundary of the infratemporal
fossa. Posteriorly they join the squamosals, which are widely
expanded to form the lateral elements of the frill. The
1 Marsh, O. C., Amer. Jour. Sci. (3), XLI, p. 171.
[December, 1903.) 4A
690 Bulletin American Museum of Natural History, [Vol. XIX,
quadrate is flattened on its ventral aspect and somewhat cylin-
drically concave on its dorsal surface. The head is elongated
transversely to a length of 147 mm., the facet which articu-
lates with the lower jaw being somewhat saddle-shaped.
The posterior end of the quadrate is embraced between the
exoccipital and the squamosal bones. The quadratojugals are
comparatively small bones lying between the quadrates and
the jugals. Dorsally they extend in thin, plate-like expan-
sions between the aforesaid bones, and in their posterior por-
tions form part of the infratemporal arcade, almost, if not
quite, meeting the forwardly extending process of the squa-
mosals. In their ventral portion where the distance widens
between the quadrates and the jugals the quadratojugals
dilate into a thick, wedge-shaped mass to fill the gap. The
greatest thickness is 89 mm.
The occipital region of the skull is rendered very massive to
support the great weight of the head, the sutures between
the various elements being closed. The occipital condyle is
almost spherical, and has a diameter of 115mm. It looks
almost directly backward and but little downward. - An-
teriorly it merges into a heavy basioccipital and laterally into
the exoccipitals, the limits of these three elements in the
condyle itself not being discernible.
The bastocctpital diverges into two stout limbs with heavy,
rugose extremities, in front of which appear the pulley-like
basisphenoid bones, the parasphenoid (‘ vomer’) arising be-
tween the limbs.
The exoccipitals run out laterally to join the quadrates and
Squamosals, overlapping the former and firmly articulating
with the latter to afford a strong brace across the entire base
of the frill. They thin away posteriorly and are, together with
the supraoccipital, overlain by the largely developed parietals
which form the median element of the frill.
The ventral aspect of the frill or crest is well shown in
Plate LIX, and is without vacuities of any sort, although
just behind the exoccipital bones the parietals are excessively
thin. Vascular impressions occur on the posterior half of the
parietal bones on either side, but there is no evidence of a
1903. | Lull,, Skull of Triceratops Serratus, 691
wide free margin sheathed with horn as in the frill of Tri-
ceratops prorsus.*
The squamoso-parietal suture is a squamous one for a short
distance backward, the squamosal overlapping; but at the
point where the suture bends outward it becomes a plain
harmonic suture having but little strength, as is evidenced
by the fact that in the specimen under consideration the
squamosal bone had slipped dorsally past the parietal on the
right side, while on the left the bones were flush with each
other. In the type specimen of Triceratops serratus in the
Yale University Museum both sutures, that on the left as well
as on the right, have slipped. The frill thus seems to have
afforded leverage to assist in moving and supporting the huge
head with its weighty armament and also to have protected
the neck against the assaults of enemies, but it seems hardly
probable in the present species that the dorsal part could have
withstood crushing blows- without injury to the frill. The
hinder margins of the parietals have decomposed somewhat
and the marginal ossicles are here wanting, though most of
them are present on the squamosals.
Dorsal Aspect.
The anterior part of the skull has been weathered off, as it
formed the outcrop of the specimen, and much of the bone has
been disrupted by grass roots even where it had not yet been
exposed by erosion.
The postjrontals, with the exception of the horn cores, are
entire and the underlying sinus is readily explored through the
large postfrontal fontanelle (the parietal or pineal foramen of
authors). This sinus is continuous with those of the horn
cores and in turn with the space within the skull behind the
orbits, but not with the brain case. It is more or less wedge-
shaped, tapering dorso-ventrally as one goes forward, the
anterior limit being just in front of the orbit. The flat roof is
formed by the overlying postfrontal and frontal bones, while
the sinus is laterally constricted into three chambers. The
anterior chamber has a rather flat floor and is separated from
1 Marsh, O. C. 1806. Dinosaurs of North America, pl. lx, fig. 4.
692 Bulletin American Museum of Natural History. [Vol. XIX,
the median chamber by vertical pillar-like bones, one on either
side, which serve also to support the antero-internal portions of
the horn cores. The floor of the second or largest chamber is
deeply excavated, and it is this chamber which communicates
with the horn-core sinuses by openings in the lateral walls.
The posterior chamber, lying just beneath the fontanelle, is
small and round. and in the specimen in question has a small
pencil-like bone running obliquely from the left lateral wall to
the floor, after the manner of a flying buttress. There is no
indication of a pineal foramen opening into the brain case
which lies directly beneath the above sinus; hence the Cera-
topsia agree with other Dinosauria in this respect. The post-
frontal fontanelle closes in old animals, as in the type skull of
Triceratops prorsus, which is that of a fully adult though com-
paratively small animal, and is thus analogous to that in the
skull of the human infant.
The loss of the frontals and nasals from our specimen
renders possible the study of the interior of the skull, the bones
of which are admirably preserved, and while the entire skull
gives an appearance of massiveness, the individual bones are
comparatively thin, but so constructed as to brace in the
most admirable manner the portions of the skull subject to
strains and impact, especially beneath the horns.
The jrall viewed from above presents much the same relative
expanse of bone as is shown in the ventral aspect except that
the squamosals now extend forward and upward to the base of
the horn cores. Anteriorly they are bounded by the jugals, the
infratemporal vacuities, and the quadrates. On one squa-
mosal, and to a less extent on the other, a ridge for muscular
attachment extends diagonally upward and backward across
the posterior portion of the bone. The parietals have the
same extent as in the ventral view except that here they
overlie the occipital bones and articulate with the postfrontals
at about the posterior limit of the horn cores. The supra-
temporal vacuities open forward beneath the postfrontals and
above the parietals into the main sinuses of the skull. Large
blood-vessels had their exit through these vacuities, their
branches being deeply impressed into the surface of the parie-
1903. | Lull, Skull of Triceratops Serratus. 693
ee ee ee
tals and to a less extent into the squamosals, thus implying a
compactly fitting integument. The base, especially of the
right horn core, is well preserved. It is extremely hollow, but
with a shelf-like circular projection of bone running around the
inner wall just above the level of the postfrontal bones with-
out, and doubtless to aid in resisting the thrust of the latter
bones when lateral pressure was brought to bear upon the
horns. Around the outside base of the horn is a horizontal
ridge which may have supported the base of the horny sheath.
The orbits are nearly circular and are surrounded by a thick-
ened ridge of bone, especially in front. The downward and
outward crushing of the left horn core has partially closed the
left orbit, adding to the sinister expression of the skull.
THE LOWER JAw.
The left mandible, which is admirably preserved, consists of
dentary, surangular, and coronoid, with a full magazine of
thirty-nine vertical rows of teeth. On the inner face is a row
of thirty-eight dental foramina, and the meckelian groove on
the inferior face is wide and deep, but was covered by the thin,
plate-like splenial which, though lying detached in the quarry,
presents a perfect contact when placed in position. Cope *
claims that in Hadrosaurus it is the splenial which contains the
magazine of teeth. Whether or not this be true of Hadrosaurus
it is certainly not true of Triceratops, in which the magazine is
contained in the dentary in the normal manner. The teeth
arise in alternate series in the successive vertical rows, only one
series being in full use at one time, though those of the
secondary series, arising between the teeth of the primary
series, show partial wear, while in the posterior part of the
jaw individual teeth of the primary set are already succeeded
by tertiary teeth. The vertical worn faces of the teeth present
the surface known to mathematicians as an hyperbolic para-
boloid or warped surface; the whole mechanism reminding one
of aslightly twisted saw with alternating higher and lower teeth.
Marsh notes the fact that in the Ceratopsia the teeth are
double-rooted, a feature almost unique among reptiles. This
1 Cope, E. D., Amer. Naturalist, July, 1883, p. 775.
694 Bulletin American Museum of Natural History. (Vol. XIX,
seems to have been brought about by the mechanical necessity
of a base widened transversely to meet a lateral strain in the
shearing process of mastication and the subsequent constric-
tion of this base into an inner and outer pillar due to the
crowding of the crowns of adjacent teeth set at a lower level.
There could have been no lateral movement in mastication,
but a chopping motion, possibly with a slight orthal movement
combined with it. The food gathered by the cutting beak
was probably chopped into short pieces by the teeth, being
kept in the mouth by the muscular wall of the cheeks. It is
doubtful whether the gape of the mouth had a posterior extent
further than the anterior end of the tooth series, as otherwise
the portions of food chopped off, falling outside of the lower
teeth, could not have been retained in the mouth.
The alveolar grooves are equally developed in the inner
surface of both inner and outer walls of the dental channel
and not in the inner surface of the outer wall only as in
Trachodon (Hadrosaurus) as shown by Lambe.! This is due
to the fact that in Triceratops the crowns of the teeth do not
form so flat a tassellated pavement when viewed from within;
their position in the jaw being more nearly vertical than in
Trachodon.
Measurements.
Lensth of skull (estimated). 4.55 oe eee ee 2160 mm.
Width across: frill. .:. 2c6.c.a2). fas Sai cee ae eae ee eee 1578
Maxillary bones, Lemp thiaen s.r sme ae ee me ee 672
> length ofmentalichann clare er 482.8
ie average width dental channel.......... 30
Premaxillary bones, width at posterior end... aes) 177
Vomer, lengthwt.: acd oilae eee ee ee eee 410
ey widthatcanterionendiian 4: een e eee 70
““_\ width of shalt. i-307 25 toe See een eee 15
Palatine. bones, lengthy... =o. 2 45 ae ee eee 293
Occipital condyle,.diameter) (37 a5 5 ee ee L205
Poramen aorta, awa Glieey lence ee en pee 47
> be evo ty... ! jaune sk ee ee 36
Basioccipital bone, widths). oe ac eee ae 280
Exoccipitals, distance from tip) tor tipi.) eee eee 790
1 Lambe, L. M., 1903, Ottawa Naturalist, Vol. XVII, pp. 136, 137; Osborn, H. F.,
and Lambe, L. M., 1902, Contributions to Canadian Paleontology, III, Part II, p. 73.
1903. | Lull, Skull of Triceratops Serratus.
Exoccipitals, greatest fore and aft width...............
ce least ay “eo “é ee
“cc “ce ec “cc
“cc cc oe “e
“cc “é “e “cc
: zs width
PHoteavempordlowacwity, lengths... 6 de. vee ee noes eats
i. i etohiGages Suey eet stares ere?
Right horn core, longitudinal diameter at base .........
a a “transverse 3 ay hm yeni te
“S (@HSMLR UKeralea Rol Aretguaen aay tenn Becidim eho ee oceno eons tc CLO
% 2 om Sol TET olen sly Ae a ed Re SERA EU eS RoR cc
cower jaw, lenoth of dentary....:...- +. 002.6022 oe 0%
r ‘““ greatest depth to summit of teeth..........
ie eI tEOl COLONO1IG a teemioe aie eee rere
ste “length of dental channel ..........---+se0-
Parietal-bones, ventral aspect, length?......-.2...s..5~.<
Leas twa Glisten
ageeNKES GT Baoosagasco as
Squamosal bones, ventral aspect, length...............
WG yer igatcun wen eenerleratere
Postirontal fontanelle, height.....0... 0.2.6 s0s%se enue
Sires (el wi ieJe a) elie elo) secs! s!\6 «ui vane serene
Ce
2
Lhe Skull of Creosaurus.
By Henry Fatrrirenp Osporn.
AUTHOR’S EDITION, extracted from BULLETIN
OF THE
American Busewum of Aatural History,
Vou. XIX, ARTICLE XXXI, pp. 697-7or.
New York, December 24, 1903.
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Article XXXI. — THE SKULL OF CREOSAURUS.
By Henry FParrrieip Osporn.
The skull of the Jurassic Carnivorous Dinosaur Creosaurus
is represented by two specimens, from the Bone Cabin Quarry,
in the American Museum Collection, Nos. 600 and 666, which
supplement each other admirably and give a very complete
knowledge of all except the palatal region. The side and
back view of No. 600 is presented in figures 1 and 2.
The chief features are: (1) the presence of three preorbital
fenestra, (2) the great elongation of the facial region, (3) the
abbreviation of the temporal region, (4) the wide extensiort
of the parietals on the occiput, (5) the deep depression of the
_-- quadrate and its rounded articular connection superiorly with
4 the squamosal.
Be Proportions. — The skull is relatively high and narrow, the
chief measurements being as follows:
4 No. 600 No. 666.
zP Length, occipital condyles to premaxille...... 81omm. 885mm.
fn Depth, parietal crest to distal extremity of
¥ ALCAN R Ea sg cla GoD CED ey ena aT 415 430
¥e Mirdch. Of occiput across pafietals...........5: : 230
z Vertical measurement of quadrate............ 205 235
}
Openings. — The openings are placed as follows: (1) the
paired narial openings, of elongate oval form, at the sides of
; the snout, bounded by the premaxille, nasals, and maxille;
ee (2) the greatly abbreviated supratemporal fenestra; (3) the
deeply extended latero-temporal fenestra (the post-temporal
fenestra is entirely closed, and there is no pineal opening);
(4) there is a slit-like foramen between the quadrate and
quadratojugal; (5) the orbital openings are deeply vertical;
; (6) the preorbital fenestra, f’, is by far the largest vacuity in
me) the skull, and is bounded by the maxillaries, prefrontals, and
j jugals; (7) a much smaller second preorbital fenestra, f*; 8
bounded entirely by the maxillaries; (8) the third preorbital
» — fenestra, 7’, lies between the premaxillaries and maxillaries,
and is very small.
[697]
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700 Bulletin American Museum of Natural History. [Vol. XIX,
The premaxillaries retain the alveoli of five teeth, and re-
semble the type premaxilla of Creosaurus atrox, a genus which,
however, has not yet been clearly separated from Allosaurus.*
The maxillaries contain the alveoli for from 16 to 18 teeth.
Length, 460mm. The masals are remarkably elongate, 470
mm. (No. 666), and gradually increase in breadth posteriorly ;
in No. 666 they are continuous back to the frontals; in No.
600 there is a fracture or suture above the preorbital fenestra
which is not observed in No. 666, and, therefore, is apparently
accidental. The frontals are extremely short, 140 mm. in No.
666, extending back to a point midway between the supra-
temporal fosse. The superior portion of the parietals is also
abbreviate, but laterally and posteriorly they form two large
transversely extended vertical plates constituting the crest or
superior and lateral portions of the occiput. The narrow
supraoccipitals apparently extend partly between the parietal
crest above; below they are in contact with the conjoined
exoccipitals. The exoccipitals unite in the median line above
the foramen magnum and extend outward in two broad wing-
like plates with which the paroccipitals are coalesced. The
bastoccipital alone enters into the prominently convex condyle.
The lateral portions of the occipital region are formed by the
posterior plates of the squamosals, which fill the bordering
angle between the parietals and paroccipitals; in side view the
squamosals are indented by a spur of the postorbito-frontals,
and send a descending process for articulation with the
ascending process of the quadratojugals. The quadrates are
vertically extended with a singly convex head above for a
jointed articulation with the squamosals, and a doubly convex
transversely extended distal extremity for union with the
articulars. The quadratojugals are placed on the outer sides
of the quadrates, and are shorter elements than the jugals.
The jugals are triradiate bones articulating with the quadrato-
jugals, postorbito-frontals, and prefronto-lachrymals, respec-
tively. The elements bounding the orbits anteriorly, uniting
with the jugals and maxillaries below and with the nasals and
1 This matter has been carefully discussed by S. W. Williston: ‘The Dinosaurian
Genus Creosaurus, Marsh.’ Amer. Journ. Sci., Vol. XI, root, p. 111.
1903. | Osborn, The Skull of Creosaurus. 701
frontals above, are considered to represent a complex com-
posed of the prefrontals and lachrymals; they are surmounted
by a stout tuberous process which is deeply excavated on the
antero-external surface; this process may have supported
something in the nature of a low dermal horn.
Below the condyles the basioccipitals descend abruptly
downward and backward; on each side of this downward
projection are the apparently inferior plates of the exoccipi-
tals. From these inferior plates two still more depressed
wings extend downward, which probably represent antero-
inferior processes of the basisphenoids for articulation with
the pterygoids.
Lower jaw. — In the lower jaw the articulars, angulars, and
dentaries are clearly determinable. The suture between the
coronoid and angular is faintly determinable posteriorly, the
coronoid being depressed. The convex element extending
along the inner side of the dentary but firmly coalesced with
it may represent a presplenial; if this is truly the presplenial
it appears to extend as far forward asthe symphysis. There
are apparently sixteen alveoli for the laterally compressed
lower teeth in the dentary.
A fuller description of these skulls will appear in a forth-
coming memoir.
AMERICAN MUSEUM OF NATURAL HISTORY,
NEW YORK CITY, U. S.A.
SUPPLEMENTS TO THE AMERICAN Museum JOURNAL.
The following Guide Leaflets to the Hall of Fossil
Vertebrates will be prepared under the direction of
Professor HENRY FAIRFIELD Osporn, Curator of the
Department of Vertebrate Paleontology :
I. The Hall of Fossil Vertebrates. Introduction and
General Guide. (Published January, 1902.)
I]. Evolution among Fossil Mammals as illustrated
by the AmgEricAN Museum Collections.
Section 1. Phe Horse.
The Camel.
pp
2. Dhe Titanenners
A. he Khinoceros:
5. Lhe. Elephant
Dib xine: Carnivora.
IV. The Dinosaurs or Giant Reptiles.
Section 1. Sauropoda.
V. Marine Reptiles.
Section 1. Mosasaurs.
bo
Ichthyosaurs.
Plesiosaurs.
G2
VI. The Fossil Fields of the West, their Topography
and Geology, with a brief account of the Field
Expeditions sent out by the American MusEuM
or NATURAL HISTORY.
February, 1902.
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AMERICAN MuseuM oF NATURAL History,
DEPARTMENT OF VERTEBRATE PALAZZONTOLOGY.
ProvisionaL GuIDE
10) “anistia’
Hall of Fossil Vertebrates.
(No. 402, FourTH FLoor, East Winc.) >
EDITION OF Ig00-Igot.
The collections in this hall are not yet complete nor permanently arranged.
We cannot therefore issue a guide-book at present.
The arrangement of the specimens is intended to
show the history or evolution of different races of
animals, chiefly in North America. All the specimens of one
race or kind of animal are placed together, the most ancient
first, the most recent last.
All the skeletons in this hall are those of extinct
animals. The Mastodon and Great [rish Deer are half-petrified
bone dug out of peat bogs. Ali the others are petrified (z. ¢.,
have been buried so long that they are converted from bone into
stone), and have been chiseled out of the solid rock. The
Megatherium is a plaster cast.
Some of the skeletons are partly restored in plaster, indicated by a red
cross (restored bones) or red lines (outlines of restored parts of bones). Bones
supplied from other specimens are marked with the catalogue number of the
specimen, or indicated by a red circle if uncatalogued.
The most ancient of the extinct animals shown here
are the Dinosaurs or great land Reptiles, Mosasaurs or great
AMERICAN MUSEUM OF NATURAL HISTORY,
marine Lizards, and other creatures of the Age of Reptiles.
These are millions of years old. Some of the Dinosaurs are the
largest known land animals, longer than the width of this hall, and
standing on all fours with the back reaching nearly as high as the
ceiling.
The rest of the specimens are mammad/s (animals which suckle
their young, including most four-footed beasts). Most of them
are hundreds of thousands of years old, ranging from perhaps
three million years down; these lived long before man had
appeared on the earth. A few, including the Mastodon, Mam-
moth, Megatherium, Irish Elk, One-toed Horse and others, are
of the latest geological age (the “ Pleistocene ”), and, while tens
of thousands of years old, were contemporaries of primitive
man.
Many of the extinct animals are allied to animals still living,
and are called by the common names of their modern relatives.
Thus we have extinct Horses, Rhinoceroses, Tapirs, Camels, etc.
Other races have died out completely and are not related to any
living animals. For these there ts no common name, and we have
to coin a name from their (Latin or Greek) scientific name, call-
ing them “ Titanotheres,” “‘ Dinosaurs,”’ etc.
The best example of Evolution is the race of extinct
ancestors of the Horse, shown in the right hand far corner of the
hall. Next best is the series of Camel ancestors, on the left hand
side near the far end, and the series of ancestors of Titanotheres
and ancestors of Rhinoceroses, skulls and skeletons ranged along
the right hand (south) side of the hall beginning at the entrance.
All these series are arranged according to age, the most ancient
(oldest or first stage of evolution, found in the lowest rock-strata)
first, the most recent (final stage of evolution, found in the upper-
most rock-strata) last.
DEPARTMENT OF VERTEBRATE PALAEONTOLOGY.
On the left side of the hall are arranged the Amblypods,
ancient hoofed animals long ago extinct, unlike any living animal,
although suggesting elephants, rhinoceroses, hippopotami and
bears, in different parts of the body. Next there are the Condy-
larths, the most ancient of hoofed mammals, then the Creodonta
and Carnivora (flesh-eating mammals), the Insectivora (insect-
eaters), Rodents (gnawers), and other small and primitive
mammals. Then come the Elephants and the various kinds of
cloven-hoofed animals, allied to modern pigs, deer, sheep and
cattle.
The Dinosaurs or Giant Reptiles are temporarily placed
at the end of the hall and in the two high cases to the left of the
centre aisle. Small models of restorations of three kinds of
Dinosaur will be found in an A-case near the end of the hall left
of the centre aisle.
The Mosasaurs or Great Marine Lizards are represented
by the skeleton on the wall of the corridor by the staircase. This
corridor will ultimately be filled with great marine reptiles.
The Water-color Restorations by Charles Knight, mainly
based on complete skeletons exhibited in this hall, show the
probable appearance of the different extinct animals, according to
our best judgment, as indicated by the characters of the skeleton,
appearance of their nearest surviving relatives, and the habits of
life for which the animals seem to have been fitted.
The Window Transparencies are enlarged from photo-
graphs of the regions where the fossils occur, and generally show
the localities where unusually fine specimens in this hall were
found.
AMERICAN MUSEUM OF NATURAL HISTORY.
The Pillar Cards and general Labels in the cases give
detailed information about each group of fossils.
The Synoptic Case in the centre of the middle aisle illus-
trates the method by which the fossils are collected and con-
veyed to the Museum.
The Charts at each side of the entrance show the order in
which the rock strata lie, one over another, and the kinds of
fossils found in each stratum.
WHAT THE EVOLUTION SERIES PROVE.
