.s

3/+3

New Series
V. 16
1890

T R A xN S A C T I 0 N S

OF TIIK

AMERICAN PHILOSOPHICAL SOCIETY,

HELD AT PHILADELPHIA,

FOR PROMOTING USEFUL KNOWLEDGE.

VOL. XVI.-NEW SERIES.

PUB LIS HE I) BY TUE SOCIETY.

V li i I a rt c 1 1) h I ii :

MacCalla <& Company, Printers,

1890.

COHSTTEItTTS OI!^ VOX-.. 2^VI.

ARTICLE I.

PAO It

A Dictionary ot the Egyptian Language. By EJward Y. McCauley, U. S. N I

'F>J^:EIT XX.

ARTICLE II.

On llie Intercentrum of the Terrestrial Vertebnita. By E. D. Cope. (With 6 cuts and a plale.) . 24;}

ARTICLE III.
A Chemical Study of Yucca Angustifolia. By H. C. De S. Al)l)ott 2!H

ARTICLE IV.

Systematic Catalogue of the Species of Vertebrata found in the Beds of the Permian Epoch in Norlli Amer-
ica. With Notes and Descriptions. By E. D. Cope. (With a cut and two plates. ) 285

ARTICLE V.
Synopsis of the Vertebrate Fauna of the Puerco Series. By E. D. Cope. (With 13 cuts and 2 plates ) 298

ARTICLE VI.
On the Shoulder-Girdle and Extremities of Eryops. By E. D. Cope. (Willi a plate.) 802

F j^:rt III-

ARTICLE VIL

The Cretaceous and Tertiary Geology of llie Sergipe-Alagdus Ba.sin of Brazil. By John C. Branncr.

(With 10 cut.i and .T plates ) :!mi

ARTICLE VIII.

Descriptions of New Species of Fossils from ihe Clinton, Lower Ilelderberg, Chemimg, and Wnverly Oroups,
found in the Collections of the Geological Survey of Pennsylvania. By George B. SimpHon.
(With 31 cuts.) ir.

ARTICLE IX.

Tlie Mammalia of the linta Formation. By William B. Scott and Flenry Fnirfleld Osborn. (With n ruin

and 5 plates.) . .... Ill

TRANSACTIONS

OF TllK

AMERICAN rillLOSOrillCAL SOCIETY.

ARTICLK I.
A DICTIONARY OF THE EGVPTIAX LANGUAGE.

BY EDWARD V. MoCAULBY, U. S. N.
Rca<l 4><'(<>l><-r 30. iSH-i.

PREFACE.

In 1880, I made a manuscript copy of Dr. Birch's Egyjjlian Ditlioiiaiy {\o\.
V, of Bunsen's Egypt). As it was inadequate for the present requirement I added
to it a list of words compiled fiom translations, lately made, of papyrus texts and
monumental inscriptions, witli the aid of Chabas' Melanges Egyj)tul<)giques, and tlic
works of Grebaut, Deveria, Goodwin, &c. Finally, I closely compared the work
with Pierret's Yocabulairc, the latest issue of the kind, resulting in the Dictionary
now laid before the Society.

1 claim for it that it contains all the words that could be ol)tained frmn the
sources I have just mentioned, and i)robal)ly all that have been delined by Egyj)-
tologists uji to the present time.

These words, and their variants, arc placed under their proper initial syml)ols
or characters, which, being carefully indexed, even the uninitiated may work dui
the meaning of a liieroglyphical text.

I have not placed any geographical m- liieological names in I lie book.
Oui' constantly increasing information on the geogi'ai>h3' and rcliL,^ion of ancient
Egypt, necessitating constant correction, I tliought it lietu-r to restrict myself
to the compilation of a book that would be of use in ti-anslating ordin;iry lin-
gual text.

E. Y. M. ('Ari.i:^ .

PiiiLADELi'iiiA. December, 1882.

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ARTICLE n.

ON THE INTEUCEXTlll'M OF THK TEUUESTKIAL VERTK13KATA.

UY E. D. COPE.
Read before the American Philosophical Society, January i, 1886.

Since the discovery of the rhachitomous Batrachiu, two difteroiit views of the
homologies of the segments which compose the l)odies of thrii- vertehne have Ijceii
maintained. According to the one first proposed, that of the writer,'" the segments in
the accompanying cuts of Trimerorhachis ins/'gnis Cope, and Eryops vmiacephalus Copi',
marked />, represent the centrum proper; while those marked /, are pieces which are

ik^i^W^ {J

Fig. 1. — Trimerorhnehh iriKif/nui ; a, occiput from behind ; 6, angle of the niaiidible, side ; r, do. from lieliind ;
(I, pnrt of vertebrnl column, lateral view, neural arclics resting on inlercentra through a loss of chorda dorsalis and
pressure ; e, do. obliquely from above ; », intcrcentra ; p, pleuroccntra. Original ; from Americau NaluralisI, 1884, p.
33. From the Permian of Texas. Natural size.

intercalated hetween the true vertebra), and arc, therefore, appropriately termed {7iter-
centra. According to the second interpretation, that of Profi's.sor Gaudry,t the three
elements J5^^ and i, together, constitute a vertebra. It follows that /' cannol then be

•American Naturalist, 1878, p. 337. Proceedings Amer. Philosoph. Society, 1878, p. ."ilO, 51.S, 532. Natumllsi,
1878, p. 633 ; 1886, 7.5.

t Bulletin de la SociStu Geologitpie, France, 1878, p. 03; Enchaincmcnts du Monde Animal, 1883, p. 371.

A. P. S. — VOL. XVI. 2e.

244

ON THE IXTERCEKTRUM OF THE

properly termed an intcrcentrnm, so it is named by Gaudry the hypocentrnm. The
question as to which of these interpretations is correct has an important bearing on
the homologies of the corresponding parts in other batrachians and in reptiles ; and

Fir.. 2. — EryopK mer/acephalui ; vertebral column from the left side, one-fdurth natural size. Original ; from
Proceedings American Philosnpli. Society, 1881.

the phylogenies of these classes cannot be determined until the question is settled.
The following pages are devoted to this su])ject.

I. — THE BATRACHIAN IXTERCENTRUM.

That the intercentrum exists is shown by the very frequent occurrence in the
Pelycosaurian reptiles of the Permian epoch, of a wedge-shaped bone between the
vertebral centra on their inferioi- side* (Plate I, Fig. 9). Apparently homologous
elements occur in the dorsal and cervical regions of Sphenodon,t and in the cervical
regions of various other lizards. Similar pieces are found in the dorsal and caudal
regions of various Mammalia, for instance, Erinaceus.J But in general they are
Avanting from Mammalia, and are better developed in the Pelycosauria than in any
other order of reptiles.

In the Pelycosauria (Clepsydrops, Dimetrodon), the intercentra of the caudal region
are continuous with, or form a wedge-shaped common head, of the chevron bones
(Plate I, Fig. 9). In the rhachitomous Eryops, in the caudal part of the column,
the i)ieces which correspond with the intercentra of the Pelycosauria in forming the
expanded heads of the chevron bones, are those which I have termed intercentra, and
which Professor Gaudry has called hypocentra (Plate I, Fig. 1). Here, as in the
dorsal region (Cut 2), the intercentra present their lateral angles upwards towards
the neural arch. The nciu-al arch rests exclusively on the pleuroeentrum, which in
turn adheres to the intercentrum behind it by its long side, and to that in front of it
by its short side or end. The diapophysis belongs exclusively to the basal part of the
neural arch in' the doisal and cervical regions, and its extremity forms a vertical

*Cope, Proceeds. Amer. Pliilos. Soc, 1878, p. 510; 1880, p. 38.
t Giinlher, Transac. Royal Society, 1867, PI. ii, Fig. 17 ; Albrecht,
|: Meyer, Neues Jahrbuch f. Mineral., Hd. II, pp. 229-30,

TEURESTKIAL VERTEBRATA. 245

articulation for a sino^le-headed lib. In the cervical re«:ion the articular surface for
the rib is continued downwards, forming a shallow groove on the posterior part of the
side of the intereentrum. This groove becomes shorter, and finally disappears from
the intercentra at some distance posterior to the ceivical region. In the cei'vical
region in some of the specimens a groove crosses the inferior side of the inter-
centrum, passing through and dividing into two parts the co.stal groove. It appears
to be a suture which cuts ott" a segment from the posterior side of tiie intereentrum.
This segment is cobssified with the inteiccntnini in most of the cervical vertebi-aa of
Ertjops viegaceplialus, and disappears so completely from the vertebric posterior to
this region, that it is impossible to say whether it is piimitively absent, or coossified.
The narrower anteroposterior diameter of the intercentra, and the absence of the
lateral costal articular groove would indicate its total absence. This element has
been observed by Fritsch in some Bohemian forms, and has been termed by him the
hypocentrum pleuralc.

In the genera described by Dr. Fi'itsch from the Permian Gaskohle of Boluinia
some conditions have been described by him which differ considerably from those
mentioned above. The figures given by Dr. Fritsch are not entirely consistent
with each other, and ap])ear to have been taken from imperfect specimens. Thus in
SparcKjmites lacertinus Fi-.,'-' a neural arch is represented as resting on an intereen-
trum, while the arch next behind it rests on a hypocentrum pleuralc, or a divided
jjleurocentrum. It is not possible to be certain whether the neural arch stands on the
centrum or intercenti'um in the caudal region of Diplovertebron, as in some figures it
covers both (PI. 52, Fig. 2), and in others only one of these elements (PI. 50, Fig.
14). In the caudal I'cgion of the Diplovertebron ]mnct(Uum,i the vertebne have dif-
ferent forms, one being produced inferiorly apparently fi>r a chevron bone, and anotlu'r
posterior to it, being without prolongation, and without chevron. It is not possible
to be sure from Dr. Fi'itsch's descri[)tions and lignres whether the chevron bones ai'e
ai'ticulated, or arc continua, in these genera.

Di-. Fritsch, however, determines the presence for the first time of the hypocen-
trum pleuralc, and describes it in the genera Chelydosaurus and Spheno.saui-ns.j:
With Gaudry, however, he regards tlu: intereentrum as the true centrum, and then
homologizes the hypocentrum pleuralc with the intereentrum of the Pelycosauria.

I have attempted to sustain my interpretation of these homologies^ by reference

• Fauna dcr Gaskolile tier I'ermiroruiations Bu;limeas, Hil. ii, II. 1, PI. .50, Kig. 16.
f Loc. cit.. PI. 50, Fig. 11. Dr. Fritsch believes llie inferior long bone to be a rib.
i Loc. cit., pp. 2.5-38.
§ American Naturalist, 1885, p. 70; Proceeds. Amer. Philos. Soc, 1878, p. 522.

246

ox THE IXTERCENTRUM OF THE

to the genns Cricotus, type of the order Embolomeri ; a form which has not yet been
found in Europe, so far as ascertainable. lu this genus the chevron bones are con-
tinua, while the neural arch is free, as in Eryops. The element which bears the
chevi'ou l)one being regarded, as in that genus, as the intercentrum, we can affirm
(Fig. 3) that the intercentrum is continued upwards to the neural canal, forming a
disk, and the pleui-ocentra are continued downwards and are united below, forming a
complete centrum. This explanation looks the more reasonable in view of the exist-
ence of the hy[)oeentrum ])leurale in the Sphenosauridjc, which if combined with the
pleurocentra would foi'm a completed centrum. It is, however, uncertain as yet
whether this is the make-up of the centrum in Cricotus, because in Eryops the hypo-
centrum pleurale unites with the intercentrum and not with the pleurocentrum. That
my determination of the homologies of the vertebral disks in Cricotus is correct is
further evidenced by the structure of the caudal region of Eryops. In E. erycholiticus
(PI. I, Fig. 1), the pleurocentra descend further than in the dorsal region, reaching
to the inferior face of the column, and sciJai'ating the intercentra from mutual contact.

Fig. 3. — Cricotus crassidiscua Co[)c, posterior part of vertebral column two-fifllis natural size; original; from

Amerifan Naturalist, 1884, p, 1)7. Fig. n, proatla.s ; h, ilo., vvilli first cervical ; e, cervicals ; d, proximal caudals

from below ; e, distal caudals left side. I'li, pubis ; U, ilium ; in, iscliium ; i, intercentrum ; ce, centrum (pleuroccM-
trum) ; r, ribs.

Fiu-ther development of their inferior portion, with truncation of inferior surface,
would represent the structure stated by Fritsch to characterize Archegosaurus, and
which is still better developed in Cricotus. The chevron bones are always continua
ill l)()lli Eryops and Cricotus. Of the latter genus I possess in my collection many
c;iu(l;il intercentra with chevrons, of all ages, and in none of them is there any sutural
articulation visible (I'l. I, Fig. G).

The correctness of this determination is further confirmed by a study of the dorsal
region of Cricotus. In passing along the caudal region towards the sacrum, the
chevron-bearing disks or intei-centra diminish in anteroposterior di;imeter both abso-
lutely and i-elatively, while the centi'a do not un<lei-go any modification. The neural

TERRESTRIAL VERTEBRATA.

24^

arch, from resting by more than luilf its base on the intereentrnm (Fig. 3) gratliiuUy
transfers itself to the centrum, at the base of the caudal series. It maintains this
position throughout the lumbar and dorsal series,* resting freely in an angular fossa
of each side of the anterior three-fourths of the superior lace of the centrum. It bears
the diapophysis with the usual articulation for a single-headed rib (Plate I, Fig. 3).
Tlu' intercentra even continue to the cervical region, wliicli terminates anteriorly in an
intercentrum, which is perhaps the body of the proatlas, the processus dentatus of the
atlas of the existing Batrachia, and the occipital condyle (without basioccipital bone)
of the Tieptilia. t Iu addition to the preceding reasons for adoi)ting my determination
of centrum and intercentrum, it may be added, that in the Cricoius lieteroclllus Cope
the dorsal intercentra are much narrowed above,J some of them so much so as to have
a sharp edge, which forms but a narrow bi'idge above the foramen (Plate T, Figs. 7-8)
chordte dorsal is.

According to Gaudry, the structure of the dorsal part of the vertebral column in
Aetinodon and Arehegosaurus is identical with that of Eryops§ (Fig. 4). This

;/<;;)

Fio. 4. — ArchegoMurus decheni GoMil. Section of dorsal vertebra', restored by Giiudry ; natural size. Let. n, </,
neural arch ; c, neural canal.

Fio. 5. — Tlie same species ; dorsal verlebne, copied from Fritscli ; nalunil size. Letter t", Interrenlrum ; p, cen-
trum ; pz, poslzygaiiO]>liysis ; n. >p., neural spine.

author represents the neural arch as resting on the pleuiocentruni. Fritscli, on the
other hand, II represents the neural arches as slightly in contact with the intercentra, and

• Proceedings Amcr. Pliilosopli. Soc, 1884, p. 29.

t Anier. Naturalist, 1884, p. 37.

X Proceeds. .\mer. Pliilos. Society, 1877, p. 186 (see C. diicophorus, wliicli was founded on such intercentra)

§ Enchalnements du Monde Animal, Fossiles Primaircs, p. 2G1, fig. 259.

I Fauna dcr Goskohle, Tab. 58, fr. 13.

248 ON THE INTERCENTRUM OF THE

the pleurocentra as between their bases, a condition of things which is probably due
to disturbance of the parts before fossilization (Fig. 5). Fritsch also represents the
neural arch in Sphenosaui-us and Chelj^dosaurus as resting on the intercentrum, in
diagrammatic manner,''' and the pleurocentra and hypocentrum pleurale as alternating
with them. As Dr. Fritsch does not give profile figures of the original specimens,
one cannot judge how much may be due here also to disturbance of the parts. Still
more open to this question are the figures of Von Meyer.f The true relation of the
parts cannot be learned from these figures and descriptions, ^nd it is probable that the
specimens at the disposal of European paleontologists generally are as yet mucli
inferior in the condition of their perservation and in size to those found in Texas.

The vertebrjB of the true Stegocephali are, so far as known, undivided and with-
out intercentra, with one exception, and that is the genus Branchiosaurus Fritsch. J
According to this author, each vertebral body is composed of thi'ce parts, a median,
an anterior, and a posterior. The median, if separated from the other two parts, would
be bi-concave, and it supports the rib. Each of the extremital segments presents a
convexity towards the median segment, resembling the epiphysis of a mammal. What
the homologues of the three jmrts may be I'craains to be ascertained.

II. — THE KEPTILIAN INTERCENTRUM.

The Rhynchocephalian genus Si)licnodon has been represented by Giinther to
have cei'vical and some dorsal intercentra.§ It therefoi'e seemed probable that its
structure might throw some light on the homologies of the batrachian and reptilian
intercentra. I therefore, dissected the vertebi-al column of a specimen in alcohol kindly
l^resented to me by my friend, Dr. James Hector, chief of the Geological iSurvey of
New Zeiiland. The attachment of the chevi'on bones does not, in the figui-e of Giin-
ther, resemble very closely that seen in Clepsydrops, but on examination I found the
relations to be structurally and homologically identical with that found in Crico-
tus. The caudal vertebne are se[)arated by a rather thick disk of intervertebral cai'ti-
lagc Avhich sun'ounds the foramen chordi\3 dorsalis, resembling closely the intercentrum
of the Embolomeri. There is an internal annular deposit of phosphate of lime, which,
if extended to the entire cartilage, would give us an embolomerous intercentrum like
tliat of Cricotus. This resemblance is a true homology, for the chevi'on bones spring
from this cartilage, although in sliglit contact with the centrum in front of it (Plate
I, Fig. 10).

* Fauna der Goskolile, pp. 25-28.

f Palnonlographica, 1858, Vol. vi.

X Fauua dur Uaskolilc, 150. i, Heft. \.

§ Transactions of the Royal Society Loudon, 18C7, iil. ii, fig. 23.

TERRESTRIAL. VERTEBRATA.

249

There can then be no donbt of the homolof^y of the cartilaginous interceutrnm of
Sphenodon with those of the Pc!3C0sauna, and of the entire centrum of Sphenodon
with that of the I'elj'cosauria. I'he division of many of the caudal centra of Sphe-
nodon and ol" Diany lizards, is evidently' not exi)laincd by the supposition that one of
the parts is an intercentiiini. That they aic halves of a single centrum is not only
rendered probable by the above determination of the intercentrum, but is supported
by the nKidifieations presented by the centra themselves. In Sphenodon, instead of
gradually losing one of the halves, the latter become, without diminution, more and
more consolidated towards the anterior part of the caudal series, and merge into the
ordinary t^'pe of vertebra, each proximal centrum representing both halves of a distal
one (Plate I, Fig. 10).

It is thus probable that each vertebra of a Lacertilian* and of a Pelycosaurian
represents one centrum, and that intercentra are present in some types, and absent in
others, except as always represented by the chevron bones of the tail. Is this the
case with the existing: Batrachia ?

Fio. ^.—Eryoiu megaccpJuilm. Vertebral column represented in Fig. 1, from below; oue-fourtli natural size.
From Permian epoch of Te.\as.

In an important memoir, Iloffmanf presents us with the results of his investiga-
tions into the homologies of tlic ril)s of the terrestrial Vertcbrata. He finds that in all
of them, excepting the Batrachia, the primitive ribs are processes of the intervertebral,
and not of the vertebral caitilage. The intervertebral cartilage in reptiles with ball
and socket joints divides, each half uniting with its adjacent body, the one to form
the ball, and the other the concave extremity of a vertebra. In the Mammalia the
halves foi-m the epiphyses respectively. In this class the head of the rib justifies this
origin, lomaining as it does articulated in a fossa which is ci|ually excavated from two
adjacent epiphyses. I have shown thai the head of tliu lil' in thr Pclycosauria is artic-
ulated with the undivided intercentrum. t The striictine in tiiis order is in conlirnia
tion of the doctrine that the elcracni 1 have called intercentrum is such in fact. The
passage of time has seen in the Keptilia in general, the same moditication in the mode
of attachment of ribs as occurs in the vertebral column of the Mammalia, etc., in i>ass-

» Baur 3how3 that in ihr Lncerlilian genera IJniplatc.s and Gecko, intercentra are present tliroughoul the dorsal,
lumbar, and sacral regions. American Naturalist, 188fi, p. 174.
t Niederl.-ondisches Archiv. f. Zoologic, 1878, p. 190.
X Proceeds. Amcr. Philos. Society, 1884, p. 43 ; Proceeds. Amer. Assoc. Adv.' Science, 1881, p. 471.

250 ON THE rNTEKCENTKUM (JF THE

ing from the ffont to the back. The iiitercentral articulation is lost, and finally the head
of the rib disappears. Only the tubercnlar attachment to the diapophysis remains,
in the Streptostyliea, and this is a simple one.

In the modern Batrachia the rib articulation is that of a double-headed rib to a
double-headed diapophysis which originates from the neural arch. There is nothing
in the history of the class to show that, as in the case of the reptiles, the head has
been transferred or lost. In recent Batrachia, elements resembling the intercentra of
Pelycosauria are unknown. The origin of the ribs of fishes and Batrachia have been
shown by Goette to be in the connective tissue of the interspaces of the myocommata,
which are opposite the middles of the vertebral centra. It follows that there is at no
time connection of the ribs with the lines of separation of the vertebral bodies.

If the intercentra become the functional centra in the Sphenosaurid.T, it is likely
that they are such in the Stegocephali, and in the modern Batrachia.*^ It follows,
then, that Batrachia, excepting Khachitomi and Embolomeri, have no centra, but in-
tercentra only. This view is confirmed by three facts, two of them already mentioned:

I. There are no intercentrum-like bodies in existing Batrachia.

II. The ribs which originate from intercentral cartilage and intercentra in reptiles,
originate from the principal vertebral bodies in Batrachia.

III. The chevron bones are continiia with intercentra in Reptilia, and with the
caudal vertebral bodies of Batrachia.

With regai'd to the last proposition I may add, that I have examined young Nec-
tuncs maculatus, of four inches in length ; of larvfe of Gurinopliilus j^orphi/viticus, of
thi-ee inches ; of Spelerpes ruher of two and a half inches ; of Amhlystoma yuactatum,
of 35mm ; and of Siyderpes hilmeatus, of 25mint length. In none of these is the
least trace of articulation of either chevron bone or of neural aix-h in any part of the
column to be discovered (Plate I, Fig. 11).

Researches into the embryology of the Urodela and Anura have not yet brought
to light any traces of the rhachitomous structure ; which is probably a case of inexact
parallelism, cceuogeny or falsification of the embryonic record — a phenomenon which
is not uncommon. There can be no doubt, however, that the entire record was i)re-
sented in the embi-yonic history of Permian land Vertebi-ata. and for a long period
subscquentl}', but that the rhachitomous stage has been, with the true centrum, lost
from tiie batrachian line at least.

As I have shown, in the Embolomeri (he intercentra tend towards reduction in
the dorsal i-egion, while the centra arc predominant. If the tendency of the evolution

* American NiitunUisI, 1880, p. 77.

t For these specimens I am greatly indebted to the Smithsonian Institution and the Secretary, Prof. Bairil.

TERRESTRIAL VERTKRRATA. 251

of the Biitrachia has been to the extinction of the centrum, the line of the Embolomeri
tends in the opposite direction, or towards the type of tlie Keptilia. Althou<;h the-
general characters of the skull of Cricotus are Bati-achiaii, the presence of a free pio-
atlas (Fig. o,n h) adds to the evidence of reptilian afHnity.* It is probable then that
we have in the' Embolomeri that order of Bati-achia from which tlie Kei)tilia were
derived, tluough intermediate forms not yet discovered.! It is also evident that the
Sphenosainid;\; cannot be referred to this order as T have proposed, but that they con-
stitute a family of lihachitomi.

We have thus clearly shadowed forth in the Permian Vertebrata the ancestry of
the existing true Fishes,^ Batrachia, Keptilia, and Mammalia.§

Tir. — ALTERNATIVE EXPLANATIONS.

I may here consider other possible ways of interpreting the homologies of the seg-
ments of the rhachitomous vertebral column, as follows: Let us first suppose with
Gaudry and Fritsch that the segments called intercentra in the figures and plates
accompanying this paper, are the true centra, and that the chevron bones are not
continna, but are originally separate, and have l)ecome coossified with the elements
with which we find them now contiinious. It will follow that the lai'ger bodies of the
vertebral column of Cricotus, Avhicli in Ihc dorsal region support the neural arch, are
intercentra, and that the centra are in llial genus in process of extinction. The same
will Ix' tiue of the so-called dorsal, and part or ail of the so-called caudal vertebra* of
Sphenodon. and of the Pelycosauriii. We thus have the Tleptilia in the |)Osition which
I have assigned to the Batrachia, the terms centrum and iutercentrum being mei'cly
reversed. As, however, these names were first a])j)lied to the Pelycosaurian reptiles
by me, and as it will recpiii-e less change of nomenclature to retain these names for the
bodies as they appear in Mammalia and Keptilia, it will be better to maintain the
proposition that the Batrachia have lost their centra, and retain only intercentra.

Another alternative is to regard the hypocentrum ])leurale as the intercentruui w itli
the chevron luiuc ll will be remembered that the iiypoceuliiiiu pleurale is described b}'
Dr. Fritsch as lying below the i)leurocentra. This is of course on the anterior side of
the "hyi)ocentium.'' It looks reasonable to suppose that in the completion (»f (he body
of which the pleurocentrimi is a J)art, the hypocentinm pleurale should be included.
Su(h might be supposed to be the case in the cenfrum of Cricotus, and of llie Pel}'-

* The iiniallanliil renlrilin is llinroiiglily ciiossific'il in all llio saliiiiiaiiiki" laiva- aliovc rcferiiil lo.
t I have suggested this view in ilie American Naturalist, \HSi, p. ;i7 ; \>i8't, p. 77. •

X .\<l(litioDa1 specimens of Didjmodus display very distiiiclly the futures I liave described, wliidi >Ir. Onrmnn
tldnks to be nccidenlal.

S A second posterior fool of C'lcpsydrops displays perfectly the iii:iminali;in characters I lijivc ascribed to It.

A. V. S. — VOL. \VI. 2f.

252 ox TIIK rXTERCEXTimM OF THE

cosauria. But on this point we have as yet no evidence. On the other hand in the
rhaehitomous Ei-yops, where the centrum is not completed, we have seen that in the
cervical region the hypocentrum pleurale probably unites with the intercentrum in
front of it. Let us sui)pose this to have been the case throughout the vertebral
column. The interpretations would be as follows :

In the caudal region of Eryops the hypocentrum i^leurale Avould bear the chevron
bone, and the present intercentrum would become the centrum. We would then have
in the caudal region of Archegosaurus and Cricotus, the remarkable state of aifairs of
two ti'ue centra present, one of which has an intercentrum coossified with it, and the
other supports the neural arch. The same would be the interpretation of the dorsal
structure in Cricotus. As there is no direct evidence of such structure in the vei'tebral
regions mentioned, noi" in the caudal region of the Pelycosauria, its assumption appears
to me to be the least probable alternative before us. Under this interpretation we have
to regard the pleurocentrum as either vanishing, or becoming coossified with the inter-
centrum (m., h^fpocentrum) in the vertebral centra of existing Batrachia, Avhich would
then consist of four elements each, viz., two pleurocentra, hypocentrum pleurale, and
intercentrum (hypocentrum), in the caudal i-egion at least. Of this no evidence can be
obtained from embryos of a length of 25mm and over. In Reptilia the centrum Avould
consist of pleurocentrum and intercentrum (hyi)ocentrum) combined, a division of which
we have no evidence in the emlirj'ology of the vertebral column in these animals. In
the divided caudal centra of the lizards one might see intercentrum (hypocentrum) and
pleurocentrum, but this supposition is not necessary to account for this structure. I
have observed above, that the evidence furnished by the vertebrse themselves is in
favor of this division having arisen in the middle of a true centrum. The chief objec-
tion to this interpretation of the reptilian centrum is, however, to be found in the
column of Cricotus, where we find the large bodies which bear the chevron bones, in
the process of extinction, leaving behind bodies which are homologous with the centra
of the Pclycosaurian reptile. And this is equally true, whether we regard the hypo-
centrum or the hypoeenti'um pleurale as the intercentrum, for it is the pleurocentra
which remain, to be the true centi'a of Cricotus.

Thus on either of the assumj>tions just mentioned, the development of the dorsal
l)urt of the vertebral column in Cricotus is in an opposite direction to that stated by
Fiitsch to characterize the Sphenosauridai. This is the main point to be proven, li'
further I have shown that the larger dorsal bodies of Cricotus are homologous with
the centra of the Pelycosauria and Lacertilia, the i)roposition remains proven that the
inferior vertebral bodies of the Jihachitoiui and the entire vertebral bodies of existing
Batrachia, are intercentra and not centra.

— -X

TERKESTRIAL VERTEHRATA. 253

In Older that the result .shall be otherwise, it will be necessary to make the
extreme assumption, that in Cricotus the two vertebral bodies represent hypocentrum
pleurale (bearing chevron), and hypocentrum, i\\v i)ic'uroccntrum having disappeared.
The reader can judge which of the alternatives is the more probable, the disappear-
ance of the insignificant hypocentrum pleurale, which only exists in distinct form in
the SphenosauridfB, or of the large pleurocentra which arc so Avell developed in tlif
"Rhachitomi, and which in the caudal region of that order are almost identical with the
bodies marked ce in Cricotus (Fig. 3). On this hypothesis we are led to the reductio
ad ahsurdiim, that the bodies which do not support the chevron bones in the caudal
I'cgion of Eryops (PI. I, Fig. 1 c), are not homologous with the bodies which do
not support the chevron bones in the caudal region of Ci'icotus (Pi. I, Fig. 2 c-5)
with which they are identical in position and connections.

Explanation of Plate I.

Figs. 1-6, two-thirds natural size; Figs. 7-10, natural size ; Fig. 11, five times nat. size.

Fig. 1. — Eryops erythrolUiem Cope (Epicordybis dim), pro.ximal caudal vertebrr. The first intercentrum is the
last one without chevron bone. From the Permian of Texas.

Fig. 2. — Cricotus hypantrifus Cope, portions of the vertebral column of one individual, ."selected from the cervical
(a), dorsal (i), and caudal (c), regions ; left side. From the Permian of Te.xas.

Fig. 3. — The centrum of the anterior dorsal vertebra of 2fc, fnim front, showing Uie liypnnlruiii and liypantra-
pophyses. (Proceedings Amer. Philos. Soc, 1884, p. 29.)

Fig. 4. — Another dorsal centrum without neural arch, from front ; a, from behind ; b, from above, showing facet
or neural arch.

Fig. 5. — Intercentrum from cervical region, showing rib atlachnient, from front ; n, from behind ; 6, from the
side ; f, from above.

Fig. 6. — An intercentrum from the caudal region ; a, from above.

Figs. 7 and 8, dorsal intcrcentra of Cricotus heterocUtus Cope, from Illinois ; collection of W. F. E. Gurley.

Fig. 7. — Lateral view.

Fig. 8. — Front view of another intercentrum, the superior part broken away ; a, lateral view.

Fig. 9. — Clepsydrops nntalis Cope, a portion of the caudal region, showing ribs and chevron bone."!, from the side >
fig. 9a from below, .showing one free intercentrum, out of its place (»)• From the Permian of Texas.

Fig. 10. — Splunodon gucntheri BuUer ; from New Zealand, caudal vertebra;, lateral view ; «, an intercen-
trum with chevron bone from front. The reticulated portions are osseous tissue.

Fig. 11. — SpeUrpes hilineata Green, larva, vertebra;, five times natural size, linear. 11«, Three cervical vcrtebrit,
including atlas and connate centrum of proatlas ; lift, a dorsal vertebra ; lie, two sacral and two anterior caudal verte-
hrrc ; lid, second caudal vertebra from behind. The caudal vcrtcbric show the continuity of the chevron bones. Speci-
men from the collection of Professor S. F. Baird.

Litters,

pra, proatlas, centrum. poz, postzygapophysis.