Each series of specimens is arranged according to the age of the strata in
which they are found. This age is known by the order in which the strata lie
one over another. In all ordinary circumstances the undeslying stratum must
have been deposited first, the overlying one afterwards. In each stratum is
found most abundantly a particular species of a race of animals and all the
fossil specimens of that race ever fougd in that stratum are like the said
species within quite narrow limits. Arranging the characteristic species from
each stratum in order of the age of the strata, we find that they show a regular
uniform change from the most ancient to the most recent. At no point in a
given series can we draw a line and say: This is, and that is not, a horse—
or a camel—or a rhinoceros. The visitor, therefore, can demonstrate for him-
self the Evolution of the race of Horses, or Camels or Rhinoceroses, within
certain limits. (Of the evolution of Man we have not satisfactory illustration
from fossils.)
It should be observed that the evolution of a race consists mainly in the
adaptation of the structure of the animals to particular surroundings and habits
of life. There is also a universal progress in intelligence, the older animals
having relatively smaller brains.
AMERICAN MUSEUM OF NATURAL HISTORY
William D. Matthew, Ph.D.
Associate Curator of Vertebrate Palzontology
SUPPLEMENT TO AMERICAN MUSEUM JOURNAL
VOL. III, No. 1, JANUARY, 1903
Guide Leaflet No. 9
American Museum of Natural History.
Officers.
President,
Morris K. JESupP.
First Vice-President, Second Vice-President,
WitiiaM E. Dopce. Henry F. OsBorn.
Treasurer, Director,
CHARLES LANIER. Hermon C. BumpPus.
Secretary and Assistant Treasurer,
Joun H. WINSER.
Scientific Staff.
Director,
Hermon C. Bumpus.
Department of Public Instruction.
Prof. ALBERT S. BICKMORE, Curator.
Department of Geology and Invertebrate Palaeontology.
Prof. R. P. WHITFIELD, Curator.
Epmunp O. Hovey, Ph.D., Associate Curator.
Department of Mammalogy and Ornithology.
Prof. J. A. ALLEN, Curator.
FRANK M. Cuapman, Associate Curator.
Department of Vertebrate Paleontology.
Prof. HENRY FAIRFIELD OsBorn, Curator,
W. D. MattHEew, Ph.D., Associate Curator.
O. P. Hay, Ph.D., Assistant Curator.
Department of Entomology.
WILLIAM BEUTENMULLER, Curator.
Departments of Mineralogy and Conchology.
L. P. GratacaP, A.M., Curator.
Department of Invertebrate Zodlogy.
Prof. HERmon C. Bumpus, Curator.
Grorce H. SHERwoop, A.M., Assistant Curator.
Department of Anthropology.
Prof. FREDERIC W. Putnam, Curator.
Prof. Franz Boas, Curator of Ethnology.
MarsHa.u H. Savitve, Curator of Mexican and Central
American Archeology.
Haran I. Smirn, Assistant Curator of Archeology.
Library.
A. Woopwarb, Ph.D., Librarian.
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ILLUSTRATIONS. OF EVOLUTION AMONG FOSSIL
MAMMALS.
A.— THE Horses.
By W. D. Mattuew, Ph.D.,
Associate Curator, Department of Vertebrate Paleontology.
As a domestic animal the Horse is to be found almost every-
where that man can live. He is spread all over the world —
from torrid to arctic climates, in all the continents, in remote
oceanic islands — he is completely cosmopolitan. But as a wild
animal the Horse is at present limited to the Old World, and is
found there only in the open arid or desert plains of Central Asia
and Africa. There are two species in Asia, the Asiatic Wild Ass
(Equus hemionus), and the little known Przewalsky’s Horse (E.
przewalskit), while in Africa there are the African Wild Ass (E.
asinus) and the several species of Zebra (E. zebra, E. burchelli,
i. quagga). In the Americas and Australia there are no true
wild horses, the mustangs and broncos of the Western Plains and
South America being feral (domesticated animals run wild) and
descended from the horses brought over from Europe by the early
white settlers. When the Spaniards first explored the New
World they found no horses on either continent. The Indians
were quite unfamiliar with them and at first regarded the strange
animal which the newcomers rode with wonder and _ terror,
like that of the ancient Romans when Pyrrhus and his Greeks
brought elephants—‘‘the huge earth-shaking beast”’*—to fight
against them.
The Horse is distinguished from all other animals now living
by the fact that he has but one toe on each foot. Comparison
with other animals shows that this toe is the third or middle
digit of the foot. The hoof corresponds to the nail of a man or
the claw of a dog or cat, and is broadened out to afford a firm,
strong support on which the whole weight of the animal rests.
Behind the ‘“‘cannon-bone”’ of the foot are two slender little
1 Macaulay —‘‘ The Battle of Lake Regillus.”
5
J
4 EVOLUTION OF THE HORSE
bones, one on each side, called splint-bones. These represent the
second and fourth digits of other animals, but they do not show
on the surface, and there is nothing like a separate toe. So that
the horse may be said to be an animal that walks on its middle
finger-nail, all the other fingers having disappeared.
The teeth of the horse are almost equally peculiar. The
molars are long, square prisms which grow up from the gums
as fast as they wear off on the crowns. Their grinding surface
exhibits a peculiar and complicated pattern of edges of hard
enamel between which are softer spaces composed of dentine and
of a material called *‘cement,’’ much like the dentine in quality
but formed in a different way. The dentine is formed on the in-
side surfaces of the enamel while the tooth is still within the jaw-
bone; the cement is deposited on the outside surfaces of the
enamel after the tooth has broken through the jaw-bone and
before it appears above the gums.
Various other peculiarities distinguish the Horse from most
other animals; some of these are shared by other hoofed animals.
The two long bones of the fore-arm (radius and ulna) are separate
in the greater number of animals, but in the Horse, and in many
other hoofed animals they are consolidated into a single bone.
The same consolidation is seen in the bones of the lower leg (tibza
and fibula). The lengthening of the foot and stepping on the end
of the toe raises the heel in the Horse, as in many other animals,
to a considerable height above the ground, where it forms the
hock joint, bending backward, as the knee bends forward. In
these as in various other ways the legs of the horse are especially
fitted for swift running over hard and level ground, just as its teeth
are for grinding the wiry grasses which grow on the open plain.
The Zebra and the Ass have the same peculiar structure of
teeth and feet as the Domestic Horse, and differ only in the color
of the skin, proportions of various parts of the body ete.
Fosstt Horsres oF THE AGE OF MAN.
The Age of Man, or Quaternary Period, is the last and by far
the shortest of the great divisions of geological time. It includes
the Great Ice Age or Glacial epoch (Pleistocene), when heavy
SKELETON OF “EQUUS SCOTTI,” FROM THE LOWER PLEISTOCENE OF TEXAS
Mounted in the American Museum
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EVOLUTION OF THE HORSE 7
continental glaciers covered the northern parts of Europe and
North America, and the Recent Epoch, of more moderate climate
during which civilization has arisen.
In the early part of the Quaternary Period, wild species of
Horse were to be found on every continent except Australia.
Remains of these true native horses have been found buried in
strata of this age in all parts of the United States, in Alaska, in
Mexico, in Ecuador, Brazil and Argentina, as well as in Europe,
Asia and Africa. All these horses were much like the living spe-
cies and most of them are included in the genus Equus. A
complete skeleton of one of them (Equus scottt) found by the
American Museum expedition of 1899 in Northern Texas, is
mounted in the large wall-case. The difference between it and
the Domestic Horse (see framed diagram of modern horse skel-
eton) is chiefly in proportions, the skull shorter with deeper jaws,
the legs rather short and feet small in proportion to the body.
In these characters this fossil horse resembles an overgrown
zebra rather than a domestic horse. We know nothing of its
coloring. It may have been striped, and in this case would have
been very zebra-like; but there are some reasons for believing
that it was not prominently striped. The bones are petrified,
brittle and heavy, the animal matter of the bone having entirely
disappeared and having been partly replaced by mineral matter.
They are not much changed in color, however, and are so per-
fectly preserved that they look almost like recent bone.
All the remains of these native horses which have been found
in America have been petrified more or less completely; this
means that they have been buried for many thousands of years,
for petrifaction is an exceedingly slow process." It serves as an
easy method of distinguishing them from bones of the Domestic
Horse, found buried in the earth. These cannot in any case
have been buried for more than four or five centuries, and have
not had time to petrify.
Remains of these fossil horses from various parts of the
United States are shown in the counter-case. One very rich
IThe so-called petrifaction which occurs in some hot springs, coating
objects dipped into them with a white, stony coat of lime is not true petrifaction.
In true petrifaction the substance of the bone is replaced particle by particle
with mineral matter.
8 EVOLUTION OF THE HORSE
locality is on the Niobrara river in Nebraska, another in central
Oregon. Many separate teeth and bones have been found in the
phosphate mines near Charlestown, 5. C.; other specimens have
come from central Florida, from southern Texas, Arizona, Kan-
sas, Louisiana and even from Alaska. They are, in fact, so
often found in deposits of rivers and lakes of the latest geological
epoch (the Pleistocene) that the formation in the western United
States has received the name of Equus Beds.
In South America, in strata of the Pleistocene Epoch, there
occurs, besides several extinct species of the genus Equus, the
Hippidium, a peculiar kind of Horse characterized by very short
legs and feet, and some peculiarities about the muzzle and the
grinding teeth. The legs were hardly as long as those of a cow,
while the head was as large as that of a racehorse or other small
breed of the Domestic Horse.
All these horses became extinct, both in North and South
America. Why, we do not know. It may have been that they
were unable to stand the cold of the winters, probably longer
continued and much more severe during the Ice Age than now.
It is very probable that man — the early tribes of prehistoric
hunters — played a large part in extinguishing the race. The
competition with the bison and the antelope, which had recently
migrated to America — may have made it more difficult than
formerly for the American Horse to get a living. Or, finally,
some unknown disease or prolonged season of drought may have
exterminated the race. Whatever the cause, the Horse had dis-
appeared from the New World when the white man invaded it
(unless a few individuals still lingered on the remote plains of
South America), and in his place the bison had come and spread
over the prairies of the North.
In Central Asia, two wild races persist to the present day;
others were domesticated by man in the earliest times, and their
use in Chaldzea and Egypt for draught and riding is depicted in
the ancient mural paintings. In Africa the larger species became
extinct in prehistoric times, as in America, but the smaller zebras
still survive in the southern part of the continent (one species,
the Quagga, abundant fifty years ago, is now probably extinct),
and the African Wild Ass is found in the fauna of the northern
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EVOLUTION OF THE HORSE II
part. The Wild Horse of prehistoric Europe, a small race,
short-legged and shaggy-haired, was domesticated by man, a fact
that is known from the rude drawings scratched on bone or ivory
by men of the Neolithic or Polished Stone Age. But the Domes-
tic Horse now in use is derived chiefly from the Asiatic race, al-
though it is probable that in some breeds there is a considerable
strain of this shaggy, short-legged European race, and it is pos-
sible also that African races may have been domesticated and to
some extent mixed with the Asiatic species. The domesticated
Ass is a descendant of the African species.
THE EVOLUTION OF THE HORSE.
The history of the evolution of the Horse through the Ter-
tiary period or Age of Mammals affords the best known illustra-
tion in existence of the doctrine of evolution by means of natural
selection and the adaptation of a race of animals to its environ-
ment. The ancestry of this family has been traced back to
nearly the beginning of the Tertiary without a single important
break. During this long period of time, estimated at nearly
three millions of years, these animals passed through important
changes in all parts of the body, but especially in the teeth and
feet, adapting them more and more perfectly to their particular
environment, namely the open plains of a great plateau region
with their scanty stunted herbage, which is the natural habitat
of the Horse.
In the series of ancestors of the Horse we can trace every step
in the evolution of those marked peculiarities of teeth and feet
which distinguish the modern Horse from an ancestor which so
little suggests a horse that, when its remains were first found
forty years ago, the animal was named by the great paleontologist
Richard Owen, the Hyracotheriuwm or “Coney-like Beast.” Its
relation to the Horse was not at that time suspected by Professor
Owen, and was recognized by scientific men only when several of
the intermediate stages between it and its modern descendant
had been discovered. Onthe other hand this first ancestor of the
Horse line is very difficult to distinguish from the contemporary
ancestors of tapirs and rhinoceroses, and indicates how all the
12 EVOLUTION OF THE HORSE
modern quadrupeds have diverged from a single type, each be-
coming adapted to the needs of its especial mode of life.
The earliest known ancestors of the Horse were small animals
not larger than the domestic cat, with four complete toes on each
forefoot and three on each hindfoot. There is reason to believe
that the still more ancient ancestors of this and all other mam-
mals had five toes on each foot. In the forefoot of the earliest
known stage we find a splint-bone or small, slender rudiment rep-
resenting the missing first digit or thumb, which no longer ap-
pears on the surface of the foot, while in the hindfoot there is a
similar rudiment representing the outer or fifth digit, but no
trace is left of the innermost or first digit. The proportions of
the skull, the short neck and arched back and the limbs of moder-
ate length, were very little horse-like; recalling, on the contrary,
some modern carnivorous animals, especially the civets (Vzver-
ride). The teeth were short-crowned and covered with low
rounded knobs of enamel, suggesting those of monkeys and of
pigs or other omnivorous animals, but not at all like the long-
crowned complicated grinders of the Horse.
Commencing with the Hyracotherium, twelve stages have been
recognized from as many successive formations, showing the
gradual evolution of the race into its modern form, and each stage
is characteristic of its particular geological horizon. Some of the
stages have been found in several parts of the world, but by far
the most complete and best known series comes from the Tertiary
Badlands of the Western States. Besides the main line of de-
scent which led into the modern horses, asses and zebras, there
were several collateral branches which have left no descendants.
Of some stages all parts of the skeleton have been found; of others
only the jaws, or jaws and feet, are known. We can mention
only the more important stages.
rt and 2.‘ Hyracotherium and Eohippus. LowER EOCENE.
The Hyracotherium is the most primitive stage known, but only
the skull has been found, so that it has not been determined
exactly what the feet were like. The teeth display six rounded
knobs or cusps on the upper molars and four on the lower ones,
™ These numbers refer to the stages in the direct line of descent of the
modern Horse; see frontispiece.
i lags bE CE Oa a) ee a
Oligocene Horses have Three Toes on each Foot.
The Side Toes Touch the Ground
BO-Ger NE
ne ft ve Four Toes in the Fore Foot,
d Three Toes in the Hind Foot.
4 AR
EOHIPPUS PROTOROHIPPUS EPIHIPPUS MESOHIPPUS MESOHIPPUS
LOWER EOCENE MIDDLE EOCENE UPPER EOCENE MIDDLE OLIGOCENE UPPER OLIGOCENE
UNDISCOVERED ANCESTOR
CRETACEOUS
EARLY STAGES IN THE EVOLUTION OF THE FEET
From the series on exhibition in the American Museum 2
by i Bre , - a
EVOLUTION OF THE HORSE 15
and these are just beginning to show signs of fusing into cross-
crests. The premolar teeth have only one main cusp, except the
third and fourth premolars (next the molars) in each jaw, which
have two and three, respectively. The only specimens which
have been found were in the London Clay or Lower Eocene of
England and are preserved in the British Museum.
The Hohtppus is much better known. It comes from the Lower
Eocene of Wyoming and New Mexico, and is very like the Hy-
FIG. 1.—UPPER AND LOWER TEETH OF “ EOHIPPUS,’? FROM
THE LOWER EOCENE OF WYOMING. NATURAL SIZE
racothertum except that the molar teeth have the cusps more
clearly fusing into cross-crests, and the last premolar is begin-
ning to look like one of the true molars. The forefoot of this
animal has four complete toes and the splint of a fifth. The
hindfoot has three complete toes and the splint of another. A
specimen of the hindfoot is shown in the series in the A-case and
many incomplete specimens, skulls, jaws etc., of several species
in the counter-case.
3 and 4. Protorohippus and Orohippus. MippLeE EOcENE.
In these animals the splint of the first digit in the forefoot and
the splint of the fifth digit of the hindfoot have disappeared, but
there are still four complete toes in the fore- and three in the hind-
foot. The crests on the molars are a little clearer and the last
premolar has become almost like the molars, while the next to
the last premolar is beginning to become so. A skeleton of
Protorohippus is mounted in the wall-case. It shows an animal
of the size of a small dog, and proportioned much like the breed
known as the whippet, of which a skeleton has been placed near by
for comparison with the Protorohtppus skeleton. The Protoro-
hippus was found by Dr. J. L. Wortman in 1880 in the Wind
16 EVOLUTION OF THE HORSE
River Badlands of Wyoming, and was described by Professor
Cope and others under the name of the “ Four-Toed Horse.”
Of Orohtppus we have only parts of jaws and teeth. A
specimen of the forefoot is exhibited in the Museum of Yale
University.
5. Epihippus. Upper Eocene. Of this stage of the evolu-
tion of the Horse only incomplete specimens have been found.
The molar teeth have the once round cusps almost completely
converted into crescents and crests, while another tooth of the
premolar series has become like the molars. The toes are still four
in the forefoot and three in the hindfoot, but the central toe in
each foot is becoming much larger than the side toes, a feature
which may be seen in the hindfoot shown in the series in the case.
(This species happens to be somewhat smaller than those found
in the Middle Eocene stage, but no doubt there were others of
larger size living at the same time.)
Paleothertum and Paloplotherium of the Upper Eocene of
Europe form a side branch of the Horse line. They were very
abundant in Europe, but have not been found in the New World.
On each foot they had three toes of nearly equal size, and the
teeth show a rather peculiar pattern. One of these animals was
thought by Professor Huxley to be a direct ancestor of the Horse,
but.1t now is considered to be merely a collateral relative. Some
species of Paleotheritum were of large size, equaltoatapir. They
were first described in the year 1804 by the celebrated Baron
Cuvier from remains found in the gypsum quarries of Mont-
martre, Paris. A large series of skulls, jaws, foot-bones etc.,
from the Upper Eocene of France, is exhibited in one of the
counter-cases.
6 and 7. Mesohippus. OLiGocENe (White River Formation).
In this stage there are three toes on each foot, a splint represent-
ing the fifth digit of the forefoot of the Eocene ancestors. The
middle toe is now much larger than the side toes, which bear very
little of the weight of the animal. Three of the premolars have
now become entirely like the molar teeth, the crests on the
crown are completely formed, and the outside crest in the upper
molars has taken the shape of two crescents. In the Middle Oli-
gocene is found Mesohippus bairdi about the size of a coyote,
Pipes sO C ENE
Pleistocene Horses have One Toe
es.others have
Pelt OR Ein &
Some Pliocene Horses |
One
MIOCENE
Miocene Horses have Three Toes on each Foot.
The Side Toes do not touch the Ground
HIPPARION
LOWER PLIOCENE PLEISTOCENE
CHIPPUS P
MIDDLE MIOCENE MIDDLE MIOCENE MIDDLE MIOCENE
LATER STAGES IN THE EVOLUTION OF THE FEET
From the series on exhibition in the American Museum
‘MODERN
Modern Horses have T each Ff
oP , 4
ee on poise
Le oat i
5
EVOLUTION OF THE HORSE 19
while in the Upper Oligocene occurs Mesohippus intermedius as
large asa sheep. Of both these animals all parts of the skeleton
are known, and a good series of skulls, feet, jaws, palates etc. is
FIG. 2.—UPPER AND LOWER TEETH OF “ MESOHIPPUS BAIRDI,”
FROM THE MIDDLE OLIGOCENE OF SOUTH DAKOTA. NATURAL SIZE
exhibited in the counter-case, besides the specimens shown in the
series of feet and in the series of skulls.
8. Anchitherium. Lower Miocene. This stage has been
found both in Europe and in America. It is much like its prede-
cessor, but is larger and has the crests of the teeth somewhat
higher and more complete. It probably is not in the direct line of
descent of the horses, but is on a side branch. A palate, jaws,
teeth and foot-bones are exhibited here.
g. Parahippus and Hypohippus. MrippLEe Miocene. In Par-
aluppus the tooth-crests are much higher, and the transverse
ridges on the upper molars are beginning to change shape so as to
become a second pair of ‘crescents inside the outer pair. Hypo-
hippus is off the direct line of descent; its teeth are like those of
Anchitherium, by which name it has been generally called, but
the animal was much larger, equalling a Shetland pony in size. A
complete skeleton of the Hypohippus is shown in wall-case 15,
and illustrates very well the general characters of the Three-Toed
Horses, although it is not in the direct line. This specimen was
found near Pawnee Buttes, Colorado, in rg01 by Barnum Brown,
of the Whitney expedition. Other incomplete specimens of
Hypohippus, Parahippus and Merychippus are shown in the
counter-case, and casts of the feet and skull in the evolution se-
ries in A-Case 49. It may be observed that in the forefoot of
EVOLUTION OF THE HORSE
20
Hypohippus small rudiments still remain representing the first
and fifth digits, but there is no splint of the fifth, asin Wesohtppus.
The second and fourth digits still touch the ground, though lightly.
mesostyle parastyle
metastyle : rie
a
metacone-~
metaconule-.
hypostylle-- --protoconule
hypocone--
B
postfossette-— --prefossette
D
1 a
i Mee
vA s
metaloph
protoloph
FIG. 3.—UPPER MOLAR OF MODERN HORSE, SHOWING EARLY STAGES
OF WEAR OF THE TOOTH. CROWN VIEW. NATURAL S'ZE
The feet of Parahippus were much like those of Hypohippus, but
the side toes were smaller.
ro and 11. Protohippus and Pliohippus. MippLe and
THREE-TOED HORSE NYPOHIPPUS,”” FROM THE MIDDLE MIOCENE OF COLORADO
le toes
Rear view of skeleton, showing small si
ae
ks, ;
i
al
EVOLUTION OF THE HORSE 29
Upper Miocene. In this stage the crowns of the upper molars
have become much longer, the two pairs of crescents on the upper
molars are complete, with two half-separated cusps within the
inner pair. And the valleys between the crests have become
filled with cement, so that with the wear of the teeth the edges of
hard enamel are backed inside by dentine and outside by cement.
In this way the surface of the tooth has a series of enamel ridges
always projecting a little above the grinding surface, because the
softer material on each side wears down into hollows, yet never
breaking off, because they are braced so thoroughly on each side.
This is a very efficient instrument for grinding hard grasses. In
Protohtppus and Pliohippus, especially in the former, the crowns
of the teeth are by no means as long as in the modern horses;
they must therefore wear more slowly or wear out at an earlier
age.
The feet in these two genera have but one toe touching the
ground. The side toes (second and fourth digits) are complete,
but much more slender than in the earlier stages and are ap-
parently useless, as they cannot reach the ground. In some
species of Pliohippus they have almost disappeared. The fore-
foot of Protohippus still retains tiny nodules of bone at the back
of the ‘‘wrist’’ (sometimes improperly called in the Horse the
‘‘knee-joint’’), which are the remains of the first and fifth digits.
Hipparion. Purocene. This genus, probably also a side
branch of the genealogical tree of the horse family, is much like
Protohippus, but larger and with more complication about the
tooth pattern. It is common in the European Pliocene beds and
has been found in America also. The feet are still three-toed,
the side toes as large as those of the older Protohtppus.
12. Equus. PLEISTOCENE and Recent. In this stage,
that of the modern Horse, the side toes have entirely disappeared
and are represented by splints on the fore- and hind-foot. No
trace remains on the forefoot of the little nodules which in Proto-
hippus represented the first and fifth digits. The crowns of the
teeth are much longer than in the last stage, and of the two half-
separated inner columns on the upper molars, one has disap-
peared, the other has increased in size and changed in form.
The skull has lengthened and the animal is much larger.
24 EVOLUTION OF THE HORSE
Hippidium. PLEISTOCENE. SouTtH AmeERIcA. The feet are
like those of Equus, except that they were short and stout. The
teeth are like those of Pliohtppus, from which it is supposed to be
descended. The skull is large and long with very long slender
nasal bones. Casts of the skull and limbs presented by the
Museo Nacional of Buenos Ayres, Argentine Republic, are ex-
hibited here.
MEANING OF THE CHANGE IN FEET AND TEETH.
Along with the disappearance of the side toes in the evolution
of the Horse there is a considerable increase in the proportionate
length of the limbs, and especially of the lower part of the leg and
foot. The surfaces of the joints, at first more or less of the ball-
and-socket kind, which allows free motion of the limb in all direc-
tions, become keeled and grooved like a pulley-wheel, permitting
free motion forward and backward, but limiting the motion in
all other directions and increasing considerably the strength of
the joint. By this means the foot is made more efficient for
locomotion over a smooth regular surface, but less so for travel-
ing over very rough ground, and it becomes of little use for
striking or grasping or the varied purposes for which the feet of
polydactyl animals are used.
The increased length in the lower leg and foot increases the
length of the stride without decreasing its quickness. The
heavy muscles of the leg are chiefly in the upper part, and to in-
crease the length of the lower part changes the centre of gravity
of the limb very little. Consequently the leg swings to and fro
from the socket nearly as fast as before, since in an ordinary
step the action of the leg is like that of a pendulum and the
speed of the swing is regulated by the distance of the centre of
gravity from the point of attachment, as that of a pendulum is
by the height of the bob. To increase the length of lower leg
and foot therefore gives the animal greater speed; but it puts
an increased strain on the ankles and toe-joints, and these must
be strengthened correspondingly by converting them from ball-
and-socket joints to “‘ginglymoid” or pulley joints. Additional
strength, likewise at the expense of flexibility, is obtained by
the consolidation of the two bones of the fore-arm (ulna and
FIG. 4.—PALATE AND UPPER TEETH OF ‘‘ EQUUS INTERMEDIUS,” FROM
THE LOWER PLEISTOCENE OF TEXAS. ONE-THIRD NATURAL SIZE
EVOLUTION OF THE HORSE 27
radius) and of the leg (tibia and fibula) into one, the shaft of the
smaller bone practically disappearing, while its ends become
fused solidly to its larger neighbor.
The increase in length of limb renders it necessary for the
grazing animal that the head and neck should increase in length
in order to enable the mouth to reach the ground. An example
of these changes is the modern Horse, in which we find the neck
and head much elongated when compared with the little H yra-
cothertum and this elongation has taken place pari passu with the
elongation of the legs. The reduction and disappearance of the
side toes and the concentration of the step on the single central
toe serve likewise to increase the speed over smooth ground. The
soft yielding surface of the polydactyl foot is able to accommo-
date itself to a rough irregular surface, but on smooth ground
the yielding step entails a certain loss of speed. A somewhat
similar case is seen in the pneumatic tire of a bicycle; a “‘soft”’
tire accommodates itself to a rough road and makes easier riding,
but a “hard” tire is faster, especially on a smooth road. Simi-
larly, the hard, firm step from the single toe allows of more speed
over a smooth surface, although it compels the animal to pick its
way slowly and with care on rough, irregular ground.
The change in the character of the teeth from “‘ brachydont”’
or short-crowned to “‘hypsodont” or long-crowned enables the
animal to subsist on the hard, comparatively innutritious
grasses of the dry plains, which require much more thorough
mastication before they can be of any use as food than do the
softer green foods of the swamps and forests.
All these changes in the evolution of the Horse are adapta-
tions to a life in a region of the level, smooth and open grassy
plains which are now its natural habitat. At first the race was
better fitted for a forest life, but it has become more and more
completely adapted to live and compete with its enemies or
rivals under the conditions which prevail in the high dry plains
of the interior of the great continents. The great increase in
size, which has occurred in almost all races of animals whose evo-
lution we can trace, is dependent on abundance of food. A
large animal, as may be shown on ordinary principles of me-
chanics, requires more food in pri yportion to its size than does a
28 EVOLUTION OF THE HORSE
small one, in order to keep up a proper amount of activity. On
the other hand a large animal is better able than a small one to
defend itself against its enemies and rivals. Consequently, as
long as food is abundant, the larger animals have the advan-
tage over their smaller brethren, and by the laws of natural se-
lection the race tends to become continually larger until a limit
is reached, when sufficient food becomes difficult to obtain, the
RESTORATION OF THE FOUR-TOED HORSE
Oldest known Ancestor of the Modern Horse; only 16 inches high
Photo from original watercolor by C. R. Knight, based on mounted skeleton in
American Museum
animal being compelled to devote nearly all its time to getting
enough to eat.
CAUSE OF THE EVOLUTION.
The evolution of the Horse, adapting it to live on the dry
plains, probably went hand in hand with the evolution of the
plains themselves. At the commencement of the Age of Mam-
EVOLUTION OF THE HORSE 20
mals the western part of the North American continent was by
no means as high above sea-level as now. Great parts of it had
but recently emerged and the Gulf of Mexico still stretched far
up the valley of the Mississippi. The climate at that time was
probably very moist, warm and tropical, as is shown by the
tropical forest trees, found fossil even as far as Greenland. Such
a climate, with the low elevation of the land, would favor the
growth of dense forests all over the country, and to such condi-
tions of life the animals of the beginning of the mammalian
period must have been adapted. During the Tertiary the con-
tinent was steadily rising above the ocean-level, and at the same
time other influences were at work to make the climate continu-
ally colder and drier. The coming on of a cold, dry climate re-
stricted and thinned the forests and caused the appearance and
extension of open, grassy plains. The ancient forest inhabitants
were forced either to retreat and disappear with the forests, or to
adapt themselves to the new conditions of life. The ancestors of
the Horse, following the latter course, changed with the changing
conditions, and the race became finally as we see it to-day, one of
the most highly specialized of animals in its adaptation to its
peculiar environment. At the end of the Age of Mammals the
continents stood at a higher elevation than at present, and there
was a broad land connection between Asia and North America, as
well as those now existing. At this time the Horse became
cosmopolitan, and inhabited the plains of all the great continents,
excepting Australia.
It is a question whether the direct ancestry of the modern
Horse is to be searched for in Western America or in the little
known interior plains of Eastern Asia. It is also unknown why
the various species which inhabited North and South America
and Europe during the early part of the Age of Man should have
become extinct, while those of Asia (Horse and Wild Ass) and of
Africa (Wild Ass and Zebra) still survive. Man, since his appear-
ance, has played an important part in the extermination of the
larger animals; but there is nothing to show how far he is re-
sponsible for the disappearance of the native American species
of horse.
30 EVOLUTION OF THE HORSE
PARALLEL EVOLUTION IN OTHER RACES.
It is interesting to observe that while the evolution of the
Horse was progressing during the Tertiary period in North
America another group of hoofed animals, the Litopterna, now
extinct, in South America evolved a race adapted to the broad
plains of Argentina and Patagonia and singularly like the Horse
in many ways (see exhibit in A-case in centre of hall). These
animals likewise lost the lateral toes one after another, and con-
centrated the step on the central toe; they also changed the form
of the joint-surfaces from ball-and-socket to pulley-wheel joints;
they also lengthened the limbs and the neck; and they also
lengthened the teeth, and complicated their pattern. Unlike the
true Horse, they did not form cement on the tooth, so that it was
by no means so efficient a grinder. This group of animals native
to South America became totally extinct, and were succeeded by
the horses, immigrants from North America, which in their turn
became extinct before the appearance of civilized man.
Many of the contemporaries of the Horse in the northern
hemisphere were likewise lengthening the limbs, lightening and
strengthening the feet, elongating the tooth-crowns to adapt
themselves to the changing conditions around them, but none
paralleled the Horse Evolution quite so closely as did the pseudo-
horses of South America. But the camels in America, the deer,
antelope, sheep and cattle in the Old World progressed on much
the same lines of evolution, although their adaptation was not to
just the same conditions of life.
AMERICAN MUSEUM OF NATURAL HISTORY
The Collection of Fossil
Vertebrates
BY
W. D. Matthew, Ph.D.
Associate Curator of Vertebrate Palaeontology
SUPPLEMENT TO AMERICAN MUSEUM JOURNAL
VOL. III, No. 5, OCTOBER, 1903
Guide Leaflet No. 12
American Museum of Natural History
Officers
President
Morris Kk. JEsSupP
First Vice-President Second Vice-President
J. Prerpont MorGAN Henry F. OsBorn
Treasurer Director
CHARLES LANIER Hermon C. Bumpus
Secretary and Assistant Treasurer
Joun H. WINSER
Scientific Staff
Director
Hermon C. BumMpus
Department of Public Instruction
Prof. ALBERT S. BICKMORE, Curator
Department of Geology and Invertebrate Paleontology
Prof. R. P. WHITFIELD, Curator E
EpmunNpD Oris Hovey, Ph.D., Associate Curator
Department of Mammalogy and Ornithology
Prof. J. A. ALLEN, Curator
FRANK M. CHapman, Associate Curator
Department of Vertebrate Paleontology
Prof. HENry FAIRFIELD OsBorNn, Curator
W. D. Matruew, Ph.D., Associate Curator
O. P. Hay, Ph.D., Assistant Curator of Fishes and Chelonia
Department of Entomology
WILLiaAM BEUTENMULLER, Curator
Departments of Mineralogy and Conchology
L. P. Gratracap, A. M., Curator
GeorGE F. Kunz, Honorary Curator of Gems
Department of Invertebrate Zoélogy
Prof. Witt1amM Morton WHEELER, Curator
Georce H. SHERWoop, A.M., Assistant Curator
Prof. J. E. DuERpEN, Honorary Curator of Coelenterates
Department of Archeology
Prof. FREDERIC W. Putnam, Advisory Curator
Prof. MARsHALL H. SavitiLe, Curator of Mexican and Central
American Archeology
Haran I. Smiru, Assistant Curator of Archeology
GeEorGE H. Pepper, Assistant in Archeology of the Southwest
Department of Ethnology
Prof. Franz Boas, Curator
Prof. Livincston FarRAND, Assistant Curator
CLARK WissLeR, Ph.D., Assistant
Department of Physiology
Prof. Ratpu W. Tower, Curator
Department of Books and Publications
Prof. RatpH W. Tower, Curator
Department of Maps and Charts
A. Woopwarp, Ph.D., Curator
The Collection of Fossil
Vertebrates
A Guide Leaflet to the Exhibition Halls
of
Vertebrate Paleontology
in the
American Museum of Natural History
By
W. De MATE HE We 2h:
Associate Curator, Department of Vertebrate Paleontology
PUBLISHED BY THE MUSEUM
AS SUPPLEMENT TO THE AMERICAN MUSEUM JOURNAL
VoL. III, No. 5, OCTOBER, 1903
Guid Leaf t No. 12
PREEA TORY NOt
THE COLLECTION of fossil vertebrates belonging to the American
Museum of Natural History comprises the extensive material col-
lected by the late Professor E. D. Cope chiefly between 1870 and 1890
and the much larger collections made by the expeditions which have
been sent out by the Museum every year, beginning with 1891. Most
of the Museum expeditions have worked in the western States.
From the beginning of the department in 1891 the collection
and exhibition of these fossils have been under the direction of Pro-
fessor Henry Fairfield Osborn, the curator. From 1891 to 1808, in-
clusive, the exploring parties in the field were under the immediate
supervision of Dr. J. L. Wortman. Since that time Messrs. Matthew,
Granger, Brown and Gidley have been in charge of the field work.
The funds necessary for sending out the expeditions and for the
purchase of the Cope Collection have been furnished chiefly by Presi-
dent Jesup and Messrs. Osborn, Whitney and Constable. The ex-
hibit illustrating the evolution of the Horse is mostly the gift of Mr.
William C. Whitney.
EDITOR.
SIVNWWYW TISSOd JO T1IVH b
ae
THE COLLECTION OF FOSSIL VERTEBRATES.
By W. D. MattuEew, Ph.D.,
Associate Curator, Department of Vertebrate Paleontology.
TABLE OF CONTENTS.
ETENENPU COMMON Ig ales ond Seer ais it 4s0 as alas » e eeeorsd Sens ole eee eee er 5
What Fossils Are. The Divisions of Geological Time. How Fossil Skele-
tons are Mounted and Exhibited in this Hall. General Arrange-
ment of the Collections.
HAG CORRIDOR== INOw4o5. Mossel Marne Repiless. 25 ee ere 10
How they come to be Buried, Fossilized, Found and Collected. Ple-
siosaurs. Mosasaurs. Ichthyosaurs. Fishes.
BASTING NOm4o0: Mossi WMammals..+-.....--.55-240seeeee eee 12
Arrangement. Titanotheres. Rhinoceroses. Horses. Primitive
Hoofed Mammals. Primitive Clawed Mammals. Elephants,
Mammoths and Mastodons. Artiodactyls. South American
Fossil Mammals. Instances of Evolution. Restorations. Trans-
parencies. Charts, etc.
RMNURTINIE STING): AOY cc oc cob ee esi & ebro cto atee as we aoe aie Stage Ole 26
Fossil Reptiles.
Dinosaurs: Amphibious, Carnivorous, Beaked. Crocodiles. Lizards.
Turtles. Primitive Reptiles.
Fossil Amphibians. Stegocephalia..........--.- cece see eee cece eee 31
Fossil Fishes. Dinichthys. Green River Fishes............-...+++++.- 32
INTRODUCTION.
WHEN we dig beneath the present surface of the ground we
sometimes find remains of ancient cities, dwellings, bones of men
and animals, buried many centuries ago under accumulations of
débris, deposits of river mud or drifted sand. From these have
been gleaned many facts concerning the early history of man-
kind of which there is no written chronicle. From the study of
these facts the science of Archeology has arisen, the science
which deals with the early history of mankind, with the evolu-
tion of civilization.
Most of the lower animals of which the archeologist finds
traces are like those now living, although a few have what Fos-
become extinct. But in those more ancient deposits _ sils Are.
which are now consolidated into clays, sandstones etc., indica-
5
6 THE COLLECTION OF FOSSIL VERTEBRATES
tions of man are not found, and the remains of lower ani-
mals which they contain are unlike any now living—the more
unlike as the rock is more ancient. These remains are called
fossils. They consist only of the hard parts of animals (bones,
shells, spines etc.). The soft parts are never preserved, and
only very rarely is some trace of skin or hair, horns or hoofs, to
be distinguished. As in the course of ages the mud or sand in
which they are buried changes to rock, so little by little the
fossils have been changed by heat, pressure and especially by
the slow infiltration of mineralized waters into brittle, stony
material, while retaining their outward form and usually their
peculiar structure. But mud and clay, in changing into rock,
settle down and contract considerably, and the fossils are flat-
tened out correspondingly, sometimes to such a degree, in the
case of a rock which has once been a soft, oozy mud, that they
suggest rather a picture or a bas-relief than the original form
of the animal. The fossil skeletons of marine reptiles and fishes
on the walls of the corridor hall and in the case opposite the
elevator have been flattened out in this manner, especially the
Ichthyosaur skeletons.
From fossils we can interpret the history of the world of life
during the long ages before man appeared. The science which
Science of deals with the ancient history and evolution of the
Paleon- animal kingdom is Paleontology (zaldaios, ancient,
tology. ovra, living beings, -Aoyéa, science). It tells us of a
long period of time before Man appeared, probably millions of
years, during which Mammals of great size and unfamiliar form
were the dominant animals—of a yet longer era before that,
during which huge Reptiles were rulers of earth, sea and air—
and of other still more ancient periods during which Amphibians,
Fish and Invertebrate Animals held sway in turn. Vertebrate
Paleontology deals only with the higher classes of fossil animals,
the Vertebrata, or those that have backbones (fish, amphibians,
reptiles, birds and mammals).
Earth-history or geological time has been divided into many
Geological Parts according to the evidence furnished by the rocks
Time. and the fossils contained therein. The principal sub-
divisions are shown in the accompanying table:
FOSSIL SKELETONS IN THE ROCK
This slab of soft chalky clay contains five skeletons of an extinct animal. One is an old male, the other
four are young
8 THE COLLECTION OF FOSSIL VERTEBRATES
GEOLOGICAL ERAS, PERIODS AND AGES.
Eras. Periods. Ages.
Age of Man
Quaternary ‘= :
50,000 years
Cenozoic
Aon ye Age of Mammals,
Tertiary ¥
2 3,000,000 years
Cretaceous
: . Age of Reptiles
Mesozoic Jurassic ge of Reptiles,
7,000,000 years
Triassic
Permian Age of Amphibians and Coal
Plants,
Carboniferous 5,000,000 years
Age of Fishes
Palzeozoic Devonian a
2,000,000 years
Silurian
Age of Invertebrates,
zs : 10,000,000 years
Cambrian
Algonkian
Eozoic (No fossils)
Archeean
The estimates in years of the geological periods given in this
table, which is arranged in descending order from the most recent
to the most ancient time, must be understood to be merely very
rough approximations. There is no known method of finding any
exact equivalent in years of any geological period, although the
relative length of each to each is much more nearly known. The
estimates given herewith are based on the careful study of the
subject made by C. D. Walcott, Director of the U. S. Geological
Survey. In concluding his discussion Dr. Walcott stated his
belief that the duration of geological time (the entire period
included in this table) might be measured by tens of millions of
years, but not by single millions or by hundreds of millions.
THE COLLECTION OF FOSSIL VERTEBRATES 9
To give the visitor a clear idea of these extinct animals, the
skeletons usually have been removed entirely from the rock in
which they were found and have been mounted as How Foaat
much as possible like skeletons of modern animals. skeletons
To mount a petrified skeleton in this manner is a very are
difficult matter, for such skeletons are rarely perfect, Mounice,
and the bone is always very brittle and more or less shattered
and crushed out of shape. In the mounted skeleton the missing
parts have been restored in tinted plaster, modeled from other
individuals or from nearly related animals in which these parts
are known. The outlines of the restored parts of bones are
marked off with red lines, while entire bones modeled in plaster
are marked with a red cross, or with a red circle if supplied from
other individuals. All the skeletons are original specimens ex-
cept the Megatherium at the far end of the hall; and all are of
extinct animals except a few which are placed with the others for
comparison. With each fossil skeleton will be found, besides a
descriptive label, a small model and a water-color restoration of
the animal, showing its probable appearance during life and
indicating its supposed habitat. The transparencies in the win-
dows show the localities where the fossils are found, chiefly in the
Bad-lands of the western States.
GENERAL ARRANGEMENT.
FOSSIL REPTILES
FOSSIL MAMMALS ano
FISHES
MARINE REPTILES] —
eee eee:
The collections are arranged to illustrate the geological his-
tory and evolution of the different groups of \ erte- Gen unten
brata, especially those of North America. They fill rangement
; of Collec-
two large halls and a corridor.
tions.
East Wing. Hall No. 406. Fossil Mammals.
10 THE COLLECTION OF FOSSIL VERTEBRATES
East Wing. Hall No. 407. Fossil Reptiles, Amphibians and
Fishes.
In addition
East Corridor, No. 405 (in which are the elevator and stair-
ways), contains fossil Marine Reptiles and Fishes of the Age of
Reptiles.
SKELETON OF THE GREAT MARINE LIZARD IN THE EAST CORRIDOR
THE EAST CORRIDOR. No. 405.
On stepping from the elevator the visitor sees before him a
case filled with skulls and skeletons of the marine reptiles and
The Preser- {shes which inhabited the great inland sea that once
vation of Spread over the center of the North American conti-
Fossilsin nent, from Canada to Mexico, The reptiles were of
the Rocks. kinds now long extinct, Plesiosaurs with long snak
g S yi
neck, short bulky body with long flippers and stubby tail, and
Mosasaurs with short neck and longer tail. Some of the fishes
were ancestors, collateral or direct, of certain modern fishes,
others belonged to groups now extinct. These animals lived and
died, their carcasses sank to the bottom of the sea, and were
buried in whatever sediment was being deposited there—soft
white ooze in the open sea, dark gray or black mud nearer the
shores. In the course of ages this ooze or mud settled gradually
and consolidated into chalk or shale. Afterwards as the conti-
nent rose above the waters and assumed more nearly its present
dimensions, the rivers flowing over the broad plains excavated
HUNTING FOR FOSSILS IN THE BAD-LANDS
Members of the American Museum Expedition of 1894 in the Uinta Basin, Utah
12 THE COLLECTION OF FOSSIL VERTEBRATES
broad shallow valleys in the chalk and shale. In the dry climate
The Find. ©! the present day the sides of these valleys often are
ing and bare rock, carved by wind and the infrequent storm-
Collecting bursts of rain into the fantastic maze of cliffs and
of Fossils. yinding cafions known as ‘“‘bad-lands.’’ Here and
there, projecting from an outstanding ledge or trailing in frag-
ments down some crumbling slope, a fossil bone may be seen by
the trained eye of the collector as he searches along the rock
exposures; and quarrying in around the bone he is sometimes
rewarded by a skull, sometimes by a string of vertebra, occa-
sionally by a whole skeleton, buried in the rock except for such
parts of it as have been weathered out and washed away.
To excavate the fossil without damaging the brittle bones,
buried as they are in a weak and shattered mass of heavy shale
or chalk, is a slow and delicate operation, requiring special
methods and considerable care and skill. Then the specimen
must be packed, and sent in to the Museum, where the rock is
removed and the specimen 1s prepared for exhibition. When the
bones are as much crushed and distorted as those represented 1n
the photograph (page ro) the matrix is removed from one side
only, and the specimen is thus placed on exhibition.
Temporarily placed in the bottom of the case is a large Ple-
siosaur skeleton, only partly removed from the rock. This
Important specimen unfortunately lacks the skull. Beside the
Specimens Jower stairway is a Mosasaur skeleton, the finest speci-
in Corridor. Box : wae
men of its kind ever found, and above it is a large fish
skeleton which was found in the same strata in western Kansas.
Beside the upper stairway are three skeletons of Ichthyosaurs,
another long extinct group of marine reptiles, of fish-like appear-
ance, paralleling the modern Whales among mammals.
EAST WING. HALL NO. 406. FOSSIL MAMMALS.