»", intercentrum. ha, hypantrum.

p, pleurocentrum. hap, hypanlrapophysis.

c. centrum. ne, nolochordal foramen.
eh, chevron hone. jf, foramina.

r, rib. m, gacral intercentrum.

d, dfapophysis. ic, sacral centrum.
pa, parapophysis.

ARTICLE III.

A CHEMICAL STUDY OF YUCCA ANGUSTIFOLIA.*

BY H. C. DE S. ABBOTT.

Read before the American Philosophical Society, December i8, 1885

This plant is well known in the West as the " soap-weed." It grows very abun-
dantly in most of the Western States and Territories. It has attracted the attention
of botanists, and it is a plant of interest on account of the many uses to which it has
been put in the countries where it is found.

The results noted in this paper are based upon a first and introductory chemical
analysis of the Y'ucca. Previously, little has been studied of its chemistry. It is
briefly mentioned in the work of a French writer. Dr. Georges Pennetier ; f also, in a
I)aper on the study of manganese found in the ash of plants in which M. Maumene
states that the ash of the Yucca contains manganese.|. He does not name what spe-
cies of Yucca was examined. The former writer gives the micro-chemical characters
of the action of iodine and sulphuric acid, dilute chromic acid, and cuprammonia on
the fibres of the Yucca angustifoUa.

The specunens of Yucca used in these analyses were of large growth and in good
condition. The entire plant was examined, and a separate study made of the bark and
wood of the root, and of the green leaf and the yellow basal part. The roots were air-
dried, freed from adherent dust, reduced to a very fine powder, and passed through a
!N"o. 80 sieve. The leaves were less finely powdered.

Dragendortf's scheme for plant analysis§ has been generally followed. Ten
grams of the air-dried powder were used for the preliminary examination of soluble
substances. For every gi-am of the powder, ten c. c. of the solvents were em-
ployed. An additional quantity of the powder was prepared for special purposes.
Five grams of the air-dried powder were dried, in a hot-air oven, at a temperature

* An abstract of this paper was read befere the Chemical Sectiou of the American Association for the Advancc-
mriil of Science, at Ann Arbor, Michigan, Angnst 38, 18S5.

fLecjons Bur les Matieres Premieres Organiqnes, Paris, 1881, p. 44f>.

X M. E. .1. Maumene, Bui. de la Soci6le Chimique de Paris. Tome xlii, p. 30.5.

§ Plant Analysis. Qualitative and Quantitative, by G. Dragendorff. Translated from the German by Henry
Greenish, London, 1884.

A CHEMICAL STUDY OF YUCCA ANGUSTIFOLIA.

255

between 100° C. and 110° C. until the weight remained constant, for the estimation of
moisture. This powder was incinerated in a covered i)orcclain crucil)le at a dull red
heat until the carbon was entirely consumed. The per cent, of total ash was deter-
mined from it.

QUALITATIVE ASH ANALYSES.

Calcium, magnesium, potassium, sodium, iron, manganese, chlorides, phosphates,
and sulphates, were found in every part of the plant.

C I. MOISTURE )

Determination of i . , ^ on the powder.

in. TOTAL ASH j ^

fi.7S per rent.
11.07

8.11
.^7.00

//.

Color <!/■ A»h.

(1.) The bark of the root

(2.) " wood "

(3. ) " green leaf

(4.) " yellow base of loiif

17.38 per cent.
1.5.75 "
5.75
10.G.S

reddish

" gray

gray
■white

I'ETKOLEUM SPIUIT EXTRACTS.

Extract (1) Bark of Boot.

The maceration was conducted in an apparatus similar to one described in Drag-
endorff's " Plant Analysis."^' A light petroleum spirit was used which boiled between
25° C. and 45° C. The extract was filtered from the jwwder-residuc. It was a clear
pale yellow-colored liquid, and slightly acid in reaction. A drop of the extract on
evaporating left a uniform spot on blue paper. The extract was evaporated at the
oidinary temperature. The residue was a solid, and it had the odor and characteristic
cry.stalline structure of fatty acid.s, suggesting the presence of a fixed oil. Its melt-
ing point was taken. 'I'hc substance melted at 00° C, and on cooling solidified amor-
phous. To determine the total amount of solids extracted, a definite volume of the
extract was evaporated, dried, and weighed.

TOTAL SOLIDS.

Petroleum gpirii residue (ir\c(\ at lOOO C 1.24 per cent of solids,

" UOOC 1.20 " " "

0.04

loHS.

The residue was identified as a fixed oil. It was soluljle in jjctrolcum spirit,
ether, benzole, chloroform, ainyl alcohol, carbon di-sulphide, and cold arpieous alka-
lies; incompletely .soluble in cold or boiling 8G per cent, alcohol, 1*5 pcM- cent, alcohol,
absolute alcohol, acetic ether, and aminoniura hydrate. No change of color was

* Page 90, Tollen's apparatus.

256 A CHEMICAL ST LTD Y OF

observed on treating the fixed oil with concentrated sulphuric acid, nor on the addi-
tion of syrupy phosphoric acid, though it was partially soluble in these acids. Phos-
phoric acid colored it yellow ; it was colored yellowish by concentrated hydrochloric
acid and nitric acid of 1.22 specific gravity. A mixture of concentrated sulphuric
acid and nitric acid of 1.22 specific gravity changed the color of the fixed oil to a red-
dish-brown; it was colored pale green by sulphuric acid of 1.634 sp. gr. and of 1.53
sp. gr. Calcium di-sulphide gave a bright green color reaction with the fixed oil, but
did not form an emulsion with it ; aqueous solutions of gold and platinum chlorides
were reduced by it. The fixed oil was saponified with . difliculty by alcoholic soda ;
but readily by boiling aqueous soda ; a white fragile soap was separated and filtered
from the liquid. The soap was decomposed by hydrochloric acid and the fatty acids
separated. The filti-ate from the soap was examined for glycerin. By the method*
used, an oily liquid was obtained, it was heated with anhydrous borax on platinum
foil, and gave the usual green-colored flame test for glycerin. The alcoholic solution
of the j^etroletmi sjnrit residue was fractionally precipitated with an alcoholic solution
of magnesium acetate, and traces of an amorphous residue were recovered.f

The petroleum sjyirit residue was digested with water containing sulphuric acid,
and examined for alkaloids which are sometimes brought down Avith fixed oils. The
usual reagents failed to detect traces of alkaloids.

:Extract (2), Wood of the Boot.

The maceration was carried out under the same conditions as in extract (1).

The extract was a clear, colorless solution, neutral in reaction. A drop of the
liquid left no uniform spot on blue paj)er. The extract was evaporated at the ordinary
temperature. The residue was light yellow- coloied, of a semi-solid consistency and
melted at 36°j^C. A definite volume of the extract was evaporated, dried, and weighed.

TOTAL SOLIDS.

Petroleum spirit reiidue dried at 100° C 0.55 per cent of solids.

" llOOQ 0.35

0.20 " " loss.

The residue was identified as a fixed oil ; associated with volatile fatty acids.
The latter were indicated by the 0.2 per cent of loss, and the disagreeable odor of the
residue which was dissipated on heating at 110° C.

The petroleum spirit residue from the extract was evaporated at the ordinary tem-
perature, dissolved with difliculty in cold 95 per cent, alcohol, and in boiling weaker

*" riant Analysis." G. Draggendurir, p. 13.
t Loc. cit., page 10.

YUCCA ANGUSTIFOLIA. 257

alcohol ; absolute alcohol hardened and discolored it. Concentrated sulphuric acid,
nitric acid, and hydrochloric acid, did not appreciably act on the residue. It was not
saponified, but slowly dissolved by boiling aqueous and alcoholic soda. The alco-
holic solution of the petroleum spirit residue Avas submitted to a fractional precipi-
tation with an alcoholic solution of magnesium acetate. The first precipitation
obtained was purified by boiling alcohol ; it was an opaque scaly crystalline solid
which melted at 85° C. The second precipitation yielded traces of a white amorphous
substance. The third precipitation resulted fi-om adding strong ammonia water to the
magnesium acetate solution, and the purified residue melted at G0° C.

Negative tests for alkaloids followed an examination of the aqueous treatment of
the petroleum spirit residue.

Extract (3), Qreen Fart of the Leaf.

The method of extraction was the same as that used in the previous extractions.
The extract was clear, pale green in coloi-, and non-fluorescent. It was colored by a
small quantity of chlorophyll, which the petroleum spirit dissolved. The liquid was
acid in reaction. A drop of it left a permanent stain on lilue paj)er when evapoi-ating.
The extract was evapoi-ated at the ordinary temperature, and the residue was a dark
greenish-yellow semi-fluid substance. The solidifying point was taken. It was found
to be about 15° C. A definite volume of the petroletim spirit extract was evaporated,
dried, and weighed.

TOTAL SOLIDS.

Petroleum ipiril residue dried at lOQO C 2.20 per cent of solids.

IIOOC 2.01

0.19 " " loss.

The pet roleu)7i sjurit residue was identified as a fixed oil with a small amount of
chlorophyll that had been brought into solution by it. It was soluble in cold 83 per
cent, alcohol, 95 per cent, alcohol, absolute alcohol, ainyl alcohol, ether, acetic ether,
chloroform, ben/ole, carbon di-sulphide and glycerin. It was also soluble in oil of
turpentine, almond oil, ammonium hydrate, mercuric chloride, and slowly soluble in
acetic acid. Concentrated nitric acid, and hydrochloric acid slowly dissolved the
fixed oil, the former colored it dark green, and on stirring the mixture the color was
changed to a brown. Concentrated sulphuric acid dissolved and changed it to a very
dark-brown color ; on adding concentrated nitric acid, the li(|uid was changed to a
leddish-brown color.

The following reactions were noted: Tlie lixcd oil cliangi'd to a hard greeiii.sh-
yellow substance on heating it with anhydrous boia.x on platinum foil. When rubbed

258 A CHEMICAL STUDY OF

on a crucible lid with powdered rosaniline, it was colored red, showing the presence of
free fatty acids. It did not emulsify with calcium di-snlphide nor with syi'upy anti-
mony chloride, but it was colored dark-green by the latter. It Avas imperfectly dis-
solved by phosphoric acid, and slowly soluble in equal parts of cane sugar and concen-
trated hydrochloric acid; more rapidly soluble in equal parts of cane sugar and niti'ic
acid. An aqueous solution of picric acid made allvaline by sodium carbonate colored
the fixed oil a light reddish-brown color, cane sugar added to the solution facilitated
dissolving it. It was instantly dissolved by equal parts of picric acid and acid ammo-
nium phosphate, and on warming with stannous chloride, leaving a tui-bid yellow-
colored liquid. It was insoluble in aqueous barium hydrate; soluble in alcoholic am-
monia with no coloration, and in sulphurous acid. It was colored brown when mixed
with sulphuric acid of 1.634 specific gi'avity, and incompletely dissolved ; it was also
colored brown by ferric chloride. On adding to the fixed oil sulphuric acid of 1.475
specific gravity, and a small quantity of zinc, hydrogen was generated, and the solubil-
ity of the oil in the acid liquid was accomjjanied by a rosy tint given to the solution.

Extract (4), Yellow Part of the Leaf.

This extract was obtained by a similar process as that used for the other petro-
leum spirit extracts. The extract was a pale yellow-colored liquid. T\\q reaction was
slightly acid. A uniform spot was left on blue paper as the drop evaporated. The
petroleum spirit was evaporated at the ordinary temperature, and a yellow-colored
residue recovered, of a semi-solid consistency and crystalline in structure. It solidi-
fied at 12° C. From a definite volume of the petroleum spirit extract, the amount of
total solids was determined.

TOTAIi SOI.TDf.

Petroleum spirit residue ilried at lOQO^C 1.1 per ceut of solids.

IIO^C 1.1

O.UO " " loss.

The residue was identified as a fixed oil. It was soluble in warm absolute alcohol,
incompletely soluble in weaker alcohol ; soluble in cold acetic ether, chloroform, ben-
zole, amyl alcohol, ether, carbon di-sulphide, and glycerin. It was saponified with
aqueous soda and a white soap separated. No reaction Avas observed with i)iciic acid
and ammonium phosphate, nor with nitric acid of 1.32 si)ecific gravity and 1.18 sj^e-
eific gravity. The fi.xed oil was soluble in jKitassio-mereuric iodide solution ; and
colored dark-brown by alcoholic ammonia. A mixture of fej'ric chloride solution and
powdered rosaniline gave a fine violet-colored reaction with the fixed oil.

An examination of the aqueous treatment of the petroleum spirit residues (3) and

YUCCA ANGDSTIFOLIA.

259

(4), for alkaloids, gave negative results. A poi'tion of the original powder, from each
of the four i)arts of the plant, was mixed with an aqueous solution of caustic soda,
and the distillate examined for volatile alkaloids with negative results.

SUMMARY I.

Petvoleum Spirit Extracts.

1. The bark of tlie root. .

2. " wood

3.

4.

-green leaf

yellow base of leaf.

Solids extracted.

1.24 per cent.
0.55

2.20

1.10 "

Character of residue

fixed oil

/ " " 1

\ chlorophyll (
fixed oil

Renetion with
litmus.

Sidling point.

slightly acid
neutral

acid

faintly acid

60° U.
3(;o C.

{semilhiid at "|
ordinary \ I
temperature j |

Solidifying point.

I f solid at ordinary
' X temperature
15° C.

120C.

The solids extracted by petroleum spirit from the four parts of the plant are iden-
tified as fixed oils*; associated with a volatile principle (0.2 per cent) in extract
(2), and with traces of chlorophyll in extract (3).

Fixed oil (1) was crystalline in structure. It was soluble in ether, chloroform,
benzole, carbon di-sulphide, and amji alcohol ; incompletely soluble iu cold or boiling
alcohol, acetic ether, and ammonium hydrate. It was colored pale green b}"^ sulphiu-ic
acid of 1.631 specific gravity, and changed to a bright-green color by calcium di-
sulphide, but formed no emulsion with it. Phosphoric acid colored it yellow. The
fixed oil was saponified, and a white soap separated. This was decomposed, and the
fatty acids recovered. Glyc(M-in was separated from the soap filtrate.

Fixed oil (2) was dissolved with difllculty in boiling 95 per cent, alcohol, and.
hardened and discolored by absolute alcohol. It was not saponified. Crystalline
solids were separated by precipitating the alcoholic solution with magnesium acetate.
They melted at 85° C, and at GO'' C, respectively.

Fixed oil (3) was soluble in alcohol, ether, chloroform, benzole, carbon di-sul-
phide, oil of turpentine, almond oil, glycerin, and slowly soluble in acetic ether. The
presence of free fatty acids was demonstrated. The fixed oil was colored dark-green
by syrupy antimony chloride ; on adding to it sulphuric acid of 1.475 specific gravity,
and a small cpiantity of zinc, hydrogen was generated, and the solubility of the oil in
the acid liquid was accompanied by a rosy tint given to the solution.

Fi.xed oil (4j was crystalline iu structure. It was soluble in warm absolute
alcohol, in cold acetic cthei", chlorofoim, benzole, amyl alcohol, ether, carbon di-sul-
phide, and glycerin. It was saponified, and a white soap separated. The fixed oil
was colored dark-brown by alcoholic ammonia, and a mixture of ferric chloride solu-
tion and powdered rosaniline gave a violet colored reaction with it.

•Fixed Oils. Science, Sept. 11, 1885.
A. P. S. — VOL. XVI. 2g.

260 A CHEMICAL STUDY OF

These fixed oils differed in their physical chai-actei's and chemical reactions.
This diffeience may he due to the presence of fi-ee fatty acids and glyceiides in vary-
ing proportions in the four parts of the plant. It is of interest to note that in the sub-
terranean part of the Yticca, the oil extracted from the bark was solid at the ordinary
temperature ; from the wood it was of a less solid consistency; while the yellow base
of the leaf contained an oil quite soft, and in the green leaf the oil was almost fluid.

Extract (2) contained an oil of low melting point. It melted at 36° C. An
alcoholic solution was fractionally precipitated with magnesium acetate, and three
members of the fatty acid series were isolated. The quantities obtained were small,
and it was impossible to do more than to take the melting point of two of the jiurified
crystalline residues. They melted at 85° C, and at 60° C. respectively. It is a well-
known fact that a mixture of fat acids in certain proportions has a lower melting point
than those of its constituents.

Alkaloids and volatile-alkaloids were not detected in the petroleum sjnrit extracts.

ETHER EXTRACTS.

Extract (1), Earl- of the Boot.

The residual powder from the petroleum spirit extraction was dried until
thoi'oughl3' freed from petroleum spirit. It was then macerated with Squibbs' stronger
ether in the apparatus already described. The ethereal extract was filtered from the
powder. It was a clear crimson-colored liquid, tinted by some red coloring matter
dissolved ; and acid in reaction. The extract was slowly evaporated at the ordinary
temperature ; white needle-shaped crystals Avere seen as the liquid concentrated. The
ethereal residue was of a resinous character. It was ruby-colored, transparent, and of
a softer consistency than ordinarj^ resin. Microscopically, the residue was identified
as a resin by its color reaction with Ilanstein's aniline violet solution.* The ethereal
residue was treated Avith petroleum spirit to remove any traces of fat that may have
been extracted with it. It was heated in a small tube, at 50° C. it experienced a
slight change, and melted at 70° C. For a determination of the total solids, a defi-
nite volume of the etlicreal extract was evaporated, dried, and weighed.

TOTAL SOLIDS.

Ethereal residue dried at 100° C 3.16 per cent of solids.

" HOC; 3.1G

0.00 " " loss.

The resin was incompletely soluble in 95 per cent, alcohol, absolute alcohol, and

♦Botanical Mici-o-Cliemistiy. PoulscuTrcIcase. Boston, 1881, p. E9.

YUCCA AXGUSTIFOLIA. 261

amjl alcohol; readily soluble in ether; not appreciably soluble in chloroform, benzole,
and carbon di-sulphide. It was dissolved by sulphuric acid to a colorless solution,
which, on warming, turned to a yellow color, and gradually darkened to a dull brown
color, fading to a pale yellow.

An attempt was made to separate the white needle-shaped crystals mentioned
above. The ethereal residue was agitated with acetic ether. The liquid was filtered
from the insoluble matter and evaporated. Traces of a resinous substance were sepa-
rated. The insoluble matter was treated with boiling ether, filtered hot, and the fil-
trate concentrated. On cooling, the white needle-shaped crystals reappeared. They
were insoluble in water and in acetic ether.

A separate portion of the ethereal extract was evaporated, and treated with warm
distilled water. The aqueous extract was made up to a definite volume, and a known
quantity evai)orated, dried, and weighed. The amount of total solids was almost in-
appreciable by weight. The aqueous extract was not colored by iron salts, and it did
not form a precipitate with alum and gelatine solution, lead acetate, potassio-mercuric
iodide, nor gold chloride solutions ; showing absence of tannin, gallic acid, and alka-
loids. The ethereal extract was directly tested for these compounds, and with nega-
tive results. A portion of the aqueous extract was evaporated to dryness, and treated
with potassa solution, and the residue dissolved Avith no coloration. Another por-
tion of the aqueous exti-act was agitated with acetic-ether, and the liquids were sepa-
rated ; on evaporating the acetic ether solution, traces of a residue were obtained
which sulphuric acid acted upon. A resinous substance separated from the gi-een-
ish-colorcd acid liquid, the former was partially disintegrated by cold water.

The specific gravity of the resin was 1.091.

Extract (2), the Wood of the Hoot.

The residual powder from the petroleum spirit extraction was macerated in
stronger ether. The ethereal extract was of a reddish-yellow coloi-; slightly acid in
reaction. It was slowly evaporated at the ordinary temperature, and as the liquid con-
centrated, white needle-shaped crystals appeared, and presented the same physical
structure as the crystals found in the ether extract (1).

The ethereal residue was identified as a resin. It was a transparent, ruby-colored
8ul)stance, and acid in reaction. It was heated to 50° C, at that temperature its color
deepened, and at 70'^ C. it melted. The specific gravity of the resin was l.OUl. A
definite volume of the el/ier extract was evaporated, dried, and weighed to determine
the amount of total solids.

262 A CHEMICAL STUDY OF

TOTAL SOLIDS.

Ethereal residue dried at 100° C 1.70 per cent, of solids.

" " 110°C 1.45 " '•

0.23 " " loss.

The resin Avas examined by Hirschsohn's scheme* with a view to classify it
■with known resins. It was imperfectly sohible in 95 per cent, alcohol and chloroform ;
soluble in ether. The alcoholic solution gave a turbidity with lead acetate, not cleared
upon boiling, and with ferric chloride formed a clear mixture. Concentrated sul^jhuric
acid dissolved the resin, leaving a dark yellow-brown liquid which faded to a dull yel-
low color. The sulphuric acid solution when mixed with alcohol, changed to a pale
gray coloi-. On addition of water to the acid solution, there was no coloration nor
separation of the resin. Alcohol containing hydrochloric acid gave no color reaction
with the resin. Bromine solution added to the chloroform-resin extract, and iodine
solution to the ether-peti'oleum-resin extract, gave no reactions. Sodium carbonate at
the ordinary temperature had no effect on the lesin, but on boiling the liquid was
colored yellow.

By the above examination, this resin was thrown out of the numerous classes of
described resins. It is proposed to name it yuccal.t

Yuccal was soluble in boiling absolute alcohol and acetic ether ; incompletely
soluble in benzole, carbon di-sulphide, alcoholic ammonia, and cold acetic ether. The
red color of the resin was removed by cold acetic ether, a transparent substance
remaining, soluble in hot acetic ether.J Yuccal Avas dissolved by potassio-mercuric
iodide. It reduced aqueous solutions of gold and platinum chlorides. A blood-red
color reaction Avas obtained by Avarming a small quantity of the resin on a crucible lid
with a crystal of ammonium molybdate and a few drops of nitric acid. On adding to
the resin mixture a few drops of strong sulphui'ic acid, and again Avarming, it Avas dis-
solved. Warm dilute nitric acid dissolved the resin, colorless ; cold nitric acid gave a
brownish-green color reaction. § Yuccal was heated on platinum foil, and as it decom-

*E. llirschsolin. AVatts Clicm. Diet. Vol. viii, Pt. ii, p. 1743.

fl suggest that in future all resins be distinguished by the terminal syllable al, for uniformity of resin nomencla-
ture. Yuccal ; Science, September 11, 1885, p. 310.

X I have examined the action of acetic ether as a solvent for resins. Cold acetic ether dissolved ordinary resin,
turpentine, styrax, tolu-balsam, mastic, elemi, Canada-balsam, Peru-balsam, copaiba-balsam, Venice-turpentine, and
incompletely spruce gum, and yuccal. In hot acetic ether spruce gum and yuccal were soluble. The following resius
were insoluble in hot or cold acetic ether, guiacam, sandarac, shellac, benzoin, olilianum, ammoniac, myrrh, galba-
num, and asafatida.

^A reddish-yellow decomposition product resulted from the action of nilric^ acid on many resins which fol-
lowed generally quite soon after adding the acid to a small quantity of the resin (0.1 gram of the resin and 5 c. c. of
nitric acid l.-l sp. gr.). But the reaction which took place varied according to the conditions, i. c, strength of acid used.

TUCCA AKGUSTIFOLIA. 2G3

posed the fumes that wore given oft' were pleasant and aromatic. Tests failed to show
the presence of benzoic or cinnamic acids.

The ethereal residue was treated with warm water, and on cooling, the liquid Avas
agitated with acetic ether, which was separated, and when evaporated yielded a small
quantity of resinous substance. The ethereal residue insoluble in water was treated
with boiling ether, and as the liquid concentrated, the white needle-shaped cr3'stals
wore seen floating in it, but on further concentration they could not be scon, and a
yellow greasy-looking mass settled in the bottom of the beaker. On driving oft' the
ether, a transjjarent and rub3'-colorod resinous substance remained. The aqueous
extract obtained in the way described above gave no coloration with iron salts, and no
precipitate with gelatine and alum solution, potassio-mereuric iodide, or gold chloride
solutions. Fehling's solution was not reduced by boiling, though the aqueous ex-
tract was boiled with acid, then rendered alkaline before adding the copper tost. The
preceding tests gave negative results for gallic acid, tannin, alkaloids, and glucosides.
A portion of the aqueous extract was acidified and agitated successively with difter-
ent solvents, for glucosides, bitter principles, and alkaloids which may be removed
from solution by this means. The acid liquid was then rendered alkaline with am-
monia, and agitated successively with the same order of solvents that wore used Avith
the aciditlod liquid. No solids wore separated b}' those methods. The ethereal residue
insoluble in Avater was treated with alcohol ; and yielded traces of a resinous sub-
stance. The residue insoluble in Avater and alcohol Avas not dissolved by ether, acids
or alkalies.

Yuccal or the ethereal residue soluble in ether and alcohol Avas saponified, and the
soap boiled Avith load acetate. The yelloAV masses Avere collected on a filter and treated
Avith boiling ether, and the filtrate Avas slowly evaporated. The residue Avas a granular
solid. This substance Avas imperfectly purified by repeated boiling Avith ether, and a
solid of ci-ystalline structure oljtainod. It gaA'c an acid reaction AviLh litmus, and a rod
color Avith concentrated sulphuric acid. Tiie acid dissolved a substance enclosing the
crystals, leaving the structure of the latter uninjured and colored. Strong nitric acid
dissoh'cd the crystals witli no coloration. They aaxm-c soluble in absolute alcohol,
amyl alcohol, benzole, chloroform, glycerin, and a solution of alcoholic soda; soluble
in potassium iodide, potassium chromate, mercurous nitrate, cobalt nitrate, potassium
ferro-and-fcrri-cyanide solutions ; insoluble in ammonia and aqueous alkalies.

llie application of heal to the resin iicid mixture, or the addition of solvents to the mixture. The more concentrated
lliu acid the more rapid was the reaction. The application of heat also hastened the change, especially if a more dilute
acid was used in the mixture. Some solvents acted like heat by increasing the energy of the reactions. Alcohol and
ether were active solvents, and the reaction was atteodcd by the escape of nitrous fumes from the combination of alco-
hol or ether and nitric acid. Chloroform and benzole were indinurent. Ainyl alcolio! acted feebly.

264 A CHEMICAL STUDY OF

Yuccal was treated with sjiirit of different strengths, as a means of separating
resin acids if any were present. It was treated with 85 per cent, spirit ; an opaque
brown substance was left undissolved, which was soluble in absolute alcohol ; insolu-
ble in ether, and colored brown by concentrated sulphuric acid. The color was not
discharged by alcohol or ether. The 85 per cent, spirit solution was evaporated, and
the residue treated with 50 per cent, spirit and a small quantity of a brown residue
was insoluble. The 50 per cent, spii-it solution on evaporating left a non-crystalline,
transparent, I'eddish-colored solid ; acid to litmus. It was colored cherry-red by con-
centrated sulphuric acid, and slowly dissolved to a yellowish-red liquid.

Extract (3), tlie Gh'een Part of the Leaf.

The residual powder from the petroleum sjiirit maceration was thoroughly dried,
and again placed in the percolator. It was treated with Squibbs' stronger ether. The
extract was a deep-green colored liquid and fluorescent. The reaction was slightly
acid. Alcohol, benzole, and petroleum spirit added to the ethereal extract did not
cause a precipitation. An amorphous and green-colored residue was obtained on
evaporating the extract. The amount of total solids Avas estimated from a definite
volume of the extract, which was evaporated, dried, and weighed.

TOTAL SOLIDS.

Mhereal residue dried al 100^ C , 1.25 per cent of solids.

" 110° C l.li '

0.11 " " loss.

The ethereal residue was brought into a state of fine division and treated with
water. The amount of total solids soluble in ether and water was 0.34 per cent. The
aqueous extract was neutral in reaction. It was faintly colored and slightly bitter to
the taste. It was not colored by iron salts nor precipitated with alum and gelatine
solution, showing absence of gallic acid or tannin. Copper solutions were not reduced,
indicating absence of glucosides, though the precaution was observed of boiling the
aqueous extract with acid and rendering alkaline before adding the copper solution.
The aqueous extract was agitated with acetic ether and a distinctly crystalline resi-
due separated. Under the microscope these crystals were white, needle-shaped and
arranged in bundles. They did not respond to tests for gallic acid. Potash solution
formed a yellow mixture Avith the crystals. The color was discharged by a drop of
hydrochloric acid. Chloroform did not dissolve any substance from the ethereal
residue. The ethereal residue was treated with acidulated water and tested negatively
for alkaloids.

The ethereal residue insoluble in water was treated with alcohol. The amount of

YUCCA ANGUSTIFOLIA. 265

substances insoluble in water, and soluble in ether and alcohol was 0.15 per cent. The
alcoholic solution was evaporated, and the residue was crystalline in structure.
Concentrated sulphuric acid imperfectly dissolved it, and gave a reddish-yellow color
i-caction ; acetic ether discolored the solution. The alcoholic residue was insoluble in
acetic ether, cold and boiling aqueous alkalies ; soluble in chloroform. It saponified
with alcoholic soda.

The amount of the ethereal reskhie insoluble in water and alcohol was 0.65 per cent.
It was not soluble in alcoholic or aqueous soda. This would indicate a resin anh}'-
dride. Concentrated sulphuric acid gave no color reaction with it ; and a mixture of
sulphuric acid and cane sugar dissolved the residue.

The ethereal residue on treating with cold ether was not entirely soluble in it. It
was soluble in chloroform, benzoic, and carbon di sulphide ; incompletely soluble in cold
alcohol, and insoluble in amyl alcohol. The ethereal residite was treated with 95 per
cent, alcohol, in which it was slightly soluble. A turbidity foi-med in the alcoholic so-
lution on adding lead acetate, ferric chloride, ammonium hydrate, and sulphuric acid ;
it did not clear up on warming. Hydrochloric acid made a muddy mixture with the
alcoholic solution. The ethereal residue yvas not entirely soluble in acetic ether ; the
latter separated coloring matter from it. The ethereal residue, insoluble in acetic ether
and freed from coloring matter (chlorophyll), was a resinous substance. It melted at
80' C. The resin was boiled with absolute alcohol, and on throwing the alcoholic so-
lution into cold water it was precipitated as a white cloud. It was not saponified.

Extract (4), Yellow Base of Leaf.
The residual powder from the petroleum spirit maceration was dried and extracted
with stronger ether. The ether extract was a turbid yellow liquid ; slightly acid in reac-
tion. On evaporating the ethereal extract Sit the ordinary temperature a reddish-yellow
granular solid remained. It melted at 79' C For the determination of total solids
extracted, a definite volume of the extract was evaporated, dried, and weighed.

I.

TOTAL SOLIDS.

Ethereal residue dried at 10(P C 1.7 per cent of solids.

110=C ^7

0.0 " " loss.

The ethereal residue was treated successively with distilled water, alcohol, and

ether.

II.

Substances souble in elher and water ^-^ P*"' '^''"''

alcohol f'

" insoluble in water and " '^

Total solids ^-^

2G6

A CHEMICAL STUDY OF

The aqueous extract gave a neutral reaction with litmus. I^egative results fol-
lowed examination for tannin, gallic acid, gliicosides, alkaloids, and anj^ compounds
containing nitrogen.

The ethereal residue (the residue insoluble in water) was an opaque reddish-yellow
colored substance, and was identified as a resin. It melted at 79° C. It was insoluble
in ether, benzole, chloioform, and acetic ether ; incompletely soluble in cold absolute
alcohol, amyl-alcohol, carbon di-suljjhide, and oil of turpentine. It was soluble in
aqueous and alcoholic soda. On boiling with them, it was saponified. Concentrated
sulphuric acid dissolved the resin and colored it a yellowish-brown. Chloroform
formed a turbid mixture with the acid solution. The action of strong nitric acid on
the resin was slow. The resin was incompletely soluble in 95 per cent, alcohol. Lead
acetate gave a cloudiness with the alcoholic solution which increased on boiling. Ferric
chloride thickened the alcoholic solution, and on boiling it gave a yellow precipitate
which was insoluble in acids, alkalies, absolute alcohol, and acetic ether. The chloro-
form extract gave no coloration with bromine solution.