The ancestors of our modern quadrupeds are to be found in
the East Wing, No. 406, together with many extinct races more
Arrangement OT less nearly related to them. All the fossil speci-
of the Fossil mens of each group of mammals are placed together
Mammals. in one alcove, where they have been arranged ac-
cording to their geological age. Thus all the fossil Horses, direct
THE AGE or MAMMALS
ok (CENOZOIC. OR TERTIARY AND QUATERNARY.)
_ WESTERN LAKE BASINS and CHARACTERISTIC MAMMALS’
THE TERTIARY FORMATIONS ARE REPRESENTED IN | IN THE SEDIME
WESTERN AMERICA BY A SERIES OF DEPOSITS | WERE SuRIER Ee MAINE Dr AES
BASINS [RNA CHARACTERISTIC. MAMMALS
ELEPHANTS. LAST MASTODONS
LAST GROUND SLOTHS. LAST SABRE-TOOTH TIGERS
CAMELS: ONE-TOED HORSES. CAVE BEARS
__*_ |PECTARIES, TAPIRS. COGS wouvEs KFotls Rovtats
150 |GROUND SLOTHS CAMELS ONE-TOEO HORSES.
FIRST HYAENAS. FIRST TRUE Cars PECCARES. PeOtuTs.
| MASTODONS, TRUE HORNLESS RHINOCEROSES
400 cast oneoponts CAMELS, THREE TOED HORSES
DEER. FIRST PRONG HORN ANTELOPES
TRUE HORNLESS RHINOCEROSES. OREOOOMTS. CAMELS:
FIRST MASTODONS, FIRST TRUE OLER
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FIRST SELENODONT ARTIODACTYLS (MOMACODON)]
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FIRST FOUR -TOEO HORSES (MYRACOTHERIUM)
FIRST TAPIRS (S¥STEMODON)
FIRST ARTIODACTYLS e* CLOVEN HOOFED ANIMALS.
PRIMATES (MONKEYS on LEMURS)
CREODONTS oe PRIMITIVE CARNIVORES,
(RESEMBLING CATS. DOGS«s0 BEARS)
FIRST RODENTS
TILLODONTS, INSECTIVORES
PRIMITIVE GROUND SLOTHS
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“WIND RIVER
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NEW MEXICO
LARAMIE
AGE OF KEPTULS
DIVISIONS OF THE AGE OF MAMMALS
Characteristic fossil mammals, and the geological formations in which they are found
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THE COLLECTION OF FOSSIL VERTEBRATES 5
or collateral ancestors of the modern Horses, Asses and Zebras,
are in one alcove, arranged in series from the most ancient to
the most recent. The most ancient and structurally primitive
groups of mammals come first, the most modern and familiar
types come last.
SoutH (Ricut) Sipe. Nortu (Lert) Sipe.
( TITANOTHERES AMBLYPODS i Primitive Hoofed
Chalicotheres CONDYLARTHS Mammals
Monkeys, Bats, Rodents, Insectivores
and other Small Primitive Mammals
Marsupials
CREODONTS t Carnivorous
CARNIVORES ) Mammals
Seals, Dolphins, Whales, ? Marine
Pee aan Sirenians etc. § Mammals
erissodactyls -
or Odd-Toed J 1) hindonts ELEPHANTS} Proboscideans
Hooted RHINOCEROSES ELOTHERES }. .
Mammals ‘Anthiacethorss Artiodactyls
Pigs and Peccaries ( a
OREODONTS * Even-Toed
CAMELS Hoofed
Palzotheres DEER ete. Mammals
HORSES Litopterna ] Fossil Mammals
Toxodontia Peculiar to South
Typotheria [deste
EDENTATA
SOUTH (RIGHT) SIDE.
The south side of the hall is entirely devoted to the PErRts-
SODACTYLS or Odd-Toed Hoofed Mammals in which the number
of toes (in the hind foot and generally in the forefoot) is either
I, 3 or 5, while in the other main division of hoofed animals, the
Artiodactyls, it is either 2 or 4; or more exactly, the axis of
symmetry of the foot passes through the central toe in Peris-
sodactyls, while in Artiodactyls it passes between two toes.
The TITANOTHERES come first in the series of Perissodactyla,
large animals which suggest rhinoceroses in general gosh ay
proportions, but have a differently shaped head and cove I.
peculiar teeth. These began as hornless animals of re
moderate size (Cases 1 and 17) and increased in size
and developed large bony horns (Cases 3, 5 and 19) before they
16 THE COLLECTION OF FOSSIL VERTEBRATES
became extinct. The Titanotheres occupy the first of the
three main alcoves into which the south side of the hall is
divided.
The second alcove is devoted to the RHINOCEROSES, which
RESTORATION OF TITANOTHERIUM, AN EXTINCT HOOFED MAMMAL OF WESTERN AMERICA
The picture shows a bull, a cow and a calf
From the original water-color, based on mounted skeleton and skulls in American Museum
of Natural History
were very common beasts in North America as well as in the Old
ann World during the Tertiary period. They also began in
Alcove 2, the Eocene as small hornless animals (Hyrachyus, Case
Rhinoce- 7), but diverged in the Oligocene into cursorial, aquatic
i and true (terrestrial) Rhinoceroses of which the two
former soon became extinct. True Rhinoceroses also became ex-
tinct in America by the Pliocene epoch, while in the Old World
several of them have survived to the present day.
Third Alcove. Horszs.‘. This fine exhibit is due chiefly to
™ For more detailed information regarding the evolution of the Horse, see
Guide Leaflet No. 7, ‘“The Evolution of the Horse.’’ Published January, 1903.
MOUNTED SKELETON OF THE TITANOTHERE, FROM THE BIG BAD-LANDS OF SOUTH DAKOTA
18 THE COLLECTION OF FOSSIL VERTEBRATES
the liberality of Mr. Wm. C. Whitney. The Evolution of the
South Horse is illustrated by a series of feet and skulls, and
Alcove 3. of complete skeletons, from the little Four-Toed Horse
Horses. of the Lower Eocene to the different varieties of the
modern animal. The construction of the modern Horse, structure
of the bones, the way in which the teeth grow, characters of the
different races of domestic Horse and of the different wild species
are shown in the end case (Case 15).
NORTH (LEFT) SIDE.
First come the AMBLYPopDS and CONDYLARTHS. These groups
of Primitive Hoofed Mammals are first found in the lowest Eocene
North strata, at the very beginning of the Age of Mammals
Alcove r. and they became extinct before the end of the Eocene
Amblypods, epoch. Like so many other races the Amblypods begin
ete: with small hornless animals (Pantolambda) and finally
develop into huge elephantine beasts (Uzntathertum) with six
horns on the skull, and great sabre-like tusks. The Condylarths
were more slender types, fitted for running. The best known
among them is Phenacodus, which is considered to represent
very nearly the prototype of the hoofed mammals, although it
was not the direct ancestor of the later groups.
The second alcove is devoted to Rodents, Insectivores, Bats,
Marsupials and other groups of small mammals, among which
North are the ancestors of the Monkeys and Lemurs and col-
Alcove 2, lateral ancestors of Man. Most of these remains are
Small small and incomplete. Here are also some very frag-
Mammals,
mentary remains of ancient and primitive mammals
which represent all that we know of the evolution of the mam-
malia during the Age of Reptiles, before the Age of Mammals
began. These teeth and jaws are of interest because they are
the oldest of mammals, from some of which are probably de-
scended all the later mammal groups.
In the third alcove are the Carnivorous Mammals, on one side
North the CREoponTms or Primitive Carnivora, on the other the
Alcove 3. True CARNIVORA (Dogs, Cats, Bears, Martens etc.), rep-
Carnivora. resented by a number of finely preserved mounted skele-
tons, and a large series of skulls, together with other specimens.
MOUNTED SKELETON OF PHENACODUS IN NORTH ALCOVE 2
This animal, although not a direct ancestor, represents the prototype of the hoofed mammals
20 THE COLLECTION OF FOSSIL VERTEBRATES
Most remarkable among extinct carnivora are the Sabre Tooth
Tigers, in which the upper canine teeth are enlarged into long,
curving, flattened, serrate fangs, most terrible weapons, effective
no doubt against the thick hides of the primitive pachyderms.
North The fourth alcove is very narrow. In it are placed
Alcove 4. @ few remains of fossil marine mammals: Seals, Ceta-
Marine ceans and Sirenians. These groups are very imper-
Mammals. fectly known as fossils.
The fossil ELEPHANTS and Mastodons are in the next broad
North alcove, about the middle of the hall. The evolution of
Alcove 5. these animals is shown by a series of skulls. The
Probos- Mastodon skeleton and the skull and fore-limb of the
cideee Imperial Mammoth from Texas, and tusk of the Si-
berian (Hairy) Mammoth are noteworthy specimens.
Beyond the Elephants are the ARTIODACTYLS or Cloven-
Hoofed Mammals. They divide into two groups, typified by the
North Pigs and the Ruminants, the latter including the greater
Alcove 6. part of modern hoofed mammals, but by no means pro-
Elotheres. ortionally common as fossils. First among the fossil
artiodactyls are the Elotheres, an extinct race of large animals
distantly related to Pigs and Hippopotami. Next are the fossil
Peccaries; then the Oreodonts, pig-like animals with the teeth
North of ruminants, very abundant in America during the
Sana middle and later Tertiary, but extinct before the Plio-
ee * cene epoch. Then come the Camels, which although
Oreodonts, Now found only in Asia and South America, originated
Camels. in North America, and afterwards migrated to these
other continents and became extinct in their native land. The
evolution of these animals is shown by a series of stages only
less complete than the stages in the evolution of the Horse.
The higher ruminants (Deer, Antelope, Sheep and Cattle) are
rather poorly represented in the collections. The Great Irish
North Deer is the most striking among the extinct species;
Alcove 8. attention is also called to the mounted skeletons of
Deer, Ante- Protoceras, a deer-like primitive ruminant of the Oligo-
eRe etC Is eene epoch, and of Merycodus, a graceful little animal
of the Miocene epoch intermediate between the Deer and the
Prong-horn Antelope.
SKULL AND TUSKS OF THE IMPERIAL MAMMOTH
yn exhibition in the American Muse
From a photograph of the specimen «
22 THE COLLECTION OF FOSSIL VERTEBRATES
The northeast corner of the hall is devoted to a number of
peculiar groups of SoutH AMERICAN Foss, MAMMALS, almost all
North extinct. During the Age of Mammals the two great
Alcove 9. northern continental areas were joined together from
pou time to time, so that there has been an occasional
American . :
Poccil interchange of animals and plants among them, the
Mammals. races developed in one continent spreading to the
other. The animals of North America therefore, although
mostly of species distinct from those of Europe and Asia, are
more or less nearly related to them. But during most of the Age
of Mammals South America was an island continent, as Aus-
tralia is still; and its extinct animals are as peculiar and as dif-
ferent from those of the rest of the world as are the living animals
of Australia different from those of other continents. It is by
no means certain where these animals originally came from, but
there is much evidence to show that both South America and
Australia were peopled from an Antarctic continent, now sunk
beneath the ocean or buried in the ice fields of the more frigid
climate of modern times.
Of these peculiar South American groups the most extraor-
dinary are the Edentates, including the Sloths, Armadillos and
Anteaters which still survive, and the huge MEGATHERIA or
Ground-Sloths and GLyproponts or Tortoise-Armadillos which
have become extinct. Others were the TOXODONTS, TYPOTHERES,
ASTRAPOTHERES and LITOPTERNA, peculiar groups of hoofed ani-
mals all now extinct. Some of the Litopterna lost their side toes
and evolved into a one-toed race curiously like the horses of the
northern hemisphere, although not at all related to them; this is
one of the most interesting examples of the parallel adaptation of
two different races of animals to similar conditions of life; the
horses in the plains and prairies of the north, the litopterna in
the pampas of the southern continent.
The best example of the evolution of a race of animals is
shown in the southeastern corner of the hall. Here is exhibited
Instances of the Ancestry of the Horse, the specimens from succes-
Evolution. sive geological strata showing how the modern Horse
has descended from diminutive ancestors with four toes on each
forefoot and three on each hind foot, and with teeth and other
= ‘ : AGE OF MAMMALS AGE OF MAN.
ESTIMATED AT S
REPTILES
EE es Se
—
PLEISTOCENE
i Pleistocene Horses have One Toe on
PLIOCENE
i Some Pliocene Horses have Three Toes. others have
rE
One Toe on each Foot
MIOCENE
Miocene Horses have Three Toes on onch Foot
The Side Toes do not touch the G
ae
OCENE
Three Toes
as Tork the Gro
nee ae
Eotene Four Tor «Ff
T r nthe Kind oot
RETA
| }
t
a
WDISCOVERED ANCESTOR * EDHIPPUS PROTOROMIPPUS CHIPPUS
re Taceoun MIDOLE Miocene MIOOLE MWS
EVOLUTION OF THE HORSE. FEET 23
24 THE COLLECTION OF FOSSIL VERTEBRATES
parts of the skeleton different from those of their modern repre-
sentatives.
Almost equally complete, although less familiar, is the series
illustrating the Ancestry of the Camel, which may be found on the
north side of the hall near the east end. These animals, like the
Horses, evolved from small and primitive ancestors to large and
highly specialized descendants, and then became extinct in their
former home, the broad and arid plains of western America, before
the advent of civilized man, but survived to modern times in other
parts of the world (Asia, Africa and South America). Less com-
plete series are the skulls and skeletons illustrating the ancestors
of Titanotheres and the ancestors of Rhinoceroses. These are
ranged along the south side of the hall beginning at the entrance.
All these series have been placed according to geological age.
The most ancient specimens, found in the lowest rock-strata,
and hence representing the earliest stage of evolution, are placed
first in the series. The most recent ones, found in the upper-
most rock-strata, and representing the final stage of evolution of
the race, are placed last. Arranging the species of a race from
each stratum in the order of the age of the strata, we find that
they show a regularly progressive change from the most ancient
to the most recent. At no point in a given series can we draw a
line and say: This is and that is not, a Horse—or a Camel—or a
Rhinoceros. The visitor, therefore, can demonstrate for himself
the evolution of the race of Horses or Camels or Rhinoceroses,
within certain limits. Of the evolution of Man we have no satis-
factory illustration from fossils.
It should be observed that the evolution of a race consists
mainly in the adaptation of the structure of the animals to par-
ticular surroundings and habits of life. There is also a universal
progress in intelligence, the more ancient animals having rela-
tively smaller brains than their successors.
The water-color restorations by Charles R. Knight, done
under the immediate supervision of Professor Osborn, mainly
Water-color based on complete skeletons exhibited in this hall,
Restora- show the probable appearance of the different extinct
Ba: animals, according to our best judgment, as indicated
by the characters of the skeleton, appearance of their nearest
THE COLLECTION OF FOSSIL VERTEBRATES 25
surviving relatives and the habits of life for which the animals
seem to have been fitted. The general proportions of the animal,
the outlines and form of head and body, and, toa great extent, the
expression of the features are usually accurately known from
the fossil skeleton. The nature of the skin is sometimes but not
often certainly known, and the coloring is always conjectural,
7
SCENE iN THE BAD-LANDS OF THE UINTA BASIN—TERTIARY FOSSIL FIELD OF
NORTHEASTERN UTAH
the paleontologist and the artist having been guided by the col-
oring of living relatives and the supposed habits of the animal.
The window transparencies are enlargements from photo-
graphs of the regions where the fossils occur, and generally show
the localities where unusually fine specimens in this window
hall were found. The expeditions sent out yearly to Transpar-
the fossil fields carry with them photographic outfits, ae
and several hundred characteristic views have been taken, from
26 THE COLLECTION OF FOSSIL VERTEBRATES
which these have been selected. The pillar cards and general
labels in the cases give detailed information about each group
of fossils. One of the cases in the center of the middle aisle
illustrates the method by which the fossils are collected and con-
veyed to the Museum. The charts at each side of the entrance
show the order in which the rock-strata lie, one over another,
and the kinds of fossils found in each stratum.
EAST WING. HALL NO. 407. FOSSIL REPTILES, ETC.
This hall forms an introduction to an earlier world, the Age
of Reptiles. These fossils are of strange and unfamiliar out-
lines, quite unlike ordinary quadrupeds; they represent an era,
long since passed away, when reptiles were the “lords of crea-
tion.’’ Chief among them were the Dinosaurs, great land and
amphibious reptiles to which the greater part of this hall is
devoted. They occupy the north, east and west sides and the
center.
The AMPHIBIOUS DINOSAURS, on the west and north sides
and in the center of the hall, were the largest of land animals,
Amphibious Some of them sixty to seventy feet in length, and of
Dinosaurs. enormous bulk. They were quadrupedal beasts, with
long necks and long tails, and comparatively long and very
massive limbs. The head was very small in proportion to
the size of the animal, and the brain inferior to that of modern
reptiles. They were cold-blooded, slow-moving, unintelligent
creatures, vast storehouses of flesh which lived and grew to huge
size with but little occasion for very active exertion amidst the
rich vegetation of the moist and tropical climate of the reptilian
era. Several incomplete skeletons of Amphibious Dinosaurs are
exhibited, besides limbs and other separate parts. The Bronto-
saurus skeletons in Case 1 (on the right-hand or south side of the
entrance) and in the center of the hall are among the largest.
The thigh bone in this animal was nearly six feet long and
weighs in its petrified state 500 to 600 pounds. The Diplodocus
(Case 2 on the left-hand or north side of the entrance) was less
robust but almost as long. This specimen lacks the fore part of
the skeleton and most of the limbs, but the tail is very perfectly
SKELETON OF BRONTOSAURUS
RESTORATION OF THE SKELETON OF AN AMPHIBIOUS DINOSAUR OF THE AGE OF REPTILES. THE SKELETON WAS 65 FEET LONG 27
TAKING UP FOSSIL DINOSAUR BONES AT ‘BONE CABIN’? QUARRY, WYOMING
Parts of several skeletons appear in the photograph
28
THE COLLECTION OF FOSSIL VERTEBRATES 29
preserved. In Case 4 are limbs and other parts of several species
of Amphibious Dinosaurs.
The BEAKED Dinosaurs (Predentata) northeast corner of
the hall, had a horny beak or bill at the front of the jaw, and
teeth at the back of it. They were most extraordinary Beaked
and bizarre animals, huge in size, although not so Dinosaurs,
large as the Amphibious Dinosaurs. Stegosaurus had eee
a series of great bony plates projecting from the back, ana aa
and stout bony spines on the tail. Triceratops had an rosaurus.
enormous skull with three great horns projecting forward, and a
strong bony frill projecting backward around the neck. Both
these were quadrupedal animals with massive limbs and elephan-
tine feet. Hadrosaurus was a bipedal dinosaur with long hind
limbs and three-toed bird-like feet, but with hoofs instead of
claws. Its beak was broad and flattened, as in the spoon-bill
duck or Ornithorhynchus of Australia.
The Carnivorous Dinosaurs are exhibited on the east side
of the hall (opposite the entrance). They were bipeds with bird-
like feet, sharp claws and large heads with sharp- cCarnivor-
pointed teeth. Some of these, the Megalosaurs, were ous
of gigantic size, much larger than any modern car- Dindeae
nivore. Allosaurus was as large as an elephant, while other
Megalosaurs were even larger. Other Carnivorous Dinosaurs,
such as Ornitholestes, were small and of slender proportions;
these probably lived on the small animals of that period—toothed
birds, pterodactyls, small reptiles of various kinds—while the
large herbivorous dinosaurians were more probably the chief
prey of the Megalosaur.
All the Dinosaurs had become extinct by the end of the Age
of Reptiles. Their place was taken by the more intelligent
and adaptable mammals, the evolution of which into the differ-
ent kinds of modern quadrupeds has been seen in the Other Rep-
Hall of Fossil Mammals. ma . Gicanat
On the south side of the Fossil Reptile Hall are Turtles,
fossil remains of four other groups of reptiles, the Pterodac-
CROCODILES, TURTLES, LIZARDS and SNAKES, which, tyls etc.
more fortunate than the dinosaurs, have survived to the present
day, though in much diminished numbers and importance.
AGE OF REPTILES ||
— (MESOZOIC)
L FORMATIONS and CHARACTERISTIC ANIMALS.
AG! oF REPTILES eee THE AGE | DURING THIS AGE THE REPTILES APPEARED,
vm D- REPRESENTED IN | FLOURISHED GREATLY, AND DECLINED AT
ITS CLOSE TO THEIR PRESENT IMPORTANCE.
THE MAMMALS APPEARED WELL DOWN IN
THIS AGE BUT REMAINED SMALL AND
SCARCE NTS iTS END.
ESTUARY AND FR
DIVIDED INTO THREE ‘GREAT PERIODS,
TRIASSIC, JURASSIC AND CRETACEOUS.
bee IN LARGE NUMBERS
| TRUE LIZARDS and SPHENGDONS
ALLIGATORS and CROCODILES
TURTLES
NUMEROUS BONY FISHES (TELEOSTS}
CARNIVOROUS DINOSAURS
JHERBIVOROUS DINOSAURS
HORNED (HERBIVOROUS DINOSAURS
J NUMEROUS SMALL MAMMALS
LAST PLESIOSAURS j
FIRST SOFT-SHELLEO TURTLES
MODERN TAILED AMPHIBIANS (SALAMAKOERS)
“EOCENE.
BIRDS, PROBABLY TOOTHED.
PTERODACTYLS, TOOTHLESS
1200)MOSASAURS and PLESIOSAURS
B700|GIGANTIC MARINE TURTLES
DOLICHOSAURIAN LIZARDS
SHARKS, CAT-FIBH, STURGEONS act Ast PIKES
ooTHED BIRDS TOOTHLESS PTERODACTYLS
JOINOSAURS
1000) MOSASAURS ani PLESIOSAURS
3000] LARGE MARINE TURTLES
BONY FISHES (reteosrs) SHARKS
GANOID FISHES.
FIRST SNAKES
| TURTLES
TRUE LIZARDS and DOLICHOSAURS
HERBIVOROUS DINOSAURS (icuanooonr
: CARNIVOROUS DINOSAURS (mecavosaunsy
rf | PTERODACTYLS, TOOTHED and TOOTHLESS
COMANCHE MOSASAURS
ICHTHYOSAURS and PLESIOSAURS
GROSODILES. TURTLES.
SHARKS and GANOID FISHES
CHIMARO!ID FISHES
(RSUPIALS, INSECTIVORES
MULTITUBERCULATES)
CARNIVOROUS DINOSAURS [CERATOSAURUS,
HERBIVOROUS DINOSAURS (ATLANTOSAURUS)
TURTLES. PTERODACTYLS
| F1RST BIRDS WITH TEETH (SAURURA)
-}ICHTHYOSAURS (TOOTHED and TOOTHLESS)
PLESIOSAURS PTERODACTYLS TOOTHED)
SMALL PRIMITIVE MAMMALS:
LONG NOSED CROCODILES (TELEDSAURS)
ICHTHYDSAURS and PLESIO
Hinst HERBIVERGUS. TINOSAURS.
ieee a CAONTROSTEAN J SHES :
REPTILE- MAMMALS (oF
TRITYLODON, MIGROLESTES)
SF RICHMOND GCOAL-BEDS FIRST CARNIVOROUS DINOSAURS |
and CONN. and NEW JERSEY i
=.