SUMMARY II.

Ethereal Extracts.

1. The bark of the root.

2. " wood "

3. " green leaf.

4. " yellow base of leaf.

Snlidft er-
tratted.

3.16 p. ct.

1.70 "

1.25 "
1.70 "

Character of
residue.

chlorophyll J

Reaction with
litmus.

Melting Specific
Poult, ^gravitft

Substances

soluble in

ether and

water.

Substnnces

soluble in

alcohol

and water.

acid
slightly "

11 (1

70° C.
70° C.

80O C.

79° C.

1.091
1.091

traces

0.34 p. ct.
0.80 "

traces

0.15 p. ct.
0.40 "

Substances

soluble inily

in ether.

O.Go p. ct.
0.50 "

The residues from the ether extracts (1) and (2) of the bark and of the wood of the
root contained I'csins which were identified as the same compound. They correspond
in color, melting point, specific gravity, solubilities, and reactions. The resin is a
transparent, ruby-colored substance, crystalline in structure, and of a softer consist-
ency than ordinaiy resin. It was examined by Plirsehsohn's scheme.* It diftered
from all described resins in its reactions Avith the reagents used to identify them. It
is proposed to name it yuccal.f

Yuccal is imperfectly soluble in 95 per cent, alcohol ; soluble in boiling absolute
alcohol, in cold ethei", and amyl alcohol ; not appreciably soluble in chloroform, benzole,
carbon di-sulphide, and alcoholic ammonia. Cold acetic ether dissolved the coloring
matter from the resin, leaving a colorless solid. Hot acetic ether dissolved it perfectl3^
Yuccal when heated on platinum foil gave ofl' as it burned a pleasant and aromatic

* Loc. cit.

fSec foot-note, ether extract (2).

YirCCA ANGUSTIFOLIA. 267

odor. Tests failed to show the presence of benzoic or cimiamic acids. A blood-red
color reaction was obtained by warming yuccal with a ci-ystal of ammonium molybdate
and a few drops of strong nitric acid. AVarm dilute nitric acid dissolved the resin, color-
less; cold nitric acid gave a brownish-green color reaction. Yuccal was mixed with
concentrated nitric acid and heated. After some time had passed, an energetic reac-
tion occurred and nitrous fumes were given off. A yellowish-brown residue was one of
the products of the reaction. This residue was almost insoluble in water, or acids. It
was soluble in alcohol and potassium hydrate.*

As the ether extracts (1) and (2) were concentrated, white needle-shaped crystals
appeared floating in the liquids, whose physical structure, and insolubility in water
and acetic ether, suggested identical substances. The crystals separated fiom yuccal
by the lead acetate method, already described, have not been sufficiently studied to
identify them with the white needle-shaped crystals in the ether extracts nor with any
class of chemical compounds. However, the absence of gallic acid, glucosides, and
alkaloids in the acpieous extracts from the ethereal residues, would show that the
crystals separated from yuccal are a constituent part of the resin.

The experiments with spirit of different strengths are only of value, so far as they
were carried out, in showing the possibility of separating the resin into distinct jiai'ts.f

Tannin was not present in these ethereal extracts.

Ethereal extract (3) was green-colored and fluorescent iVom the chlorophyll of the
leaves. On evaporating, the ethereal residue was amorphous and of a green color.
The aqueous extract obtained from treating this ethereal residue was neutral in reaction
and bitter to the taste. It contained neither gallic acid, tannin, noi- glucosides. It was
agitated with acetic ether, and the solvent removed a solid, which undei- the micio-
scopc ])roved to be white needle-shaped crystals arranged iu bundles. Potash solu-
tion formed a yellow-colored mixture with the crystals, hydrochloric acid discharged
the color. The sulyect has been too little studied to state definitely if these crystals
are or are not identical with the ciystals found iu the ethereal extracts {1) and (2). But
it should be noted, that unless the crystals from Ethereal extract (3) arc brought into
aqueous solution mechanically b}' some compound not present in ethereal extract.'f^l), and
(2), the indications are in favor of tlio crystals from (3) not being identical with them;
foi' the crystals from (1) and (2) were insoluble in water and not removed by acetic
ether.

* See fool-note 3, ether extract (2).

f The amount of material on wliicli these experiments, as well as others described in this paper, were tried, was
too small in quantity for me to obtain more conclusive results. The facts which have been ascertained will serve as a
guide in future investigations.

A. P. s. — VOL. x\i. 2n.

268 A CHEMICAL STUDY OF

The ethereal residue was treated with acidulated water and tested negatively for
alkaloids.

The ethereal residue insoluble in water was a mixture of two resins (I) and (II).
The one (I) was dissolved by absolute alcohol, the other (II) was mostly soluble in
ether. The alcoholic residue was crystalline. It was insoUiblc in acetic ether, but
was saponified with alcoholic soda. The ether residue was a resin anhydride ; it was
insoluble in alcohol, and in alcoholic or aqueous alkalies.

The amorphous and green-colored ethereal residue was not entirely redissolvcd by
cold ether. It was soluble in chloroform, benzole and carbon di-sulphide ; incompletely
soluble in cold alcohol, and insoluble in amyl alcohol. It was slightly soluble in 95 per
cent, alcohol and in acetic ether. 1 he latter separated the green coloi'ing matter fi'om
it. The resinous mass insoluble in acetic ether melted at 80° C. It was not saponi-
fied. This resinous mass insoluble in acetic ether is a mixture of the two resins just
described (I) and (II). It was noticeable that the resinous mass was not saponified.
Resin (I) was saponified. Resin (II) did not saponify, and as this resin exceeded in
amount by 0.5 per cent, resin (I), it would show that a certain percentage of resin
anh3'dride in a mixture of two resins forbids the saponification of the mixture.

It was not determined if the crystals dissolved by water and separated by acetic
ether were a part of resin (I) or resin (II) or an independent compound.

Ethereal extract (4) Avas a turbid yellow liquid. On evaporating, a i-eddish-yellow
granular solid i-emained. The extract from the aqueous treatment was tested Avith
negative results for tannin, gallic acid, glucosides, and alkaloids. The ethereal residue
insoluble in water was identified as a i-esin. It was soluble in ether, benzole, chloro-
form, and acetic ether; incompletely soluble in cold absolute alcohol, amyl alcohol,
cai'bon di-sulphide, and oil of tui-pentine. It was saponified. A resin was extracted
by boiling absolute alcohol from the residual powder of the leaves (the yellow base)
which was identified as the same resin, and the name of pyrophreal* was proposed
for it.

I. Resins (1) and (2) are identical substances (yuccal).

II. Ethereal residue (3) is a mixture of two. resins, and a ci-ystalline principle
soluble in water.

III. Resin (4) pyrophteal is identical with a resin found in alcoholic extract (4).

ALCOHOLIC EXTRACTS.

Extract (1), Barh of the Hoot.

The residual powder from the ether extraction was dried, and replaced in the
percolator. The maceration was conducted at the boiling temperature of alcohol.

* Pyroi)liu'!il. Science, September 11, 1885.

TUCCA ANP.USTIFOI.IA. 2G9

Squibb's stronger alcohol was used. A dark red-colored liquid was extracted. It
was neutral in reaction with litmus. The alcoJiolic extract was evaporated in a current
of carbonic acid. The residue was non-ci-ystalline and of a I'cd color. A definite volume
of the alcoholic extract was evaporated, dried until the weight remained constant, and
the residue incinerated in a weighed platinum dish and the ash estimated.

I.

TOTAL SOLIDS.

Akoliolk residue dried at lOOO C 9.25 per cent.

" " " lioo C 0.25

" fisli 0.2

The alcoJiolic residue was treated with distilled water, and a definite volume of
the extract was evaporated, dried, and weighed. The alcoholic residue insoluble in
water was treated with water containing ammonia (one part in fifty). This ammoni-
acal extract was evaporated with excess of acetic acid, and the residue rinsed with a
little water on a filter, dried, and weighed. The dried aqueous extract insoluble in

ammonia was then estimated.

II.

Distilled water residue ;'.'-'2 per cent.

Ammonia" " •">.-t3 "

Insoluble '• " O.fiO "

Total solids 9.25

The aqueous extract from the alcoholic residue was studied as follows : It was
not colored by a ferroso-ferric salt nor precipitated by gelatine and alum solutions,
showing absence of gallic acid and tannin. A portion of the aqueous extract was
acidified with sulphuric acid and agitated successively with petroleum spirit, benzole,
chloroform, and amyl alcohol. The acidified liquid was rendered alkaline by ammonia
and agitated with the solvents in the same order. Petroleum spirit removed from the
acidified solution traces of an amorphous residue, soluble in sulphuric acid and caustic
soda. Benzole and chloroform separated no substances from the solution. As the
amyl alcohol solution was evaporating white needle-shaped crystals were seen fioating
in the liquid. On drying the residue they were decomposed and melted, leaving a dai k-
colored liquid. Several attempts were made to dry these crystals, without success.
A few of the crystals were recovered from the solution, and tested for alkaloids; no
reactions were obtained with the usual reagents for them.

Glucose was estimated from the aqueous extract. The licpiid was heated over a
water bath with Fehling's solution, and the pi-ecii)itated red cui)rous o.vide was thrown
upon a weighed filter, dried, and incinerated. The glucose was estimated gravimetri-

270 A CHEMICAT. STUDY OF

cally by calculating the amount of cupric oxide. It yielded 0.619 per cent.' A portion
of the aqueous extract was boiled with acid, neutralized, and heated over a water bath
with Fehling's solution to calculate, by difference, saccharose or other reducible com-
l)ounds, and b}' this method 0.18 per cent, was obtained.

The alcoholic extract was described as being deeply colored. This coloring prin-
ciple-" Avas completely precipitated by sub-acetate of lead. The lead precipitate was
collected on a filter, suspended iu water, and decomposed by sulphuretted hydrogen,
filtered, and the filtrate freed from all odoi-. It was allowed to evaporate slowly
over sulphuric acid. The I'esidue was a brownish-gi-ay mass, interspersed with
fine crystals which radiated from a nucleus. The mass was weighed and gave 3.27
per cent, of solids. Another portion of the alcoholic extract was agitated with water
and acetic ether. The coloring matter was taken up by the acetic ether, and on
evaporating a red-colored substance was recovered. It was dried and weighed, yield-
ing 2.2 per cent. This red-colored residue was perfectly soluble in cold water. This
solution was tested with the following reagents : It gave with potassium bichromate
a creamy-colored precipitate ; ferric chloride, a yellowish-green precipitate ; ferrous sul-
phate a reddish-brown precipitate ; stannous chloride, no precipitate, a yellow cloudy
liquid; alum, a cloudy solution; neutral acetate of lead, a slight precipitate. The
red color of the colorino- matter was brought out on addition of alkalies. It was
destroyed by acids.

Extract (2), Wood of the Root

The residual powder from the ether treatment was dried and macerated with
Squibb's stronger alcohol. The alcoholic extract was neutral in reaction ; when warm
it was a clear reddish-golden-colored liquid. On cooling, a creamy- white solid settled
at the bottom of the flask. This substance was soluble iu water, and was identified
as saponin by the usual tests for it. A definite volume of the alcoholic extract was
evaporated in a current of carbonic acid, dried, and weighed. The residue was incin-
erated iu a weighed platinum crucible for the ash determination.

TOTAL SOLIDS.

Alcoholic residue dried at 100° C 14.3 per cent.

" " " IIOOC 14.3

" " ash 00.1

The alcoholic residue was treated with cold water in which it Avas soluble. A
cloudy solution was formed, and on shaking, it became frothy, and presented the
appearance of an emulsion. It was allowed to stand for several daj's to see if the
resinous matter separated, but the emulsion was permanent, as no separation had

* A red crystalline coloring matter. Science, September 11, 1885.

YUCCA ANGUSTIFOLIA. 271

taken place. The craulsificd li(jiiid was agitated witli acetic ether, and this solvent
readily separated most of the resin from the aqueous i)ortion. The water extract was
then evaporated to di-yness and redissolved in watei'. Gelatine and alum solution did
not precipitate the extract, showing absence of tannin ; no coloration w ilh iron salts,
absence of gallic acid; negative results followed tests lor alkaloids; the aqueous
extract was boiled witli ijotash and no ammonia fumes were formed; addinjr irold
chloride, and potassio-mercuric iodide solutions to the extract gave no iireciijitate. A.
measured portion of the aqueous extract was acidified with sulphuric acid, and
agitated successively w'ith petroleum spirit, benzole, and chloroform. The solvents
were evaporated; peti'oleum si)irit removed 0.01 per cent of a resinous substance,
imperfectly soluble in cold and boiling aqueous alkalies, dissolved by sulphuric acid
with a red coloration; chloroform left a brownish residue which, on weighing, yielded
0.4 {)er cent. This residue was moistened with a few drops of concentrated sulphuric
acid, and changed to a red-violet color characteristic of saponin.

A certain portion of the aqueous extract w^as rendered alkaline, and heated over a
water bath with Fehling's solution. The i)recipitated co])per was collected on a
weighed filter, dried, and incinerated, and the glucose estimated gravi metrically from
it. It yielded 1.592 per cent. .Vnolher portion of the aqueous extract was acidified,
boiled, and potash added until the solution was alkaline to litmus jxaper, then the
li(^uid was mixed with Fehling's solution and heated ovei' a water bath. The per-
centage of saccharose or other substances which reduced the copper was calculated by
dilference. It amounted to 0.929 per cent.

The resin separated by acetic ether was an opaque substance, greenish-yellow in
coloi-, and insoluble in ether. The resin was dissolved in water and frothed on shak-
ing. The emulsion in this ease was not quite so permanent, as a slight resinous sedi-
ment settled after a time, possibly due to changes in the resin through oxidation.

Extract (3), the Green Part of the Leaf.

The dried residual powder was raacei'ated by the aid of heat with Squibb's
stronger alcohol. When wai-m the alcoholic extract was clear, but on cooling the solu-
tion became cloudy, and a creamy-white fine i)recipitate settled. The alcoholic extract
was neutral in reaction. It was evaporated in a current of carbonic acid, dried, and
weighed. A certain part of the residue was incinerated and the ash determined.

TOTAL SOLIDS.

Alcoholic retiduc dried at 100° C 3.80 i)cr cent.

" liooc :i.80 ••

ash 0.15

272 A CHEMICAL STUDY OF

The alcoliolic residue was ti-eated with cold distilled water. It had a slightly acid
reaction with litmus. An emulsion was formed on the addition of water to the alco-
liolic residue. A measured quantity of it was evaporated, dried, and weighed. It
amounted to 3.4 per cent., 0.4 per cent, of the alcoholic residue was insoluble in water.
Tannin, gallic acid, and alkaloids were tested for and with negative results.

The liquid fi-om the aqueous treatment of the alcoholic residue was rendered alka-
line, and boiled with Fehling's solution, and there was no reduction. Boettger's bis-
muth test was also tried and with negative results. The aqueous portion was boiled
with acid and examined in the usual way for glucosides; the results were negative.

One volume of the aqueous solution was mixed with three volumes of stronger
alcohol. It was placed on ice, and after some time a white precipitate formed. The
precipitate was collected and dissolved in water. It frothed on shaking. On addition
of a concentrated solution of caustic baryta, a creamy-white precipitate of saponin-
baryta was obtained. Sulphuric acid gave the usual red-violet color reaction with the
precipitate from the alcoholic aqueous solution.

The method of successive agitation of an aqueous extract with solvents already
described was followed. Petroleum spirit on evaporating left a resinous substance.
The residue separated by chloroform from an acidified solution was a brownish-
colored substance. It was soluble in water, and fi-othed on shaking. It was colored
i-ed-violct by sulphuric acid, and the aqueous solution was precipitated by barium
hydrate. Chloroform sejiarated a brownish solid from an alkaline aqueous solution.
It was precipitated by barium, colored red-violet by sulphuric acid, and its aqueous
solution frothed on shaking. This brownish i-esidue was identified as saponin.

Extract (4), tJie Yellow Base of the Leaf.

The residual powder, dried from all traces of ether, was macerated with hot alcohol.
The alcoholic extract was a currant-colored liquid, and slightly acid in reaction. The
liquid became clear on standing, and a creamy-white solid, identified as saponin,
scpai'ated fi'om it. The alcoholic extract was evaporated, dried, and weighed, and the
ash of the residue was estimated.

TOTAl, SOLIDS.

AlcohoUr residue dried at 100"^ C 4.30 i)or cent.

" llOOC 4.30 ■■

ash 0.05

The alcoholic residue was treated with cold distilled water. The solution was
slightly colored, and faintl}' acid in reaction. The absence of gallic acid, tannin, and
alkaloids, Avas determined by negative results with iron salts, gelatine and alum solu-

TUCCA ANGUSTIFOLIA.

273

tion, gold chloride, ;uul potassio-mercuric solutions. Acetate of" lead caused no pre-
cipitation. Fehling's solution detected a trace of glucose.

An imperfect emulsion formed on adding water to the alcoholic residue. Upon
standing, the resin settled ; the licpiid was filtered several times, and the greater part
of the resin collected. It was an opaque reddish-3-ellow-colored substance. It had
the same melting point (79° C), solubilities, and physical appearance as the resin of
ether extract (4). The resin was examined by llirschsohn's scheme. It differed in
character from the many resins described by that author, and it is proposed to name
it pyrophajal.*

P^'rophical was slightly soluble in ether, and 9j per cent, alcohol ; soluble in ben-
zole, chloroform, and acetic ether ; incompletely soluble in cold absolute alcohol,
amyl alcohol, carbon di-sulphide, and oil of turpentine. It was saponified with aqueous
and alcoholic soda. The ethereal resin solution was cloudy. The alcoholic resin solu-
tion gave a precipitate with lead acetate which did not disappeai- on boiling; ferric
chloride and aqueous ammonia formed turbid mixtures with it. The chloroform resin
solution was not affected by bromine solution. The petroleum-ether-resin solution
turned to a turbid mixtuie on adding iodine solution. Alcohol containing hydro-
chloric acid was not colored by the resin. Sulphuric acid and alcohol gave a turbid
brown mixture with it, and sodium carbonate solution was colored pale brown when

cold or on wai'minir.

SUJIMARY III.
Alcoholic Extracts.

^UiU extracted.

1. The Imrk of the root 9.2.") p. ct. 0.20 p. ct. ash

2. " wood " ....

'A. " green leaf

4. " yellow base of leaf. .

14.:!0

3.80

! 4.30

O.IO
0.15
0.05

Character of
reiidue.

I C red color- |
} ing mutter >
( crystalline j

resin, saponin

.resins, "

Iresin, " I

Reaction wUh
litmus.

neutral

slightly acid

iQunnlitaliie cili-
Quantitative c.ili.' '""""" "/lacclia.
mution o/gluciac. roseor other redu-
cible compounds.

O.GlOp. Ct. 0.180 p. Ct.

1..593
none
traces

0.029
none
traces

Extract (1).

My attention was not directed to the presence of saponin in extract (1), for the
characteristic properties which it imparted to extracts (2), (8), and (4), were absent;
but, it was evident that saponin was ijrcsenl in the bark, for on boiling the latter in
distilled water, the presence of the compound was indicated. The .solution frothed on
shaking, and by adding a concentrated solution of caustic baryta, sapoiiin-baryta was
precipitated.f

• Pyroplucal. Science, September 11, 188.5.

\ Saponin in the bark of Yucca anguiU/olia. Science, September 11, 18H5.

274 A CHEMICAL STUDY OF

A coloring matter* contained in tlie bark was extracted, and imparted to the
alcoholic extract a bi-illiant red color. It was precipitated by sub-acetate of lead, and
tlie lead precipitate suspended in water and decomposed by sulphuretted hydrogen.
The lead sulphide filtrate was evaporated over a water bath until the odor of sul-
l)huretted hydrogen was expelled, and the concentrated liquid was placed over sul-
phuric acid to evaporate slowly. A crystalline residue was obtained. On addition of
alkalies to the colorless lead sulphide filtrate the red color of the original solution was
developed. Acid discharged the color. Acetic ether took up the red colored sub-
stance. The acetic ether residue was a red uniform solid, and soluble in water. It
was precipitated from the aqueous solution by sub-acetate of lead, potassium bichro-
mate, ferric chloride, ferrous sulphate, and it was clouded by alum, and stannous
chloride solutions.

Tannin, gallic acid, and alkaloids were absent.

Amyl alcohol separated from the acidified aqueous extract white needle-shaped
crystals. It was not determined if these crystals were the same as those of the color-
ing matter.

Extracts (2) ,'(3), r/juZ (4).

Extracts (2) and (3) when warm were clear, and on cooling a creamy-white solid
separated. Extract (4), if warmed, was turbid, and as the liquid cooled, a creamy-white
substance remained at the bottom of the flask, and the supernatant fluid became clear.
This creamy-white substance was identified in each of the extracts as saponiu.f

The results following an aqueous treatment of alcoholic residues (2) and (3), were
noticeable. The residues were dissolved, and by shaking the mixtures, emulsified.
This emulsion was permanent, as no resinous matter separated on standing several
days. The emulsion was agitated with acetic ether, and by this means, most of the resin
and saponin were separated from the aqueous portion. The saponin was removed
mechanically with the resin, as it is almost insoluble in acetic ether. The resin-sapo-
nin mass was insohible in ether, soluble in water. The solution frotlied on shaking
and emulsified, but the emulsion was not so permanent as in the first case, for a resi-
nous sediment settled after a time. Chloroform separated saponin from an acidified
acjueous solution, and also from an alkaline aqueous solution of the residues; and the
red-violet saponin reaction with concentrated sulphuric acid was obtained.

The solubility in water of the alcoholic residues (2), (3), and (4), and the result-
ing emulsion were unusual, and explicable by the facts collected from a series of ex-

* A reil crystiillino coloring matter. Sfieiioc, Sci)tember li, 1885.

•f Saponin iu the wood of Uie root ami leaves. Science, September 11, 1885.

TUCCA ANCrUSTI FOLIA. 275

periinents with resins and saponin, since I had successfully emulsified resins with
aqueous antl alcoholic saponin solutions.*

By hot alcoholic treatment the ?/i<cca yielded a residue of saponin and resin which
became emulsified on the addition of water, giving results identical witli those of the
resins above described.f

Extract (4) contained a resin. It was an opaque reddish-yellow colored sub-
stance, and it differed, by its reactions, from the many resin classes given in Ilirsch-
sohii's scheme. It is proposed to name it pyroph;eal.:|; A resin having the same melt-
ing point, solubilities, physical appearance, and chemical reactions, was discovered in
the ethereal extract (I). It was identified as the same compound for which the name
pyrophiTial is proposed.

Tarmin, gallic acid, and alkaloids were not detected in extracts (2), (3), and (4).
In extract (3) glucose was not found.

Tlie Solids of the Alcoholic Extracts.
I. A red coloring matter (crystalline).
II. A new resin (yuccal).§

III. A second new resin (pyropha3al).§

IV. A mixture of a crystalline resin and a resin anhydride.
V. Saponin. §

VI. Glucose, and saccharose or other rcducil)le compounds.
VII. Ash.

AQUEOUS EXTltACTS.

Extract (1), 13or1c of the Hoot.

The residual powder was thoroughly dried from alcohol. It was returned to the
percolator, and cold distilled water added until a definite amount had been used.
The aqueous extract was dark colored, and of a faintly acid reaction. A certain (piaii-
tity of the extract was evaporated, di'ied, and weighed. Fr(jin a known weight of the
aqueons residue, the ash was calculated. The incineration was conduced in a covered
porcelain crucible of known weight.

•The same kinds of resins were used in Uieso experiments as in those with whicli I determined tlie solubility of
resins in acetic elher. See footnote (2), ethereal extract (2).

fit was not until a later date following the lime of these experiments that I found a reference to suponin-rcsin
emulsion in L'OfBcine ou Repertoire General do Pharmacie Pratique, par Dorvault. Iluilii-me Edition, Paris, 1872, p.
81G. Also, refer to examination of the Turra ani/ii»lifoUn, hy H. C. I)c S. .Miliolt, published in the Medical and
Surgical Reporter, Philadelphia, September 12, lSH."i, |i:if;e liOI.

JPyrophtcal, loc. cit.

§ Science, September 11, 1885, page 210. Abstract of a paper on the chemical Btuiy of Yueea anguitifolia, hy
H. C. De 8. Abbott.

A. P. S. — VOL. XVI.' 2r.

276 A CHEMICAL STUDY OF

TOTAL SOLIDS.

Aqueous residue dried between 100° 0 and IW^ C 4.00 per cent.

ash 2.05 "

Oum.

One volume of the aqueous extract was mixed with two volumes of Squibb's
stronger alcohol. The mixture was kept in a cool place for twenty-four hours, and the
precipitate which had formed was collected on a weighed filter, Avashed with 06 per
cent, alcohol, dried, and weighed. The precipitate and filter were incinerated in a
weighed porcelain crucible, and the weight of the filter being deducted, the percentage
of ash was determined.

Weight of precipitate by stronger alcohol yielded 2.0 per cent.

" " ash yielded 0.3 "

Another poi'tion of the aqueous extract was precipitated by stronger alcohol, and
the precipitate consisted of gum and albuminous substances. It was incompletely
soluble in water. The soluble matter was gum, it was recovered from solution by
evaporating the liquid to dryness. The gummy residue was almost completely solu-
ble in cold water. It was precii^itated from a concentrated aqueous solution by
stronger alcohol ; basic acetate of lead precipitated it as a fiocculent precipitate. Borax
did not thicken the gum solution, and ferric chloride and sodium chloride solutions did
not precipitate it. The gum was boiled with dilute acid, and heated over a water bath
with Fehling's solution which it reduced. A few drops of hydrochloric acid and
stronger alcohol were mixed with the concentrated gum solution for the separation of
arabin. It was not separated.

Carhliydrates.

The filtrate and wash alcohol from the gum precipitate were mixed, and evapo-
rated to a syrupy consistency at a temperature of 70° to 8U° C. The concentrated sohi-
tion was treated Avith four volumes of stronger alcohol, and the resulting precipitate
of carbhydrates rapidly filtered off. It was soluble in water. It was not precijji-
tated from aqueous solution by basic acetate of lead, and by this means it was dis-
tinguished from vegetable mucilage. The carhhydrates were boiled with dilute acid,
and the solution was rendered alkaline, and heated over a Avater bath Avith Fehling's
solution. The latter was reduced. The percentage of carbhydrates as estimated,
amounted to 0.2 per cent. An aqueous carbhydrafe solution Avas mixed Avith a solu-
tion of barium in 40 i)er cent, alcohol. It yielded no precipitate.

TUCCA ANGUSTIFOLIA. 277

Carhhy (Irate Filtrate.

The carhlnjdrate filtrate was concentrated at a low temperature in a ciu-rent of
carbonic acid until the alcohol was dissipated. The residue was examined for glucose,
organic acids, saponin, and tannin. Traces of glucose were detected qualitatively by
Fehling's test; the amount of cui^rous oxide present was too small to estimate gravi-
metrically. A i)ai-t of the carhhydrate filtrate residue was boiled with 83 per cent,
alcohol, and filtered Avhilc hot. On cooling, a precipitate formed. This precipitate was
identified as saponin. It was almost insoluble in stronger alcohol. Baryta-water pre-
cipitated it from aqx;eous solution. Its aqueous solutions frothed on shaking. When
agitated with chloroform and on evaporating the chlorofonn solution, a light-colored
residue was obtained. A few drops of concentrated sulj^huric acid mixed with it gave
a reddish-A-iolet color reaction. Another portion of the carhhydrate -filtrate residue was
precipitated with neutral acetate of lead and filtered. The precipitate was suspended
in water, decomposed by sulphuretted hydrogen, and the lead sulphide filtrate evapo-
rated over a water-bath to expel all odor of sulphuretted hydrogen. The liquid was
cooled and lime-water added until the reaction was alkaline to litmus. A turbidity
forinc<l when the lime-water was added to the filtrate, and was not entirely cleared
on the addition of dilute acetic acid. A neutralized portion of the lead sulphide
filtrate gave a yellow precipitate with a ferrous salt. Oxalic acid by these tests was
indicated, and possibly other vegetable acids were present in the filtrate. The carh-
hydrate filtrate residue was examined for tannin, and with a negative result. Calcium
oxalate was separated.

Extract (2), the Wood of the Root.

The powder used in the alcoholic maceration was thoroughly dried, and replaced
In tlir poicolator. A measured quantity of cold distilled water was allowed to percolate
slowly tlirough the powder. The extract was colored, and slightly acid in reaction. A.
definite volume of the extract was evaporated, dried, and weighed. A known weight
of the residue was incinerated in a weighed covered porcelain crucil)le, and the ash
determined. The ash was white and incompletely soluble in water.

TOTAI, SOLIDS.

Aqueoui residue dried between lOO^ and llO'^ (' 12.10 per cent.

" ash 1.74

Oum.

A certain quantity of the a/piexnis extract was mixed with two volumes of stronger
alcohol (Squibb's). The mixture was allowc'l to stand ini- twenty-fnui- hours, and the

278 A CHEMICAL STUDY OF

precipitate which formed was collected on a weighed filter. It was dried, and weighed.
The precipitate and filter were incinerated in a weighed poi'celain covered crucible, and
the percentage of ash calculated.

Weight of precipitate b}' stronger alcoliol yielded l-'^O per cent.

" ash yielded 0-34 "

CarhJiydrates.

The filtrate and wash-alcohol fi-om the gum precipitate were concentrated at a low
temperature, and the residue was mixed with four volumes of stronger alcohol. The
precipitate was rapidly filtered off, and the percentage of carNiydrates calculated
gravimetrically in the usual way, from the amount of cupric oxide reduced from Feliling's
solution. It yielded 2.75 per cent.

Carhhydrate Filtrate.

A portion, representing a certain volume of the aqueovs extract, of the carhhydrate
■filtrate was evaporated, dried, and weighed. It yielded 7.65 per cent, of the total solids
of the aqueous extract residue. A lalo^yll weight of the carhhydrate filtrate residue was
dissolved in water and heated over a water-bath with Feliling's solution, and the
amount of glucose present estimated gravimetrically from the weight of the cupric
oxide. It was estimated as 4.47 per cent. Another portion of the carM?/(7r«fe^^^rafc
residue was boiled with 83 per cent, alcohol. A precipitate formed on cooling, which
was collected on a weighed filter, dried, and weighed. It yielded 1.98 per cent. The
83 per cent, alcohol precipitate was identified as saponin by the usual tests. The carh-
hydrate -filtrate residue was precipitated b}'- acetate of lead and the precipitate examined
qualitatively for organic acids. The lead precipitate was decomposed by sulphuretted
hydrogen and filtered, and the filtrate concentrated over a water-bath, and mixed
with lime-water until turbid. The turbidity did not clear on adding dilute acetic acid.

The aqueous extract was examined for tamiin, and with negative results. Calcium
oxalate was present.

Extract (3), the Qreen Fart of the Leaf.

The residual powder was dried from all traces of alcohol, and cold distilled water
was allowed to slowly percolate through the powder. The extract was colored, and
slightly acid in reaction. A definite volume of the extract was evaporated, dried, and
weighed, and tlic ash calculated from incinerating a known weight of the residue.

TOTAL SOLIDS.

A'pieons residue dried between 100° C and 1 10° 4.n.T per cent.

" ash 0.40 "

TUCCA ANGUSTIFOIilA. 270

Gum.

One volume of the aqiceoics extract was mixed with two voUimes of stronger
alcohol. The precipitate was collected after twenty-four hours, washed with 66 per
cent, alcohol, dried, and weighed. The precipitate and filter were incinerated, and the
ash estimated.

Weight of precipitate by stronger alcohol yielded 0.775 per cent

ash yielded 0.125

CarhJiydrates.