WEALD EN
POTOMAC
*: LM PPER
ca
Cc
=
>
a
2
bid
| WUDDLE
| LOWES *
oS
LPL
-|FIRST YELEOST on BONY FISHES
Xx TRIASSIC 3000 SHARKS, CHONDROSTEAN and LUNG FISHES
6000|pLesiosaurs (NOTHOSAURS)
FIRST ICHTHYOSAURS (MIXOSAURUS
PLACODONTS
LARGE AMPHIBIANS (LABY-
RINTHODONTS)
FIRST PLESIOSAURS (NOTHOSAURS)
LOWS SO MIMOLE ,
FIRST REPTILES (COTYLOSAURS,
.| PROGANOSAURS and PELYCOSAURS,
| PRIMITIVE AMPHIBIANS ‘STEGOCEPHALIA
SHARKS. LUNG FISHES.
‘ PRIMITIVE AMPHIBIANS ‘STEGO-
= se ae CEPHALIA. moony SMALL SPErES)
ARBMEROUS e ae : : PRIMITIVE. SHARKS and LUNG FISHES
HOMDROSTEAM and CROSSOPTERYRIAN FISHES
WD COAL PLANTS
Al
AGE OF AMPHIBIANS
DIVISIONS OF THE AGE OF REPTILES
Characteristic fossil reptiles, amphibians and fishes, and the formations in which they are found
30
THE COLLECTION OF FOSSIL VERTEBRATES 31
Crocodiles in their palmier days were of world-wide distribu-
tion and comprised marine as wellas fresh-water types. Turtles
are among the commonest of fossils in the Bad-lands and some
of them of very large size. Lizards and snakes, the only common
reptiles of modern times, are very rare and fragmentary as
fossils, and little is known about them.
Besides these surviving groups, several extinct groups of rep-
tiles are shown on the south side of the hall. The BeLoponts,
of the dawn of the Reptilian Era, were partly intermediate be-
tween Dinosaurs and Crocodiles. The still older PELycosauRs
were remarkable for an enormous rigid bony fin on the back;
among the contemporary THERIODONTs there existed perhaps the
remote ancestors of the Mammals. The PreRopacty_s or Fly-
ing Reptiles were the most extraordinary of reptiles, tailless, with
batlike wings, supported on the enormously lengthened little fin-
ger, and with a spread in the largest species of twenty feet from
tip to tip. The RHYNCHOCEPHALIANS are an interesting group
of very primitive reptiles, of which a single species, the Tuatara,
still survives in New Zealand.
Fossit AMPHIBIANS.
The Age of Reptiles was preceded by an Age of Amphibians,
when the dominant animals were allied to modern Frogs, Toads
and Salamanders, but had the skulls covered by a aymorea
solid bony roof and the bodies by more or less scaly Amphibi-
armor. These Armored Amphibians have been called ans (Stego-
Stegocephalia (oréyn, “epadr = deck-head) or Laby- cepnae
rinthodonts (Ae Bvprv 0s, odovs = labyrinth-tooth, from the com-
plicated fluting or infolding of the enamel on the teeth). Some
of them, like Eryops, were large animals with heads eighteen
inches long and a foot wide; others resembled colossal tad-
poles; but the majority of them were quite small animals, either
proportioned like salamanders or else long and eel-like with
minute limbs or none at all.
These fossil Amphibians are the most ancient of fourfooted
animals, and are not far removed from the central type from
which all the higher vertebrates are believed to be descended.
They are exhibited near the middle of the south side of the Hall
of Fossil Reptiles.
32 THE COLLECTION OF FOSSIL VERTEBRATES
FossiIL FISHES.
Some of the finest specimens of fossil fishes in the collection
are exhibited in the corridor hall. Others are placed in the
southwest corner of the Fossil Reptile Hall. These range from
the exceedingly ancient and archaic types, such as the huge
Dinichthys of the Age of Fishes, older even than the fossil Am-
phibians, to more modern and familiar types such as the fossil
Perch and Herring of the Green River Tertiary formation.
AMERICAN MUSEUM OF NATURAL History.
DEPARTMENT OF VERTEBRATE PALAZONTOLOGY.
Revised List
OF
Casts, Models, and Photographs
OF
Fossil Vertebrates.
New YORK:
Published by the Museum.
19O4.
SUPPLEMENT. TO BULLETIN AMERICAN MUSEUM OF
NATURAL HISTORY, VOL. XX, 1904.
The Knickerbocker Press, Ae
iF
ee |
_ AMERICAN MUSEUM OF NaTurat History.
DEPARTMENT OF VERTEBRATE PALEONTOLOGY.
' REVISED LIST: OF CASTS, MODELS, PHOTOGRAPHS,
AND RESTORATIONS OF FOSSIL VERTEBRATES.
This Department of the American Museum of Natural
History was established in May, 1891, as the Department of
Mammalian Paleontology, and was extended to include Verte-
_ brate Paleontology in 1896. The object of the Trustees is
to procure a representative collection of the American Fossil
Vertebrates from the successive geological horizons of the
West for purposes of exhibition, research and publication.
The collections are being made readily accessible to students,
and placed upon exhibition as rapidly as they are prepared.
Such duplicate original specimens as are available for purposes
of exchange are offered to other museums.
In order to publish as widely as possible the discoveries of
scientific and popular interest and to place the knowledge of
them within reach of those unable to visit this Museum, a
series of plaster casts of specimens and models, and of photo-
graphs of skeletons and restorations has been prepared for
"sale to, or exchange with, other museums, colleges and high
— schools.
t. Plaster Casts of Specimens. — These are from sharp
gelatine moulds, of plaster mixed with gum-arabic, strength-
ened by embedded rods or wires, carefully finished and ac-
curately colored by a skilled artist, so as to be as nearly as
| possible fac-similes of the original specimens. The price is
> based on the actual cost of making, and includes coloring
and packing, but not transportation charges. The packing
a 1 Revised to January, 19004.
I
2 Department of Vertebrate Paleontology.
is done in the department, as carefully as possible, but the
more delicate casts and models cannot be absolutely guaran-
teed against breakage.
2. Models of Extinct Animals. — These are plaster casts of
reduced models (from 1 to 3 feet in length) made by the animal
painter and sculptor, Mr. Charles Knight, under direction of
Professor Osborn. ‘The casts are made in the same manner
as those of specimens and colored under Mr. Knight’s personal
supervision.
3. Photographs of Mounted Skeletons of Extinct Animals. —
These are bromide enlargements, 18 x 22 inches, mounted on
card, with descriptive labels, from photographs by Mr. Ander-
son of mounted skeletons in this Museum. These enlarge-
ments are not available for purposes of exchange and are sold
at $4.00 each.
4. Photographs of Restorations of Extinct Animals. — These
are bromide enlargements, 18 x 22 inches, on card, with de-
scriptive labels, from photographs of the series of water-color
restorations by Charles Knight. These restorations are drawn
from the models above mentioned, and in almost every case
based upon a careful study of the complete mounted skeleton,
made by Mr. Knight under direction of Professor Osborn,
and with advice and assistance of other specialists in Verte-
brate Paleontology. The bromides are sold at $4.00 each
and are not available for exchange.
The models and photographs are copyrighted and are sold
upon the express understanding that they are not to be copied
or adapted for publication.
Orders should be addressed to the CuRATOR.
Payments, either by draft or by postal money-order, should
be made to the order of Treasurer of the American Museum
of Natural Htstory.
HENRY FairFIELD OsBoRN,
Curator.
W. D. MattHEew )
OPS Elay
Associate Curators.
BASHFORD DEAN )
List of Casts, Models, and Photographs. 3
I. — Casts OF ORIGINAL FossiIts AND TYPEs.
Casts Nos. 1 and 2. Coryphodon testis (Cope).
Fore and Hind Feet.
Am. Mus. No. 258.
This large species of Coryphodon was the first described
from this country, and is second only to C. anax in size. The
specimens consist of the left fore and hind feet of the same
animal. The casts consist of the 52 podial elements colored
Fig. 1. Coryphodon testis. No. 258. Fore and hind feet. One-sixth natural size. Lower
Eocene. Wyoming.
and mounted in the natural position upon plaster blocks
imitating the matrix. Each piece is complete and can be
studied separately.
E 7 Price, $20.
Coryphodon radians Corr, OSBORN & WORTMAN, Fossil Mammals of the
Wahsatch and Wind River Beds, Bull. Am. Mus. Nat. Hist., 1892,
Vol. IV, p. 119; OsBorn, A Mounted Skeleton of Coryphodon
radians, ibid. Vol. X, pp. 81-91.
Coryphodon testis (COPE) OSBORN, Evolution of the Amblypoda, Bull.
Am. Mus. Nat. Hist., 1898, Vol. X, pp. 189-205.
No. 3. Palzosyops major Leidy.
Fore Foot.
Am. Mus. No. 1544.
Cast from the foot of the complete skeleton mounted in the
American Museum. This is the characteristic Titanothere of
4 Department of Vertebrate Paleontology.
the Middle Eocene Period (Bridger and Washakie). The foot
bones are not cast separately.
The especial interest of the Titanothere fore foot is that it
is essentially paraxonic (Artiodactyl) although belonging toa
member of the Perissodactyla, with a typically Perissodacty]
(mesaxonic) pes. Several other Artiodactyl characters are
found in this family, supporting Cope’s contention that the
Perissodactyla and Artiodactyla should be united in a single
order (Diplarthra).
Price, $5.
Nos. 4 and 5. Diplacodon emarginatus Hatcher.
Front of Skull and Lower Jaw.
Cast, by courtesy of the Princeton University Museum,
from the type specimen described by Hatcher.
It shows the intermediate stage in the development of
the horns between the ancestral Titanotheres of the Middle
Eocene, hornless or with very rudimentary horns (Pal@osyops
and Telmatotherium), and the horned species (Tztanotherium) of
the Oligocene. The greatest diameter of the horns is antero-
posterior, and both the nasals and frontals enter to some
degree into the base. Diplacodon is found in the Upper or
True Uinta Beds of the Upper Eocene.
Price, $15.
HatcHeEr, On a New Species of Diplacodon, Am. Nat., 1895, p. 1084,
pl eRsorvitd: |
No. 6. Dromatherium sylvestre Emmons.
Type Lower Jaw.
Cast, by courtesy of the Geological Museum of Williams
College, from the original.
This classic specimen is the better preserved of the two jaws
found by Prof. Ebenezer Emmons in 1854 in the Triassic Coal
Beds of Egypt, North Carolina. It is distinguished from the
jaw of a reptile by apparently consisting of a single bone, while
the teeth begin to show two fangs and multiple cusps prophetic
of the mammal type, the Protodonta of Osborn.
Price, $3.
Emmons, American Geology, Part VI, p. 93.
Oszorn, Proc. Acad. Nat. Sci. Phila., 1886, p. 3509.
List of Casts, Models, and Photographs. 5
No. 7. Micronodon tenuirostris Osborn.
Type Lower Jaw.
Cast, by courtesy of the Philadelphia Academy of Natural
Sciences, from the original there preserved.
This specimen, referred by Emmons to Dromatherium
syluestre, was shown by Osborn in 1886 to present important
differences, and made the type of a new genus and species.
The specimen is so small that the cast is not very perfect in
detail.
Price, $3.
No. 8. Periptychus rhabdodon (Cope).
Brain Cast.
Am. Mus. No. 366s.
Periptychus, the most abundant quadruped of the Basal
Eocene Torrejon Formation, presents a very small brain re-
sembling that of a rabbit.
Price, $1.
No. 9. Pantolambda bathmodon Cope.
Brain Cast.
Am. Mus. No. 3057.
The brain of Pantolambda was very similar to that of its
contemporary Periptychus, and gives us the early Amblypod
type, ancestral to that of Coryphodon and Uintathertum.
Price, A igh
10. Giraffe-Camel. Alticamelus altus (Marsh).
Hind Limb.
Am Mus. No. 9109.
Upper Miocene (Loup Fork) of Colorado, Am. Mus. Expedition, 1898.
— Cast from the hind limb of the specimen in the American
Museum Collection. This animal, although a true Camel,
6 Department of Vertebrate Paleontology,
simulated the Giraffe in the extreme elongation of the neck
and legs, and equalled it in size, the hind limb being over
seven feet long. It is a most instructive example of parallel-
ism, as well in its points of difference from the Giraffe as in
the resemblances.
Price, $12.
MatTTHEW, Mem. Am. Mus. Nat. Hist., Vol. I, Pt. vii, 1901, p. 429, pl.
RoR
11. Dissacus saurognathus Wortman.
Lower Jaw,
Am. Mus. No. 2454. Type specimen.
Dissacus was the largest and one of the most characteristic
Creodonts of the Torrejon horizon of the Basal Eocene. It is
believed to have been the direct ancestor of the great Mesonyx
mn
Fig. 2. Diéssacus saurognathus. Lower jaw. Basal Eocene (Torrejon or Upper Puerco)
New Mexico. Three tenths natural size.
of the Middle and Upper Eocene. This very perfect lower
jaw, sixteen inches long, is the best specimen of the large
species.
Price, $8.
MatTTHEw, Bull. Am. Mus. Nat. Hist., 1897, p. 286, fig. o.
List of Casts, Models, and Photographs. 7
12. Polymastodon tadensis Cope.
Lower Jaws,
Am. Mus. No. 968.
A pair of complete rami of the lower jaw. The Multituber-
culates, a characteristic group of Mesozoic time, reached their
culmination in the Puerco or lower horizon of the Basal
Eocene, in this species, about the size of a Beaver. The true
position of this group of pseudo-rodents is uncertain; they
are generally considered, though on very insufficient evidence,
to be related to the modern Monotremata.
‘ Price, $6.
OsBporn & Earve, Bull. Am. Mus. Nat. Hist., 1895, p. 13, fig. te.
13. Polymastodon attenuatus Cope.
Upper Teeth,
Am. Mus. No. 970.
The incisors of one side and premolars and molars of both
sides, set in plaster.
Price, $3.
Osporn & Earte, Bull. Am. Mus. Nat. Hist., 1895, p. 13, fig. rb.
14. Palzonictis occidentalis Osborn & Wortman
Lower Jaws and Front of Skull.
Am. Mus. No. 110. Type specimen.
From the Suessonian (Wasatch) of Wyoming.
This rare and fine specimen, found by Wortman in 1891,
supplies another proof of the homotaxy of the Wasatch with
the Suessonian of Europe, where the genus is represented by
the less perfect specimens preserved in the Paris Museum.
| Price, $12.
Ossorn & WortMaN, Bull. Am. Mus. Nat. Hist., 1892, p. 104, pl. iv.
8 Department of Vertebrate Paleontology.
15. Carnivorous Dinosaur, Allosaurus.
Hind Limb Complete.
Am. Mus. No. 290.
Upper Jurassic (Atlantosaurus or Como beds) of Wyoming, Expedition
of 1808.
The limb is seven feet two inches high, as mounted in the
natural position, and is almost absolutely complete, the distal
Fig. 3. Alosaurus hind limb,
from Upper Jurassic of Bone
Cabin Quarry, Wyoming.
parts of the fourth digit being perfect, the proximal splint only
wanting. Each bone is cast separately—twenty-four pieces
ia cally
Price, $40.
OsBorn, Bull. Am. Mus. Nat. Hist., 1899, p. 161, figs. 1-4a.
16. Patriofelis ferox (Marsh).
Fore and Hind Feet.
Am. Mus. No. 1507.
Original from the Bridger or Middle Eocene of Wyoming, Expedition
of 1893.
Typical Creodont or Primitive Carnivore foot. Taken from
the mounted skeleton in the American Museum Collection.
List of Casts, Models, and Photographs. 9
Displays the broad, spreading foot with blunt, hoof-like claws
and very limited play on the joints, common to most of the
larger Creodonts. The hind foot is nine inches long.
Price, $ro.
OsBorn, Patriojelis and Oxyena Re-studied as Terrestrial Creodonts
Bull. Am. Mus. Nat. Hist., XIII, 1900, pp. 269-279. .
Wortman, Osteology of Patriofelis, Bull. Am. Mus. Nat. Hist.
VI, 1894, pp. 129-164. :
Fig. 4. Patriofelis ferox (Marsh). Fore and hind feet. One-fourth natural size.
17-20. Type Specimens of Eocene Tapiroids.
PRICE
17. Desmatotherium guyoti, upper jaws, type.--.----- $1.00
18. Dilophodon minusculus, lower jaw, type..------- I.00
19. Isectolophus latidens, jaw fragments, types....-.. 1.50
20. Isectolophus annectens, jaw fragments, types....-. 1.50
Four little-known Tapiroid mammals from the Bridger and
Uinta Eocene of Wyoming. ‘The originals are in the Prince-
ton Museum.
Scott, Desmatotherium and Dilophodon, Contributions from the Mu-
~ seum of Princeton College, 1883, p- 51, pl. vill.
Scotr & OsBorN, The Mammalia of the Uinta Formation, Trans.
Amer. Philos. Soc., 1889, pp- 519, etc.
10 Department of Vertebrate Paleontology.
21. Heptodon calciculus Cope.
Palate and Complete Lower Jaws.
Amer. Mus. No. 204.
Primitive Lophiodont from the Wind River Eocene of Wy-
oming. Probably ancestral to the Colodons of the American
Oligocene.
Price, $7.
OsBorN & WorTMAN, Fossil Mammals of the Wasatch and Wind River
Groups, Collection of 1891, Bull. Amer. Mus. Nat. Hist., IV, 1892,
1s, 1A 7fe
22. Colodon dakotensis Osborn & Wortman.
Upper Jaws.
Am. Mus. No. 1212.
White River Oligocene, 5. Dakota.
Price, $3.
23. Colodon occidentalis (Leidy).
Lower Jaws and Fore Foot.
Am. Mus. No. 6583
White River Oligocene, S. Dakota.
This rare Lophiodont is known only by fragmentary speci-
mens. The group is intermediate between Tapirs and Horses
in some respects, between Tapirs and Rhinoceroses in others.
It left no descendants.
Price, $5.
OsBorN & WorrmMan, Perissodactyls of the Lower Miocene White
River Beds, Bull. Amer. Mus. Nat. Hist., VII, 1895, p. 362, fig. 7.
WortTMAN & Earte, Ancestors of the Tapir from the Lower Miocene of
Dakota, Bull. Amer. Mus. Nat. Hist., V, 1893, p. 174, figs. 6 and 7.
List of Casts, Models, and Photographs. II
24. Systemodon primevus Wortman.
Palate and Lower Jaw.
Am. Mus. Nos. 144 and 131.
Suessonian (Wasatch Beds) Wyoming.
Systemodon is the starting point, as far as known, of the
Tapir line, and strikingly resembles in most respects the con-
temporary ancestors of the Horse (Hyracotherium, etc.).
Price, $4.
OsBoRN & WorRrMAN, Fossil Mammals of the Wasatch and Wind River
Beds, Collection of 1891, Bull. Amer. Mus. Nat. Hist., IV, 1892,
Be rage fis. 17;
WormTMAN, Species of Hyracotherium, etc., Bull. Amer. Mus. Nat. Hist.,
Niliie=r896; p. $7, fg."3:
25. Protapirus validus Hatcher.
Skull.
Type specimen (Original in Princeton Museum).
Protoceras Beds, White River Formation (Oligocene), S. Dakota.
Price, $7.
Hatcuer, Recent and Fossil Tapirs, Amer. Jour. nci., [| 1896, pi 169;
pleas.
26. Protapirus obliquidens Wortman & Earle.
Lower Jaws and Fore Foot.
Am. Mus. Nos. 661 and 662.
Protoceras Beds, White River Formation (Oligocene), S. Dakota.
Protapirus is intermediate between the modern Tapirs and
their primitive Eocene ancestors (Systemodon and Isectolo-
a) Price, $6.
WortMan & Ear e, Ancestors of the Tapir from the Lower Miocene
~ of Dakota, Bull. Amer. Mus. Nat. Hist., V, 1893, p. 165, figs. 2,
3, 4-
12 Department of Vertebrate Paleontology.
27. Oreodon culbertsoni Leidy.
Fore and Hind Foot.
Am. Mus. No. 1287. Cast from the mounted skeleton in the American Museum.
Oreodon Beds, White River Formation (Oligocene), S. Dakota.
Shows a very primitive Artiodactyl foot, the fore foot still
retaining a rudimentary thumb, and the lateral digits being
less reduced than in any of the more recent Artiodactyls.
Price, $6.
Scott, W. B., Beitrage zur Kenntniss der Oreodontidz, Morph. Jahrb.,
1890, Pp. 328,334} pl. xvi, figs: Sy, 28:
28. Hyznodon horridus Leidy.
Fore and Hind Foot.
Am, Mus. No. 1375. Cast from the mounted skeleton in the American Museum.
Oreodon Beds, White River Formation (Oligocene), S. Dakota.
Hyenodon was the latest and most specialized of the Creo-
donts or Primitive Carnivores, and the feet are adapted for
running, although not so well as in the modern swift-footed
Carnivora.
Price, $7.
Scott, W. B., The Osteology of Hyenodon, Journ. Acad. Nat. Sci.
Phila Vol wexe
29. Series of Fossil Horse Feet.
Illustrating the Evolution of the Horse.
The Horses afford the best illustration of the evolution of
a race of animals during geological time, commencing in the
Lower Eocene as small fox-like animals with several toes on
each foot, and ending with the modern Horses, Asses, and
Zebras. At least ten intermediate stages are known in the
direct line of descent, besides several stages of collateral
branches, leading into types which have left no modern de-
scendants. Those of which casts are now ready are:
List of Casts, Models, and Photographs. 13
Stage b.—Hyracotherium craspedotum. Fore and hind feet. Middle
Eocene (Wind River), Wyoming. Four toes and a rudi-
ment of a fifth (1st digit) on the fore foot, three toes on
the hind foot. Size of a fox. Price, $4.
Stage e.—Mesohippus bairdi. Fore and hind feet. Lower Oligocene
(White River), S. Dakota. Three toes on fore and hind
foot, the fourth (5th digit) represented on the fore foot by
a short “‘splint-bone.’”’ Lateral toes much reduced, but
still touching the ground. Size of a prairie wolf.
Price, $4.
Stage 7’.—Mesohippus intermedius. Fore and hind feet. Upper Oli-
gocene (White River, Protoceras Beds), S. Dakota. Like
M. bairdi, but larger. Price, $5.
Stage 7”—Mesohippus copei. Hind foot. Upper Oligocene (White
River, Protoceras Beds), S. Dakota. Larger than M.
intermedius. Price, $3.
Stage h”—Hypohippus equinus. Fore and hind feet. No. 8407
Middle Miocene (Loup Fork), Colorado. Larger than any
of the preceding species; size of Shetland pony. Fifth
digit of the fore foot reduced to a little nodular rudiment.
Side toes still touch the ground. Price, $12.
Stage i!”.—Merychippus sejunctus. Fore and hind feet. Middle Mio-
cene, Colorado. Fifth digit of fore foot a small nodular
rudiment, side toes much reduced, not reaching the
ground. Size of 3-months-old colt. Price, $10.
Stage j”.—Neohipparion whitneyi. Fore and hind feet. Upper Mio-
cene (Loup Fork), S. Dakota. Feet and limbs elongate,
proportions like those of the deer. Side toes more re-
duced than in Merychippus, size somewhat larger.
Price, $12.
30. Series of Fossil Horse Skulls.
Illustrating the Evolution of the Horse.
(Stages corresponding with those in No. 29.)
Those now ready are:
Stage c.—Protorohippus venticolus. Crushed skull and jaws. No,
4832. From mounted skeleton in Cope Collection. Mid-
dle Eocene (Wind River), Wyoming. Short-crowned
f primitive pattern. Size of fox.
teeth of pri I ethe wk
14 Department of Vertebrate Paleontology.
Stage e—Mesohippus bairdi. Skull and jaws. No. 1477. From
Lower Oligocene (Oreodon Beds, White River), Nebraska.
Short-crowned teeth of more horse-like pattern. Size of
prairie wolf. Price, $8.
Stage h” —Hypohippus equinus. Skull and jaws. No. 8407. From
Middle Miocene (Loup Fork) of Colorado. Teeth much
like those of Mesohippus. Size of Shetland pony.
Price, $15.
Stage 2’”’ —Merychippus sejunctus. Skull and jaws. No. 8291. Cope
Collection. Middle Miocene (Loup Fork) of Colorado.
Long-crowned teeth with cement. Pattern intermediate
between Mesohippus and Equus. Size of 3-months-old
colt. Price, $15.
31. Pantolambda bathmodon Cope.
Fore and Hind Feet.
From mounted skeleton in American Museum. No. 2540.
Torrejon Formation (Basal Eocene) of New Mexico.
Primitive ungulate foot, with five toes, first digit semi-
opposable, plantigrade step, wide, loosely joined wrist and ~
ankle-bones (giving much flexibility in all directions at the
expense of power), and other characters now preserved chiefly
among arborealmammals. The foot approaches nearly those
of the more ancient unguiculates or clawed animals. Panto-
lambda was ancestral to the ancient Amblypoda (Coryphodon
and Utintathertum), but only a collateral ancestor of the more
recent hoofed animals.
Price, $5.
Ossorn, Evolution of the Amblypoda, Bull. Am. Mus. Nat. Hist., X,
1898, pp. 183-188, figs. 9, Io, 12.
32. Series of Fossil Camel Feet.
Illustrating the Evolution of the Camels and Llamas.
Although now found only in the desert regions of Asia,
Africa, and South America, the Camel family was of North
American origin, spreading to other continents only in the
Pliocene epoch, and becoming extinct in its original home
List of Casts, Models, and Photographs. 15
during the Pleistocene. During the Tertiary the race evolved
from small deer-like animals, no larger than jack-rabbits, with
sharp, pointed hoofs and separate metapodial bones, to their
present size and proportions, the metapodials becoming solidly
fused together and a large pad on the foot supporting the
weight of the body. Collateral lines of descent also existed,
one ending in an animal of singularly giraffe-like proportions,
although a true Camel (Giraffe-Camel, see also No. 10, on
page 5).
Stage a. — Protylopus petersoni. Hind limb. No. 2564. Upper
Eocene (Uinta), Utah. Separate metapodials, lateral
digits represented by short splints, pointed hoofs. Size
of jack-rabbit. Price, $5.
Stage b. — Poébrotherium wilsonit. Fore and hind feet. No. 1364.
Lower Oligocene (White River), S. Dakota. Separate
metapodials, lateral digits represented by small nodular
rudiments, pointed hoofs. Size of vicufia.
Price, $5.
Stage e’— Protolabis montanus. Fore and hind feet. No. 9108.
Middle Miocene (Loup Fork), Colorado. Separate meta-
podials, lateral rudiments fused, hoofs pointed. Size of
llama. Price, $6.
Stage 7”.— Alticamelus altus. (Giraffe-Camel.) Hind limb. No.
gtog. Middle Miocene (Loup Fork), Colorado. United
metapodials, greatly elongated, hoofs considerably re-
duced. Size of small giraffe. Price, $12.
33. Protoceras celer Marsh.
Fore and Hind Foot.
Protoceras Beds, White River Oligocene, S. Dakota.
Protoceras is the largest of a pec uliar group of primitive
ruminants characteristic of the White River Oligocene. The
fore foot has four fully functional digits, but in the hind
foot the lateral digits are reduced to small splints. The
animal was somewhat smaller than the musk deer, much less
16 Department of Vertebrate Paleontology.
gracefully proportioned, the skull remarkably specialized in
the male.
Price, $5.
Scott, Osteology and Relations of Protoceras, Jour. Morph., Vol. XI.
OsBoRN & WortTMAN, Characters of Protoceras, etc. Bull. Amer.
Mus. Nat. Hist., IV, 1892, pp. 351-372.
34. Type Specimens of Fossil Equide.
PRICE
Equus complicatus Leidy. Upper molar. Pleistocene......... $2.00
“« excelsus Leidy. Upper jaw. ees PG te ieee 4.00
‘* occidentalis Leidy. Upper molars. pei LS eae nein 2.00
S| pectinatus Cope. Upper teeth asenics, ua enn) seni er 5.00
Hipparion speciosum Leidy. ee teeth.» Miocene. eke 3.00
ie affine Leidy. eR a era 3.00
zs occidentale Leidy 0 SO Se ees 3.00
gratum Leidy * Ape PE Seren oo 25 2.50
montezsume Leidy. Upper and lower tooth. Mio-
COTS sce d cs 55 wa ety te toc ee So asta ees fect a a 2.00
Merychippus mirabilis Leidy. eee TENE Miocene ty accor 3.00
= insignis Leidy. oie teeta eee Saeed 3.00
Parahippus cognatus Leidy. Upper teeth, PSs treme tact 2.50
Parahippus (Desmatippus) crenidens Scott. Upper and lower
jaws. Miocenes...% 622 ee peter et ee aerator oF 6.00
Anchippus texanus Leidy. Upper molar. Mliocene.......... 1.50
Hypohippus affinis Leidy. a Sat, i besetcer ses inet 1.50
Hypohippus (Anchitherium) equinus. Upper and lower jaws.
MIOCENE... wis'ein'ae sts, fi sie cues eye pein etal te te hee te ite kena eat 6.00
Mesohippus bairdi Leidy. Figured skull and jaws. Leidy,
1860; pl. xx. Oligocene. ci Gigwise ee ee eee 5.00
The patterns of the molars in the above type specimens
have been copied with especial care and accuracy on the
casts.
35. Phenacodus primevus Cope.
Fore and Hind Feet.
Wasatch Formation, Lower Eocene, Wyoming.
The mounted skeleton of Phenacodus in the Cope Collection
(now in the American Museum) is well known by descriptions
List of Casts, Models, and Photographs, EF
and figures in all textbooks of geology and paleontology.
Phenacodus is the type of the Condylarthra, a very archaic
order of ungulates retaining the primitive form of wrist and
ankle articulations, five toes on each foot, and pig-like teeth,
and considered to represent very nearly the central prot-
ungulate type from which all hoofed animals are descended.
The size of P. primevus was somewhat less than that of a pig.
Price, $ro:
Copz, E. D.,Tertiary Vertebrata, Rep. U. S. G. S. Terrs., Vol. III,
B’k I, p. 435, and plates.
Ossporn, H. F., Remounted Skeleton of Phenacodus, Bull. Am. Mus.
Nat. Hist., X, 1898, pp. 159-164.
36. Euprotogonia puercensis (Cope).
Hind Foot.
Torrejon Formation, Basal Eocene, New Mexico.
Euprotogonia was the ancestor of Phenacodus, and has a still
more primitive type of foot. The five toes are tipped with
narrow, claw-like hoofs, and in other characters this foot is
Fig. 5. Luprotogonia puercensis.
Hind foot. One-half natural size.
intermediate between the hoofed animals (Ungulata) and
clawed animals (Unguiculata), and indicates the derivation
18 Department of Vertebrate Paleontology,
of both from a common ancestor. This species was about
the size of a cat, and the cast is taken from an incomplete
skeleton in the American Museum.
Price, $4.
MarrHew, Revision of the Puerco Fauna, Bull. Amer. Mus. Nat. Hist.,
DEL 189%, ps SOs; a. Eee
OsBorN, Remounted Skeleton of Phenacodus, Bull. Amer. Mus. Nat.
Hist., X, 1898, p; 150.
37. Stereosternum tumidum Cope.
Skeleton.
Upper Permian, near Sao Paulo, Brazil.
This beautifully preserved skeleton is one of the most an-
cient and primitive reptiles known, and represents a distinct
and very primitive order of Reptilia, the Proganosauria,
with many characters connecting it with the Amphibia.
Skeleton in block, about two feet long, showing all parts
except the left fore and hind feet, and with the outlines of
the body indicated. By courtesy of the Director of the Geo-
logical Survey of Brazil.
Price, B25:
Cope, Contribution to the Vertebrate Paleontology of Brazil, Proc.
Am. Phil. Soc., XXIII, 121, 1885; The Carboniferous Genus
Stereosternum, Amer. Nat., 1887, 1109.
OssBorn, Memoirs Am. Mus. Nat. Hist., Vol. I, Pt. VIII, p. 481, figs.
15-109, and pl. xl, Nov., 1903.
II.— Casts or MopEets or EXTINCT VERTEBRATES BY
CHARLES KNIGHT.
Originals presented by $. Pierpont Morgan, Esq.
No. 1. Three-horned Frill-necked Dinosaur. Triceratops
prorsus Marsh.
The original model was made by Mr. Knight for the National
Museum, under direction of Mr. Lucas, and is based on parts
of several skeletons there preserved.
List of Casts, Models, and Photographs. 19
Triceratops was one of the last and most remarkably
specialized of the Dinosaurs. It was herbivorous, quadru-
pedal, with elephant-like legs and feet. The skull was of
huge size, 6-8 feet long, with long, powerful horns projecting
forward, and a great bony frill projecting backward and com-
pletely covering the neck. One-tenth natural size. Length
of base, 18 inches.
Price, $20.
No. 2. The Duck-billed Dinosaur. Hadrosaurus mirabilis
Leidy.
The model of this remarkable Dinosaur is based on an almost
complete specimen in the Cope Collection, now in this Museum,
which will shortly be placed on exhibition as a mounted
skeleton. It was found in the Laramie Cretaceous by Messrs.
Wortman and Hill, and described by Professor Cope (under
the generic name Diclonius) in 1883. The animal was thirty
feet in length, with a long neck, flattened, duck-like bill,
numerous small teeth, small fore limbs, and heavy hind limbs
and tail. It was probably of amphibious habits, feeding on
soft water-plants. It was covered with a thick, rhinoceros-like
hide, parts of which are preserved in Professor Cope’s speci-
men. One-eleventh natural size; length of base, 12 inches.
Cope, Proc. Acad. Nat. Sci. Phila., 1883, p.97; Am. Nat., 1883, p. 774.
Price, $10.
No. 3. Leaping Carnivorous Dinosaur. Dryptosaurus (Le-
laps) aquilunguis Cope.
The first Dinosaur described by Professor Cope was Lelaps
aquilunguis from the New Jersey Cretaceous beds. The model
is based upon this fragmentary skeleton and upon the restora-
tion by Professor Marsh of the allied form Ceratosaurus.
The Carnivorous Dinosaurs, although smaller than some of
the herbivorous kinds, were animals of great size. This species
20 Department of Vertebrate Paleontology.
was about fifteen feet long. According to Professor Cope’s
views, by which Mr. Knight was chiefly guided in making this
and the following restoration, the animal was an active and
powerful leaper, and the model, representing two fighting
Lelaps, carries out this conception. One-seventh natural
size. Length of base, 18 inches.
Cope, Proc. Acad. Nat. Sci. Phila., 1866, p. 275; Proc. Amer. Philos.
Soc., XXX, p. 240, May, 1892.
Mars, Dinosaurs of North America, pp. 157-163, pl. xiv.
Price, $20 (2 figures, complete).
No. 4. Naosaurus claviger Cope.
From the Permian Beds of Texas. This belongs to the prim-
itive Reptilian order Pelycosauria of Cope, but represents a
highly specialized side-branch, related to the Khynchocephahia
or Proganosauria as shown by Baur and Case. The precise
object of the extraordinary rigid fin on the back is not known;
it was humorously suggested by Cope that it might have been
used as a sail. It was, perhaps, chiefly ornamental. Differ-
ent species of Naosaurs reach from three to ten feet in length,
and the dorsal fin reaches a length even greater than that
shown in the model. One-fifth natural size. Length of base,
12 inches.
Cope, Proc. Amer. Phil. Soc., 1878, p. 510, and subsequent papers.
Baur & CasE, Morphology of the Skull of the Pelycosauria, Anatom.
Anzeiger, Bd. XIII, p. 109, Jena, 1897.
Price, $12.
No. 5. Cervalces americanus (Harlan).
This Pleistocene American Elk was of the size and propor-
tions of the living Moose, but had horns almost as large as
those of the Extinct Irish Deer of Europe, and expanded in
three planes of growth at right angles to each other. The
mhodel is based on a remarkably perfect skeleton found in New
Jersey and mounted in the Princeton University Museum.
List of Casts, Models, and Photographs, 21
This skeleton was fully described by P
rofessor Scott, in 1885,
as possessing characters intermediate between those of the
deer and moose. One-fourth natural size. Length of base,
24 inches. |
scott, Proc. Acad. Nat. Sci. Phi
No. 6. Long-horned Titanothere. Brontotherium platyceras
(S. & O.,
Based on the mounted sk I : ~
1 y ry] L110 O tne [ ita
Museum, and on the monograj
by Professor Osborn.
22 Department of Vertebrate Paleontology.
The model represents a charging Titanothere, and illustrates
a chief use of the great horns developed in these animals.
One-eighth natural size. Length of base, 25 inches.
Price, $20.
No. 7. Series of Heads of Titanotheres.
This series of five heads represents the evolution and poly-
phyletic development of the Titanotheres during the Lower
Oligocene.
Brontotherium platyceras,
Megacerops robustus,
Titanotherium ingens,
Symborodon acer,
Diplacodon emarginatus.
eBooe
One-sixth natural size.
Price, $5 each; for the set, $20.
No. 8. Restored Head of Dinocyon. Dinocyon gidleyi Matthew.
Upper Miocene Epoch, Texas. One-fourth natural size.
Based on the skull and jaws in the American Museum.
This gigantic dog equalled the polar or Kadiak bear in size,
and resembled them in many external characters. One-
fourth natural size.
Price, $5.
No. 9. Sabre-tooth Tiger. Smilodon.
This restoration model is based on a complete mounted
skeleton of S. necator in the American Museum.
Smilodon, of the Pleistocene epoch, was the latest and
largest of the Sabre-tooth Tigers, equalling a polar bear in
size, and provided with canines which projected seven inches
fromthe jaw. It ranged through the New World from Canada
to Patagonia, and probably preyed on the great ground sloths
or other gigantic herbivora of the Tertiary Period, to pierce
whose thick hide and heavy fur the enormous canines were
well adapted. One-sixth natural size. Length of base, 14
inches.
Price, $15.
List of Casts, Models, and Photographs.
to
Ge
III. — PHotoGrapus oF MounTED SKELETONS.
Bromide enlargements from the original negatives. Size,
IS in. % 22 in,
These photographs are taken from the fossil vertebrates
in the American Museum, as an exhibition of the Succession
of Vertebrate Life in North America. Other skeletons are in
various stages of preparation, photographs of which will be
added to this series.
The skeletons show the actual form, proportions, and atti-
tudes of these extinct animals as nearly as they can be de-
termined or the preservation of the specimen permits.
The photographs also exhibit the different methods em-
ployed in mounting, each method being adapted to the special
condition of the fossil. For example, Cenopus tridactylus,
No. 11, and Tylosaurus, No. 14, are cases where a LOW
RELIEF mounting was necessary. Amynodon, No. 18, is an
instance of the HIGH RELIEF mounting. In some cases the
matrix is wholly or partly the original rock in which the
fossil was found; in others it is a close imitation of it, made
partly by mingling the rock with plaster, thus giving the
visitor and student a truer conception of the original em-
bedding. Other animals are mounted in FULL RELIEF, like a
recent skeleton, with the aid of concealed, or partly concealed,
iron rods. Except when abundant duplicate material is
available, each bone is made easily removable for purposes
of study, fastened to its mountings by wire ties or by con-
-cealed screws and bolts. The mounting is under the direction
of the Curator and of Mr. Adam Hermann, Preparator.
Price of the photographs, $4 each.
1. Metamynodon planifrons Scott G Osborn.
Am. Mus. No. 546.
This aberrant rhinoceros of the Oligocene epoch was dis-
covered by Garman and described by Scott and Osborn in
1887. The mounted skeleton is a composite from materials
obtained bv the Amer. Mus. Expeditions of 1892 and 1894 in
24 Department of Vertebrate Paleontology.
the Big Badlands of South Dakota. Its length is 9 ft. 7 inches.
The general impression is of a very broad, flat skull, with
formidable canine tusks; small, but prominent and greatly
elevated eye-sockets, and a very broad chest. The fore and
hind limbs are quite powerful, but the metapodials are rather
slender, especially in the manus. This animal is widely
separated from the true Rhinoceros by its four completely
functional digits on the fore foot and by the strong develop-
ment of the canines.
Osporn & WoRTMAN, Perissodactyls of the Lower Miocene, White
River Beds, Bull. Am. Mus. Nat. Hist., VII, 1895, pp. 373-375,
Pllc
2. Brontotherium, sp.
Am. Mus. No. 518.
This skeleton represents the largest size and the last stage
in the evolution of the Titanotheres, and was discovered by
the American Museum Expedition of 1892 in the Upper
Titanotherium Beds of South Dakota, absolutely complete as
far back as the border of the pelvis. The hind limbs, belonging
to different animals, but of the proper proportions, were secured
in the same regions of the West in the summer of 1894.
The complete skeleton, about 7 feet 7 inches (M. 2.312)
high; 13 feet 8 inches (M. 4.17) long, and 3 feet 10 inches
(M. 1.17) broad, probably belongs to an adult female, as we
judge from the imperfect development and ossification of the
horns, which in males of this period are very long and power-
ful. An interesting feature of the skeleton is an exostosis and
false joint in the centre of the seventh rib, undoubtedly an
after-result of fracture.
OsBorN & WorRTMAN, loc. cit., pp. 346-352, pll. viii, ix.
3. Hyrachyus agrarius Leidy.
Am. Mus. No. 5065.
" This is the original skeleton discovered by Professor Cope
himself in his explorations in the Bridger Basin (Middle
List of Casts, Models, and Photographs. 25
Eocene) in 1873, and was mistakenly referred by him to H.
eximius. It was mounted as found, with an incomplete skull,
and so figured by Cope in ‘The Tertiary Vertebrata,’ The
American Museum Expedition of 1893 secured a complete
skull belonging to this species, and of the proper size, which
has been affixed to the skeleton. In order fully to expose the
bones, and correct several errors in the original mounting, the
entire animal was taken apart and remounted, as here photo-
graphed. The animal was about as large as a sheep, and is
the oldest known type of Rhinoceros, more directly ancestral to
the Hyracodon or Cursorial Rhinoceros of the Oligocene.
Cope, Tertiary Vertebrata, U. S. Geol. and Geog. Sur. Terr’s, F. V.
Hayden in charge, Final Report, Vol. IV (188s), PP. 657-677, pill.
liv, lv, lva, etc.
OssBorn & WortMaAn, Perissodactyls of the White River Beds, Bull,
Am. Mus. Nat. Hist., VII, 1895, pp. 367-371.
4. Patriofelis ferox (Marsh).
Am. Mus. No. 1507.
This animal was originally described by Leidy from a frag-
ment of the lower jaw. The American Museum Expedition
of 1893 procured the complete skeleton, represented in two
different animals, in which the skull alone was in an imperfect
fragmentary condition, and the teeth, unfortunately, entirely
wanting. This animal was as large as a jaguar, and exhibits
short, powerful, highly flexed limbs, widely spreading clawed
feet, heavy backbone, very deep sagittal crest and small
‘brain case. It is a typical Middle Eocene Creodont, with
highly specialized cutting teeth.
WormtaN, The Osteology of Patriofelis, Bull. Am. Mus. Nat. Hist., VI,
1894, pp. 129-164, pl. i.
OsBorn, Oxyena and Patriofelis Re-studied as Terrestrial Creodonts.
Bull. Am. Mus. Nat. Hist., XIII, 1900, pp. 269-279, pll. xviii, xix.
5. Protorohippus venticolus (Cope).
Am. Mus. No. 4832.
This is the famous skeleton described by Cope in ‘ The
Tertiary Vertebrata,’ as the four-toed Lower Eocene Horse.
26 Department of Vertebrate Paleontology.
It was found by Dr. J. L. Wortman in the Wind River Beds of
Wyoming (Middle Eocene). Since its purchase by the Ameri-
can Museum the entire skeleton has been taken apart and re-
mounted as a walking animal; in the original mounting the
animal was represented as pacing. The skeleton is far from
perfect, the limbs upon the left side being largely restored,
while those upon the right side are complete. The ribs are
entirely restored, as is the pelvis. In the remounting of the
skeleton these missing parts were studied from the correspond-
ing bones in the well-known form Mesohtppus.
Cope, Tertiary Vertebrata, pp. 635-647, pl. xlixa, b and c.
WortMAN, Species of Hyracotherium and allied Perissodactyls from the
Wahsatch and Wind River Beds of North America, Bull. Am. Mus.
Nat. Hist., VIII, 1896, pp. 81-110.
6. Hoplophoneus primevus Leidy.
Am. Mus. No. 1406.
The skeleton was procured by the American Museum Expe-
dition of 1894 in the Oreodon Beds of South Dakota, and is one
of the most complete fossil skeletons ever found. The only
parts missing were some of the processes and spines of a
few vertebre. This animal is characterized by powerful
canines protected by a heavy flange upon the lower jaw, and
is considered the ancestor of the great Sabre-tooth Tiger
Smilodon.