The gum fdtrate and wash alcohol were concentrated at a low tempe'rature, and
the residual liquid mixed with four volumes of stronger alcohol. The resulting precipi-
tate was rapidly filtered and collected. The percentage of carlliydrates was estimated
gravimetrically from a copper solution in the usual way. It amounted to 0.525 per

cent.

Carhhydrate Filtrate.

The fiUrafe was evaporated to dryness. A part of the residue was boiled with 83
per cent, alcohol. On cooling, a iirecijjitate formed. It was identified by the usual tests
as saponin. Another portion of the residue was precipitated with ])asic acetate of lead.
The lead precipitate was decomposed by sulphuretted hydrogen, the solution fil-
tered, and the filtrate evaporated over a water bath until all odor of sulphuretted
hydrogen was dissipated. A part of the lead sulphide filtrate was mixed with lime-
water, and a precipitate fornied not completely dissolved by acetic acid. The remain-
der of the lead sulphide filtrate was allowed to evaporate over sulphuric acid. The resi-
due consisted of a mass of fine crystals radiating from a centre.*

The crystals gave a very acid reaction when placed on moistened blue litmus
paper. They turned black and left a residue when heated on |il,ilimim foil, and the
residue was sloAvly dissolved by nitric or hydrochloric aciil. Tlic quantity of residue
was very small, and no effervescence was observed. The melting point of the crystals
was taken. A small quantity Avas placed in a tube Avith thin walls, and gradually
heated ; at 150° C. the substance sublimed, leaving a Avhite, cloudy stain on the inner
surfiice of the tube; at 190° C, this cloudy stain changed to a pale green spot, and
with increasing temperature to 210° C, no further change was noted. Dry sodium
carbonate was added to an aqueous solution of the crystals, and a slight effi-rvesccncc
Avas observed. Some iron A\-as scparate<l Avhich possibly Avas in combiiKiiion with the

• The material nsed in this analysis of the green part of the leaf was quite dry and powdered readily. A previous
examination of the/r«*A Uaten gave more satisfactory quantitative resulLs. A gum was extracted which promises to ho
of interest for a future study, and the crystals separated from the lead sulphide filtrate arc to be further investigated.

280 A CHEMICAL, STUDY OF

ciystallinc principle. jSTegative results followed tests for formates, acetates, malates,
tarti-ates, citrates, phosphates, oxalates, alkaloids, and glucosides.

The amount of glucose present in the aqueous extract was too small to determine
quantitatively. iN^egative results for tannin.

Extract (4), the Yellow Base of the Leaf.

The residual powder from the alcoholic maceration was dried and replaced in the
]iercolator. Cold distilled water was allowed to percolate slowly through the powder.
The aqueous extract was slightly acid in reaction. A known measure of it was evapo-
rated, dried, and weighed. The ash was determined from a part of the aqueous extract
residue.

TOTAL SOLIDS.

Aqueous residue dried between 100° aud 110"^ C 11.35 per cent.

ash 3.10

Oum.

One volume of the aqueous extract was mixed with two volumes of stronger
alcohol. The precipitate was collected, washed with 66 per cent, alcohol, dried, and
weighed. The ash was calculated from incinerating the precipitate, and deducting the
filter.

Weight of precipitate by stronger alcohol j'ielded 3.850 per cent.

ash yielded 0.G76

Carhhydrates.

The filtrate and wash alcohol from the gum precipitate were concentrated at a Ioav
temperature. The residual liquid was mixed with four volumes of stronger alcohol, when
a precipitate formed, and was rapidly filtered off. The carNiydrates were dissolved in
water, hoiled with dilute acid, and the liquid rendered alkaline and heated over a Avater
l)ath with Fehling's solution. The amount of carhhydrates was estimated gravimetric-
ally in the usual way. It gave 2.95 per cent.

CarhJiydrate Filtrate.

The filtrate residue was examined for glucose, and traces of it were present. The
filtrate residue was precipitated with acetate of lead, and the lead precipitate was dis-
solvofl in watci- and decomposed by sulphuretted hydrogen. The lead sulphide fil-
trate was tested qualitatively for organic acids, and a turbidity formed on adding to the
filtrate lime-water. It was not completely cleared by acetic acid.

Negative results followed tests with alcoholic methyl-violet solution for mineral
acids. The aqueous extract contained no tamiin. Calcium oxalate was determined in it.

\UCCA ANGUSTIFOLIA. 281

AQUEOUS MACERATIOX AT A TEMPERATURE OF 50° C. TO 60° C

The BarJc of the Hoot (1), the Wood of the Boot (2).

The powder (1) used in tlic cold water extraction was macerated with distilled
water heated between 50° and 60° C. The warm aqueous extract (1) was cooled and
mixed with three volumes of stronger alcohol. A precipitate formed, it was dried,
weighed, and the percentage estimated. It yielded 0.03 per cent.* The precipitate
was dissolved in warm water. On evaporating the filtrate, a >\hitr residue was oh-
tained. It was stained yellow by iodine.

The i)owder (2) from the cold water treatment Avas macerated in the Avarm water.
The tcann aqueous extract (2) Avas a dark colored liquid, indicating a coloring matter.
A certam measure of the extract Avas evaporated, and the solids estimated. It
amounted to 4 per cent. The percentage of solids precipitated from the extract by
stronger alcohol A\'as 0.25 per cent.

QUANTITATIVE ESTLAIATIOX OF SAPONlX-f

The tAA'O methods of Christophsohn and Otten for the rpuuititatiA'c estimation of
saponin Avere adopted. The Avood of the root Avas examined.

A. — Ten grams of the original poAvder Avere boiled Avith distilled Avatcr. The sapo-
nin was precipitated by baryta-water. After A\-eighing, it was ignited, and \\\v baryta
estimated as carbonate, calculated into oxide and deducted from the weight of the
saponin-baryta, the dilfcrence being llie Aveight of saponin.

B. — The saponin-baryta Avas decomposed by a(i<l and the weight of the sapogenin
was ascertamed and calculated to saponm.

Several estimations Avere made on tAvo specimens of the Yucca, collected at difter-
ent times of the year.

Mean percentage, A 8.9o per cent.

B 10.40t "

SUMMARY IV.

Aqueous Extracts.

SoliOi
extracted

Ath.

Qum.

Ash.

Qlucosc.

Saponin.

1. The bark of the root

2. " wood "

3. " green leaf

4. " yellow base of the leaf. .

4.00 p. ct.
12.10 '•

4.a5 "
11..35 "

2.fi5 p. ct.
1.74 •■
0.40 "
3.10 "

2.00 p. ct.
1.70 "
0.77 "
3.85 "

0.20 p. ct.
0.34 "
0.12 •'
0.67 "

traces

4.47 p. ct.

traces

A. 1 B.

8.!i.j p. ct.^ i6.40'p.

ct.

* Examination for Inulin, page 87, Plant Analysis, G. Dragendorff. English translation,
t Loc. cit., page 68.

i Examination of the Tueca angutlifolia, by II. 0. Du S. Abbott. The Jleaitiil and Surgical Reporter, Sept.
12, 1885, p. 301.

282 A CHEMICAL STUDY OF

The aqueous extracts contained gum, albuminous substances, carbhydrates, glu-
cose, sajwnin, organic acids, calcium oxalate, and no tannin, mineral acids, nor alka-
loids. Arabin was not separated from gum (1). Calcium oxalate was brought into
aqueous solution possibly by means of the organic acids or saponin. Needle-shaped
crystals were found in extract (3). They did not respond to tests for formates, ace-
tates, malates, citrates, tartrates, phosphates, oxalates, glucosides, and alkaloids.

Aqueous extracts of nO° C. to GO °C. from the bark and wood of the root contained
Inulin.

DILUTE CAUSTIC SODA EXTKACTS.

Extract (1), the Barl: of the Roof.

The residual powder insoluble in water was suspended whilst moist in a dilute
soda solution (0.1 to 0.2 per cent). After twenty-four hours the mixture was filtered.
One volume of the filtrate was acidified with acetic acid and mixed Avith three volumes
of 90 per cent, alcohol, and allowed to stand in the cool. The precipitate was collected,
washed Avith 75 per cent, alcohol, dried, and weighed, deducting ash. It consisted of
mucilaa'inous substances and albuminoids.

Weight of precipitate by 90 per cent, alcoliol yielded 0.85 per ceut.

ash yielded 0.2.5

Lassaigne's test showed the presence of albuminous substances.

The filtrate and wash alcohol from the 90 per cent, alcohol precipitate was evapo-
rated to dryness, and weighed, deducting the amount of soda acetate. It gave 0.24
per cent. The residue soluble in water was mixed with acetate of copper solution. A
very small quantity of alltuminous substances was precipitated by the reagent.

The allouminoids of the bark Avere estimated from the total nitrogen in one gram
of the original poAvder. It yielded 4.75 per cent of albuminoids.

The powder insoluble in dilute soda solution Avas Avashcd Avith distilled Avater.
The liquid was deeply colored. It Avas evaporated, and the amount of solids estimated.
It gave 1.3 per cent.

Extract (2), the Wood of the Root.

The poAvder insoluble in Avater Avas treated in the same Avay as in extract (1).
The filtered solution Avas mixed Avith 90 per cent, alcohol in the manner described. The
precipitate was estimated, deducting ash.

Weight of precipitate by 90 per cent, alcohol yielded 2.170 per cent.

ashyielded 0.25G

The filtrate from the 90 per cent, alcohol precipitate was treated Avith Avater, and
the soluble matter precipitated by copper acetate. The precipitate was collected, dried,

YUCCA ANGUSTIFOLIA. 283

■weighed, and ignited, the resulting oxide of copijer being deducted. It yielded 0,101 per
cent, of albuniinoid.s. A current of Avashcd cai'bonic acid was passed through the
dilute soda extract to determine the presence of globulin (vitcllin. myosin), and with
negative results. The albuminoids were determined from the total nitrogen in the
powdered wood. It amounted to 4,75 per cent. The total albuminoids in the leaves
gave 9,02 per cent,

DILUTE IIYDKOCIILORIC ACID KXTRACTS.

Extract (1), the BarJc of the Hoot.
The i)owder insoluble in ililute so<1a was washed with water and suspended in
water containing 1 per cent, of hydrochloric acid. The absence of the blue color which
the stai-ch granules assume when treated with iodine solution was determined by exam-
ining the bark under the microscope ; and consequently it was not looked for in the
extract. A qualitative test showed the presence of calcium phosphate, and calcium
oxalate, A measured quantity of iXw filtrate was neutralized with ammonia and mixed
with three volumes of 90 per cent, alcohol. The precipitate was collected on a weighed
filtei-, washed with GO per cent, alcohol, dried, and weighed. It was incinerated, and the
ash deducted from the precipitate.

The precipitate yielded rj.CO per cent.

ofash 0.98

Organic substance 4.23 "

The filtrate from the 90 per cent, alcohol precipitate was evaporated. The residue
was composed of ammonium chloride, from the reagents employed, and an organic sub-
stance having an odor like gum benzoin. It was agitated with ether and on evapo-
rating the solvent a white residue with an odor like benzoin was obtained. Sulphuric
acid gave a red color with it. The amount of this substance was calculated. It
gave 0.4.J per cent.

Extract (2), the Wood of the Boot.

The insoluble powder rrom llie diliile .soda maceration was washed with distilled
water, and suspended in water containing 1 per cent, of hydrochloric aci'l. The same
means were used as in extract (1) to determine the absence of starch in the wood of the
root. Parabin was also absent. Calcium oxalate was detected by qualitative tests.
A similar method, as employed for its estimation in extract (1), was used to determine
it quantitatively.

The precipitate yielded 0,"0") per cent.

" " ofash 0.1.5.') "

Organic substance 0. loO "

A. P. S.— VOL. XVI, 2 J.

284

A CHElvnCAL STUDY OF YUCCA ANGUSTIFOLIA.

Extract (3), (he Qreen Part of Leaf.

The powder used in the dilute soda maceration was Avashed with distilled water
and suspended in water containing 1 per cent, of hydrochloric acid. Iron and calcium
phosphate wei"e detected in the extract. The leaves were examined under the micro-
scope, and a blue color was developed by an aqueous solution of iodine, indicative of
starch granules. Starch was also present in the yellow base of the leaves.

TOT.Vr, QUANTITATIVE RESULTS.

(I.) The bark 0/ the
root.

(2.) The wood of the
root.

(3.) The green part of
leaf.

(4.) The yellow base of
leaf.

06.78 per cent.

17.38

01.24

03.16

09.25

04.03

01.09

01.30

05.05

11.67 per cent.

15.75

00.55

01.70

14.30

16.10

02.41

08.11 per cent.

05.75

02.20

01.25

03.80

04.35

37.00 per cent.
10.63

2 Total ash

3. Petroleum spirit extract

4. Ethereal extract

5 Alcoholic '*

01.10
01.70
04.30

11.35

8 W^asli ret^idiip ...

00.30

1

49.88

62.78

25.46 per^cent.

66.08 per cent.

Total albuminoids estimated on powder.

Quantitative saponin determination
on powder by A and B \

04.75

04.74

08.95 per cent.

B.
10.40

09.62

In my paper on The Chemical Study of Yucca angustfolia, read at Ann Arbor,
Mich., I stated what methods I had employed to separate sa])onin, and the properties
of the compound as observed in that plant. Since which time a further study of it
has induced me to withhold the notes used at Ann Arbor, from this, and to offer them,
with those collected later, in a separate and more complete publication.

I am indel)ted to Dr. F. M. Endlich, for his courteous consideration and kind-
ness in selecting and forwarding the fine specimens of Yucca which were used in these
analyses, and which were grown in the neighborhood of Lake Valley, jN^ew Mexico.
"Within a few weeks I have received, in addition, several hundi-ed pounds of the plant
from Dr. Endlich..

The investigations described in the preceding pages Avere conducted in the
chemical laboratory of the Philadelphia College of Pharmacy, from February to
August, 1885.

ARTICLE IV.

SYSTEMATIC CATALOGUE

OP THE

SPECIES OF VERTEBRATA FOUND IN THE BEDS OF THE PERMIAN EPOCH IN NORTH AMERICA

WITH NOTES AND DESCRIPTIONS.

BY E. D. COPE.

Read May 7, 1886.

PISCES.

SELACHII.

TnoR.\CODUS Cope ; Proceeds. Academy Philiulelphia, 1883, p. 108.

1. T. emydinus Cope ; 1. c, 109.
Janassa Munster.

2. J. gurUiana Cope ; Proc. Anier. Phil. Soc, 1877, p. 191. Eastern Illinois.

3. J. stri'jiliim Cope ; Amer. Naturalist, 1881, p. 163 ; S. Uiujuaformis Cope ; Proc. Amer. Phil. Soc, 1877, p. 53,

not of older authors. Eastern Illinois.

4. J. ordiana Cope. Texas.
ORTBACANTnns Agass.

5. 0. gracilis Newberry. Eastern Illinois.

6. O. (juadriseriatus Cope ; 1. c, p. 192. Eastern Illinois.

DiDYMODUS Cope ; Proc. Acad. Phila., 1883, p. 108 ; Proc. Amer. Phil. Soc, 1884, p. 573.

7. D. texensis Cope ; D. fcompressus Newb., Cope, 1. 1. c. c. Texas.

8. D. platypternus Cope ; 1. c Texas. *

DIPNOI.

Ctejjodus Agass.

9. C'.fossatut Cope ; Proc. Amer. Phil. Soc, 1877, p. 51. Eastern Illinois.

10. C. gurleian-u» Cope ; 1. c, p. 55. Eastern Illinois.

11. 0. periirrion Cope; Proc. Amer. Phil. Soc, 1878, p. 537. Texas.

12. C. parrectus Cope ; 1. c. Texas.

13. C. tabasentit Cope ; Proc. Acad. Phila., 1888, p. 110. East Illinois.

14. C. dialophm Cope ; Proc. Amer. Phil. Soc, 1878, p. 528. Texas.

15. C. pusHlus Cope ; Proc. Amer. Phil. Soc, 1.^77, p. 191. Eastern Illinois.
Pttonodcs Cope ; Proc Amer. Phil. Soc, 1877, p 193.

10. P. vinilocii Cope ; Proc. Acad. Phila., 1870, p. 410 Eastern Illinois.

17. P, paucicrittalus Cope ; Proc. Amer. Phil. Soc, 1877, p. 54. Eastern Illinois.

286 CATALOGUE OF THE SPECIES OF VERTEBEATA FOUND IN THE

Gnathorhiza Cope ; Proc. Amer. Phil. Soc, 1883, p. 639.

18. O. serrata Cope ; 1. c. Texas.
Ceratodds Agas.9.

19. C.favosus Cope ; Proc. Amer. Phil. Soc, 1884, p. 28. Texas.

TELEOSTOML*

ECTOSTEORHACHIS Cope ; Pal. Bull. No. 32, 1880, p. 19.

20. E. nitidus Cope ; 1. c. Texas.

21. E. ciceroniua Cope ; Proc. Amer. Phil. Soc., 1883, p. 628. Texas.

BATRACHIA.

GANOCEPHALA.

Trimerorhachis Cope ; Proc. Amer. Phil. Soc, 1878, p. .'321 ; 1880, p. 54.

22. T. insignis Cope ; 1. c, p.. 524. Texas.

23. T. bilobatus Cope; 1. c, 1883, p. 029. Texas.

RHACHITOMI.

Zatkaciiys Cope ; Proc. Amer. Phil. Soc, 1878, p. 523 et iufra.

24. Z. serratvs Cope ; 1. c et infra. Texas.

25. Z. apicalis Cope ; Amer. Naturalist, 1881, p. 1020, New Jlexico.
Ekyops Cope ; 1. c, 1877, p. 188.

26. K megaceplmlus Cope ; 1. c Hhachiiomus valens Cope ; 1. c, 1878, p. 526. Texas.

27. E. erythrolilicus Cope ; I. c, 1878, p. 515 [EpicordyUs] ; Traus. Amer. Phil. Soc, 188G, P!. I, fig. 1. Texas.

28. E.ferrieolus Cope ; 1. c, 1878, p. 521 (Parioxys). Texas.

29. E. reticulatus Cope ; Aracr. Naturalist, 1881, p. 1030. New Mexico.
AcHELOMA Cope ; Proc. Araer Phil. Soc, 1882, p. 455.

30. A. cximmiiisi Cope ; 1. c, 450. Texas.
Anisodexis Cope ; 1. c, 1882, p. 459.

31. A. imbriearius Cope ; I. c Texas.

STEGOCRPHALI.

DiPLOCAULUS Cope ; Proc Amer. Phil. Soc, 1877, p. 187 ; 1883, p. 451.
33. D. salamandroides Cope ; 1. c, 1877, p. 187. Eastera Illinois.

33. D. magnuornis Cope ; 1. c, 1882, p. 453. Texas.

EMBOLOMERI.

Cricotus Cope ; Proc. Acad. Phila., 1876, p. 405 ; Proc. Amer. Phil. Soc, 1884, p. 29.

34. e. heUroditus Cope ; Proc. Acad. Pliila., 1870, p. 405 ; Proc Amer. Phil. Soc, 1884, p. 29 ; Trans. Amer.

Phil. Soc, 1880, p. 247, PI. I, figs. 7-8. U. discophorua Cope ; Proc Amer. Phil. Soc, 1877, p. 186.
Eastern Illinois.

35. C. gibsoni Cope ; Proc. Amer. Phil. Soc, 1877, p. 185. Eastern Illinois.

30. C. crassidiscus Cope; Proc Amer. Phil. Soc, 1884, p. 29 ; C. netcroeliim Cope ; 1. c, 1878, p. 523 ; Araer.

Naturalist, 1884, p. 39. Texas.
37. C. hypantricus Cope ; Proc. Araer. Phil. Soc, 1884, p. 30 ; Transac Amer. Phil. Soc, 1880, p. 253, PI. I, figs.

2-0. Texas.

* Owen ; llyopnmata Cope ; Tclcostei el Qanoidei pars Miiller.

BEDS OF THE PERMIAN EPOCH IN NORTH AMKHICA. 287

KC.I'TII.IA.

TIIEKOMOUrUA.

Clepi<i/(lropidie.

? LTSOiinoriius Cope ; Proc. Amer. Phil. Soc, 1877, p. 187.

38. L. triearinatus Cope ; 1. c. Eastern Illinois.
Arcii.«:obelus Cope ; Proc. Araer. Phil. Soc, 1877, p. 1".)2.

39. A. teUicittui Cope ; 1. c. Eastern Illinois.
CLErsYDROPS Cope ; Proc. Acad. Phila. , 1876, p. 404.

40. C. eolletUi Cope ; 1. c. p. 407. Eastern Illinois.

41. C. vinsloBii Cope ; Proc. Amer. Phil. Soc., 1877, p. (ii. Eastern Illinois.

42. C. pedunculatus Cope ; 1. c, p. 63. Eastern Illinois.

43. C. nalalis Cope ; Proc. Amer. Phil. Soc, 1878, p. 509. Texas.

44. C. macrospondijlu$ Cope ; I.e., 1884, p. 3-5. Texas.

45. (7. Uptocephaliti Cope ; 1. c, 1884, p. 30. Texas.

DlMETRODON Cope ; Proc Amer. Phil. Soc, 1878, p. 512 ; 1. c, 1880, p. 42 et infra.

46. D. gigas Cope ; 1. c, 1878, p. 513 ; 1. c. 1880, p. 44. Texas.

47. D. inemou Cope ; 1. c. Texas.

48. D. rectiformii Cope ; 1. c, p. 514. Texas.

49. B. semiradicatua Cope ; Bull. U. S. Geol. Surr. Terrs., 1880 (81).
Naosaurus Cope ; Amer. Naturalist, 1886, p. 545 et infra.

50. N. eruciger Cope ; Dimetrodon cruciger ; Proc. .Vmer. Phil. Soc, 1880, p. 44 ; Amer. Xaturalist, 1878, p. 830.

Texas.

51. -V. datiger Cope ; Amer. Naturalist, 1886, p. 545 el infra. Texas.

52. A', microdu* Cope ; 1. c, 1886. p. 545 ; Edaphosaurus microdus Cope ; Proc. Amer. Phil. Soc, 1884, p. 37.

Texas.
TuEROPLECRA Copc ; Proc. Amer. Phil. Soc, 1878, p. 519, 1880, p. 40.

53. T. retrotersa Cope ; 1. c. Texas.

54. T. uniformin Cope; 1. c, 1878, p. 519 ; 1880, p. 40. Texas.

55. T. trianguliita Cope ; 1. c, 1878, p. 520. Texas.

56. T. obtusidena Cope ; 1. c, 1880, p. 41. Texas.
Embolopiiorcs Cope ; 1. c, 1878, p. 518.

57. E. fritillut Cope ; 1. c Texas.

58. E. dollotianus » Cope ; Proc Amer. Phil. Soc, 1884, p. 43, I'l. I, figs. 4-5. Texas.
Edaphosaurus Cope ; Proc. Amer. Phil. Soc, 1882, p. 448.

59. E. pogonias ; \. c, 449. Texas.

Parioiichidx.

Pariotichus Cope ; Proc. Amer. Phil. Soc, 1878, p. 508.

00. P. hraehyops Cope ; 1. c Texas.

61. P. mtgalopt Cope ; 1. c, 1883, p. 030, Texas.
EcTOCYNODOH Copc ; 1. c, p. 509.

02. E. aguli Cope ; 1. c. 1882, p. 451. Texas.

63. E. ordinatui Cope ; I. c, 1878, p. 508. Texas.

04. E. incititut Cope, infra. Texas.
Pastylcs Cope ; Bull. U. S. Geol. Surv. Terr., 1881 (80).

65. P. eordnlus Cope ; 1. c. Texas.

•Dedicated to Dr. L. DoUo, the distinguished paliconlologist of the Royal Museum of Brussels.

288 CATALO&UE OF THE SPECIES OE VERTEBRATA POUND IN THE

Bolosauridse.

BoLOSADRDS Cope ; Proc. Amcr. Phil. Soc, 1878, p. 50C.

G6. B. striatus Cope ; 1. e. Texas.
CuiLONYX Cope ; 1. c, 1883, p. 631.

67. C. rapidens Cope ; 1. c. Texas.

Incertee sedis.

Metarmosadkus Cope ; Proc. Amer. Phil Soc, 1878, p. 516.

68. M.fossaius; 1. c. Texas.

Diadectidas Cope.
Pal. Bull. No. 33, 1880, p. 8.
DiADECTES Cope ; Proc. Amer. Phil. Soc, 1878, p. 50.j.

69. D. sideropelicus Cope ; 1. c Texas.
Empedias Cope ; Proc. Amer. Phil. Soc. ; Empedocles Cope ; Proc. Amer. Phil. Soc, 1878, p. 516 ; 1880, p. 634.

70. K phaaeoUnus Cope ; Pal. Bull. No. 32, 1880, p. 9. Texas.

71. E. alaius Cope ; 1. c. Texas.
73. E. laiibueeaius Cope ; 1. c. Texas.

73. E. molaris Cope ; Pal. Bull. No. 32, 1880, p. 10. Texas.

74. E.fissuaCoye; Proc Amer. Phil. Soc, 1880, p. 684. Texas.
Helodectes Cope ; Pal. Bull. 11, No. 33, p. 11.

75. E. paridens Cope ; 1. c. Texas,

76. H. isaaci Cope ; 1. c, p. 13. Texas.

SYNOPSIS OF THE SPECIES.

Pl.SCES.

Selachii,

Dipnoi,

Teleoaioim,
Batraciiia.

Oanoce.2:>liala,

RhachUomi,

Stegocephali,

Emholoma-i,
Keptilia.

Theio:norpht,

Total,

The only catalogue of this fauna hitherto published appeared in the American
Naturalist for February, 1881, p. 162. In that list fifty-one species were enumerated.

CTENODUS VABASENSIS Cope ; Proc. Phila. Academy, 1883, p. 110.

Two fine teeth received from Mr. "W. F. E. Gurley, from Danville, Illinois,
probably belong to this species, their anterior crests being perfect. They are proven
to be produced forwards as in C. porrectus Cope. One of the specimens agrees Avith

Gen.

Specu

5

8

4

11

1

2

1

2

4

8

1

2

1

4

15

39

33

76

BEDS OF THE I'ERMIAlSr EPOCH IX XOHTII AMEUICA.

289

the type in tlie number of its crests, while the second, which is a httle larger, has them
7{ instead of 6{-.

ERYOPS PLATYPUS Cope. Ichlhyeanthut platypus Cope ; Proc. Amer. Phil. Soc, 1877, p. 574.

A reexamination of the type specimen of this species from
the Coal Measures of Ohio, preserved in the museum of Cohimbia
College, Xew York, enables me to refer this species to the Rhach-
itonii. The neural spines are distinct, showing that it belongs,
probably, to the Eryopidne, As the skull is not preserved I can-
not determine the genus positively, but refer it for the present to
Eryops. I append a figure of the posterior foot, which displays
the characters of the tarsus of this group for the first time. The
number of tarsals is as in a Theromorph reptile, except that two
elements represent the cuboid bone, as in the reptile Stereosternum
tumidwn Cope ; giving five elements in the distal tarsal row.
There is but one centrale and no intermedium. Two fragments
of caudal vertebra) adhere to the specimen (marked cv in the accompanying cut).
The lettering of the cut is as follows: Fi, fibula; F, ni)ulare ; T, tibiale ; c, centrale;
i-v, tarsalia.

Z.\TKACHYS SERRA.TUS Cope; Proceedingsof the Amer. Philos. Soc, 1878, p. 523 ; American Naturalist, 1884, p. 36.

This species has been thus far certainly known from a portion of the maxillary
bone only. Analogy of general characters led me to associate with it a second species
under the name of Z. apicalis. This form was clearly rhachitomous, so that in the
American I^Taturalist, as above cited, I referred the genus Zatrachys to the family of
the Ei"yopid:Tc.

A skull of the Z. serratus having come to hand, I am able to give some of its
characters. These indicate that the position assigned to it as above is correct, and
that it represents a genus dificrent from any of the others of the fiimily so far as our
present understanding of the characters goes.

There are two approximated occipital condyles. There is no distinct basi-occipital
bone distinguishable, and it is possibly wanting. The palatopterygoid arch is convex
outwards, keeping near the maxillary bone, and separated from the parasphenoid by a
wide foramen. The intercalare forms a prominent angle on each side of the cranial
table. The occipital asjiect of the skull displays fontanelles between two ascending
portions of the exoccipitals, one of which bounds the foramen magnum, and the other
the intercalare. The latter has a superior ;iiul inferior posterior angle which are sepa-

290 CATALOGUE OF THE SPECIES OF VERTEBRATA FOUND IN THE

rated by a notch. The posterior part of the quadratojugal arch forms a wide roof or
ledffe overhang-ino" the lower iaw. The teeth are small, acute, and close together. The
median and anterior teeth are unknown, as the anterior half of the skull is-wanting.
The angle of the lower jaw projects little or not at all beyond the cotylus. The
condyle of the quadrate is nai-row and little distinguishable. Between the intercalare
and the transverse process of the parasphenoid, is the deep tympanic chamber. It
is traversed on one side of the specimen by a curved club-shaped bone, with the larger
end truncate and internal. Its tissue is very spongy. It may be the columella auris,
but its slender external extremity appears to be continuous with the os intercalare.
This may, however, be due to the mode of preservation.

The surface of the skull has a dense reticulated ffl3ulpture, which is in places radi-
ated. It is especially pronounced along the external border of the quadratojugal arch
where it develops nodules which are arranged in a serrate manner. Thei-e is a pro-
nounced fossa in front of each orbit, which is bounded within by a convex i-idge extend-
ing forwards from the orbit. Between these ridges is another deep fossa of the middle
regions, whose posterior border is in line with the anterior edge of the orbits. The
cranial wall is here very thin. On each side of the supraoccipital bone a sharp process
projects backwards and inwards forming a short horn. I do not imagine that the
value of this character is more than specific. The external face of the mandible is
sculptured, the sculptured surface presenting an obtuse angle upwards, and leaving
a narrow smooth face anterior and posterior to the angle.

iVeaiuremeiits. M.

Width of skull posteriorly, .138

" between intercalare bones inclusive, .062

" " occipital processes " .019

" " orbits, .039

(anteroposterior, .017

Diameters of orbits - „„„

(transverse, .OM

Elevation of occiput from foramen inclusive, .010

Width of parasphenoid behind transverse processes, .024

This cranium presents a ciu'ious mixture of defective and excessive ossification.
Its form is more depressed than any others of the family. I await the discovery of
its anterioi' regions with interest.

ECTOCYNODON INCISIVUS, sp. nov.

A. nearly complete but somewhat distorted cranium represents this species. It
presents the generic characters of roofed temporal fossae, sculptured cranial bones
without lyra, and an elongated tooth near the middle of the maxillary series.

BEDS OF THK I'ERMIAX EI'OC'II IN XOIITH AJIERICA. 291

The muzzle is quite piomincnt, a cluiracter somewhat exaggerated in the speci-
men by pressure. The nostrils are large, lateral in dii'ection, and situated close to the
end of the muzzle. The orbits are subround, of medium size, and look mainly up-
wards in the present condition of the specimen. Oue of tlie most important peculiar-
ities of the species is the disproportionately large size of the first or anterior incisor or
l)rcmaxillary tooth. The crown is conical and nearly straight, with an acute apex
slightly posterior to the central point. Its section at the base is slightly angulate.
The two other prcmaxillaiy teeth arc much smaller, the third quite minute and with
a sharp apex.

Thei'p are three maxillary teeth separated by rather wide interspaces anterior to
the large tooth which gives character to the genus. Tlie latter is abruptly large, but
not equal in dimensions to the large first incisor. Posterior to it the maxillary teeth
are closely placed, and with obtuse crowns. They commence very small, and increase
in size posteriorly. At a point where the palatine or ectopterygoid, as the fact may
be, joins the maxillary, the tooth-bearing surface is wide, and supports four rows of
small obtuse-crowned spaced teeth of equal size. This dental patch is triangular, with
its long angles extending anteriorly and posteriorly. The latter angle terminates a
little posterior to the middle of the orbit. The teeth have a small axial pulj) cavity,
and the dentine is perfectly simple.

The head sculpture is well defined, and is reticulated in pattern.

Measurements. M.

Length from end of muzzle lo posterior border of orbit, .054

Transverse diameter of orbit, .016

" " inlerorbital space, .020

Length from end of muzzle to orbit, .034

Vertical diameter of nostril, .008

Vertical depth of maxillary in front, .018

Length of first premaxillary tooth, .006.5

Transverse diameter of do., .0038

Distance between first incisor and large maxillary teeth, .013

Distance from large maxillary tooth to posterior angle dental p;itch, .024

Width of dental patch, .010.5

Elevation of a posterior tooth, .0015

This species is intermediate in size between the E. ordmatiis, wliicii is small, and
the M Of/uti, which is large. In its disproportionate inecpiality in the size of its
teeth, it differs from the latter; while the former has larger orbits and a difierent sculp-
ture, besides having half the linear measurements. The sculpture of the JEl ordtnatiis
is in parallel ridges, enclosing minute deep punctiform pits between them.

A. p. S. — VOL. XVI. 2k.

292 CATALOGUE OF THE SPECIES OF TERTEBKATA FOUXD IX THE

The characters of this species confirm the propriety of my proj)osition that Paii-
tylus, Ectocynoclon and Pariotichus are members of a single family which difters from
the Clepsydropidae in the overroofing of the temporal fossa.*

DIMETRODON SPECIES.

My last account of this genus was jniblished in the Proceedings of the American
Philosophical Society, 1880. p. 42. Since that time additional material has enabled
me to develop more fully the characters of this singular type.

Hibs. — In this genus and in I^aosaurus the sacral i-ibs are present as in Batrachia.
They are short, and vertically compressed, forming a wedge-like. body.

Vertebral. — I have at various times described the extraordinary development of
the neural spines of the dorsal vertebrse in the genus Dimetrodon, which belongs to
the ClepsydropidcTB, one of the carnivorous families of the Saurian order Thei'omorpha.
The dentition of these animals is of the most formidable character, consisting of com-
l^ressed finely serrate teeth on the maxillary and dentary bones mingled Avith huge
conic tusks on the middle of the maxillary anterior end of the dentary, and occupying
the entire alveolar face of the preraaxillary. The huge neural spines formed an ele-
vated fin on the back. In a medium-sized specimen of Dimetrodon mcisivus, where
the vertebral body is 35 mm. in length, the elevation is 900 mm. or twenty and a half
times as great. The apex of the spine in this species is slender, and apparently was
flexible. Its utility is difficult to imagine. Unless the animal had aquatic habits,
and swam on its back, the crest or fin must have been in the way of active movements.
Accordingly the spines are occasionally found distorted by union of surfaces of frac-
ture. The limbs are not long enough nor the claws acute enough to demonstrate
arboreal habits as in the existing genus, Basiliscus, where a similar crest exists.

Stermcm. — A singular bone which I can only regard as this element accom-
panies a fragmentary skeleton of the D. incisious or D. gigas ; and other examples
occur with other specimens.

The anterior two-fifths of the bone is nearly square, and slightly concave above,
with three angles, one at each side and one anterior ; the rest contracts posteriorly
into a long, narrow, flattened shaft, which constitutes three-fifths of the length.
This portion is depressed, so that the transverse section is lenticular. The lateral
edges are acute, and without articular facets of any kind. The distal extremity is
first grooved, and then fissured, each half terminating in an obtusely narrow apex
which is applied to the other half. The surface of this part of the element is longi-
tudinally grooved both above and below.

* See Proceedings Aiiier. Philos. Soc., 1883, p. 031.

BEDS OK THE PERMIAN EPOCH IX NORTH AMEUICA. 293

Clavicles. — The clavicles in the genus Dimeti-odon are well developed elements.
They consist of a vertical narrow and a horizontal expanded portion. The anterior
border of the bone is rounded ; the internal border is serrate or semidigitate. If the
latter unites -with the epistenunn l)y suture it must be by a very open one. This por-
tion is more expanded than in the monotreme mammalia; while the episternum is
more produced posteriorly. Tlie tyjic of structure of this [lart of the scapular arch is
less mammalian than is liiat of Empedias above referred to, but is not far removed
from the latter.

Posterior foot. — A posterior foot of a species of Pelycosaurian, from Xe Mexico,
displays the characters more perfectly than any specimen in my collection. It con-
firms the inferences I have derived from the posterior foot of Clepsydrops natalis Cope,
as to the mammalian affiliations of this order of reptiles. Thus it has the same number
of tarsal bones distributed in the same manner. That is an astragalus and a calca-
neum in the proximal series ; then a navicular distad to the astragalus, which is
succeeded by three cuneiform bones. Distad to the calcaneum is but a single bone,
the cuboid. The specimen described differs in some important particulai's from tliat
of the species above mentioned. Thus the astragalus and navicular together areas
long as the calcaneum, while in the C. natalis, the calcaneum and astragalus have
equal lengths. It is possible that there are but four digits in the posterior foot ; at
least I can only find one metatarsal in connection with the cuboid. The internal edge
of the astragalus is broken away, so that the presence of the spur or a homologous
digit caimot be substantiated.

As the asti'agalus of Dimetrodon is closely similar to that of Clepsydrops, the
species now described does not enter that genus. The rather numerous vertebi'te
which accompany it reselnble, on the other hand, those of Theropleura, and it may be
that they belong to a species of that genus. By comparison of the i)late, with the cut
given under the head oi Enjops plati/jrus, the dill'erence between that type and this
may be perceived.

The lettering of the ligure is as follows: ^s., Astragalus ; 6\<., Calcaneum ; Cn.,
Cuboid ; Wa., Xavicular ; Mic, Entoeuneiform ; Msc, Mesocuneiform ; JEcc, Ecto-
cuneiform ; i-iv. Metatarsals.

N.\OS.VURUS CLAVIGKU Cope. American Xiitumli.sl, June, 188G.

Char. gen. — Xaosaurus differs from Dimetrodon only in the presence of transverse
l)rocesses on the neural spines.

The above named very peculiar si)ecies is congeneric with the Saurian described
under the name of Dimetrodon cruciger Cope. The neural spines are not quite ho ele-

294 CATALOGUE OF THE SPECIES OP VERTEBRATA FOUND IN THE

vated as in the D. incisious, but they are moi'e robust, and have transverse processes
or branches which resemble the yard-arms of a ship's mast. In a full-sized individual,
the longest cross-arms, which are the lowest in position, have an ex^janse of two hun-
dred and sixty millimeters, or ten and a quarter inches, while the spine has about the
height of five hundred millimeters (19.75 inches), the body being 60 mm. long. The
animal must have presented an extraordinar}' appearance. Perhaj^s the yard-arms
were connected by membrane with the neural spine or mast, thus serving the animal
as a sail, with which he navigated the waters of the Permian lakes.
The thi-ee sj^ecies of Kaosaurus differ as follows:

I. Neural spines distally cylindric.

Distal transverse processes represented by tuberosiiies, iV, cruciger.

II. Neural spines distally dilated and compressed.

Palatine teeth small, widely spaced, iV. elamger.

Palatine teeth large, closely packed, N. microihis.

The skull. — One of the best preserved skeletons of the iV". claviger includes a skull,
but the extremity of the muzzle is unfortunately wanting. The median line rises for-
wards so that the convexity of the top of the muzzle is higher than the posterior parts
of the skull, whose profile descends rapidly. This throws the orbit far back and gives
the animal a peculiar appearance.

The orl)it is nearly round, the superciliary border being arched. Anterior to it
is a large anteorbital fossa bounded by a longitudinal ridge above. Above the'ridge
is a longitudinal groove, which is separated from that of the opposite side by a narrow
ridge only. The quadrate bone is large and laminiform, and is truncate above, having
a good deal the shape of the corresponding bone in a fish. The parietal buttress is
produced downwards and backwards, and is in contact with the superior third of its
posterior border. Beneath and within it is a narrow opisthotic. The pterj^goid is
largo, and is distally vertically compressed. Anteriorly it becomes flattened so as to
be horizontal, and is studded with small conical teeth rather distantly placed.

The transverse series of palatine teeth on a massive Z-shaped bone, seen in the
D. inciswus, is not preserved in this specimen, but the explanation of the structure is
furnished by a specimen of the Dimetrodon semiradicatus Cope, of which a second
specimen has been found by Mr. Cummins. Here the palatine bones with their teeth
are preserved. They are not so massive as in the D. incisivus. Posteriorly they jjass
into the longitudinally flattened i)art without interruption by suture, so that I suspect
that this part is to be rel'erred to the palatine rather than to the pterygoid bone. It
is studded with small teeth, but they are not nearly so numerous as in the N. cruciger.

BEDS or THE PEKMIAN EPOCH IN NORTH AMERICA. 295

These specimens show that the species I named Edaiihosaurus microdus* must be
placed in I^aosaarus, where it represents the second species with transverse processes
on the dorsal neural spines. The teeth of the palatine patch in this species are larger
and more closely i)laced than in the D. cruciger. The distinction between the two
species in the form of the apices of the neural spines, to which I referred in my descrip-
tion (1. c.) holds good ; but the N. clavif)er has them dilated antei-oposteriorly nearly
as in the D. inicrodiis.

Vertebne. — A large series of these is preserved, and they show many interesting
characters. The intercentra are not distinct in the anterior part of the column, but
are separated posteriorly and in the sacrum. The centra are compressed and have
an acute inferioi* heel. The neural spines are moderately compressed below the first
transverse processes ; above this point they are antei'opostei'iorly oval in section. The
distal half is compressed. They expand to a point below the apex, where the antei'ior
edge extends obliquely backwards to the summit. A short corresponding oblique
edge truncates the posterior superior angle. The medullary cavity of the spine is
not closed at the apex. On several of the vertebras the lowest transverse process is
double, but the sides of the same vertebra differ from each other in this respect in some
instances.

The two sacral vertebras are not coossilied, and the zygapophyses are well de-
veloped and distinct, as are the intercentra. The latter are flat, and but little
developed in the upward direction. The neural spines are rather elevated and slen-
der. They are compressed without cross-pi'ocesses, and the apex of the spine has
small tubercles.

JRibs. — The ribs ai'c long and well curved, and are moderately compressed on
their proximal half, and cylindric for their distal. The head is w^ell distinguished
from the tubercle as in the manner of a mammal. That is, the tubercular surface
is sessile on the convexity of the rib, and not licdunculate. In this respect these ribs
diffei" from the usual form of two-headed reptilian ribs. The head is so long on the
anterior dorsal vertebra", as to articulate with the posterior edge of the vertebra in
front of the one with which its tubercle articulates. It becomes shorter on the pos-
terior parts of the column, articulating with the edge of the rib which supports the
tubercle. On the caudal sei'ies the head is reti-acted so as to be close to the tuber-
cular articulation, \yhich is the most extensive, and which is deeply notched on one of
its faces. This gives the appearance of a three-headed rib in this genus and in Dime-
trodon.

* Proc. Amer. Pliilos. Soc, 1884, p. 37.

29G CATALOGUE OF THE SPECIES OF VERTEHKATA FOUND IN THE

Besides the skull, the vertebrge and the ribs, I cannot yet positively identify the
skeleton of this species.

Measurements. M.

No. 1. (Skull.)

Depth of muzzle at sixth tooth from last, •1'''2

Length of skull posterior to same tooth, -SSS

(■anteroposterior, -057

Diameters of orbit -< . , „-„

(vertical, •"O"

Vertical depth from line of eyebrow to end of quadrate, .200

Width of condyles of quadrate, -050

( vertical, .038

Diameters of crown of sixth tooth X anteroposterior, .014

( transverse, .011

Thickness of maxillary bone at sixth tooth, .032

No. 3. (Vertebra', probably of No. 1.)

\ anteroposterior, .064

Diameters of dorsal neural arch, with zygapophyses -, , ' . ^

(transverse (posterior)

Diameters of neural spine near base \

(transverse,

( anteroposterior,
Diameters neural spine at process <

.036

f anteroposterior, . 033

base -!

037

No. 3. (Vertebrae)
anteroposterior, .039

Diameters centrum dorsal vertebra •< transverse, .034

I. vertical, -035

Expanse prezygapophyses do., .041

" diapophyses do., .083

Elevation of neural spine to first process, .034

Transverse diameter of process, -025

.030

(transverse, .038

Length of the two sacral vertebra;, .084

Elevation of neural canal of do., .010

" " " spine of do., .084

Length of rib on outside of curve, .360

" head of rib, .045

Transverse diameter of rib just beyond tubercle, .017

Anteroposterior do., .020

?SPUR OF A PELYCOSAURIAN.

In Plate 111, fig. 7, is represented the basal part of a bone which has the form
and characters of a mctapodial of one of the Clepsydropida\ It diftbrs from all of
these which have come under my notice in the l)urred or serrate character of the bor-
der of the proximal extremity. I do not know of any genus which is likely to have
such a mctapodial of the usual series. The lirst of the series, which articulates with

/

>K!'

■°==--l'_

v., , . JJ,' , p A'Sj*

>«;i--

A"

\^,---./

■■?'/'■

f':Cm

^■'

-J^'-

1 -2. NAOSAURUS CbAVIGER Va . 3.N. CRUCIGER %. 4.N .MICRODUS 'A. 5 DIME TROD ON %.
6 THEROPLEURA ? 'V*. 7. METAPODIAL ? 8. PLATYPUS.

BEOS OF THE PEliMIAN Kl'OCU IN NOUTU AMERICA. 297

the distill pait of the astragahis has, however, not yet been discovered, and this bone
may be that missing piece. Some probability attaches to this identification from the
fact that the corresi)onding element in Ornithorhynchus (PI. Ill, fig. 8), is burred in
the same manner at its proximal extremity. The present piece is, however, longer
than the bone of Ornithorhynchus, and has much more the usual character of a meta-
tarsal. It is not jjcrforate at the base, and has only the usual medullary cavity. It
is flat on one side, and convex on the other. It must, however, be regarded as proba-
ble that fi'om a more or less normal metatarsal, the basal bone of the spur of Ornitho-
rhynchus has been derived ; the spur proper representing one or more phalanges.

That the posterior foot of Vertebrata includes six toes is maintained by Baur.
This is confirmed by the presence of a digit within the hallux in various Batrachia
Anura. In Rana catesheiana this digit has three segments, a metatarsal and two
phalanges, the former resting directly on the astragalus. This digit appears to have
been present in the ClepsydropidiE.

EXPLANATION OF PLATES.

PLATE II.

Figs. 1-2. Niiosaurus clntiger Cope. From the Permian formation of Texas.

Fig. 1. Posterior part of skull, left side ; one-half natural size. Restored from skulls of N. crticiger and Dimelrodoii
incmtm. Pa, Parietal bone ; J, jugal ; QJ. Quadralojugal ; Opo, Opisthotic ; Sq. Squamosal ; Q, Quadrate ; Pt, Ptery-
goid.

Fig. 2. Part of palatopterygoid arch of the same from below ; three-quarters natural size.

Fig. 3. Naosaurus mierodus, part of palatopterygoid arch from below ; three-quarters natural size.

Figs. 4-5. Ectocynodon incisirms, skull ; from the Permian formation of Texas ; natural size.

Fig. 4. From above.

Fig. 5. From bclow.

PLATE in.

Figs. 1-2. Naosaurus claviger, dorsal vertebras ; one-half natural size.

Fig. 1. Front view.

Fig. 2. Right side.

Fig. 3. Naosaurus cruciger, neural arch and spine, from front ; one-half natural size ; centrum in outline.

Fig. 4. Naosaurus mierodus, distal view of distal end of ndiiral spine ; three-fourths natural size.

Fig. .5. Dimtlrodon? sp., sternum, inferior face.

Fig. (i. Theropleura sp. ?, pari of posterior foot ; three-fourths natural size. As, Astragalus ; Ga, Calcancum ; Na,
Navicular ; C«, Cuboid ; Ece, Ectocuueiform ; .)/, Mesocuneiform ; Enc, Entocuneiforra ; phalanges partly restored.

Fig. 6a. Calcancum, anterior view.

Fig. 7. Supposed first metatarsal of Pelycosaurian, twice natural size ; 7 and la, opposite lateral views , b,
proximal end.

Fig. 8. Ornithorhynchus anatinus, internal digit or spur, from Owen, Anatomy of Vertebrata, ii, flg. 199 ; a, meta-
tarsal ; J, phalange : c, metatarsal, proximal view.

ARTICLE V.

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

BY E. D. COPE.

Read before the American Philosophical Society, January 20, 1888.

The Puei-co formation rests on the Laramie in IS'orthwestern New Mexico and
Sonthwestern Colorado, and is largely covered by the Wasatch Eocene in both
regions. It was discovered by the writer in 1874 at its eastern outcrop of about 500
feet thickness, and was identified by Endlich and Holmes in Colorado in 1876, where
the thickness reaches 1000 to 1200 feet. On the San Juan river its thickness is 700
feet, while at its western outcrop south of that river, its thickness is 800 or 900 feet.
"While the 'formation possesses lithological peculiai-ities, no clue to its importance in
geologic chronology was known until the discovery of vertebrate remains was made
in 1880, by Mr. David Baldwin. With the evidence derived from this material, the
writer has been able to interject into the series of epochs of geological time a period
which must have possessed many peculiarities, and which differed in such important
essentials from those which preceded and from those that followed it, that an immense
interval between them is proven to have existed, such as had not been previously sus-
pected. The rich fauna which it contains displays characters which indicate other
discoveries yet to be made befoi'c connections with other epochs both prior and subse-
quent can be known.

The vertebrate fauna includes up to the present date one hundred and six known
species. Four species of Mollusca have been discovered, which have been determined
by Dr. C. A. White of the U. S. National Museum. They are Unio rectoides White ;
Helix adijns White ; H. nacimientensis White, and Pupa leidyi Meek, The first
named is found in the Wasatch, and the last in the ? Bridger ; the two other species
are peculiai*. Besides these the only other indication of organic life at that period is
petrified wood of undetermined trees, which is quite abundant.

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE I'UEIICO t^EKllJrf. 299

The character of the vertebrate fauna is indicated by the following table :

Reptilia 12

Crocodilia 3

Testudinata 5

Rhynchoct'phalia • ;!

Ophidia 1

Avcs , 1

Mammalia 93

? 3Iarsupialia 11

Bunotheria 53

Tivniodonta 3

Creodouta 49

Taxeopoda 28

Quadrumana ?4

Condylartlira 24

Amblypoda 2

Total 10(1

In 1874 the wi-itei* advanced the proposition that the ancestors of modern pla-
cental Mammalia would be found to be " plantigrade pentadactyle bunodouts." This
anticipation was partly realized in the fauna of the "Wasatch epoch subsequently dis-
covered, but is completely so in the characters of the Mammalia of the Puerco epoch.
All the placentals, and probably the implacentals also, were " plantigrade pentadactyle
bunodouts."' More than this the placentals nearly all present the primitive type of
lentition of the maxillary series, since the superior molars are nearly all of the tritu-
bercular type. But foui- species out of the eighty-two placentals are quadrituber-
cular. In the inferior molars the tubei-culo-sectorial, or quinquetuberculai- type of
dentition, is extensively prevalent, but not so generally so as the superior tritubercu-
lar. Thus of the eighty-two placentals sixty-four present the primitive type.

Ill its relations to other fauna?, the Puerco is totally distinct as to species, No
determined species came to it from an earlier epoch, and none continued after i(. Ol"
genera not widely distributed in time, one of lizard-like Khynchocephalia, Champso-
saurus, comes over from the Laramie, with a genus of tortoises, Compsemys. An-
other genus of tortoises, Chelydra, probably commences at this epoch, to conliniic
through the European Miocenes to the present time, since it still exists in jMorlh
America. Among Mammalia, two genera only coiitinue later. Didymictis is found
in the Wasatch and Bridger formations, and Chriacus in the AVasatch. iS'ot only
this, but the entire family of tlic I'eriptychidai ceased at the close of the Puerco.
The same is true of the Amblypod family Pantolambdidic. One of the most import-
ant features of the fauna is, however, the presence of eleven siJecies of the ?Marsu-

c:

300 STKOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

pialia Multitubereulata, a suboi-der whidh commenced in the Triassic age, and which
terminated its existence so far as the ISTorthern hemisphere is concerned with the
end of the Puerco epoch. This series of animals gives a Mesozoic character to the
fauna, which is not necessarily counterbalanced by the characters of the remaining
types. The placentals are in all probability those which existed during the latter
part of Mesozoic time, and the absence of some of the forms of the Eocene increases
the weight of the impression thus produced. Thus two orders universally present
in the Eocenes, the Perissodactyla and the Rodentia, are wanting from the Puerco.

In conclusion it may be safely asserted that in the Puerco fauna we find the
ancestors of the species of Eocene and of later times. In the Tajniodonta we get
ancestors of Tillodonta and probably of Rodentia and Edentata. In Creodonta we
get the ancestors of the Carnivora, in the flxmily of the Miacidse. In the Condylar-
thra we get the ancestors of the Diplarthra and Amblypoda, and in the Puerco Ara-
blypoda the ancestors of those of the following epochs. Hence the investigation of
this fauna possesses an especial interest for the mammalogist and for the evolutionist,
as well as for the geologist proper.

I give first a list of the species, and then give descriptions of new species, with
the osteology of such as the material permits.

In describing the dentition I have, after consultation with ray friend Professor
W. B. Scott, of Princeton, followed the method of enumeration of premolar teeth
inti-oduced by Kowalevsky, and adopted b}^ Schlosser. In this method the premolar
teeth are counted from behind forwards, so that the one usually enumerated as num-
ber four becomes number one, and vice versa.

The only catalogue of the Vertebrata of the Puerco which has appeared was
published in the Proceedings of the American Philosophical Society for 1882, begin-
ning at page 461. Since that time the following publications relating to that fauna
have appeared :

First Addition to the Fauna of the Puerco Eocene. By E. D. Cope, loc. cit., 1883, Jan., p. 545.

Second Addition to the Knowledge of the Fauna of the Puerco Epoch, loc. cit., 1883, Dec, p. 309.

On Some Fossils of the Puerco Formation. Proc. Acad. Nat. Sci. Phila., 1883, p. 168.

Tlie Tertiary Vertebrata of the West. Report U. S. Geol. Surv. Terrs., P. V. Ilayden, Vol. Ill, Feb., 1885.

On Some New T:cniodonta of the Puerco. Amcr. Naturalist, 1887, p. 469.

The Marsupial Genus Chirox, loc. cit., 1887, p. r)G6.

The present enumeration brings together all the species hitherto described, and
adds a number of ncAV ones. The whole number, it will lie observed, reaches 106,
which are referred to thirty-four genera.

Information at present available indicates that there is some faunal difference

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUKRCO SERIES.

301

between the lower and upper beds of the formation. I give a list of species which
have been found in the lower part of the formation only, and leave for future research
to determine whether they occur in the upper part of the formation or not :

Maksupialia.

Creodonta.

CONDYLAUTUUA.

Jfeoplagiaulax amerkanita .
Polymastodon taoensis.

" attemifitus.

" latimolis.

nemiganus otariiden».
Onychodectes tisonensis.
Miodtenus pentacus.

" bathygnaOtus.
crnssiaisptK.

" coryphieus.

" gaudrianvs.

" jUholianu».

" turgiduneulus.
Chriacus prisevs.

" hyattianus.

" ruetimeieranus
Trmodon bicuhninatus.
Periplychus brabensis.
' ' coarctatus.

Eetoconus ditrigonus.

All but one of the species of MiocUenus belong to the section Sareothraustes,
which has five cusps on the inferior true molars.

IJKI'TIIJA.

Croeodilus sp.
Croeodilits sp.
Crocodilui sp.

Plfiitomenus f communis Cope.
Chelydra cra»»a Cope, sp. nov. infra.
Compsemys sp.
Emys sp.
Trionyx sp.

CROCODILIA.

TESTUDINATA.

RIIYNCIIOCEPnATJA.

Cfiampsoiaurut auntralis Cope ; Anier. Nnlumlist, 1881. p. 090 ; Teitiiiry Vcrlobmta, p. 107, PI. -Will 1), figs. 1-4.
CliampfOMttriin piierceMi* Cope; Proc. Amor. Phil. Soc, 1881, p. 19."> ; Tcrliftry Vertcbniln. p. 107, PI. XXIII b,

figs. .")-10.
Champtomurui mponcn»i» Cope ; loc. cit., 1*91, p. 190, p. 109, PI. XXIII b, ngs. 11-22.

302 SYNOPSIS OF THE VERTBBEATE FAinSTA OP THE PUERCO SERIES.

OPHIDIA.

Belagras pi-isHformis Coiie ; Proc. Amer. Phil. Soc, 1883, p. 545 ; Tertiary Vertebrata, 1885, p. T31, PI. XXIV g,
fig. 2. Several individuals.

AAT2S.

Fragments of bones of a few undetermined species of birds have been found.

MAMMAXTA.

? MARSUPIALIA.

It has been recently discovered that the genus Ornithorhynchus possesses, in an
early stage, molar teeth exceedingly similar to those of the genus Ptilodus. I sus-
pect foi- this reason, that the Multituberculata belong to the Monotremata rather than
to the Marsupialia. This fact, if demonstrated, will account for the taxonomic isola-
tion of this group from the known Marsupialia, though some Monotreme has prob-
ably given origin to the latter. (See Amer. ]S"aturalist, Feb., 1888.)

Multituberculata.

PtUodus medimvus Cope ; Amer. Naturalist, 1881, p. 933 ; Tertiary Vertebrata, p. 173, PI. XXIII d, fig. 1. Five indi-

viduals.
Ptilodus trovesshrtianus Cope; loc. cit., 1882, p. 686 ; Tertiary Vertebrata, p. 737, PI. XXV f, flg. 19. Two individuals.

Additional and much more perfect specimens of the lower jaws of this species conflrm its characters as origi-
nally defined.

Neoplagiaulax americaims Cope ; Amer. Naturalist, 1885, p. 493. One individual.
Neoplagiaulax molestus Cope ; loc. cit., 1886, p. 451, et infra. Two individuals.

Chirox pUeatus Cope ; Proc. Amer. Phil. Soc, 1883, p. 331 ; Amer. Naturalist, 1887, p. 566, flg. Two individuals.
Polymastodon laiimoUs Cope ; Amer. Naturalist, 1885, p. 385. One individual.
Polymastodon taoenais Cope ; loc. cit., 1883, p. 684 ; 1884, p. 688, figs. 3-4 ; Tertiary Vertebrata, p. 732, PI. XXIII c,

figs. 1-6. Tmniolabis scalper, Amer. Naturalist, 1883, July ; Tertiary Vertebrata, p. 193, PI. XXIII e, fig. 7.

Catopsalis poUux Cope ; Amer. Naturalist, 1883, p. 685 ; Tertiary Vertebrata, 1885, p. 734, PI. XXIII c, figs. 1-5.

Twenty individuals.
PoUjmastcdon attenuaius Cope ; Amer. Naturalist, 1885, p. 494. One individual.
Polymastodon JiMideiis Cope ; Amer. Naturalist, 1884, p. 688. Catopsalis Jissidcns Cope ; Proc. Amer. Phil. Soc,

1883, p. 333. One individual.
Polymastodon foUatus Cope: Amer. Naturalist, 1884, p. 688, fig. 5. Catopsalis foliatus Cope; loc cit., 1883, p. 41G ;

Tertiary Vertebrata, 1885, p. 171, PI. XXIII d, flg. 3. One individual.

BUNOTHERIA.
Tseniodo7ita.

Psiitacotherium aspasiiv Cope; Proc Amor. Phil. Soc, 1883, p. 103 ; Tertiary Vertebrata, p. 196, PI. XXIV c, flgs.

3-4. Two individuals.
PsiitacotJieritm multifragum Cope ; Amer. Naturalist, 1883, p. 156 ; Proc. Amer. Phil. Soc, 1881, p. 191 ; Tertiary

Vertebrata, p. 196, PI. XXIV c, fig. 3 ; Amer. Naturalist, 1888, p. 5, fig. 1. Throe individuals.
Psittacotherium megalodus Cope ; Amer. Naturalist, 1887, p. 469. One specimen.

SYNOPSIS OP THE VEKTE15UATE FAUNA OF THE PUERCO SEIilES. 303

Creodonta.

nemiganus vulluosus Cope ; Amer. Naturalist, 1883, p. 831 ; Tertiary Vertcbrata, 1885, PI. XXIII c. Four specimens.

Bemiganua otariidens Cope ; Amer. Naturalist, 1885, p. 492 ; et infra. One specimen.

Conoryetes comma Cope ; Proc. Amer. Phil. Soc., 1881, p. 48G ; Tertiary Vertcbrata, 1885, p. 198, PI. XXIII c, figs.

1-5; XXV c, figs. 3-4. Ilexodon mokstus Cope; Amer. Naturalist, 1884, p. 705, fig. 3. Seven indiviiluals.
Onyehodectes tisonensis Cope, gen. et sp. nov. infra. Three individuals.

^fioel<Bnu» antiquus Cope. Sarcothraiistes (intiqiius Cope; Proc. Amer. Phil. Soc., 1881(2), p. 193 ; Tertiary Vertc-
brata, 1885, p. 347, PI. XXIV d, figs. 19-33. One specimen.
Mioelanus eonidens Cope. I'nisodon conidens Cope ; Proc. Acad. Pliila., 1882, p. 297 ; Tertiary Vertcbrata, 1885, p.

274, PI. XXIII d, figs. 9-10. Diacodon conidens; Amer. Naturalist, 1884, p. 3.j0. Three specimens.
3fiocl<enus bat/iygnathus Cope, sp. nov. infra. One specimen.
Mioelanus cragsicusph Cope. Conoryetes crassicuspis Cope ; Tertiary Vertcbrata, p. 201, 1885, PI. XXXIII e, fig. G.

Three specimens.
MiocUenus coryp/uvus Cope. Sarcothraustes coryplimus Cope; Amer. Naturalist, 1885, p. 380. Kine specimens.
Mioclcmus pentacus Cope, sp. nov. infra. Seven individuals, from the Lower Puerco.
MioeliFnus gaudrianus Cope, sp. nov. infra. One specimen.
MiocUenus lydekkerianus Cope, sp. nov. infra. Three specimens.
Jlioekmus filhoUanus Cope, sp. nov. infra. Four specimens.
JliocUenus i/iterruptus Cope. Deltatherium interruptum Cope; Proc. Amer. Phil. Soc, 1883, p. 403; Tertiary Vcrle-

brata, 1885, p. 383, PI. XXIII d, fig. 13. One specimen.
Mioel(enus acolytus Cope. Eyopaodus acolytus Cope ; Proc. Amer. Phil. Soc, 1882, p. 402; Tertiary Vertcbrata,

1885, p. 238, PI. XXXIII d, figs. 5-C Five specimens.
Mioelanus aasurgens Cope. Tri'isodon assurgens ; Proc. Amer. Phil. Soc, 1883, p. 311. Two specimens.
Mioelanus levisanus Cope. Tri'isodon levisanus Cope; Proc Amer. Phil. Soc, 1883, p. 446; Tertiary Vertcbrata

1885. p. 273, PI. XXIV f, fig. 3. Six specimens.
Mioelanus heilprinianus Cope. Triisodon heilprinianus Cope ; Proc. Amer. Phil. Soc, 1880, p. 193 ; Tertiary Vertc-
brata, 1885, p. 278, PI. XXIII d, fig. 11. One specimen.
Mioelanus ruslicus Cope. Triisodon rusticus Cope ; Proc Amer. Phil. Soc, 1882, p. 300. One specimen.
MiocUenus subtrigonus Cope ; Proc. Amer. Phil. Soc, 1881, p. 491 ; 1883, p. 555 ; Tertiary Vertcbrata, 1885, p. 338,

PI. LVII f, fig. 5, LIV f, fig. 4. Seventeen specimens.
MiocUenus cuspidalus Cope ; Proc. Amer. Phil. Soc, 1883, p. 312. One specimen.
MiocUertus protogonio'ides Cope; Amer. Naturalist, 1882, p. 833; Tertiary Vertcbrata, 1885, p. 340, PI. XXV f, fig. 17.