Riccs, Restoration of Hoplophoneus occidentalis Leidy, Kan. Univ.
Quar., V, 1896, pp. 37-52, pl. i.
7. Palzosyops major Leidy.
Am. Mus. No. 1544.
Pal@osyops was one of the first fossils found in the Eocene
deposits of the Western States. Materials for this mounted
skeleton were secured with great difficulty by the American
Museum Expeditions of 1893 and 1895 in the Bridger and
Washakie Beds of Wyoming. The skull, fore limbs and foot,
and a large part of the vertebral column and ribs belong to a
single skeleton, and the parts associated with these were put
List of Casts, Models, and Photographs. 27
ee ee
together after being positively determined as belonging to this
species, and as representing individuals of corresponding age,
The skeleton is thus believed to represent accurately the
original, and there is little restoration, except in the head of
the femur. This species is distinguished by the convexity
of the forehead, and by the low, flattened cones of the molar
teeth, and is not directly ancestral to the later Titanotheres.
Ear e, On the genus Pal@osyops, etc., Jour. Acad. Nat. Sci. Phila., 1892
8. Phenacodus primevus Cope.
Am. Mus. No. 4369.
This is the famous skeleton of Phenacodus found by Dr.
Wortman in the Lower Eocene of the Big Horn Basin,
Wyoming, and described by Cope in ‘Tertiary Vertebrata’
and other publications. It here has an entirely different ap-
pearance, and gives us a different conception of the animal
from that presented in its original mounting. The skeleton
was very largely embedded in the rock, and it required four
months of continuous work to remove the limbs, ribs and
vertebre preparatory to remounting. The animal as figured
in “Tertiary Vertebrata’ appeared like a plantigrade; it was,
however, restored by Sir William Flower in his book upon the
Horse as a digitigrade. As here shown, the animal was as
digitigrade as the Tapir; the hind limbs are much longer and
more powerful than the fore; the tail is of great size. The
head, on the other hand, is extremely small, and out of all
- proportion to the body. It is thus a microcephalic type, in
contrast with.Coryphodon, which is macrocephalic.
Cops, Tertiary Vertebrata, pp. 428-463, pll. lviib—Ivin.
Ossorn, Remounted Skeleton of Phenacodus, Bull. Am. Mus. Nat.
Hist., X, 1898, pp. 159-164.
9. Coryphodon testis Cope.
Am. Mus. No. 2865.
Coryphodon is the characteristic large mammal of the Lower
Eocene or Wasatch. Its remains are usually found scattered,
*OZIS
Jeanqeu yaj}[aA\y-2uO
*ALOISIF{ [eANIENE JO UlNasNT ULOLIAUIW ayy UI SHAHuZLg SNporvUaY JO UOJa[ayxs pajyunoy 2 “Big gz
List of Casts, Models, and Photographs. 29
and it was not until 1806 that the Museum Expeditions
enabled us to mount the complete skeleton. The animal ex-
hibits a curious mixture of primitive and specialized charac-
ters, the former being dominant in the limbs and body, the
latter appearing in the skull and feet. The flat-topped skull
with slight rudiments of frontal and parietal horns, the spread-
ing, hippopotamus-like front teeth, the short elephantine feet
and broad pelvis, are the most noticeable specialized charac-
ters. Coryphodon was nearly related although not directly
ancestral to the Uintatheres (Dinoceras) of the Middle Eocene
or Bridger Beds.
The skeleton as mounted is composed of a number of in-
dividuals of the same species, size and age, collected by the
Museum Expedition of 18096.
OsBorNn, Coryphodon radians Cope, A Complete Skeleton of Corypho-
don, Bull. Am. Mus. Nat. Hist., X, 1898, p. 81.
Coryphodon testis OsBorn, Evolution of the Amblypoda, ibid., pp.
189-204.
See also Marsu, Am. Jour. Sci., 1893, p. 321.
10. Teleoceras (Aphelops) fossiger (Cope).
Am. Mus. No. 2604.
This Rhinoceros was the most abundant large animal of the
Upper Miocene of Western America. Its bones are found in
some localities (such as the famous quarry at Long Island,
Phillips Co., Kansas) in great numbers and remarkably well
preserved, but the individuals are not distinguishable. This
_ skeleton, made up of many individuals of appropriate size, is
mounted with concealed steel rods traversing the bones and
only two visible main supports. The male probably bore a
small horn, which was rudimentary or absent in the female.
The animal had much shorter legs and a heavier body than
any modern Rhinoceros, the principal measurements being:
height at shoulder, 4 ft. (M.1.22); at pelvis, 3 ft. 8 in. (M. 1.12);
total length, 1o ft. 3 in. (M. 3.12); greatest girth or chest
circumference, 9 ft. 2 in. Its size was about that of the
largest living species, except in the height. The neck was
30 Department of Vertebrate Paleontology.
short, the head rather small, the body extremely thick and
barrel-like, the belly almost reaching the ground.
OsBorn, Bull. Mus. Comp. Zodl., XX, p.92; Bull. Am. Mus. Nat. Hist.,
LOGS s pees
Wiutston, Restoration of A phelops fossiger, Kans. Univ. Quar., 1894,
p. 280, pl. viii.
11. Cznopus tridactylus (Osborn).
Am. Mus. No. 538.
This skeleton was found by the American Museum Field
Party in 1892, and was the first skeleton mounted by the
Department; although very complete it is considerably
crushed, and has been mounted in low relief. The skeleton as
it lay in the rock was doubled over on itself; it has been
straightened out and a missing fore limb modelled in plaster,
otherwise it is in the position that it had in the rock. The
matrix is chiefly plaster, colored and chipped to imitate the
surface of the gray sandstone original. Length from tip of
nasals to bend of tail, 7 feet g inches. The animal was about
the size of the living Sumatran Rhinoceros, but was hornless,
or with rudimentary paired horns on the male.
OsBorRN & WortMAN, Fossil Mammals of the Lower Miocene White
River Beds, Bull. Am. Mus. Nat. Hist., VI, 1894, pp. 206-207, pl.
ili.
OsBorRN, Mem. Am. Mus. Nat. Hist., Vol. I, Pt. IV, Oct., 1899.
12. Phenacodus wortmani Cope.
Am. Mus. No. 4378.
This small species of Phenacodus was found by Dr. Wort-
man in the same beds as P. -primevus. This skeleton was
mounted in the original matrix in the Cope Collection, but has
been chiselled out of the rock and remounted in low relief, the
missing parts of the bones being restored in plaster. Each
bone is easily removable for separate study.
Cope, Tertiary Vertebrata, p. 464, pl. xxixe.
List of Casts, Models, and Photographs. 3!
13. Canopus occidentalis (Leidy).
Am. Mus. No. 1132.
This characteristic Rhinoceros of the Middle Oligocene of the
West is ancestral to C. tridactylus of the Upper Oligocene, and
thus leads to the Dicerathere line. C. occidentalis was horn-
less, somewhat smaller than the living Sumatran Rhinoceros
and more lightly proportioned. This skeleton is mounted in
full relief. .
Ossporn, Mem. Am. Mus. Nat. Hist., I, Pt, III, April, 1808.
14. Tylosaurus dyspelor (Cope).
Am. Mus. No. 221.
Upper Cretaceous (Niobrara) of Kansas.
Tylosaurus dyspelor was one of the largest of the Mosasaurs,
great marine lizards, distantly related to the living Monitor
Lizard (Varanus), but adapted to strictly aquatic life. This
fine skeleton, found by Mr. W. G. Bourne, is twenty-eight feet
long and exceptionally perfect, even the cartilages being pre-
served. The skeleton is mounted in the original matrix and
position.
OssBorn, Mem. Am. Mus. Nat. Hist., I, Pt. 1V, October, 1899.
15. Protoceras celer Marsh.
Am. Mus. No 1236.
Upper Oligocene (White River, Protoceras Beds), S. Dakota.
Protoceras was the largest of a group of Primitive Ruminants
found in the White River Beds, which had two toes in the
hind foot, four in the fore foot. The proportions are nearly
those of the smaller deer, but with shorter limbs and neck.
The male skull bore several bony bosses comparable to the
bosses on a giraffe skull instead of true horns or antlers; the
32 Department of Vertebrate Paleontology.
female was hornless. This skeleton is that of a female, and
is mounted in high relief.
Marsu, Am. Journ. Sci., 1891, Jan., p. 81; 1893, Nov., p. 407; 18097,
Sept., p. 165. .
OsBoRN & WorTMAN, Characters of Protoceras, Bull. Am. Mus. Nat.
Hist., [V, 1892, p. 351.
Scott, Osteology and Relations of Protoceras, Jour. Morph., XI, p.
393°
16. Oxyzna lupina Cope.
Am, Mus No. 107.
Lower Eocene (Wasatch), Big Horn Valley, Wyoming.
Oxyena was a typical Creodont or Primitive Carnivore.
Large head, with small brain and long, powerful jaws, short,
bandy legs, short, subplantigrade feet, and long tail, are the
most marked primitive characters.
This skeleton was pieced together with great labor out of
two fragmentary specimens, one found by Dr. Wortman in
1881 and described by Cope in ‘ Tertiary Vertebrata’ (Speci-
men No. 2), the other found in 1891 and described by Osborn
and Wortman in the American Museum Bulletin for 1892. On
comparing these two specimens it was found that they be-
longed to the same individual, and that the skeleton was
nearly complete, though broken into innumerable fragments.
The skeleton is mounted in full relief, every bone being re-
movable.
WortTMAN, Restoration of Oxyena lupina Cope, etc., Bull. Am. Mus.
Nat. Hist., XII, 1899, p. 139.
17. Equus scotti Gidley.
17a. Equus scotti and Protorohippus venticolus.
Am. Mus. No. 10612.
Pleistocene, Rock Creek, Llano Estacado, Texas.
This skeleton is one of eight found together by J. W. Gidley
of the American Museum Expedition of 1899. It shows as
List of Casts, Models, and Photographs. 33
mounted the zebra-like proportions of this native American
wild horse, especially seen in the large head and deep jaws,
short barrel, small limbs and feet.
GipLEy, A new Species of Pleistocene Horse from the Staked Plains of
Texas, Bull. Am. Mus. Nat. Hist., XIII, 1900, pp. 111-116; and
Revision of American Species of Equus, ibid., 1901, XIV, pp. 91-
I4o.
18. Amynodon intermedius S. & O.
Am. Mus. No. 1931.
Upper Eocene (Uinta) Utah. Found by A. O. Peterson, American
Museum Expedition, 1895.
Amynodon was a collateral ancestor of the large Aquatic
Rhinoceros, Metamynodon, of the Oligocene epoch, and is
largely intermediate, both in size and in characters of the
skull and skeleton, between it and the Middle Eocene Rhi-
noceros. This skeleton is mounted in high relief.
19. Mastodon americanus (Kerr).
Pleistocene Epoch, Newburgh, New York.
Remains of Mastodon have been found in numerous locali-
ties scattered all over the Eastern States and Mississippi
Valley, and mounted skeletons are to be seen in several
American and European museums. ‘This skeleton was found
in a peat bog near Newburgh, N.Y., and is one of the largest;
size, 9 feet high, 18 feet long as mounted. The Mastodons
differed strikingly from the Mammoths in proportions, as
well as in the number and character of their teeth, form of
skull and curve of tusks.
20. Platygonus leptorhinus W7/liston.
Pleistocene Epoch, Kansas.
This skeleton is one of nine found together in Western
Kansas and described by Prof. Williston in 1894. The pro-
portions and pose of the skeleton at once suggest the living
pecearies (Dicotyles), from which Platygonus differed chiefly
in its larger size, more specialized feet and teeth,
34 Department of Vertebrate Paleontology.
21. Oreodon culbertsoni Lezdy.
Oligocene Epoch, Big Badlands, S. Dakota.
Oreodon was first described by Joseph Leidy in 1851, and is
the most abundant and characteristic fossil of the Big Bad-
lands. This species is of the size of a peccary, and the skeleton,
a composite of two very perfectly preserved individuals ob-
tained by the American Museum Expedition of 1894, shows
somewhat similar proportions, but is much more primitive in
characters, and in most respects has departed comparatively
little from the old ancestral type of the Artiodactyla.
22. Hyznodon horridus Leidy.
Am. Mus. No. 1375.
Oligocene Epoch, Big Badlands, South Dakota.
This finely preserved skeleton was found by the American
Museum Expedition of 1894, and is all one individual, the
few missing parts being restored in tinted plaster. Hyaenodon
is the best-known and the most highly developed of the
Creodonta, and is found both in Europe and America, This
species is about the size of the Tasmanian Wolf (Thylacinus),
which it resembles to a striking degree in proportions of
limbs and feet and in many characters of the skull.
23. Pantolambda bathmodon Cope.
Basal Eocene, San Juan Basin, New Mexico.
Pantolambda is the most ancient mammal of which the
entire skeleton is known. This mount is a composite of -
several incomplete skeletons obtained by the American
Museum Expedition of 1896 in the Torrejon (Upper Puerco)
horizon of Northwestern New Mexico. It exhibits the
short, crooked legs, five-toed, plantigrade feet, long, heavy
tail, arched back, primitive skull, with heavy jaws and small
brain case, and many other characters which were the common
heritage of the early mammals from their reptilian ancestors.
OsBorn, Evolution of the Amblypoda, Bull. Am. Mus. Nat. Hist., X,
1898, pp. 183-188.
List of Casts, Models, and Photographs, 35
ee eee
24. Hypohippus equinus (Scott).
Middle Miocene, Pawnee Buttes, Colorado,
This three-toed Horse is not in the direct line of descent
of the modern horses, but on a somewhat conservative side-
branch. The skeleton is as large as that of a Shetland pony
and is mounted in full relief in a walking pose. The head,
limbs, and feet are in exceptionally fine preservation. The
side view shows best the general proportions of the animal;
the view from behind, which best displays the small lateral
digits, can also be supplied. H ypoluppus is closely related to
Anchitherium of the Lower Miocene, from which it is probably
directly descended. This skeleton was found by Mr. Brown
of the American Museum Expedition of rgor.
25. Ichthyosaurus quadriscissus Quenstedt,
Jurassic Period, Holzmaden, Wurttemberg.
This fine skeleton was presented to the American Museum
by the Royal Natural History Museum of Stuttgart, through
Prof. Eberhardt Fraas. It is of especial interest as showing
that the Ichthyosaurs were viviparous instead of egg-laying
reptiles. The skeletons of seven young (unborn) Ichthyosaurs
can be seen, partly in, partly washed out of the body-cavity.
26. Portheus molossus Cope.
Upper Cretaceous (Niobrara) of Kansas. Found by C. H. Sternberg.
This great fish is characteristic of the marine Upper Creta-
-ceous chalk formation of Western Kansas. It was related
to the modern Tarpon of Florida but of gigantic size, this
skeleton being 18 feet in length. The head and tail are
especially fine; the ribs and fins are mostly restored in plas-
ter. It is much flattened and is mounted in low relief.
27. Hippopotamus liberiensis orton.
The Pigmy Hippopotamus of the West Coast of Africa is a
rare species, almost extinct. This skeleton is semi-fossil, and
J . . .
was presented by the Museum of Christiania, Norway.
36 Department of Vertebrate Paleontology.
28. Ornitholestes hermanni Osborn.
Upper Jurassic (Como), Wyoming.
This beautiful little skeleton, about seven feet in length,
represents a little-known group of Carnivorous Dinosaurs
adapted for swift running and the seizing of a light and agile
prey. The tail is extremely long, slender, and whip-like, the
hind limbs long and the feet like those of a bird, while the
small fore limbs are modified into remarkable prehensile
organs, the first and second digits greatly elongated and
opposed, with large curved claws, the third digit small and
slender, and the fourth atrophied.
Ossorn, Bull. Am. Mus. Nat. Hist., XIX, 1903, pp. 459-464.
29. Mesohippus bairdi Lezdy.
Middle Oligocene (White River), South Dakota.
This classic species represents an early stage in the evolu-
tion of the Horse. It is smaller than the modern Dorcas
Gazelle, and has three toes on each foot, the lateral toes
slender but reaching to the ground. Vestiges of the first and
fifth digits are still preserved in the fore foot. Mesohtppus is
somewhat peculiar in the unusual length of the hind limbs as
compared with the fore; in other respects it is intermediate
between the four-toed horses of the Eocene and the modern
horse.
Scort, W. B., On the Osteology of Mesohippus and Leptomeryx, Jour.
Morph., V, 1891, pp. 301-406.
Farr, Notes on the Osteology of the White River Horses, Proc. Amer.
Phil. Soc., XX XV, 1896, p. 147—175-
30. Merycodus osborni Matthew.
Middle Miocene, Pawnee Creek Bed, Colorado.
Merycodus is a collateral ancestor of the modern Pronghorn
Antelope of Western North America, but is distinguished by
large, branching, deciduous antlers like those of the Deer.
List of Casts, Models, and Photographs. oy
aw ee ee
The bony rudiments of lateral toes, present in nearly all deer
but absent in all antelopes, are still preserved in Merycodus,
but exceedingly minute, much smaller than in the deer. The
animal was of about the size of the Dorcas Gazelle.
MattueEw, W. D., A Complete Mounted Skeleton of Merycodus, Bull.
Am. Mus. Nat. Hist., XX, 1904, pp. —.
See also Scort, W. B., Bull. Mus. Comp. Zoél., XX, 1890, p. 82 (Cosoryx
furcatus).
ITV.—PHOTOGRAPHS OF RESTORATIONS, BY CHARLES KNIGHT.
Presented by $. Pierpont Morgan, and exhibited in the Depart-
ment of Vertebrate Paleontology. Bromide enlargements
jrom the original negatives. Size, 18 in. % 22 in.
These photographs are from a series of large water colors
executed for the American Museum by the animal painter,
Mr. Charles Knight, under direction of Prof. Osborn, with
suggestions and criticisms by other paleontologists, the object
being to increase the popular interest in these extinct animals,
and to give a fuller and truer idea of their anatomy and
external form than is afforded by the skeleton.
The artist begins each study by preparing a number of
models in wax, based upon the actual proportions and muscu-
lar indications of the skeleton, and by a series of preliminary
- sketches, representing different attitudes, habits, and sur-
roundings. Thus the position of all the joints and angles of
the feet and limbs is true to life. The lips, nostrils, and gape
of the mouth are determined by comparison of the length of
the nasals, size of the anterior nares, character and position
of the teeth, with similar parts in the remotely related living
forms. The eyes are carefully located and proportioned. Upto
this point the animal is a fairly correct representation of the
original. On the other hand, the shape of the ears, the color
and epidermic characters of hair and hide are largely imagina-
tive, except in so far as they are suggested by relationship
28 Department of Vertebrate Paleontology.
to modern allies, as of Protorohippus to the Horse, or of Ceno-
pus, Metamynodon, and Hyracodon to the Rhinoceros.
These restorations are copyrighted and the photographs are
sold with the understanding that they are to be used only for
exhibition purposes, and are not to be copied or adapted for
publication.
Price of the photographs, $4 each.
1. Patriofelis, Middle Eocene Creodont.
This animal is based upon studies of the skeleton mounted
in the American Museum Collection (No. 1507). The follow-
ing quotation from an article by Dr. Wortman explains the
picture: ‘‘The broad, flat, plantigrade feet, with their spread-
ing toes, suggest at the first glance their use for swimming.
The eversion of the feet, together with the general clumsiness
of the limbs point, moreover, to the fact that the animal was
not an active runner. . . . He was perhaps not as expert
a swimmer as the seals are now, but was sufficiently active
in the water to capture turtles.” This is perhaps the least
original and successful of the restorations, being modelled
somewhat too closely upon the Otter. More recent studies by
Osborn have cast much doubt upon the aquatic habits of the
animal as indicated by Wortman.
Gavinally reproduced by Osborn in ‘ Prehistoric Quadrupeds of the Rockies,’ Century
Magazine, September, 1806.
WortMan, The Osteology of Patriofelis, Bull. Am. Mus. Nat. Hist.,
VI, 1894, pp. 119-64, pl. i.
OsBorRN, Oxyena and Patriofelis Re-studied as Terrestrial Creodonts,
Bull. Am. Mus. Nat. Hist. XII, 1900, pp. 269-279.
2. Protorohippus venticolus, Four-toed Lower Eocene Horse.
The studies for this animal were based upon skeleton No.
4832, mounted in the American Museum Collection. The very
primitive characters of this early stage in the evolution of the
Horse are especially seen in the short neck and legs, the
heavily muscular part of the limb extending much further
down in proportion to the lower leg and foot. The arched
liad.
List of Casts, Models, and Photographs. 39
back and short head are likewise striking characters. The
striping of the neck and fore quarters is based upon the fact
that the young of all modern species of Equide show a more
Fig. 8. Restoration of the Four-toed Eocene Horse Protorohifjus. Height at withers
sixteen inches.
or less marked tendency to striping in these parts. The ani-
_mal in life was about 4 hands, or 16 inches, in height at the
withers.
Originally reproduced by Lucas in McClure’s Magazine. 1890.
Cope, Tertiary Vertebrata, U. S. Geol. & Geog. Sur. Terrs., F. V. Hay-
den in charge, Final Report, IV (1885), pp. 635-647, pll. xlix,
GD iG.
3. Cznopus, A Hornless Upper Oligocene Rhinoceros.
The study is based upon the complete mounted skeleton,
No. 538, in the American Museum, of Cenopus tridactylus.
This animal compares in its dimensions and in the general
characters of its skeleton with the Two-horned Rhinoceros of
4O Department of Vertebrate Paleontology.
Sumatra; and Mr. Knight’s studies for this picture were
practically based upon that living type.
OsBorn & WortTMAN, Fossil Mammals of the Lower Miocene White
River Beds, Bull. Am. Mus. Nat. Hist., VI, 1894, p. 206, pl. iit.
4. Protoceras, Six-horned Upper Oligocene Ruminant.
The characters of this animal are known from complete
female skeletons in the American Museum of Natural History
and the Princeton Museum, and a very complete series of
male and female skulls, secured by the American Museum
Expeditions of 1892 and 1894. As exhibited, there is a wide
contrast between the male and female skulls, the latter being
entirely devoid of horns. The very marked recession of the
nasals has suggested the presence of a long, somewhat ex-
panded upper lip, as in the modern Saiga, while the coloring
of the animal is based upon that of the North American
Pronghorn Antelope.
Originally reproduced by Osborn in the Century Magazine, September, 1896.
Osporn & WorTMAN, Characters of Protoceras Marsh, Bull. Am. Mus.
Nat. Hist., IV, 1802, pp: 351537 5-
5. Metamynodon, Aquatic Middle Oligocene Rhinoceros.
The study is based upon the mounted skeleton, No. 546, in
the American Museum, and represents the general conception
of the animal as aquatic. The opinion as to the aquatic .
habits of Metamynodon was very largely due to the studies
made for this drawing in which it became apparent that the
very elevated position of the orbits, the widely expanding
ribs, and the four complete spreading toes in the fore foot,
strongly suggested the general build of the Hippopotamus.