Four specimens.
MiocUenus floverianus Cope, sp. nov. infra. One specimen.
Mioclienus corrugatus Cope, Proc. Amer. Phil. Soc, 1883, p. 500; Tertiary Vertcbrata, 1885, p. 341, PI. XXIV f, fig.

5 ; XXrV g, fig. 8 be (" M. ferox "). Six specimens.
MiocUenus ferox Cope ; Proc. Amer. Phil. Soc, 1883, p. 547 ; Tertiary Vertcbrata, 1885, p. 328, PI. XXFV t, fig. 6 et

seq. Four specimens.
MiocUenus opisthacus Cope; Amer. Naturalist, 1882, p. 833; Proc Amer. Phil. Soc, 1883, p. 312. HemithUeus opis-

thaeus Cope ; Tertiary Vertebrata. 1885, p. 407, PI. XXV f. figs. 8-9. Twelve specimens.
MiocUenus turgidus Cope ; Amer. Naturalist, 1881, p. 489 ; loc cit., 1881, p. 830 ; Tertiary Vertcbrata, 1885, p. 325,

PI. LVTI f, fig. -.i-A ; XXV e, figs. 19-20. Twenty-five specimens.
Mioelanus zitlelianus Cope, sp. nov. infra. One specimen.
Mioelanus tnrgidunculus Cope, sp. nov. infra. Three specimens.
Mioelanus minimus Cope, Proc. Amer. Phil. Soc, 1882, 468 ; Tertiary Vertebrata, 1885, p. 337, PI. XXV e, figs.

22-4. Five specimens.
Tricentes bueculentua Cope ; V roc. Amer. Phil. Soc, 1883, p. 310. lliorUnnus burculenlus Cope ; Proc. Amer. Pliil.

Soc, 1883, p. .5.55 ; Tertiary Vertebrata, 1885, p. 341, PI. XXIV g. fig. 2. Three specimens.
A. P. S. — VOL. XVI. 2m.

304 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUEItCO SERIES.

Tncentes crassicoUMeus Cope ; Proc. Amer. Phil. Soc, 1883, p. 315. One specimen.

Tricentes inicquideiu Cope ; loc. cil., 1883, p. 317. Two specimens.

Chriacus truncatiis Cope; Proc. Amer. Phil. Soc, 1883, p. 313. Six specimens.

Chriacus pelvidens Cope ; Proc. Amer. Phil. Soc, 1883, p. 314 ; Tertiary Vertebrata, 1885, p. 740. Lipodectes pelvidens
Cope; Amer. Naturalist, 1881, p. 1019. Pelycodus pelvidens Cope ; Tertiary Vertebrata, 1885, p. 325, PI. XXHI d,
flgs. 7-8. Six specimens.

Chriacus siinj>lex Cope ; Proc. Amer. Philos. Soc, 1883, p. 314. One specimen.

Chriacus priscus Cope, sp. nov. infra. Sis specimens ; Lower Puerco.

Chriacus schlosseriamis Cope, sp. nov. infra. Fonr specimens ; Upper Puerco.

Chriacus haldwini Cope. DeUatherium haldioini ; Proc. Amer. Phil. Soc, 1883, p. 403 ; Tertiary Vertebrata, 1885, p.
383, PI. XXIII d, fig. 13. Three specimens.

Chriacus ki/aUiantis Cope ; Amer. Naturalist, 1885, p. 385. Loxolophua adapimis Cope ; loc. cit., p. 380. Four speci-
mens ; Lower Puerco.

Chriacus ruetimeycra/ius Cope, sp. nov. infra. One specimen.

Chriacus stenops Cope, sp. nov. infra. Three specimens ; Upper Puerco.

Chriacus inversus Cope, sp. nov. infra. One specimen.

DeUatherium fimdaminis Cope ; Amer. Naturalist, 1880, p. 338 ; Tertiary Vertebrata, 188.5, p. 278, PI. XXIH e, figs.
8-11 ; XXV a, fig. 10 ; XXV d, fig. 3. Lipodectes penetrans Cope ; Amer. Naturalist, 1881, p. 1019. Twenty-
three specimens.

Triisodoii hiculminatus Cope, sp. nov. infra. Two specimens ; Lower Puerco.

Triisodon quivirensis Cope ; Amer. Naturalist, 1881, p. 607 ; Tertiary Vertebrata, 1885, p. 370 ; PI. XXV c, fig. 3. One
specimen.

Dissacus navajovius Cope ; Amer. Naturalist, 1881, p. 1019 ; Tertiary Vertebrata, West, 1885, p. 344, 741. Eight in-
dividuals.

Dissacus carnifex Cope ; Amer. Naturalist, 1883, p. 834 ; Tertiary Vertebrata, 1885, p. 345, PI. XXIV g, figs. 3^.
One individual.

Didymictis haydenianus Cope ; Proc. Amer. Phil. Soc, 1883, p. 464 ; Tertiary Vertebrata, 1885, p. 306 ; PI. XXIII e,
flgs. 12-13. Three specimens.

Didymictis primus Cope; Proc. Amer. Phil. Soc, 1883, p. 309. Three specimens.

? Lemuroidea.

Mixodectes pungens Cope ; Proc Amer. Phil. Soc, 1883, p. 447 ; Tertiary Vertebrata, 1885, p. 341, PI. XXIV f, fig. 1.

Two specimens.
Mixodectes crassiusculus Cope; Proc. Amer. Phil. Soc, 1883. p. 447; Tertiary Vertebrata, 1885, p. 343, PI. XXIV f,

fig. 2. Three individuals.
Indrodon malaris Cope ; Proc. Amer. Phil. Soc, 1883, p. 318. One specimen.

CONDYLARTHRA.

Ilaploconus corniculatus Cope, sp. nov. infra. Six specimens.

Jlaploeonus liiieatus Cojie ; Amer. Naturalist, 1883, p. 417 ; Tertiary Vertebrata, 1885, p. 417, PI. XXV e, fig. 1-4.

Twenty-four specimens.
Ilaploconus angvstus Cope ; Tertiary Vertebrata, 1885, p. 410, PI. LVII f, fig. C Mioclirnus angustus Cope ; Amer.

Naturalist, 1881, p. 831. Two specimens.
Haploconus xiphodon Cope : Proc. Amer. I'hil. Soc, 1882, p. 406 ; Tertiary Vertebrata, 1885, p. 430, PI. XXV e, figs.

5-6. Two specimens.
Haploconus cntoconus Cope ; Amer. Naturalist, 1883, p. 686 ; Tertiary Vertebrata, 1885, p. 431, PI. XXV f, figs. 4-5.
flaploconus cophater Cope. Anisonchus cophater ; Proc. Amer. Phil. Soc, 1883, p. 331. Three specimens.

SYNOPSIS OF THE VERTEBRfVTE FAUNA OF THE PUERCO SERIES. 305

Anisonehu$ mandibularia Cope. Mioclaniis mandibtilaris Cope ; Amer. Naturalist, 1881, p. 831 ; Tertiary Vertebrata,
1885, p. 389, PI. LVII f. flg. 7. Two specimens.

Aniionehus sectorim Cope ; Proc. Amer. Phil. Soc, 1881, p. 488 ; Tertiary Vertebrata, 18a5, p. 41u, PI. XXV c,
figs. 5-6 and 8. Jliodanus teetorius Co^c ; Amcv. Naturalist, 1881, p. 831. Thirteen individuals.

Anitonehm contferut Cope ; Amer. Naturalist, 1883, p. 833 ; 1884, p. 80S, fig. 13 c ; Tertiary Vertebrata, 1885, p. 409.
Four specimens.

Anisonchus giUianus Cope ; Proc. Amer. Phil. Soc, 1883, p. 407 ; Tertiary Vertebrata, 1885, p. 411, PI. XXV f, figs.
10-11. ITaploconvs gillianus Cope ; Amer. Naturalist, 1883, p. 68G. Eight individuals.

Anisonchiis agapetiUus Cope ; Proc. Amer. Phil. Soc., 1883, p. 320. Si.\ specimens.

Zetodon gracilis Cope ; Tertiary Vertebrata, PI. XXIX d, explanation, and fig. 9. Two individuals.

Hemithlceus apiculatus Cope. Anisonchus apicutatiis Cope ; Tertiary Vertebrata, PI. XXV e, flg. 7. Eight indi-
viduals.

Hemithlaus k&iealeMkianvs Cope ; Amer. Naturalist, 1883, p. 833 ; Tertiary Vertebrata, 1885, p. 405, PI. XXV f, flgs.
6-7. Sixteen specimens.

Periptyc/iiis brabensis Cope, sp. nov. infra. Fourteen individuals ; the Lower Puereo.

Periplychus coarclatus Cope ; Tertiary Vertebrata, PI. XXIX d, explanation and figs. 7-8. Five individuals ; from the
Lower Puereo. ^

Periptychus carinidens Cope ; Amer. Naturalist, 1881, p. 337 ; Proc. Amer. Phil. Soc, 1881, p. 484 ; Tertiary Verte-
brata, 1885, p. 403, PI. XXV a, fig. 16 ; XXIII d, flgs. 14-15 ; XXIV g, fig. 5. Five specimens.

Periptychus rkabdodon Cope; Tertiary Vertebrata, 1885, p. 391, PI. XXIII f, XXIII g, figs. 1-11 ; LVII, figs. 1-3;
Amer. Naturalist, 1884, p. 801, figs. 1-3 and 6-9. Catathlmis rhabdodoii Cope ; Amer. Naturalist, 1881, p. 830 ;
Proc. Amer. Phil. Soc., 1881, p. 487. One hundred and thirty-eight individuals ; from tlie Upper Puereo.

Ectoconut ditrigonua Cope ; Amer. Naturalist, 1884, p. 796, figs. 4-5. Periptychus ditrigonus Cope ; Tertiary Verte-
brata, laS.'), p. 404, PI. XXIII g, flg. 13 ; XXIX d, flgs. 3-6. Thirty-two individuals ; from tlie Lower Puereo.

Protogonia zuniensis Cope. Phenticodas ziiniensis Cope ; Proc. Amer. Phil. Soc., 1881, p. 4!)3 ; Tertiary Vertebrata,
ISaj, 491, PI. LVII f, flg. 10. Three individuals.

Protogonia caleeolata Cope. Phenacodus calceolatus ; Proc. Amer. Phil. Soc, 1883, p. 559 ; Tertiary Vertebrata, 1885,
p. 487. One individual.

Protogonia plicifera Cope : Amer. Naturalist, 1883, p. 833 ; Tertiary Vertebrata, 1885, p. 424, PI. XXV f, figs. 3-3.
Eight individuals.

Protogonia puercensis Cope. Phenacodus piiercensis Cope ; Proc. Amer. Phil. Soc, 1881, p. 492 ; Tertiary Vertebrata.
1885, p. 488, PI. XXV e, flgs. 12-13 ; LVII f, figs. 8-9. I^otogonia subguadrata Cope ; Proc. Amer. Phil. Soc
1881, p. 492 ; Tertiary Vertebrata, 1885, p. 436, PI. LVII f, figs. 11-13. Twenty-five individuals.

AMBLYPODA.

Pantolambda bathmodon Cope ; Amer. Naturalist, 1882, p. 418 ; Proc. Amer. Phil. Soc, 1883, p. 558 ; Tertiary Verte-
brata, ^15, p. COl, PI. XXIX b, XXIX c. Seven individuals.
Pantolambda eavirietus Cope ; Tertiary Mammalia, PI. XXIX d, explanation and flg. 1. Four individuals.

DESCKirTIONS OF .SPKCIKS.

TESTUDINATA.

CHELYDRA Schw.

(J/iar. gen. — Marginal l)onc.s of the bridge imitetl wilii Uie eo.'^tals Iiy siini)le
gompliosis, and with the phistron by conii)omid gomphosiss. Bridge of phistron iiar-

306 SYNOPSIS OF THE VERTEBKATE FAUNA OF THE PDERCO SERIES.

row, with an iiitermarginal series of scuta, and without fontanelle. Surfaces not
sculptured.

Chelydra crassa sp. nov. Dermatemys sp. Cope ; Proc. Amer. Phil. Soc, 1883, p. 461.

This species is represented in my collection by fragments of two individuals.
Of the typical specimen there are preserved, two vertebral, nine marginal, and three
plastral bones; of the second, three vertebral bones. The specimens indicate
an animal of the average size of the existing snapping turtle, Chelydra serpentina.

The bones of both carapace and plastron are relatively much thicker than the
corresponding parts of the snapping tortoise, equaling in this respect the largest
existing sj^ecies of Emys. The bridge of the plastron is not so slender as in C. ser-
pentina. The vertebral bones have a median keel-like angle, which becomes at the
anterior part of each vertebral scutum a pi-ominent rib. This results from the abrupt
depression of the suiface on each side immediately posterior to the transverse dermal
suture. In the lai'ger specimen this suture is deeply notched anteriorly, and its
anterior border is so prominent posteriorly as to give an imbricate appearance, the
anterior vertebral scute rolling over the posterior by an obtuse border. The marginals
of the bridge are very massive, and the pit for the process of the costal is at one side
of the middle, and is nearly round. It is flat in the C. serpentina. The pits for the
plastral fingers are three, on the inner inferior edge of each marginal, and are directed
obliquely. The external foce of the marginals is distinguished by a rabbet, the in-
ferior margin of which projects as a ridge beyond the external face. Inferior face
convex. No other except the fine mutual sutures on the marginals of the bridge.
The free marginals, of which I have two, and one of them the anal, have no gomphosis
nor suture with the costals or pygal, being held in place by the integuments and by
the mutual marginal sutures. The dermal scuta are well marked, the marginals having
theii- bounding suture below that of the marginal bones. Surface of the shell every-
where smooth.

Measurements.

No. 1. Inch.

AntcropnsUTior dianictev of vertebral bone 8

r anteroposterior 83

Diameters of marginal of bridge I transverse (greatest) 35

[ vertical 98

anteroposterior 13

s ver

[ transverse 43

( intoriorh' 1 00

anteroposterior { ,

i medially 4d

transverse (witliout gomphosis) 1-90

vertical (at middle) 23

Diameters of anal marginal i vertical 35

Diameters of hyosternal ■

SYNOPSIS OF THE VEBTEBR^VTE FAUNA OF THE PUERCO SERIES. 307

Measurements.

No. 2. Inch.

r anteroposterior 1.13

Diameters of vertebral bone ■! transverse 1.20

I. vertical 40

This is the oldest species which can be icferred to this genus.

■! .MAKSUPIALIA.

All of the Puerco implacentals belong to the suborder Multituberculata, and to
three families, which are dclincd a>J follows :*

Premolars present, compressed and trenchant in form Phir/iaulaeida.

Premolars present, molariform, dillerent from true molars in form Chirogida.

Premolars wanting or rudimental Polymasiodontida:.

To the first belong, in the Puerco fauna, Ptilodus and Neoplagiaulax ; to the
second, Chirox ; and to the third, Polymastodon.

NEOPLAGIAULAX Leinoine.
Bulletin of the Geological Society of France, 1881, November.

I have referred the American species of PlagiaulacidsTi with but one, the first
inferior premolar, to the above genus. Those with two premolars I have referred to
Ptilodus (October, 1881).

Neoplagiaula.\ molestus Cope ; Amer. Naturalist, 1886, p. 4.51.

This species was establi.shed on a separate first inferior premolar. T now
describe a mandibular ramus whicii probably belongs to it. It has sullered the loss
of the crown of the molar, and the greater pan of that of the i)remolar teeth. The
latter has its base oblique in both the vertical and horizontal directions ; its posterior
base rounded. True molars small, and on the inner side of the coronoid process.
Incisor one, rodent-like. Angular infiection of the mandible well developed, but the
posterior bordei- has 1)een broken away. Its form is lobust, especially at the exteiiial
base of the large premolar tooth, where a longitudinal swelling rises posteriorly, and
disa])pears in the base of the coronoid process. Below and posterior to this protuber-
ance the mas.seter fossa extends, having an oblique boundary below in the externo-
superior face of the external infiection of the base of the ramus. The internal infiec-
tion commences more posteriorly, and projects inwards at a right angle to the vertical
plane. Anterior to these infiections the inferior outline of* the i-amus is gently con-
vex downwards in conformity to the arc required by the alveolus of the hu-ge

*Amer. Natunilist, 18S4, p. 687 ; 1887, p. !;07.

308 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

inferior incisor. The latter issues from the jaw at a considerable distance in advance
of the first premolar, leaving a wide diastema, which has a rather wide and obtuse
superior surface. The alveoli of the true molar are within the base of the coronoid
process, and indicate a tooth of much smaller size than the premolar. The symphysis
is short, and is indicated almost exclusively by rugosities of the inferior and anterior
border of the extremity of the ramus. The articulation has been a loose one by
ligament.

MeasuremcnU. M.

Length of fragment of ramus 057

" " diastema 013

anteroposterior 013

Diameters p. ni. i , . , „„_

transverse posteriorly 005

Length of base of m. i (estimated) 006

Depth ramus at diastema , 016

" << «' posterior base of p. m. i 031

( vertical 007

Diameters of incisor I

> transverse 004

The tooth from which the species was originally characterized has the following
characters : Length of base one-third greater than in Neoplagiaulax americanus, and
there are fifteen keel-crests on the side of the crown, while there are but seven in the
N. mnericanus. The outline of the crown is elongate and moderately convex, and
less elevated than in the known species of Ptilodus. The irregularity in the outline
of the base of the crown is less than in the other species, and the diameter of the
roots is subequal. The anterior base of the crown is not excavated for the second
premolar as in Ptilodus. Length of base of crown, 16 mm. ; elevation at mid-
dle, 8 mm.

In size this species is about equal to the P oly mastodon foUatus.

CREODONTA.

Important additions to the knowledge of this suborder have been made recently
by Professor W. B. Scott* and Dr. Max Schlosser.f The former has determined the
presence of the siibcylindric postzygapophyses in Didymictis, and I have ascertained
their existence in Esthonyx and Dissacus. Scott shows that in HyjTJuodon the char-
acters of the carpus and of the brain coincide with those of this group. He also
finds an os centrale in that genus and in Mesonyx, and Osborn has found it in
Esthonyx.

In the following pages considerable additions to the general osteology of Hemi-

* On Some New and Little known Creodonta ; Jour. Acad. Phil., 1880, p. 15.5.

\ Die Affen, Lemuren, Chiroplcrcn, Insectivoren, Marsujiialia, Crcodouten und Caniivoren des Europiiischen Ter-
tiars ; Wien, 1887. Alfred Holder.

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE rUEItCO SERIES. oOO

ganus, Onychodectes, Miocljv3nus, Chriacus and Dissacus are made, and they throw
considerable light on the phylogeny of the various genera. It is apparent on all
grounds that MioclaMnis is the central type of Creodonta for the Puerco Epoch, and
that all other types of the suborder may be traced to it as derivative, either by pro-
gressive 01" retrogressive specialization. Thus the line which our present knowledge
ends with Hemiganus, is one of simplification and reduction of the molar dentition at
the expense of increase in the development of the canine dentition, as takes place in
the line of the seals. In the line of the Mcsonychid:c another kind of simplification of
the molar dentition of the inferior series appears, and the feet undergo specializa-
tion, apparently in accordance with the needs of a terrestrial life. A third line
leading towards the Oxya'uidiB commences with the genus Chriacus, which is easily
modified into Stypoloj)hus on the one hand and Deltatherium on the other, the latter
having some special characters of its own, and forming probably the end of its line.
The line which gives us Miacida', and in Miocene times the Carnivora, probably
diverged from a Miochenus, but from a species of an earlier horizon than the Puerco,
since two species of Didymictis are already contemporai'ies with the Miochenus of
that age. These relations may be displayed as follows :

Carnivora Oxymnidm

/

Didymictis Deltatlicrium Stypolophus

Miacis Chriacus

Mioclicnus
Triisodon Onycliodecles

Dissacus Conoryctes

PachysDna Ilemiganus

/
Mesonyx

In the reference of these genera to families, lack of information as to ceitain
parts of the structure forbids liiial C()iK;lusi()ii>. Leaving aside the JNEiacida' and
0.xytcnida\ we find that tliu I'roviverridiu (Schlosser, Leptictida-, I'iv. II, (Jopf ) cm-

310 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

braces Deltatheiium, Mioclfenns, and probably Triisodon and Onychodectes, of the
Puerco genera. Dissacus is on the bonndaries of the Mesonychidjie, but the nngues
are unknown, and the astragalus is not trochlear. That Hemiganus can be referred
to the Proviverridse is improbable. The characters of a family Hemiganida? would be :
Sujjerior molars with confiuent roots, and trituhercular crowns; inferior molars quadri-
tubercular ; astragalus not trocJilear ; ungues much compressed. • The sole character
separating this flimily from the Proviverridjie is the confluence of the roots of the
superior true molars. It is approached in this respect by the genus Conoryctes,
which will, however, remain in the Pl•oviverrid^e as an aberrant form, connected with
Miocla^nus by Onychodectes. The tendency of this group is to resemble the Tseni-
odonta, but whether there is any affinity involved in this resemblance does not yet
appear.

The line of the Mesonychidaj shows definite changes in the following respects :
First, loss of the internal cusps of the inferior molars. The diminution of these
cusps is seen most prominently at first in the genus Triisodon, and more distinctly in
the later T. quivirensis than in the earlier T. hiculminatus. It has progressed far in
Dissacus. Second, in the reduction of the number of the digits ; the hallux has
become very small in Dissacus and is gone in Mesonyx. Third, in the production of
a distinct facet for the cuboid bone on the extremity of the astragalus. The contact
of the astragalus and cuboid is seen in many if not all Creodonta, but the navic-
ular and cuboid surfaces are not distinguished from each other. In Mesonyx and
Pachysena it is well known that they form distinct facets separated by an angle, as in
Perissodactyla. This structure appears in a primitive condition in Dissacus, the
angle being obtuse and rounded. It is an excellent illusti'ation of the origin of a
zoulogical character. The thoroughly diplarthrous type of the astragalus in Mesonyx
is associated with an ungulate form of ungual phalanges, and I suspect that the
resemblance is not accidental. The form of the distal end of the diplarthrous astra-
galus is probably due to impacts combined with energetic flexion and extension.
But why the structure should appear in Mesonyx and not in Phenacodus is, as yet,
an unanswered question.

Although it is now evident that the Pinniped Carnivora cannot be derived from
the genus Mesonyx, as is truly maintained by Scott, it appears to me that they have
been derived, like other Carnivora, from some form of Creodonta. The retrograde
development of the molar teeth probably passed through stages like those of Dissacus
and Hemiganus, and the ungual phalanges might have been easily derived from such
as are possessed by Piichyjena.

SYNOPSIS OF THE VEIiTKUKATE FAUXA OF THE TUEKCO SEKIES. ."Jll

HEMIGANUS Oope.
Amcr. Naturalist, 1882, p. 831 : Tertiary Vertebrata, 1885, PI. XX HI c, figs. 7-13 ; Amer. Naturalist, 1885, p. 4!1l'.

The claws are large and compressed like those of a prehensile-footed carnivore.
The astragalo-tibial articulation is nearl}' flat. The femur is ver}- robust, and has a
low third trochanter, as in Bunotheria generally. The vertebr;e of the neck are short
and wide. The jaws liave a very lai'ge and wide coronoid process, as in Calamodon,
and the horizontal rami are very robust. Only one true molar (the tirst) is preserved,
and it has the crown worn. Its outline is subround, with a notch on the internal side.
There are probably but two true molars ; they have two roots. There are at least four
premolariform teeth, and their crowns are short, obtuse cones, with a low heel-like
expansion at the inner side of the posterior base, and have but a single root. They
resemble very nearly the teeth of some of the eared seals. There is a robust canine
tooth in the upper jaw, which is not separated from the premolars by a diastema.
There is at least one superior incisor, but the exact number is unknown. There is a
large tooth on each side of the symphysis of the lower jaw, but in the specimens it
is not in place. It has enamel on the anterior face only, and its apex is worn ti'ans-
versely. The wear descending passes to one side of the middle line. It evidently
has a median position, and may be therefore an incisor. Its form reminds one of that
of the second inferior incisor of Calamodon, but the enamel-face is much shorter.

Should the lai'ge inferior teeth be canines, the mandibular dentition will greatly
I'csemble that of the seals, as does that of the maxillary bone. The absence of })ost-
orbital angles resembles the condition in the Phocidaj. The wide vertical coronoid
process and the flat vertical angle are as in Calamodon. The sagittal crest is ele-
vated, and the brain-case very small.

This genus resembles in several respects the Tseniodonta, and confirms the pro-
priety of the union of that group with the Creodonta into the ordei' Bunotheria.

The typical species, H. vuUuosus, was an animal of probably the size of a grizzly
bear. A second and considerably smaller species is described below.

IIemioanus OTAKiiDENS Cope ; Aiiicr. Naturalist, 1885, p. 495. llujus operis, Plates IV and V.

Only one individual of this species has been found, but it is represented by many
parts of the skeleton. It was a plantigrade beast of about the size of a black bear,
of robust proportions, and with a wide head with an exceedingly short thick muzzle,
armed with some formidable teeth in front. These, with its sharp claws, made it the
most formidable animal yet known of the Piierco fauna, excepting its larger and
more powerful congener, the H. vultuosus.

The nares are well roofed by the nasal bones, which border the premaxillarieR

A. I'. S. — \()L. XV I. 2n.

312 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE rUERCO SERIES.

above to the line of the front of the second superior incisor, by a wide sutural sur-
face. The superior process of the premaxillary bone is short, not extending posterior to
the vertical line of the posterior face of the superior canine tooth. A small foramen,
perhaps the infraorbital, issues above the second tooth posterior to the canine. Ex-
terior to the third tooth that follows the canine, the external face of the maxillary
bone spi-eads outwards as though forming the malar process, and that this is the case
is rendered probable by its smooth superior surface, which is the inferior orbital bol-
der. Just anterior to the orbital border, a large foramen from the maxillary antrum
perforates the maxillary bone. The two teeth in the maxillary bone are injured, but
the anterior has a conical crown and a single root, while the crown and Ixise of the
second are widened a little transversely. I can find no superior true molars in the
collection.

The mandibular rami are remarkable for the shortness of the dentary portion, and
the elevation and width of the coronoid process. Tlie condyle is elevated above the
alveolar border of the lower jaw, when the inferior border of the ramus is horizontal.
The ramus increases in depth anteriorly, as in Tseniodonta, to accommodate the large
anterior teeth. The inferior border is straight and compressed, and the posterior
border is gently concave to a short rectangular angle, which does not extend poste-
riorly to the line of the base of the condyle. It is therefore much less prominent
than in Creodonta generally, resembling in this respect the Tseniodonta. There are
four alveoli for single-rooted molars, and apparently another one in front of the ante-
rior one of the four. This would give seven molars, the first true molar having the
form of a premolar ; but the distribution of the teeth is not quite certain. As already
described, the heel of these premolariform teeth is partly internal.' The first true
molar may be one of these simple teeth. The second has two roots, and the crown
is about as wide as long. The crown consists of an anterior portion, which is slightly
elevated above a posterior heel. The superior face of the crown is worn by mastica-
tion so that its construction is not evident, but there is no trace of a division between
fourth and fifth tubercles, so that I suspect that the latter did not exist. It is not
j)robable that there were well-marked cusps on the heel.

The parietal region of the skull is very much compressed, and the sides slope
regularly upwards to the elevated sagittal crest. The temporal ridge is an oblique
angular line of the surface, and the frontal regiou is flat. No other parts of the skull
are preserved.

Measurements of Skull. M.

DepUi of maxillary bono at ]). in. iv 038

" " nasal " "canine 031

Length of maxillary boiiu to orbital border 045

SYNOPSIS OF THE VERTKBRATK FAUNA OF THE PUERCO SERIES. 313

Measurements of Skull. M.

{anteroposterior (oblique) 015
transverse (oulique) <Ji-

Lengths of p. m. iii and iv Old

" " seven inferior molars 053

I anteroposterior 085
tn»nsverse 072
vertical 0'"'0

{to condyle 070
l\-r
to angle 0^><'

f anteroposterior 054

Diameters of coronoid \ „ , .

[ vertical 044

Width of condyle 017

Depth of ramus at coronoid 081

' m. iii 034

" " " "p.m.iv 042

Vertical depth of sagittal suture 027

Only cei'vical vertebriB are preserved. These have small anteroposterior diame-
ter, and their transverse exceeds their vertical diameter. In general .they resemble
those of Periptychus. The atlas is peculiar in the small anteroposterior diameter of
the paradiapapohysis, whose base is perforated by an anteroposterior canal. It sends
upwards a vertical keel to opposite the middle of the facet for the axis. The axis has
a cylindric and rather slender odontoid process whose superior extremity is obliquely
beveled on a curve. Its articular surface is continuous with the large atlantal facets
laterally and inferiorly. The longitudinal axis of the cervical centra is oblique to the
horizontal, showing that the head was elevated above the body. The llooi- of the
neural canal is pierced by a foramen of considerable size on each side. A posterior
(? seventh) cervical has a greater anteroposterior diameter than the two which pre-
cede it, and the vertical diameter is relatively greater. The posterior aiticular face
of all three is slightly concave.

JUeasuremenis of Vertebrw. ^^f-

Anteroposterior diameter of atlas 020

Width of axis at atlantal facets 0;!8

Length of odontoid process "1 1

I anteroposterior 0085
vertical 010
transverse 02C

f anteroposterior Oil

Diameters of centrum cervical ? v | vertical 015

(. transverse 029

f anteroposterior 01.)

Diameters of centrum cervical ? vii J vertical 010

I. transverse Oi.8

314 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

The anterior limb is represented by parts of both uhite, part of one radius, and
a metacarpal of the pollex. More than half of one ulna is preserved. The olecranon
api)ears to be short and terminating in an acute apex at the basal border ; but it may
have been broken off. The humeral cotylus is oblique, extending backwards and out-
wards, and inwards and forwards. The i^ostei'ior border is elevated into a ridge,
which is convex forwards. The anterior marginal ridge is limited to the external
part, and it extends outwards, downwai'ds, and then backwai'ds, overhanging the
internal face of the ulna. The radial facet is flat, and slopes gently, and not steejily
inwai'ds, it is bordei'ed on the outer side by a low ridge, external to which is a longi-
tudinal groove. Both do not extend far distad, and the superior edge of the shaft of
the ulna is narrow and convex. The inferior edge is similar except below the humeral
cotylus, whei'e it is transversely flattened, the inferior face turning upwards on the
inner side posteriorly. The internal side of the shaft of the ulna is concave, and the
extei'nal side is convex. The head of the radius is a transverse oval, with subequal
bi-oadly rounded extremities. The superior border is openly shallowl}^ excavated,
while the inferior is obliquely beveled for the ulnar face. A short tuberosity projects
longitudinally from the middle of the ulnar facet. A metacarpal, supjjosed to be that
of the pollex, is quite short and robust and has a pi'oximal excavation of the internal
side for the trapezium. This concave facet extends half its length. The distal end
is a subround convex facet which presents outwards. It has neither median keel nor
groove. It indicates a robust digit.

Measurements of Anterior Limb. M.

Length of fragment of ulna 117

DepUi at miildle of glenoid surface 020

Width" " ■' ■• " 027

Depth at coronoid 0;!2

Ilc'iilh at middle of sliaft 0215

I,. , , , , r fvertical 017

i'lami'lers head oi railuis \

[ transverse 02G

,,. , , r. T n,- r • , , f vcrtical 012

Uiamelers shalt radms .04.) from pro.ximal end -

(^ transverse 013

Length of metacarpal of polle.x 020

TV , r . 1 r . f , (^''^'■li<"il 013,5

Diameters distal facet of do. \

[ transverse 013.5

The femur lacks the distal extremity so that it is not possible to determine its
exact length. Its proximal portion is robust, and about as large as that of a fully
grown pig. That the animal is not fully grown is shown by the fact that the epi-
physis of the head is not united, although it is preserved. The projection of the
great trochanter is about equal to that of the head, and is robust, and truncate both

SYNOPSIS OP THE VERTEBRATE FADNA OF THE PUERCO SERIES. 315

proxiraally and externally, and incloses a considerable trochanteric fossa. The head
has a large fossa lor the round ligament which is near the neck, from which it is sep-
arated by a low boi-der. The little trochanter is quite prominent, and is reverted, but
it is not connected with the great trochanter by a ridge. The thii-d trochanter is
well developed, and has a wide external surface, whose anterior edge is recurved for-
wards. Its upper portion overlaps the line of the infeiior edge of the little trochanter,
being higher up than in Pachyania. The middle of the shaft is somewhat depi'essed,
and its margins are rounded.

The proximal part of the left tibia, and the distal part of the right, give the char-
acters of that element. The proximal part is laterally crushed. It is evident, how-
ever, that the crest is large and obtuse at the apex, and that the spines are low ridges.
The e.xterno-posterior border forms a roughened lidge for 35 mm. below the intei-nal
femoral surfiice. and ceases rather abruptly below. At or near the middle the
shaft is normally somewhat compressed, and slender. The malleolus is very promi-
nent, and terminates in an apex in its internal plane. The astragalar surface is l)ut
little o])lique; the libular articular surface is large.

One of the metatarsals shows that the foot was short, since it is neither the lirst
nor the fifth. Its proximal face is concave in the transverse direction, but nearly
sti'aight anteroposterioi'ly. The arc of the phalangeal face is less than half a circle,
and is slightly concave in transverse section. It is divided medially at its inferior
foui'th by a short, narrow, and low trochlear keel. Inferior border of phalangeal face
Ijrominent and o])enly emarginate. An ungual phalange is preserved, but whether
of the anterioi- oi' posterior foot I do not know. Its apex is lost. It is strongly
compressed, and has a narrowly rounded supei-ioi" border. The phalangeal
cotylus is deeply excavated, and is rather narrow, and has a weak median
keel. The superior ])rocess ovei-hangs the phalangeal surface rathei* further than the
inferior. The tuberosity for the llexor tendon is a longitudinal oval, Avith sui'face
transversely convex, which gradually ascends to the narrow but flat inferior surface
ol" what lemains of the phalange. The large nutritive foramen enters above its
middle.

Measurements of Posterior Limb. M.

I.cugtli of f'omur preserved 123

Widtli at liead 0(!1

Length pro.ximad of little trochanter, inclusive 004

" third " " 085

^. f nnteroposlcrii)r 010

Diameters .shaft of femur \

I transverse 029

^. . , ( anteroposterior 023

Diameters shaft of iibia •(

transverse 010

316 STKOPSIS OF THE VEKTEBRATE FAUTSTA OF THE PUERCO SERIES.

Measurements of Posterior Limb. At.

( anteroposterior 025

Diameters distal end of tibia -^

i transverse OdU

Lengtli of plialange 024

anteroposterior 013

Diiuneters of distal end -J

transverse .011

Depth of ungnis

f at tendinous insertion 019

I anterior to tendinous insertion 013

I at phalangeal surface 009

Width of unguis [ at tendinous tuberosity 006

[ anterior to tendinous tuberosity 005

CONORYCTES Cope.

Proe. Amer. Pliil. Soc, 1881, p. 486; Tertiary Vertebrata, 1885, p. 198. Hexodon Cope; Amer. Naturalist, 1884,
p. 794.

Additional material representing the type species of this genus enables me to
give the chai'acters more fully than hitherto. The C comma experienced an early
obliteration of the details of the structure of the crowns of the molar teeth through
the thinness of the enamel layer and the mastication of hard substances, so that they
are seen in but few specimens.

The superior true molars, and the first premolar, have two external conical cusps,
and an internal triangular table, whose inner angle is produced downwards to a line
with the apices of the internal cusps. The inferior premolars and the first premolar
have the anterior part much elevated above the posterior. The former consists of a
large external and a small internal cusp joined to near their summits, except on the
first premolar which has but one anterior cusp, which is simple acute cone. A rudi-
mental lifth cus]) is present on the true molars.

The ma.stication in this genus is aftected by a scooping action of the keels of the
inferior molars against the internal table of the superior molars by a motion which is
partly transverse, as in Onychodectes.

There are probably four infei'ior i)remolars in this genus, but the anterior two
have but one root each, and are close together. My supposition that some specimens
had but three premolars led me to propose the genus Hexodon, which is now aban-
doned.

The position of the genus is doubtful, owing to the absence of the ungual pha-
langes. It is probabl}' Ci'eodout rather than Condylarthrous, for two reasons; one is
the close resemblance of the dentition to those of Onychodectes and Hemiganus,
between which it takes a natural position. The other is, that it displays no resem-
blance to any of the Condylarthra in the details of its structure.

But one species is known to me.

SYNOPSIS OF THE VERTEBKATE FAUNA OF THK I'UEKCO SERIES. 317

CoNORTCTES COMMA Copc ; loc. cit., Hexodon mole»tu» Cope ; loc. cit., fig. 3.

This animal was about the size of a wolverine, of which species one is romindcd
by its robust characters. Ft liad an elevated sagittal crest and a strong iiiion. In a
series of teeth which arc but little worn the following characters may be discerned.
The crown of thi- infeiior canine has a flat inntr face, beyond which llie antciior sur-
face extends inwards, forming a rib-like border. The enamel on the internal and [)os-
terior faces extends but a short way from the apex, and is thin, while on the convex
anteroexternal face, it extends below the usual position. It thus approaches the condi-
tion seen in Ileniiganus. A similar state of affairs is seen in the molars, where the
enamel is extended much further on the external face of the inferior and the internal
face of the superior molars than is usual, approaching the genera mentioned and also
the TjeniodonUi in this respect. There are no cingula on any of llu' molars of either
series excepting on the external side of the superiors ; and there it sends out a process
or cusp between the two external cusps. The crowns of the inferior molars are notched
at the junction of the antei'ior and posterior parts. The notch is the section of a
vertical groove from the base of the crown on the external side, and of a very short
superficial one of the internal side. The fifth cusp is median, and about opposite the
rim of the heel in elevation. The grooves of the first premolar are similar to those
of the ti'ue molars. There is no anterior basal cusp. The heel is large and has a
raised border on the posterior and inner sides, and an external median lateral conic
cusp. This when worn joins the cuived crest, forming a comma-shaped figui'c.

A si)ecimen represented by fragments includes a canine chai-acteristic of the
genus, but of such relatively smaller size, that I suppose the animal to be a female.

Seven individuals have come under my observation.

ONYCIIODKCrKS Cope, gen. nov.

Superior molars ti'itubercular, the external cusps distinct ; the internal with the
intermediate confounded in a prismatic form with flat grinding surface, and whose
internal angle rises claw-like to an elevation equal to that of the external cusps, and
without cingula or api)endicular cusps. First premolar with but one external and
one internal cusps. Inferior molars seven, the true molars Avith five cusps, the ante-
rior triangle distinct. Last inferior molar with a heel; canine large.

Caudal vertebne robust. Iliiini rather slender, flat-triangular in section, and
with a small anterior-inferior spine. Scapula with coracoid hook, and abruptly rising
spine. Astragalus with unequal trochlear ridges, the internal the lower. Internal
face oblique, but less so than in the species of Miochenus where it is known, and not

318 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE RUERCO SERIES.

produced farther posteriorly than the external face, which is vertical. Head depressed,
convex, and without angles. Cuboid with a small external distal facet.

This genus is intermediate in character of teeth between Conor3'ctes and Mio-
ela^nus. The molai-s are those of the former as to the internal portion of the crown.
The external cusps are moi-e those of MiocijBiius, and there is but one external cusp
of the first premolar, while there are two in Conoryctes. It is in the i-emarkable
table-like form of the interior part of the crown and the hoof-like production of the
internal angle, that Onychodectes differs from Miochenus.

But one species is known to me.

Onychodbctics tisonensis sp. nov.

Two individuals certainly represent this species in my collection, both of which
include superior molars, while a third, which includes two mandibular rami, belongs to
it. Of the former the typical and most important specimen includes the following
parts. Both maxillary bones with the posterior five. molars; the left mandibular
ramus with all the alveoli, and the second true molar in place; the glenoid extremity
of the scapula; the left ilium; the right astragalus and cuboid.

It is characteristic of the superior molars that the external cusps have a lenticular
section, and not a triangular or a round one as in the species of Mioclajnus and Chri-
acus. The external cusp of the first premolar is large and elevated, and has the same
fore and aft lenticular section with obtuse euttino^ edges. The internal table of the
crown is of parabolic outline and its edges are right angles. The sides ascend per-
pendicularly to the alveolar border without the least trace of cingulum or other irreg-
ularity. The ci-own has a weak external cingulum, which does not support any
cusps. The posterior of the external cusps of the third molar is well developed, and
nearly in longitudinal line with the anterior.

The anterior triangle of the second inferior molar has a broadly rounded external
a[)ex, and it is a little elevated above the heel. The latter has two internal marginal
cusps, but its summit is so worn that the form of the surface cannot be further deter-
mined. N^o cingula. The manner of mastication is such as to wear the crown
obliquely from within outwards in conformity with the form of the inner table of the
su])erioi- molars. The anterior triangle fits, as usual, between two adjacent superior
molars, and the claw-shaped internal border of the superior molar worked, scoop-like
across the heel, the inferior molar moving from without inwards. The motion was the
same as in Conoryctes. In neither genera do I possess the glenoid surface for the
mandibular condyle, but it is highly probable from the evident lateral movement of
the lower jaw that neither genus possessed a preglenoid crest as is found in Mioclsenns.

SYNOPSIS OF THE VKKTEBUATE FAUXA OF THE PUERCO SEUIBS. 319

The mandibular ramus is slender and moderatclj^ stout, especially so at the ante-
rior base of the eoronoid process. It follows that the anterior border of the masseteric
fossa is well marked, but there is no distinct inferior border. The angle is prominent,
straight, and compressed; apex lost. The dental foramen is below the middle of the
base of the eoronoid; and in lino with the alveolar border. The symphyseal surface
is smooth. The fourth premolar is close to the canine, and to the third, and has one
root. Premolars all closely contiguous. From the appearance of the alveoli the last
infei-ioi- molar is of reduced size. Enamel everywhere smooth.

Measurements ; from one individual. M.

Length of posterior five molars 031

" " true molars 0155

(anteroposterior 005
vertical, outside 004
tjansverse 006

(anteroposterior 0055
vertical, iusiile 005
transverse 0075

(anteroposterior 0043
vertical, inside 0043
transverse 006

Length of inferior molar scries 041

" " true molar series 031

( anteroposterior 007

Diameters m. ii < „„,_

i transverse ou4o

Depth ramus at m. iii 015

p. m. iii 014

anteroposterior 016

transverse Oil

anteroposterior 013

transverse 009

Diameters glenoid cavity of scapula
Diameters peduncle of ilium

( anteroposterior 031

Greatest diameters of astragalus < „,_

^transverse 017

Width of trochlea 0135

" head Oil

Length of cuboid 012

The specimens are all from the lowest beds of the Puerco.

MIOCLiENUS Cope.
I'roc. Amer. Phil. Soc, 1881, p. 489; 1883, p. 547 ; Tertiary Vertebrata, 1885, p. 334.

An examination of my material of species allied to the types of this genus results
in the following conclusions :

The fifth or anterior inner cusp of the inferior molars in this genus, displays
various conditions of development down to absence. In the species which I formerly

A. p. S. VOL. XVL 2o.

320 SYNOPSIS OV THE VERTEHRATE FAUNA OF THE PtJERCO SERIES.

referred to Sarcothraustes it is well developed, although small and conic in shape,
and occupies an elevated position on the inner side of the front of the crovi^n, close to
the fourth cusp. In the M. interrnptus, it is less distinctly developed, and is wanting-
from the last molar in some of the specimens. In M. acolytus the fifth cusp adheres
closely to the fourth, and is in some specimens wanting. In the M. suUrigonus the
fifth cusp is indifferently absent, or present in small development on the second and
third molars. In nearly all the species of the genus as here constituted the fifth cusp
is present on the first inferior true molar ; but in the M. turgidus and species most
nearly related, it is absent. The species of the latter type differ from the forms allied
to M. coryplums and M. ferox in their robust premolars, but the M. opistliacus fur-
nishes a passage between the two. It does not seem practicable to divide the genus,
as I once proposed,* on the presence or absence of an interior cusp of the third supe-
rior premolar. While this cusp is present in the M. turgidus, it is wantingf in the
nearly allied M. zittelianus, and the M. opistliacus.

In the species referred to the second section of the genus (Goniaeodon, type M.
levisanus), the fifth cusp is quite distinct, but is median in position and near the base
of the crown (except in M. Iieilprinianus), forming an anterior angle in the outline of
the crown. These species I have referred sometimes to TriTsodon and sometimes to
Diacodon, but I think I have now found their proper position.

With six new species now added, the total of those embraced in the genus is
twenty-four.

Parts of the skeleton of M. antiquus, M. coryphcms, M. levisanus, M. foverianus,
M. corrugatus and M. ferox are preserved, and do not present any but specific differ-
ences. I must here correct an error into Avhich I fell in describing the M. ferox (Ter-
tiary Vertebrata, p. 331) in calling the fibula the radius (Plate XXIV f, fig. 11). The
distal part of this element is very robust in that species.

I. Inferior true molars with anterior triangle of three cusps [Sareothraustes).
Length of inferior premolars, .009 ; depth of ramus at p. m. iv, .055 ; superior true molar ii, .023 by .016, M. antiquus.
Inferior true molars, .052 ; premolars, .043 ; infer, m. iii, .015 ; depth of ramus at m. i, .052; length sup. true molars,

,040 M. conidens.

Inferior true molars, .037 ; premolars, .047 ; depth ramus at p. m. iv, .041 M. bathygnathua.

Last inferior true molar, .012 ; depth of ramus at m. ii, .036 M. crassiewpis.

Inferior molars, .037 ; :n. iii, .012 ; premolars, .037 ; sup. molars, .030 ; depth ramus at m. iii, .037 M. eoryplmus.

Inferior molars, .030, wide; premolars, .030, close together, the first subconic ; ramus at ni. i, .020 3J. peiitiicus.

Superior molars, .030 ; depth ramus at inferior m. ii, .035 .V. gaudriaims.

Inferior true molars, .019; premolars, .018, the anterior spaced, the first anteroposteriorly short ; depth ramus at m. 1,

.013 M. ikterruptus.

Inferior true molars, .018 with anterior elevated ledge and basin-like heel ; depth at m. i, .014 31. lydekkerianm.

* Proc. Amer. Phil. Soc, 1883, p. 312. f Genus Oxyctenus Cope, 1. c.

SYNOPSIS OF THE TEUTEHHATE FAUXA OF THE PUEItCO SEHIES. 321

Inferior molars, .017 ; Ihe first small, the last large, .00G5 ; depth of ramus at m. i, .013 M. jUhoUanus.

Inferior molars, .011 ; fifth lobe small, heel with one cusp ; depth ramus at m. ii, .007 M. aeolytus.

II. Inferior true molars wilh anterior inner tubercle submcdian and depressed ^ Goniaeodon).

Inferior true molar i or ii, .01 1 ; fifth cusp elevated, inner edge of heel crenate M. heilprinianut.

Inferior true molars, .0115 ; anterior paired cusps much elevated ; last molar elongate, .007 il. asaurgens.

Inferior true molars, .023 ; last inferior and superior molars reduced, the inferior, .007 M. levisaniu.

Inferior true molars, .040; list molar large,. estimated, .013 ; ramus deep ; fifth cusp low, lieel not crenate inside

J/, rusticus.

III. Inferior true molars (except first) without anterior inner fifth tubercle (JUioclainus).
A Premolars of smaller diameters than true molars.

a Heels of inferior molars supporting cusps.

,9/9 Superior molars subtriangular, rounded at inner angle.
Superior true molars wide, .017, wilh small tubercle between e.vternal cusps, and strong posterior internal ciugulnm ;

inferior molars, .023 ; depth of ramus at m. i, .012 3t. suhtrigonvs.

Superior true molars, .019 ; external cusps close together and without intermediate cusp .)/. cuspidatus.

Superior true molars wide, .021 ; inferior molars, .025 ; premolars, .035 ; depth of ramus at ni. i, .015, M. protogonioides.

Inferior true molars, .028, the anterior with a strong anterior fifth cusp M. flovcriatius.

aa Heels of inferior molars bounded by a curved crest (anterior inferior true molar with median anterior
fifth cusp).

Enamel wrinkled ; length of ramus, .110 ; depth at m. i, .020 ; true molars, .035 ; premolars, .035 JU. corrugatus.

Enamel roughish ; length of ramus, .160; depth do. at m. i, .034; true molars, .038; premolars, .041 M. ferox.

A A Superior or inferior premolars of equal or greater transverse diameter than the true molars.
a Heels of inferior molars supporting cusps.
Last superior premolar with large internal cusp ; superior molars, .013 ; inferior molars, .010 ; premolars, .019 ; depth

ramus at m. i, .010 ; last molar large M. opiathaeua.

aa Heels of inerior molars supporting a curved crest from an external cusp (last molars small ; no fifth
cusp on inferior m. i).
Cusps obtuse, premolars swollen ; superior molars, .018 ; premolars, .039 ; superior ji. m. iii and iv, wilh internal

cusps ; depth of ramus at m. i, .012 if. turgidvt.

Like last but smaller, and internal cusp wanting on superior p. m. iii, and rudiniental on p. m. iv ; superior molars,

.015 ; premolars, .022 M. siiulianus.

Much smaller; inferior molars, .0125 ; depth of ramus at m. i, .007 ; exterior cingulum of superior molars strong ;

premolars in both jaws large ; heel of lower molars with high edge M. turgidiinculua.

Still smaller ; superior molars, .008, with strong external cingulum ; premolars not enlarged ; inferior m. iii not much

smaller than m. ii M. aeolytui.

Small ; superior molars, .010 ; without external cingulum ; intermediate tubercles very indistinct ; inferior ni. iii much
smaller than m. ii ; premolars small M. minimus.

JIlOCL.KSnS BATnTC.SATIIUS Sp. nOV.

Established on a nearly entire left mandibular ramus which supports the true
molar teeth and the second premolar. The species is distinj^uishcd at first sight by
the relatively large size of the i-amus, as comi)ared with the dimensions of the true
molais. The latter do not exceed in size those of the M. ferox, but the jaw is that
of an animal as large as the M. conidens.

The ramus is of compressed form, and is rather thin posteriorly. It is deepest
below the first true molar. Symphyseal sutuic very coarse. Coronoid process large.

322 SYNOPSIS OF THE VERTEBRATE PAUNA OF THE PUERCO SERIES.

The condyle is broken off, but it was probably elevated a little above the line of the
molars. Masseteric fossa well defined anteriorly, but not inferiorly. The angular
region is perfectly flat and straight. The dental foramen is below the middle of the
coronoid process, and just below the line of the bases of the crow^ns of the molars.
Mental foramina two, in horizontal line.

The inferior canine was large and compi-essed; part of its alveolus only remains.
The molars and premolars are small. The crowns of the fourth and thii'd premolars
are lost, but judging from the extent covered by the two roots of the third, it is as large
as or a little larger than the second. The second has a simple slightly compressed
conical crown, with a small posterior transverse heel. The first premolai* is a large
tooth, and judging from the alveoli, of greater anteroposterior extent than any of the
true molars. The true molars are somewhat worn with use. The anterior part of the
crown is more elevated than the posterior, offering in this a resemblance to the M.
conidens, and a contrast to the M. corrugatus and M.ferox. The fifth cusp is en-
tirely on the inner side of the crown, and is close to the fourth, which it did not equal
in elevation when unworn. The heel of the third molar is short so that the length of
the crown does not exceed that of the second true molar, which is the most robust of
the series. There is a cingulum on the external base of the true molars, and another
on the internal base of the anterior half of the crown of the same. The enamel is
minutely rugose.

Measurements. M.

Estimated length of ramus to below condyle 185

Depth at m. i 044

" m. iii 040

Width at base of coronoid in front 015

Length of molar series 081

" " true molars 037

Diameter of canine alveolus (greatest) 020

^. , , f anteroposterior 0145

Diameters base of p, m. i .^

I transverse 009

f anteroposterior Oil

Diameters m. i \

transverse 0095

{anteroposterior,
transverse

, ,,„„_ 0125

Diameters m. u {

010

^. , ... f anteroposterior 0125

Diameters m. lu ^ '

( transverse 0095

From the lowest bed of the Puerco of ISTew Mexico. D. Baldwin.

If the skull of this animal was proportioned as in allied forms it was about the
size of that of the black bear ( Ursus americanus).

SYNOPSIS OF THE VEWTKBRATE FAUXA OF TTIE PUERCO SERIES. 323

MioCLiENns CKA8S1CUSPIS Cope. Conovjjcles crassicuapia Cope ; Terliuiy Vcrtebrata, 1S85, p. 201, PI. xxiii e, flj;. U.

Better specimens of the Conoi'yctes comma and of this species show that there is
nothing in common between them. The M. crassicusj)is is allied to the M. cori/pJui'us,
but has a deeper ramus of the mandible and was probably a considerably larger animal.
The last inferior true molar is relatively of smaller size than in that species. Parts
of lower jaws of three individuals are in my collection. These show that the fifth
cusp is well developed in all of the true molars, and that the anterior part of the
crown is higher than the posterioi-. The type and a second specimen are from the
Lower Puerco ; the exact horizon of the other is unknown.

Mioci..«Nus coKYPHiEus Cope. Sarcothraustes cor yphceus- Cope ; Amei. Naturalist, 1885, p. 38G.

Nine individuals represent this species in my collection. The best of these in-
cludes numerous fragments of a skull with superior molar teeth, and a part of the
mandible with the second true molar in a worn condition. The mandibular rami of
other individuals furnish the entire inferior series.

The crowns of the supei'ior molars support two external conical cusps which stand
close together, but are entirely distinct, and have a circular section. There is a
single internal conical cusp flattened on the external side. The entire crown is sur-

T'" /'<"

f a I /iijii

Mioelimuseoryphmis.—Fig. 1. Higlit squamosnl bone, from witliin. Fig. 2. Occipital from fiont. Fig. :!. Left supe-
rior molars, lacking p. m. ii, iii anil iv. Fig. 4. Inferior molars lacking m. i. and p. m. iv. All from the typical
individual except p. m. i, ii, and iii. Figs. 1-2, seven-ninths natural size ; :5-4, about natural size.

rounded by a well-developed cingulum, which is especially jirominent round the exter-
na! anterior cusp of the second and third true molars. Tlie posterior external cusp ol
the last true molar is rudimental, and is situated well within the external liiif on (be
posterior border. The fourth premolar has a single external cusp, and liii- oin.i;iilinn
is wantins: on the anterior and interior sides. The outline of the Ijasc of the crown

324 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

of this tooth is subtriangular ; that of the fii-st and second true molars is a half ellipse ;
while that of the last true molar is a transverse oval, as in the two species mentioned
above. In this last respect it diliers from the species of Mesonyx and Dissacus,
where that tooth has a triangular base. Enamel delicately wrinkled where unworn.
In the M. cmtiquus and M. conidens, the first and second true molars have a triangu-
lar outline, and there is no internal cingulum.

The occiput of this species rises into an elevated transverse crest with an oval
outline, like that of the Dinocerata. This is divided in front by an elevated sagittal
crest. The brain cavity is very small. There is a preglenoid crest.

Measurements of Superior Molars. M.

Length ot true mnlais 0*51

f anteroposterior 010

Diameters of p. m. i v \ „, „

I transverse "'-^

f anteroposterior Oil

Diameters of m. ii -,' „,„

I transverse (Jtu

f anteroposterior 008

Diameters of m. iii -^ „,^

I transverse uio

Elevation of occipital crest 058

The ramus of the mandible has a low and elongate form, and is not very robust.
The inferior outline rises below the coronoid process, and the angular region is nar-
row and compressed. The condyle is produced posteriorly, and its articulai" face is
recurved backwards, and downwards, contracting to an obtuse angle. Its radial line
is directed nearly posteriorly. The posterior border of the ramus is concave imme-
diately below it. The dental foramen is below the middle of the coronoid process.
The mental foramen is single and is below the p. ra. iii. The symphyseal sutui-e is
coarse.

The canine tooth is of moderate size (crown broken oft'). The fourth premolar is
one-rooted. The third and second are two-rooted. They are of equal size and small ;
much smaller than the p. m. i. Their crowns are low, compressed-conic, with a small
anterior cingulum and a short carinate heel. The p. m. i is as long anteroposteriorly
as any of the molars. Its crown consists of a large compressed-conic cusp directed
somewhat obliquely backwards, and a heel supporting two closely-appressed conic
cusps. The internal of these is a little posterior to the external ; the latter is slightly
extended towards the principal cusp. There is a low anterior basal cusp. The ante-
i-ior part of the true molars is more elevated than the posterior. It consists of three
tubei'cles as already described. The heel supports three cusps ; a large external,
which has a crescentic section ; a minute median, and an anteroposterior interior,
which is of medium size, and little elevation. The third true molar dift'ers in having

{anterop
transve

SYNOPSIS OF THE ^■E1:TEB1IATE FAUNA OF THE rUEKCG SEKIES. 325

the median tubercle as large as the external, thus developing a liocl. There is a low-
basal ciuoiiliim on all the true molars on the external side, but none on the inner side.
The external side of" the heel of tlie i). m. i has a low ciniiuhiin. The enamel is
minutely rugose, but is worn nearly smooth in old examples.

Measurements of Ramus. M-

Total length . . .loO

Length of dental series 088

" moiar " 073

" " true molar series 037

p.m.! 0013

anteroposterior 0105

I'erse 008

f anteroposterior 0135

Diameters m. li { .._

[^ transverse OO'J

Depth ramus at m. i 028

m. iii 028

" " from summit of condyle 036

Width of coronoid process at anterior base 015

The mandible from which the above description is taken is that of a diil'erent
individual whose ci'anial characters are given on a previous page.
All the specimens are from the Lower Puerco beds.

MiOCLiESDS PENTACUS sp. nov.

Represented by mandibular rami of seven individuals, which display all the molar
teeth excejDt the fourth, which is represented by its alveolus. The character of these
teeth is a good deal like that of the Protogonia jniercensis, but the absence of superior
molars prevents my learning whether the species is one of the Phenaeodontiche or
not. If it is, it belongs to a genus of that fltmily hitherto undescribed, since the
first premolar is absolutely simple, or without internal cusp. This fact prevents my
reference of the species to the genus Chriacus, to which the details of the structure
of the true molars have much resemblance. My reference of this species to Mio-
clajnus is therefore purely provisional. The veiy robust form of the lii-st premolar
resembles that of the Ilaploconus eiituconus rather than that of any species of Mio-
clajnus.

The inferior molars are robust and Phcnacodus-like, but the lifth cusp is well
developed in all of them, forming an anterior tiiangle. The three cusps of this tri-
angle are only connected at their bases. The heel is wide and supports a large exter-
nal marginal cusp and a curved raised margin behind and within, which is nolehed
at two points so as to produce two nai-row areas on wear. On the third true molar
these are represented by distinct but rather small tubercles. The second true molar

326 SYNOPSIS OF THE VERTEBKATE FAUNA OF THE PUERCO SERIES.

is distinctly lai'ger than the first or the third. The external tubercles of all of the
true molars stand within the base of the crown, so that their external faces are unu-
sually oblique. They have a crescentic section, and the posterior sends its anterior
ridge obliquely to the base of the posterior of the anterior internal cusps, enclos-
ing a basin as in Chriacus. There is a delicate cingulum I'ound the base of the
crown on all except the internal sides. The first premolar is a robust tooth, with a
wide narrow heel, and a small anterior basal tubercle, besides the principal subconic
cusp. A delicate extei-nal basal cingulum. The third premolar has a minute heel,
and no external or anterior cingulum. Fourth premolar one-rooted. A short space
between third and fourth premolars. Enamel minutely rugose, but polished.

The ramus is not deep, but is robust. Its inferior outline rises posteriorly from
below the anterior part of the third true molar. The anterior masseteric ridge is
prominent. The coronoid process rises gradually from the third true molar, without
cari-ying that tooth with it. The symphysis extends posteriorly to the line of the
posterior border of the third premolar. Mental foramina below the second and fourth
premolars.

Measurements. M. •

Total length of inferior molar series 060

Length of true molars 030

f anteroposterior 008

Diameters p. m. i -,'

(^ transverse 0065

^. f anteroposterior 0096

Diameters m. i <

(^ transverse 0083

^. .. f anteroposterior 0105

Diameters in. n <

j^ transverse 0098

... f anteroposterior 0105

Diameters m.m-^

[ transverse 0083

Depth ramus at p. in. i 019

" " " m. iii 031

The seven individuals represented in my collection are all from the Lower
Puerco, and the si)ecies is probably confined in its range to that horizon.

MiOCLiENUS G.\DDIIIANUS Sp. noV.

In this species we have a form which in various respects suggests approach to
Onychodectes. It is represented in my collection by several fragments of the skull
of an individual of medium dimensions in the genus. Both maxillary bones are pre-
served with most of the molar teeth, and a i)art of the mandible with the last
molars.

The first superior pi-emolar has the principal cusp an acute cone with the section
but little compressed. It is joined to an internal cusp whose apex is crescentic.

SYNOPSIS OF THE VEUTEBRATE FAUNA OF THE PUERCO SERIES. 327

Externally it is bounded by a strong cingulum whose anterior and posterior extremi-
ties rise into strons: basal tubercles of erescentic section. The first true molar is
considerably smaller than the second, and has a strong postei-ior inner cingular tuber-
cle, l)ut no internal cino-uluni. The second true molar has the structural characters
of the fiist. Its external cusps are conic, and are little compressed. There is a
strong external cingulum, but no internal cingulum. The third true molar has a very
small anteroposterior diameter, and has a very strong external cingulum which in-
creases the transverse diameter of the croxAm. There are delicate anterior and poste-
rior einornla, which are continuous round the internal base of the crown. In the
lower molars the third is considerably smaller than the second, and its keel is but
little produced. The anterior part of both molars is elevated above the posterior
part.

The ramus mandibuli is characterized by the rapid increase of its depth ante-
rior to the base of the coronoid process, and by the distance of the m. iii anterior to
the base of the coronoid process. The anterior border of the masseteric fossa is not
very strongly marked. The calcancum has the form usual in Creodonta. The sus-
tentaculum and internal facets are well spread apart, and the cuboid facet is triangular
and oblique to the long axis of the hone.

3/easurements. ^f-

Length of bases of posterior six superior molars 015

true molars 021

( anteroposterior 007

Diameters of p. m. i < ._„

i transverse Uw

Diameters of ra. 1- (transverse) 008

( anteroposterior 0075

Diameters of m. ii { ,.,..

) transverse "lOo

( anteroposterior 005

Diameters of m. iii { ,,._

^ transverse "UU

Length of inferior m. iii 0085

anteroposterior 009

,.„j:

Diametersinferiorro.il s „._

^ transverse OOu

Depth of ramus at base of coronoid 020

" front of m.ii 025

Length of calcancum 025

" " free calcar 014

This species has molar teeth about the size of those of the M. protogonioides.
In that species also, the second superior true molar is larger than the fii'st, but the
third is relatively larger than in the M. (/mcflricmtcs, having two well-developed
external cusps. The ramus mandibuli of the M. protogoiiioides does not increase in
depth anteriorly, and the inferior m. iii is larger. There is nn fifth cusp on the molars.