This aquatic life is, moreover, consistent with the presence in
the same beds of two other types of Rhinoceros, it being
improbable that the three types were all land animals.
Originally reproduced by Osborn in the Century Magazine, September, 1896.
OsBorn & Worrman, Perissodactyls of the Lower Miocene White
River Beds, Bull. Am. Mus. Nat. Hist., VII, 1895, pp. 373-375
joullls, Seng ssc
——e_
List of Casts, Models, and Photographs. 41
6. Elotherium, Giant Upper Oligocene Suilline.
These animals have been depicted from very complete re-
mains in the American and Princeton Museums, and represent
the largest type of Oligocene Elothere, namely, the E. ramosum
of Cope, or E. tmperator of Marsh. The great flanges below
the cheeks for the attachment of the masseter muscles pre-
sented peculiar difficulties, and it is possible that they are
not here exhibited with sufficient expansion. The head is of
enormous size, but the chest is small, and the limbs are com-
paratively tall and stilted, as here represented. The very
remote relationship to the Pig and to the Hippopotamus is
suggested in the bristles and in the form of the lips. Elo-
thertum is in a general way a sort of dry-land hippopotamus,
as Metomynodon is an aquatic rhinoceros; in the one case the
dry-land type, in the other, the aquatic type, has become
extinct.
Originally reproduced by Osborn in the Century Magazine, September, 1896.
Marsu, Am. Jour. Sci., XLVII, 1894, p. 408, pl. ix.
7. Megacerops, Long-horned Lower Oligocene Titanothere.
This group of male, female, and young is based upon skulls
belonging to different sexes and ages, in the American Museum
Collection, as well as upon the complete skeleton, No. 518,
mounted in the American Museum. There is no doubt that
“the females had smaller skulls, with imperfectly developed
horns and narrow zygomatic arches, while the males of the
same species had extremely long, recurved horns. The ex-
tension of the premaxilla and the overhanging of the nares
by the rudimentary nasals indicate that there was not space
for a proboscis, but rather a prehensile upper lip, which sug-
gests the same structure in the Rhinoceroses.
Originally reproduced by Osborn in the Century Magazine, September, 1896.
OsBorn & WortMan, Perissodactyls of the Lower Miocene White
River Beds, Bull. Am. Mus. Nat. Hist., VII, 1895, pp. 346-352,
pill. viii, ix.
42 Department of Vertebrate Paleontology.
8. Uintatherium, Six-horned Middle Eocene Amblypod.
The male and female here represented are based upon
skeletal material as yet unmounted in the American Mu-
seum Collection, also upon the reconstruction published by
Professor O. C. Marsh. The structure of the male skull, as
here represented, corresponds with the largest type ever
found, that of the Uintathertum cornutum skull, discovered by
Professor Cope himself, and now in the American Museum.
There is no indication of a proboscis, nor is there any reason
to believe that the bony protuberances bore horns, many of
them expanding at the tip, as is never the case with true
horn-cores. The tusks, both in male and female, are found
chiefly worn upon the posterior surface, a fact which suggests
that they were used upon branches in drawing leaves and
shrubs into the mouth, as represented in the female figure in
the rear. Their chief function was probably as weapons of
defence or attack, the exceptionally wide gape of the jaw, as
indicated by the position and form of its muscular attach-
ments, giving room for their use in a very effective manner.
Originally reproduced by Osborn in the Century Magazine, September, 1896.
Cope, Tertiary Vertebrata, pp. 569-587, pll. xxxvii—xlii.
MarsH, The Dinocerata, U. S. Geol. Sur. Mon., X, pp. 1-243, pll. i-lvi,
1886.
9. Hyracodon, Cursorial Oligocene Rhinoceros.
This restoration is based also upon material in the American
Museum and Princeton Collections. The general conception of
Hyracodon as a running type was outlined by Scott in 1883.
These animals thus presented the widest contrast to the
swimming Rhinoceroses, or Metamynodon, on the one side,
and the low-land or swamp Rhinoceroses, the Aceratheres, on
the other. The upper molar teeth, however, closely resemble
those of the Rhinoceros, and form the motive for the re-
semblance to this animal, which is given in the face. The
trunk and limbs, on the other hand, resemble very closely
List of Casts, Models, and Photographs. 43
So a ee
those of the primitive horses. Hence the motion character-
istic of the galloping horse is given to the skeleton.
Originally reproduced by Osborn in the Century Magazine, September, 1896.
Scott, Osteologie von Hyracodon Leidy, Festsch. f. Gegenbaur, 1896,
PP. 353-383, pll. i-tii.
10. Mesonyx, Omnivorous Middle Eocene Credont.
This huge animal is represented preying upon the skull of an
Uintathere, in order to give some conception of its size. The
skull, as represented in the American Museum Collection, No.
1892, is extremely large and armed with very blunt teeth,
wearing down in old age, indicating that the animal was omniv-
orous, or lived partly upon turtles or decaying animal food.
The form of this body is derived from a complete skeleton in
the Princeton Museum, which has been figured by Scott. It
slopes backwards, the posterior quarters being rather small;
the tail is extremely long and powerful, the general propor-
tions resembling somewhat those of the Tasmanian Wolf.
Originally reproduced by Osborn in the Century Magazine, September, 1896.
Scott, New and Little-known Creodonts, Jour. Acad. Nat. Sci. Phila.,
IX, 1887, pp. 155-185, pll. v—vii.
Wortan, Studies of Eocene Mammalia in the Marsh Collection, Part
I, Carnivora, Am. Journ. Sci., XII, 1901, pp. 285, 377, 421; SANG
1902, Pp. 39.
11. Palaosyops, Eocene Hornless Ancestral Titanothere.
This animal is based upon the complete skeleton mounted in
‘the American Museum, and is represented as having somewhat
the habits of the Tapir, living in low, marshy land, and feeding
entirely upon the softer kinds of leaves and grasses, since its
teeth are entirely unadapted to hard grasses or the more
silicious plants. According to the studies of Earle, the ani-
mal was devoid of a proboscis, but had an elongated, pre-
hensile upper lip. The slender fore feet are very similar in
proportions to those of the aquatic Rhinoceroses.
Originally loaned for reproduction in Harper’s Magazine, 1897.
Ear.e, On the genus Palgosyops, etc., Jour. Acad. Nat. Sci. Phila.,
IX., 1892, pp. 267-388, pll. x—xiv, and restoration, p. 314.
44 Department of Vertebrate Paleontology.
12. Mastodon americanus, North American Pleistocene
Elephant.
This animal is executed from the skeleton in the American
Museum of Natural History, and it is characterized by being
partly hairy, in contrast with the extremely hairy northern
variety of the Mammoth, and in view of the probability that
all the more primitive elephants were coated with hair. It
has distinctly the low, flat skull of the African Elephant,
whereas the Mammoth had the high, prominent forehead
characteristic of the Indian Elephant. The skeleton is also
distinguished by enormously large, projecting feet, larger than
in any existing species, and by the relative shortness of the
limbs, also a primitive feature. This restoration, therefore,
while based upon studies of the elephant, exhibits the charac-
teristic proportions which distinguished the Mastodon ex-
ternally.
Originally executed for reproduction in Harper’s Magazine, 1897.
13. Dryptosaurus (Lelaps), Carnivorous Cretaceous Dinosaur.
In this and the four following restorations Mr. Knight was
guided largely by ideas and sketches given him by Prof. Cope
shortly before the latter’s death. They are based upon speci-
mens in the Cope Collection now in the American Museum, and
upon restorations by Professor Marsh of related types.
This Dinosaur, represented in the photograph as engaged in
combat, was of considerable size, about fifteen to twenty feet
in length, and well armed in teeth and claws. Prof. Cope con-
sidered that the long hind limbs and heavy tail indicated great
leaping powers, in spite of its large size, and the restoration is
made in accordance with this idea. Dryptosaurus was allied
to Ceratosaurus of Marsh and to the European Megalosaurus.
Cope, Proc. Acad. Nat. Sci. Phila,, 1866, 275; Proc. Am. Phill Soc,
XXX, p. 240, May, 1892.
Marsu, Dinosaurs of North America, pp. 157-163, pl. xiv.
14. Elasmosaurus, Long-necked Cretaceous Plesiosaur.
Elasmosaurus was one of the largest of the marine reptiles
which inhabited the Cretaceous seas. It was related to the
——e ee
List of Casts, Models, and Photographs. 45
ee a eee
European Plesiosaurus, which it resembled in shape, except in
the flattened tail. It is represented in the restoration as cap-
turing a young Portheus, one of the large, bony fishes of the
period. A skeleton in the Museum of the Philadelphia
Academy of Natural Sciences, and another more complete in
the Cope Collection (but both lacking the skull), served as the
basis for the restoration. The Plesiosaurs differed greatly in
the length and flexibility of the neck, as well as in size.
Cope, Cretaceous Vertebrata, U. S. Geol. & Geog. Surv. Letts:, 2 We
Hayden in charge, Final Report, II, pp. 75-88.
Dames, Abh. k. preuss. Akad. Wiss. Berlin, 1895, pp. 75-80; Natural
Science, Jan., 1898, p. 48.
15. Agathaumas, Three-horned Cretaceous Dinosaur.
These herbivorous Dinosaurs were of great size and had
large heads armed with three horns, and a great projecting
crest or frill protecting the neck. The body may have been
covered with small bony plates (scutes). This restoration is
based on a reconstruction of the skeleton of Triceratops
prorsus by Prof. Marsh. Agathawmas sphenocerus Cope,
shown in the picture, is distinguished by the large anterior
median horn and small posterior paired horns.
Cope, Amer. Nat., 1886, p. 716; 1892, p. 768.
Marsh, Dinosaurs of North America, Ann. Rep. U.S. Geol. Surv., 1896.
16. Naosaurus and Dimetrodon, Spine-backed Permian
Lizards.
These ancient reptiles represent an early specialized branch
of the primitive Rhynchocephalians. Their most remarkable
character is in the enormously elongated dorsal spines of the
vertebre, with (in Naosaurus) or without (in Dimetrodon)
transverse bars of bone. The restoration is based on a num-
ber of incomplete skeletons in the Cope Collection.
Cops, Proc. Amer. Phil. Soc., 1878, p. 512. .
Baur & Case, Morphology of the Skull of the Pelycosauria, Anatom.
Anzeig., Jena, 1897, XIII, p. 109.
46 Department of Vertebrate Pal«ontology.
17. Hadrosaurus, Duck-billed Cretaceous Dinosaur.
Drawn from the unusually complete skeleton of Hadro-
saurus (Diclonius) mirabilis Leidy, in the Cope Collection now
in the American Museum. This very specialized genus is
found in the Upper Cretaceous of New Jersey and Wyoming.
It was herbivorous, and probably amphibious, with long neck
and heavy hind quarters. It had a broad, duck-like, horny
bill, and back of it a magazine of numerous small, rod-like
teeth, not less than a thousand in each jaw, set on end in
several close-set rows, and wearing to a tesselated-pavement-
like grinding surface. The length was about thirty feet.
The skeleton of a nearly related genus, Claosaurus, is
mounted in the Yale Museum.
Cops, Proc. Acad. Nat. Sci. Phila., 1883, p. 97; Amer. Nat., 1883, p.
774-
18. Siberian Mammoth or Hairy Elephant.
Unlike the living elephants, the Mammoth ranged into arctic
countries, and is here restored with its appropriate environ- _
ment, taken from the region of the Taku Glacier, Alaska. It
was a contemporary of primitive man, and its enormous size is
therefore fitly indicated by the contrast with human figures
in the background. In this instance the color and texture of
the hide is certainly known, from the mammoth carcasses
which have been found frozen in the palgwocrystic ice of
northern Siberia, and parts of which are still preserved in the
St. Petersburg Museum.
19. Cervalces, Pleistocene American Elk.
Cervalces was as large as the Moose, but is distinguished by
its magnificent antlers, spreading in three directions, outward,
upward. and forward, and attaining a size and complexity
unequalled by any living species. The fine skeleton in the
Princeton Museum served as a basis for this restoration, the
superficial characters being studied from the Moose.
Scott, Proc. Acad. Nat. Sci. Phila., 1885, p. IgI.
List of Casts, Models, and Photographs, 47
20. Condylarth or Primitive Hoofed Mammal Phenacodus.
Lower Eocene Epoch.
This is drawn after the famous skeleton discovered by Wort-
man and described originally by Cope, and as remounted by
Osborn.
Phenacodus belongs to the group of Primitive Ungulates
from which all the modern hoofed animals are descended. It
is in many respects like the clawed animals (Unguiculates)
from which the hoofed animals (Ungulates) branched off.
Five toes on each foot, pig-like teeth, arched back, short legs,
narrow chest, lank sides, long tail, all are characters of the
primitive clawed animals still retained by Phenacodus, but lost
by modern hoofed animals.
OsBorn, Remounted Skeleton of Phenacodus primevus, Bull. Am. Mus.
Nat. Hist., X, 1898, p. 159.
Cops, Tertiary Vertebrata, pp. 428-463, pl. lvii b-Iviii.
21. Hornless Amblypod Coryphodon.
A short-footed, hoofed animal of the Lower Eocene Period.
This remarkable animal was related to the huge Uintatheres.
It had five very short toes on each foot. The anterior teeth
suggest those of the Hippopotamus, but the grinding teeth are
entirely different. The neck is short, the body rather long,
with many archaic characters. The legs are short and clumsy,
the upper joint disproportionately long. The brain is re-
markably small. At the sides of the top of the skull are bony
projections prophetic of the posterior horns of the Uintathere.
Coryphodon was probably an amphibious animal living in
the marshes and rivers bordering the ancient Wasatch Lake.
Its large tusks were used presumably in rooting up water-
plants, and were also formidable weapons of defense against
the larger flesh-eating animals of that period. Its general ap-
pearance was unlike that of, any modern animal, for, although
it combined some characters of such dissimilar beasts as the
Bear, Elephant, and Hippopotamus, it had many peculiari-
ties of its own.
OsBorn, Bull. Am. Mus. Nat. Hist., X, 1898, pp. 81-91.
g
48 Department of Vertebrate Paleontology.
22. Primitive Sabre-tooth Tiger Hoplophoneus.
Oligocene Epoch.
The most striking difference between Sabre-tooth Tigers and
the great living cats is in the long, flattened sabre-like upper
canine teeth, which in Smilodon (Restoration No. 31) were
seven inches long. These teeth could pierce the hides of
rhinoceroses and other thick-skinned animals common in
America in the Oligocene Period, against which the shorter
fangs of modern lions would be ineffective. The legs were
shorter and more muscular than those of the larger modern
cats, the animal more powerful, but by no means as swift-
footed.
Scorr & OsBorn, Bull. Harv. Mus. Comp. Zodl., 1887, p. 153, pl. i.
Riccs, Restoration of Hoplophoneus occidentalis, Kans. Univ. Quar.,
V, 1896, pp. 37-52, pl. 1.
23. Short-legged American Rhinoceros Teleoceras.
Upper Miocene Epoch.
Teleoceras, the last known survivor of the Rhinoceros race
in America, was also the largest; and its fossil remains are so
abundant in certain localities as to indicate that it lived in
great herds upon the plains, like the Bison in more modern
times.
The body was as long and heavy as that of the living Indian
Rhinoceros, but the legs were so short that the belly nearly
reached the ground, giving the animal the squat proportions of
the Hippopotamus. The male had a small horn on the end
of the nose; the female was hornless.
OsBorn, Complete Skeleton of Teleoceras fossiger, Bull. Am. Mus. Nat.
Hist., X, 1898, pp. 51-59, pll. iv, iva.
Wituiston, Restoration of A phelops jossiger, Kans. Univ. Quar., 1894,
p. 280, pl. viii.
Scott & Ossorn, Bull. Harv. Mus. Comp. Zoél., pp. 92-99, pl. it.
24. Amphibious Dinosaur Brontosaurus.
Upper Jurassic Period.
In the Reptilian Age, preceding the Age of Mammals, great
reptiles were dominant on land and water. The Dinosaurs, or
List of Casts, Models, and Photographs. 49
Giant Reptiles, were the chief inhabitants of the lowlands and
marshes. Preéminent in size among these were the Sauropoda,
or Amphibious Dinosaurs, a herbivorous group, mostly of
gigantic size, from forty to seventy-five feet in length,—the
greatest of land animals living or extinct, and exceeded in
size only by the modern Whales.
The long neck and small head enabled them to lie con-
cealed in marshy bayous and lagoons, the body generally im-
mersed, the head foraging for food without easily attracting
the attention of the great carnivorous reptiles which lived at
this time. The long and massive tail was useful both as a
support and a propeller.
Brontosaurus was among the largest of the Sauropoda,
seventy feet in length and about eighteen feet in height to the
arch of the back. The thigh-bone is six feet long, and weighs
in its petrified condition 500 to 600 pounds.
Marsu, Dinosaurs of North America, Sixteenth Ann. Rep. U. S. Geol.
Surv., 1896, pp. 166-175, pl. xlii (Brontosaurus excelsus).
OssBorn, Bull. Am. Mus. Nat. Hist., X, 1808, p. 210.
25. Great Marine Lizard or Mosasaur Tylosaurus.
Upper Cretaceous Period.
This restoration is made from the complete skeleton, thirty
feet long, in the American Museum.
The Mosasaurs were carnivorous Sea-Reptiles abundant in
the warm, shallow seas of the Upper Cretaceous Period, but not
yet found earlier or later. Large flat head, short neck, four
paddle-limbs like the flippers of whales, vertically flattened,
swimming tail. Length of different species, from twelve to
forty feet. Their nearest living ally is the Monitor Lizard
(Varanus) of the Nile Valley.
Ossorn, A Complete Mosasaur Skeleton, Osseous and Cartilaginous,
Memoirs Am. Mus. Nat. Hist., I, Pt. IV, Oct., 18909, pll. xxi—xxiii
and text illustrations. oom
Witutston, Kansas Univ. Quar., VIII, 1899, pp. 39-41; Univ. Geol.
5
Surv. Kansas, IV, Pt. V, pp. 83-221, pll. x-Lxxii.
50 Department of Vertebrate Paleontology.
26. Great Irish Deer Megaceros.
Pleistocene Epoch. Europe.
Drawn from the skeleton mounted in the American Museum.
Megaceros exceeded any living deer in the spread of its
. antlers, in some cases ten or even eleven feet from tip to tip.
In size it is about equalled by the Moose, but the proportions
were somewhat different, approaching those of the Fallow
Deer (Cervus dama) to which it was more nearly related. The
most abundant and complete remains have been found in the
bogs of Ireland, but the animal ranged all over Western
Europe. It is not found in America.
27. Primitive Mastodon Trilophodon.
Miocene Epoch,
Based upon a skull and incomplete skeleton in the American
Museum, and upon the restored skeleton of T. angustidens by
Prof. Gaudry. These Miocene ancestors of the Mastodons and
Elephants were much less specialized than their later de-
scendants. ‘This is especially seen in the small tusks in both
upper and lower jaws, the upper ones curving down, the
lower ones straight, and both with an external or anterior
enamel band like the incisors of rodents,—in the shorter
trunk, as indicated by the characters of the skull, in the limbs
of moderate length and toes much less reduced than in the
Pliocene and Pleistocene proboscidians.
28. Great Carnivorous Dinosaur Allosaurus.
Upper Jurassic Period.
This great carnivorous reptile was a contemporary of the
huge Sauropoda. That it preyed on their carcasses is certain,
for the bones are often found fossil with scorings and scratches
on their softer surfaces which might well have been made by
this animal, and its broken-off teeth, still more frequently
found close by, suggest that it was more greedy than prudent
in its feasts. But the Allosaurus was likewise well adapted in
List of Casts, Models, and Photographs, 5!
ee a ee ae y
teeth and claws to attack a living prey, and it is quite probable
that this was the enemy from whose attacks the huge Bronto-
Saurus sought refuge by habitually burying his great unwieldy
body beneath the water, as indicated by the skeleton structure
and illustrated in our restoration (No. 24) of that beast.
In the Allosaurus restoration Mr. Knight has represented
him as preying upon the carcass of a Brontosaur, using the
powerful bird-like hind feet in somewhat the same manner as
does a vulture or other bird of prey.
29. Amphibious Dinosaur Diplodocus.
(In preparation.)
30. Ichthyosaur, or Great Fish-Lizard.
Jurassic Period.
The discoveries of Ichthyosaurs, with the outlines of body
and tail preserved, in the great slate-quarries of Holzmaden in
Wurttemberg, as described by Professor Fraas in 1892, have
considerably altered our conceptions of this classic fossil
reptile. We now know that it had a high, triangular back fin
and broad, forked tail, like that of the shark, except that the
vertebral column ran into the lower wing instead of the upper
wing of the tail, The Holzmaden specimens, worked out with
wonderful skill and care by Herr Hauff, give the outlines of
almost all parts of the body, so that there is little (except as
to color) left to conjecture or inference in this restoration.
Mr. Knight has associated with the old Ichthyosaur a little
‘school of new-born young whose proportions and size are
taken from the seven little Ichthyosaur skeletons contained
within the. body-cavity of the large skeleton in the American
Museum.
31. Great Sabre-tooth Tiger Smilodon.
Pleistocene Epoch.
Based on the complete skeleton in the Cope Collection,
found in the Pampean formation near Buenos Aires, and now
mounted in the American Museum.
Smilodon may well be considered the most terrible of all
52 Department of Vertebrate Palaeontology.
beasts of prey. It equalled in size the largest Polar Bear, and
was probably of the most savage and ferocious disposition,
fearing nothing, and accustomed to prey on the largest and
best-defended of the great herbivora. It did not indeed equal
the modern large cats in activity, but it far surpassed any of
them in strength, especially in the fore limbs and neck. The
extraordinarily powerful attachments for the sterno-mastoid
and cleido-mastoid muscles indicate the tremendous force
with which the great seven-inch upper canines could be
driven into the flesh of the prey, while the extremely wide
gape of the jaw, indicated by the form and arrangement of
its muscular attachments, gave free play for these powerful
weapon-teeth.
The relatively small and low hind quarters and short, small
tail give the Smilodon an appearance quite unlike the modern
great cats, and the characters of skull and neck vertebre indi-
cate that it carried its head low, and was little able to raise it.
In PREPARATION.
32. ‘Oxyena lupina.
33. Platygonus leptorhinus.
34. Oreodon culbertsoni.
35. Poébrotherium wilsoni.
36. Hyenodon horridus.
37. Cynodictis gregarius.
38. Dinictis squalidens.
39. Pantolambda bathmodon.
40. Hypohippus equinus.
41. Elephas imperator.
42. Ornitholestes hermanni.
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