A. p. S. — VOL. XVT. L'P.

328 STNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

This sijecies is dedicated to my distiiiguislied friend Prof. Albert Gaudry, Pro-
fessor of Palaeontology in the Jardm des Plautes, Paris. The type specimen comes
from the Lower Puerco.

MiocL^NUS INTERRUPTUS Cope. DelUitherium inlerruptum Cope, Proc. Arner. PUilos. Soc, 1883, p. 4G3 ; Tertiary
Vertebrata, 1885, p. 283, PI. XXIII d, fig. 13.

The type and only specimen of this species is imi^erfect, but presents the char-
acter which is shared by no other in this genus, of interspaces between the second,
third, and fourth inferior jDremolars. The fourth premolar is not present in the speci-
men, whence my former reference of the species to the genus Deltatherium, but the
appearance of the surface of the ramus leads me to suspect that the tooth has been
lost by accident. The species is further peculiar in the very small anteroposterior
diameter of the first premolar, in which it differs from the other species where this
tooth is known. The fifth cusp is well-developed on the m. i, but the other inferior
molars, with the superior dentition, are unknown.

MlOCL^NUS LTDKKKERIANDS Sp. nOV.

This species is characterized by the presence on the inferior molar teeth of a
distinct curved ledge in front of the two principal anterior cusps, which terminates in
a more or less distinct fifth cusp at its internal extremity. The ledge with its curved
anterior edge is unusual in this genus, though more oi- less developed in some of them,
especially the M. opisthacus. The present species is much larger than the latter, and
exceeds the M. suMrigonus also, but is smaller than the M. jjrotogomoides.

The heel is rather wide and the edges elevated and with only an external cusp-
like elevation, giving a basin-like surface. The last molar is not enlarged or reduced,
but is narrowei- than the m. ii. The crowns are without cingula, and the enamel is
obsoletely plicate. No premolars preserved.

Measurements. M.

Length of true molars OlS

" " la.st true molar 007

-,. .. f anteroposterior OdCi

Diameters m. n -^

I transverse 00()

_ . f anteroposterior OO.'io

Diameters m. \ \

[^ transverse OO.j.i

Deplli of ramus at m. i 014

Poi'tions of three mandibular rami represent this species. It is dedicated to
Dr. Kichai-d Lydekker, the distinguished curator of vertebrate palajontology in the
British Museum.

SYNOPSIS Ob' THE VERTKBRATE FAUNA OF THE PUKRCO SERIES. 329

MlOCI,.«NUS KILIIOI.IANLS Sp. nOV.

Parts of both mandibular rami of two iiulividuals icprcsont this siiecies in
my collection. One of these presents the five posterior molars in perfect preservation.
The characteis ally the species to some of those of Chriaeus, but the simple first
inferior premolar distinguishes it from that genus. After the 3f. minimus and the
M acolytus this is the smallest species of Mioclasnus, and its inferior molars have the
cusps more acute and elevated than in any other.

The molars are distinguished from those of several othei" species, including Chri-
aeus schlosseriamts, in that they increase in size, regularly posteriorly. The first is
thus much smaller than the third, a pi-oportion which is reversed in many other spe-
cies. The two anterior cusps are joined much above the level of the heel, which
would be thus much below them, but for the fact that its border is much elevated,
leaving the median surface a deep basin. The fifth cusp is small and is elevated and
close to the foui'th, and is connected with the anterior extei'nal by a cui-ved crest.
The edge of the heel is developed into three small cusps, one median and two
lateral. The tliiid true molar is quite elongate, and has six well-developed cusps in
all. TIk' Hist premolar has a short base, and the principal cusp is truncate behind,
and has a shar[) edge in front. The heel is very short and terminates in a small acute
cusp, and there is no anterior basal cusp. The second premolar has a small cuspiform
heel. There is a weak cingulum on the first premolar and first true molar, but only
traces remain on the m. i and ii. Enamel smooth.

Measiurements. M.

Length of true molars 017

„. , ( iinteropostcrior 0045

Diiimeters of m. i

^ transverse 00:!

^. „ ( auteroposlerior 0052

Diameters of m. ii <

I transverse 0035

^. , ... (anteroposterior 006

Diameter of m. iii <

( transverse 0035

Depth of ramus at front of m. i 012

The two specimens of this species were found in the lowest beds of the Puerco
formation by Mr. David Baldwin. It is dedicated to Dr. Henri Filliol, the distin-
guished French paUoontologist, whose discoveries in the field of extinct ^lammalia
have been so numerous and important.

MiocL^NUs PROTOoosioiDES Cope ; Amcr. Naturalist, 1882, p. 833; Tertiary Vertebrata, 1885, p. 31(), I'l. XXV f,
fig. 17.

This species has been known hitherto by the last two supeiioi- molars only.
At present I have in addition, a fragment of a maxillary bone with the first and sec-

330 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

ond true molars, and a set of jaws of a single broken skull, and also a left mandibular
ramus containing all but the canine and fourth premolar teeth. The species is dis-
tinguished in the ui)per series by the relativel}^ considerable development of the
second and third superior molars, the third having two well-developed external cusps.
Also by the strong development of the intermediate tubercles, and the distinctness
of the posterior inner tubercle of the first and second true molars. Though distinct,
it is only a small acute elevation of the jDosterior cingulum. The latter passes
round the internal base of the second and third ti'ue molars, but not the iirst.

The ramus of the lower jaw is of moderate and uniform depth throughout, and
not convex below, as in M. gaudrianus. The second and third true molars have no
trace of the fifth cusp ; the anterior part of the fii'st is lost. There is a weak median
posterior tubercle on the first and second true molars, which is developed into a very
large obtuse heel in the third, so that this tooth is longer than either of the other true
molars, and reaches to the base of the coronoid process. The first premolar is not
relatively so large as it is in some of the species above described. Its principal cusp
is but little compressed, and the heel is short and transverse, and has the inner ex-
tremity almost cuspidate. A small acute anterior basal cusp. The second premolar
has anterior and posterior basal cingula oi\\y. The third is a little smaller than the
second. The only cingulum of the inferior molars is at the outer base of the anterior
half of the crown. This half is a little higher than the posterior half

Measurements of Mamus. M.

Length of true molar series 0245

f anteroposterior 0075

Diameters m. ii \

I transverse 0065

^. f anteroposterior 0095

Diameters m. m-^

i transverse 000

Depth of ramus at end of m. iii 019

" " " " " m. i OIG

MiOCLiENUS FLOVERIANUS Sp. nov.

This species is indicated by parts of two mandibular rami which display four of
the molar teeth, with which were found the following bones of the skeleton : several
vertebi'aj, parts of both humeri, distal end of radius, greater part of right ilium.
These all appear to be parts of the same skeleton.

The first true molar has a well-developed fifth cusp which is median in position
and larger and lower than the fouith cusp, from which it is well separated in position.
There is no trace of fifth cusp on the other true molars, but there is a median poste-
rior of small size on the m. ii. The last inferior molar is not reduced as in some spe-

SYNOPSIS OF THE VKKTEBKATE FAUNA OE THE PUEECO SERIES. 331

cies, but is smaller than the penultimate, and has but a short heel. The first pre-
molar is about the length of the first true molar. It is robust, and the cusp is a little
compressed, but is flattened behind. The heel is very short and is convex posteriorly.
A very small antciior basal cusp. The anterior part of the crown of the m. i
is higher than the posterior, which only supports one tubercle, the external. The in-
equality between the anterior and posterior tubercles of the third true molar is
unnoticeable. Ko cingula on molars or premolars.

There are two lumbar and one caudal vertebra}. The centra ai'c depressed by
pressure ; arches lost. The caudal vertebra is one of the proximals. It is robust, and
indicates a well-developed tail. The head of the humerus is extended in the direc-
tion of the greater tuberosity. The latter is large and truncate both proximally and
externally ; at the posterior external angle is a distinct fossa m. teretis. The bici-
pital groove is wide. The lesser tuberosity is bounded by an angle, which rises prox-
imad and towards the greater tuberosity. The deltoid ridge extends to below the
middle of the shaft. The internal condyle is large and compressed so as to be verti-
cal ; it is pierced vertico-obliquely by the large entepicondyUu- canal. The external
epicondyle is very little jjrominent, and is mostly occupied by its fossa. The trans-
verse extent of the condyles is not very great. The lateral borders of its posterior
groove are sharp. The convexity of the roller is pronounced but not approaching an
intertrochlear crest. The distal extremity of the radius is wider than deep and the
outer side is truncate, so that the section of the shaft is a triangle. At the extremity
the ulnar side is grooved, and above the prominent superior border of this groove is
another, which is bounded above by a short robust Hat ridge. The outlines of the
scaphoid and lunar fossae are distinct except where they pass into each other.

The ilium is robust, and at the peduncle the anterior or inferior face is nearly as
wide as the interior. At the sacral articulation it is moderately expanded. The an-
terior inferior s[)ine or tuberosity is moderately prominent, and is situateil a consid-
erable distance proximad of the acetabulum, opposite the narrowest part of the
peduncle.

Measurement'. M-

Length m. i 009

Width " posteriorly OOO.'i

{antero])oslerior 000
AA-r
transverse 0()i.>

f iinteropostcrior 000

Diameters m. iii -!

h

riinsverse .

.007

f anteroposterior 009

i^fiamciers p. m. i { „nr

I transverse ""■'

Length of centrum of lambar vertebra 022

332 SYNOPSIS or the vertebrate fauna of the puerco series.

Measurements. M.

[ long, with great tuberosity 021

Diameters of Iiead of humerus i , , ,, „ ., ^r,

(^ short, " small " fli

Least diameter of sliaft do OOo

Width of distal end do 035

" " condyles in front 015

f transverse 0105

Diameters distal end of radius J. . .„„

I vertical ouy

Ilium anterior widtli of peduncle 010

" external " " " Ol'l

This species is one of medium size in the j^enus. It is quite of the type of the
M.ferox and M. corrugatus. Its inferior molar teeth are considerably larger than
those of the M. prolog onioides, and the last one is shorter in relation to its length.
It is dedicated to Professor W. H, Flower, Director of the Natural History Depart-
ment of the British Museum.

MiocL^NOS CORUUGATUS Cope ; Proc. Amer. Phil. Soc, 1883, p. 560; Tertiary Mammalia, 1885, p. 341, PL XXIV f,
fig. 5 ; XXIV g, fig. 8.

The discovery of nearly entire mandibular rami of this species and of M.ferox
enables me to determine the distinctive characters of the two species better than
heretofore. While the true molar teeth in the two are of nearly equal size, or at least
within the range of variation not unusual in a variable species, the mandibular ramus
of the M. corrugatus is very much smaller every way than that of the M. ferox.
The shortening influences the length of the premolar series, which is much shorter
in the M. corrugatus, the teeth being crowded, while in the M.ferox they are (the sec-
ond, third and fourth) separated by interspaces. An astragalus which accompanies
the jaws of 3£ corrugatus is identical in character with that of the M. ferox but of
smaller size.

MiocLJENUs OPisTiiAcns Cope ; Amer. Naturalist, 1883, p. 833. Hemiihlmus opisihaeus Cope ; Tertiary Vertebrata,
p. 407, PI. XXV f, figs. 8-9.

The typical specimens of this species include parts of four mandibular rami with
teeth, and a single superior molar, which is that of a species of Hemithljeus. The
association of this superior molar with the mandibles cannot be demonstrated. I
have subsequently received a mandibular ramus with nearly complete dental series,
associated with a portion of a skull which contains the true molars and the first pre-
molar in a perfect state of [)reservation. Everything inspires the belief that these
are parts of the same animal. In addition I have parts of both rami of a sixth indi-
vidual which support most of the teeth ; and a maxillary bone with m. ii, m. i and p.
m. i, with parts of mandibles of a seventh, and mandibles of five other individuals.

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUEKCO SERIES.

333

This species exhibits characters intermediate between the types witli robust pre-
molars, and those with pi-emolars of narrower and more angular form. The second
superior pi'cmolar has no internal cus|) and is trenchant, but the first has a very large
internal cusp, whose base is as large as that of the external one, and which has slight
anterior and posterioi- cingula, but no internal or external ones. It resembles the cor-
responding tooth of some of the PcriptychidiB. The true molars arc those of this
genus, resembling those of the M. auhtrigonus. I'he external cusps arc distinct and
not compressed. The crown is surrounded by a cingulum except on the internal side.
The cingulum rises into a small cusp between the external cusps, and into a small
but distinct posterior internal cusp. In the inferior series the first and second pre-
molars ai'e enlarged and have longitudinally oval bases ; both have posterior heels,

.If II'

ai Jit-

J al III

' J ar Jir

Fig. 5. Fig. G.

Fig. 5. JUiocUenua opisthacus molars of both series | natural size : the cusps are lettered ; ae, anterior external ; pe,
posterior external ; ain, anteri(jr intermediate ; j>in., posterior intermediate ; ai, anterior internal ; pi, posterior in-
ternal ; 5, fifth cusp.

Fig. fi. Chriacui priseut | natural size; teelh of both jaws.

and the first has a small anterior one ; both are like those of Anisonchus. The in-
ferior true molars illustrate the transitional condition of the fifth cusp in this genus.
A narrow crest descends inwards from the anterior external cusp, but terminates
below the level of the fourth, or ojjposite inner cusp, and there is no liltli cusp, or but
a rudiment in the m. iii of one of the specimens.

Measurements of Fifth Specimen. M.

Length of superior true molars 014

( anteroposterior OOf.'i

Diameters of superior p. m. i <

^transverse (i07

( anteroposterior 00(i

Diaractera of superior p.m. ii (spec. No. 7) <

^ transverse 001

Length of inferior molars (except p. m. i v) 0;i;c>

true molars 01.5.'i

p. m. i 0O53

Width of inferior p. m. i 00!{

Length of inferior m. iii OOG

Width " " " 003

Depth of ramus at m. i 010

334 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

MiOCLiENUS ZITTELIANDS Sp. nOV.

This species is represented by a skull which lacks the occipital region. It is
enclosed in a silico-calcareous concretion, and the teeth have been cleanly exposed by
weathering, but the surface of the skull is not cleaned. Large parts of both mandi-
bular rami with their teeth are preserved.

The dental characters resemble those of the M. turgidus, but differ in the absence
of intei-nal cusps of the superior premolars. The dimensions are smaller, especially
those of the premolar teeth. The last molar in both jaws is much smaller than the
others, as in M. kmjidus, and tiie intermediate tubercles in the superior molars are
present, but small. The external cingulum is distinct on the superioi- true molars,
but wanting on the premolars. The same is true of the anterior and posterior cin-
gula, while internal cingula are absent from all the teeth. The heels are rudimental
on the first and second inferior premolars.

Measurements. M.

Length of superior molar series 036

" " " true molars 014

( anteroposterior 00o5

Diameters superior p. m. i < „„„

( transverse UU i

C anteroposterior 0U55

Diameters superior ni. ii < „._„

I transverse 00,3

Width between p. m. ii on palate 0075

" " m. ii on palate 015

From the Upper Puerco. Dedicated to Professor Karl Zitlel, of the University
of Munich, and Director of the Museum.

MlOCL^NUS TUnniDDNCULUS sp. nov.

This species is primarily indicated by a portion of a left maxillary bone which
supports the first two true molars, and the first premolar. There is a mandibular
ramus supporting the corresponding teeth of the inferior series, which probably be-
longs to the same species, and the posterior parts of two other rami, each supporting
the last two molars, are also referred to it.

The peculiarity of the superior molars consists in the strong external cingulum
and intermediate tubercles, and the large size of the first premolar, and especially of
its internal cusp, in which respects it resembles that of the MloclcBnus ojnstJiacus. It
has no cingula whatever, except a trace at the anterior and posterior external angles.
There are anterior and posterior, but no internal cingula of the true molars. In the
M. minimus the molars have no cingula, except faint traces of the anterior and pos-
terior, and the intermediate tubercles are wanting.

SYNOPSIS OF TIIK VERTEBRATE FAUNA OF THE PUERCO SERIES. 335

The inferior molars are referred to this species because of the large size of the
premolars, which approach in dimensions those of the M. zittelianus. The first has a
low wide heel, and no anterior basal angle. The fifth cusp exists as a cusp on the m.
i and ii. The curved border of the heel in these teeth is more elevated than in the
M. minimus on the inner side, so as to enclose a dee[) basin, which is much better
defined than in M. minimus. In the third and fourth individuals, where the last inferior
molar is preserved, this tooth is smaller than the m. ii, and both of these teeth have
the fifth cusp present, but small, and appressed as a small twin on the anterior face of
the fourth. This occurs also in the M. acolytus, and I do not believe that the char-
acter is constant.

Measurements. -'/.

LengUi of superior molars (last estimated) Oil

( anteroposterior ." 005

Diameters m. ii < „r,rr

I transverse W)jo

anteroposterior 004

Diameters m. i < [.^..^

transverse W4o

(a.

Diameters p. ni. i ■

I antoropostt-rior 0045

' transverse 000

Length inferior true molars (sp. 2) 0125

" p.ni.i 005

Depth ramus at m. ii 0075

The typical specimen is from the lowest beds of the Piierco. D. Baldwin.

MiocL.KNUS ACOi-YTCs Copc. Ili/opsodus acoli/lus Cope ; Proc. Amer. Pliil. Soc, 1883, p. 4(13 ; Tertiary Vcrtebrala,
1885, p. 238, PI. XXIII d, figs. 5-fl.

Additional specimens of this species show that it differs fi'om the M. minimus in
the presence of a strong external cinguliiin of the superior true molars, and the less
reduced size of the last inferior molar. A single specimen, figured as above, displays
the first two true molai-s and the fii-st premolar above, and both the last inferior molars
of the same skull. From the M. turr/idunculus this species is easily distinguished by
the small first and second premolars in both jaws. A damaged skull, without lower
jaw, shows this charactei-, as well as the type, and also that the fourth superior pre-
molar has but one root, and is of small size; also that the canine is well developed
and has a vertical direction. The second superior premolar has an internal cingulinn,
but no cusp.

Measurements of Skull. M.

Length of superior dental series 0353

true molars 009

Width between superior canines Oil

Five specimens ; all lioni the Upper Puerco except one, and the specific refer-
ence of this one is uncertain.
A. p. s. — VOL. xvr. 2q.

336

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

The tritubercular superior molai's and simple premolars show that this species
cannot be referred to the genus Hyopsodus.

The number of the superior premolars of the allied M. mmimus is unknown, so
that it may be a Tricentes. In this genus it can be only compared with the T. in-
cequidens (Fig. 8), which is of about the same size. But in that species the last
molai- is still more reduced than in M. minimus.

Pig. 7. ~ "^

Fig 8.
Fig. 7. Tricentes hucculcntus f natural size. Fig. 8. Tricentes inmquideiis f natural size.

CHRIACUS Cope.
Proc. Anier. Phil. Soc, 1883, p. 314 ; Tertiary Vertobrata, 1885, p. 740.

In the absence of sufficient knowledge of the skeleton in this genus I (ind its
distinction from Mioela>nus to rest on dental characters. It differs from that genus in
the presence of an internal cusp of the first inferior premolar, which is however but lit-
tle developed in some of the species. The molars of the inferior series may be usually
easily distinguished from those of Mioclfenus in the well-developed fifth cusp, and
in the trihedral cusp of the external side of the lieel, which with the raised internal
border gives the heel the basin-like character which is seen in Pel3'codus. The char-
acter of the inferior molars graduates into that of Miockenus through such species as
M. suhtrigonus and C. scldosseriamis.

The species are difficult to distinguish with tlie material in my possession. I
recognize ten from the Puerco and one from the Wasatch. The former are distin-
guished as follows. Owing to the absence of corresponding parts I am obliged to
compare the superior and inferior dental series in separate tables :

Species with Sxtperior Molars.
I. Posterior cingulum and its interior cusp large, forming a prominent single of the crown.

Larger ; true molars .0185 C. pclvidcns.

Snniller ; true molars .014 (Fig. 9) C. iruncatus.

n. Posterior cingulum and cusp small or insignificant.
«. Superior molars wider than long.

Largest ; molars .01SJ3 ; depth of ramus at m. i .013 C. jrrisctis.

Medium ; molars .015 ; depth oi ramus .011 G. schloaserianus.

Smallest ; molars .010 ; depth of ramus .007 C. simplex.

aa. Superior molars longer than wide.
Medium ; molars .0165 0. ki/ntlianiis.

SYNOPSIS OF THE VEUTKBKATK FAUNA OF TUK TUERCO SERIES. 337

Species with Inferior Molars only.

I. Size large ; premoUrs unknown.

Inferior true molars .0285 ; depth ramus at m. i .015 C. riielimei/eranus.

II. Size smaller ; premolars not spaced.

Inferior true molars .023, last large ; ramus .018 C. pelcidens.

Inferior true molars .017, the last small, the first not reduced ; ramus .011 C schlosserianus.

III. Size smaller ; spaces between p. ms. ii, iii and iv.

True molars .018, fifth cusp very distinct from fourth ; molar series .043 C. baldwini.

True molars .024, fifth cusp divergent, connected with fourth by a crest : molar series .0.50 O. stenops.

IV. Smaller ; third premolar larger than second.

Depth of ramus at p. m. i .010 C. inversus.

_ ^ CD «» \^ -

Fig. 9. '

Fig. 10.
Fig. 9. Chriams truncatus f natural size. Fig. 10. Indrodon malaris J natural size.

CnniACus priscus sp. nov. Fig. fi.

This species is represented primarily b}' a fragmentary skull of wliicli there
remain, the superior walls of the brain case, a part of the left maxillary bone, Avhich
supports the postei-ior five molars ; part of the right maxillary with three molars, and
part of the right mandibular ramus with the anterior two true molars. There are asso-
ciated as cospecific with this individual, a second, represented by superior and inferior
molars ; a third represented by superior molars onljs and two others represented
by mandibular rami only ; in all, five individuals. All are from the lowest beds of
the Puerco.

The superior molars are of the same size as those of the O. pelvidens, but are
very dilferent in form. The absence of the stiong internal angles of the crown at
the inner extremities of the anterior and jjosterior cingiila, is one character. The
straight outline with an open margination of the external side of the crown in C. jx'l-
videns is in strong contrast with the two convexities, each following an external cusp
which form the outline in the C. jfrif^cus. The first premolar has a small internal cusp
in the C. priscus ; a large one in the C. pelvidens. It is represented by an angular
cinguliiin in the C. priscus. In this species the cinguliim does not extend round the
inner base of the crown, except weakly in the last molar.

The inferior molars have a relatively greater transverse diameter than those of
any other s])ecies of the genus. They also differ from those of the (J. pelvidens and
C. stenops., which they resemble in size, in the nearly equal elevation of the anterior
and posterior cusps, and in the absence of an external basal cingiiluni. The lift h

338 STNOPSIS OF THE VERTEBRATE FAUNA OF THE PUEKCO SERIES.

cusp is a small cone, and is not spread away from the fourth and connected with it
by a crest, as in 0. steno'ps. In specimens with the last inferior molar preserved, that
tooth is seen to be of average proportions.

The brain-case is long, narrow and rather low, and the sagittal crest is low and
thin. It does not in the least resemble that of Adapis, but is rather that of a Creo-
dont. Post orbital region lost.

3Ieasurements. M.

Length of five superior molars 031

anteroposterior 0055

Diameters p. m. i , „„__

transverse 0055

f anteroposterior 007

Diameters m. ii^^ „„„_

(^ transverse 0085

( anteroposterior 0055

Diameters m. iii -

I transverse 0875

f anteroposterior 007

Diameters inferior m. ii < „„„

I transverse 00G5

f anteroposterior 006

Diameters m. i \

I transverse 006

Depth of ramus at m. i 013

Cheiacds sculosserianus sp. nov.

Parts of four individuals represent this species. Three of these are from the
Upper Puerco ; the exact horizon of the fourth is probably the same, as those from
the Lower Puerco are especially marked in my collection. The typical specimen in-
cludes jjarts of both maxillaries with molars ; both mandibular rami, one nearly per-
fect and sujjporting the last foyr molars ; parts of both humeri, an ulna and part of
the right astragalus. The species is smaller than the C. i^viscus, and has about the
dimensions of the MioclcBuns suhtrigonus.

The posterior internal cingular cusp of the second superior molar is better devel-
oped than in the Cjiviscus, but nothing like that seen in the C. pelvidens and C trun-
catus. The cingulum continues round the internal side of the crown in the second
and third molars (first injured). External cingulum present. Last molar of rela-
tively small size, about as in Mioclcenus turgidus, but larger than in Tricentes ince-
quidens. Internal cusp of p. m. i small. The inferior series is distinguished by the
small size of the third. The anterior part of the crown of the inferior molars has
a distinctive character. The fifth cusp is not distinctly defined, but the anterior part
of the crown forms a three-side prism, the anterior edge, oi- that connecting the ex-
ternal anterior cusp with the lifth, being vei-y well defined. On wearing, the summit
forms a narrow V with the apex external. External cingulum complete on the
second true molar only. The first premolar is of moderate size, and has an anterior

SYNOPSIS OF THE VERTEBRATK FAUNA OF THE PUEKCO SElilES. 339

cusp of an elevation equal to the internal cusp. There are no interspaces between
the first, second and third premolars, and probably not behind the Iburlh, as the jaw-
is shallow ami tapers rapidly. It is, however, deeper than in the C. simplex. Mental
foramina below the second and fourth premolars.

There is nothing noteworthy in the humerus. The astragalus has but a short
anteroposterior trochlear surface, and it extends well on the neck, indicating a planti-
grade foot. It is slightly concave transversely. The malleolar face is oblique, and
projects posteriorly beyond the fibular face, which is vertical. The base of the pos-
terior notch bridges the minute median foramen. The ulna is much compressed, and
has a large olecranon. The humeral cotylus is diagonal to the long diameter of the
shaft, and has a marginal llange behind on the interior side, and in front on the
exterior side. The radial face is absolutely flat and is directed inwards at an angle
of thirty degrees from the vertical. It indicates a flat head of the radius, without
rotar}" capacity. Below the superior border of the ulna, on the external side, is a deep
gi-oove which extends to the edge of the flange of the humeral cotylus.

Meaturements. M.

Length of superior true molars 015

{anteroposterior 0062
transverse 0035

( anteroposterior 000

Diameters of m. ii {

[^ transverse 008

! anteroposterior OO.M
transverse 004

Length of inferior molars less lu. i 02!)

" " " true molars 017

molar iii , OO.W

f anteroposterior (H)T)T)

Diameters m. ii {

(^ transverse 004.')

Depth of ramus at m. i 010

Width of distal condyle of humerus in front Oil

{anteroposterior OOG
transverse 007

f at olecr.inon 009

Depth of ulna i at cotylus .0005

I at coronoid 00!)

Width of astragalus 008

Width of trochlea 005

Height of trochlea, external side 0045

This species is dedicated to my friend Dr. Max Schlosser, of Munich, whose
work.s on the extinct Mammalia are among the most important of modern times.

340 SYKOPSIS OF THE VEUTEBKATE FAUKA OF THE PUERCO SERIES.

Chriacus ruetimeteranus sp. nov.

This is the largest species i-efeired to the genus, and is the only one as yet I'epre-
sented by a single individual. I possess part of the left mandibular ramus which
exhibits the true molars, and the roots of the first and second premolars.

The last molar is as large as the second, and its anterior cusps are not opposite
to each other as in other species of Chriacus, but the external is in advance of the
internal. The fifth cusp is not elevated as in other species, but is represented by an
internal angle of an interior ledge, which is quite wide. The anterior cusps of the
m. i are opposite, those of m. ii are injured. The heel has the external cusj) of cres-
centic section, and the internal elevated border. ISTo cingula. Enamel obsoletely
coarsely rugose. The heel of the m. iii is well developed, as are also the adjacent
internal and external marginal cusps.

Measurements. M.

f anteroposterior 009

Diameters m. i \ .„„

(^ transverse OUo

f anteroposterior 010

Diameters m. iii - ..._

I transverse UOoo

Depth of ramus at posterior end of m. i OKi

" m. iii 031

From the lowest Puerco beds. Dedicated to Prof. L. Euetimeyer, the distin-
guished palaeontologist of Basle.

CniiiACUS BALDwiNi Cope. Deltatherium haldwini Cope; Proc. Amer. Phil. See, 1883, p. 463; Tertiary Vertebrata,
1885, p. 283, PI. XXIII d, fig. 13.

Besides the type of this species, I have obtained since its description, the greater
|)art of the mandibulai' ramus with teeth with several parts of the skeleton, of one
individual, and both mandibular rami with most of the teeth of a third individual.
Parts of the rami of a fourth individual may belong to the species.

The complete rami show that their anterior portions are produced, and that the
premolar teeth in front of the second are spaced. The second and third are close
together. The true molars themselves are a good deal like those of the M. snhtri-
gonus with fifth cus}). llie internal cusp of the first premolar is rudimental in this
species. 'I''he premolars of the M. suUvigoii.us are not spaced. The fifth cusp is small,
distinct, and not involved in a crest. There is a small median cusp of the heel.
The first premolar has a distinct heel, llie euaniel of the molars is coarsely rugose,
a surface which long use does not entirely smooth. !No cingula in one specimen, but
in the ty])e and second specimen there is a rather weak one on the external side.
The mandibular condyle is entirely above the level of the molars in sp. No. 3. The
last inferior molar is not of reduced size. Kamus slender.

SYNOPSIS OP THE VEKTEBRATE FAUNA OP THE PUERCO SERIES. 341

The femur of the second specimen is nearly perfect. Tlie trochanters are well
developed ; the great extending proximad to the head, and connected by a curved
ridge with the less. Third trochanter superior in position, its superior i)art overlap-
ping the line of the inflMior part of the lesser. Head without fossa or notch lifia-
menti ten's. liotular face elevated, wide, with equal l^oiders and sliirht concavity.
Condyles well separated.

Measurements of JVo. S. M.

Length of inferior molar series 043

•• " " true molars 0205

" " " " m. iii '. 007

Depth of ramus at m. i 015

p. m. iv 009

Length of femur (greatest) 098

Width at head 022

" " middle of shaft (crushed) Oil

" " condyles 017

CHRIACCS 8TENOP8 Sp. nOV.

Four mandibular rami of three individuals represent this species. Two of the
laiiii arc entire to the base of the canine tooth. These show interspaces between the
second and third and third and fourth premolars, not seen in such s]iecies as <'. pelvi-
dens and C. ,scJilof;seri(iiiUf!. It resembles in this respect the (J. haldw/'iti, but diU'crs
from it in characters already mentioned, and in its superior size. The internal cusp
of the first inferior premohir is also much better developed.

The antei'ior cusps of the true molars ai-e more elevated than the posterior, but
the diirerence is not so great as in some other species. The fifth is a prominent
angle, nearly in line with the fourth, and connected with it by a crest. 'I'he iieel has
an external angular cusp with crescentic section, well distinguished from an internal
curved border which is connected with a small median posterior angle. This be-
comes a prominent cusp in the third molar, which is of average size. Molars with an
external eingulum. Enamel coarsely and obsoletely rugose in unused specimens. Tiic
first premolar has a well-developed ti'ansverse heel, and anterioi' l)asal cusp. The
second has the same, but of i-educed proportions.

In a specimen (No. 3) cleaned by weathering, the posterior border of the sym-
physis is opposite the posterior border of the p. m. iii ; the two mental foramina are
below p. m. ii, and the space between p. m. iii and iv.

Measurements No. 1. M.

Length of dental series including canine 055

" molar series Oil)

" " true molar series 024

342 SYNOPSIS OP THE VERTETJRATB FAUNA OF THE rUERCO SERIES.

Measurements No. 1. ^•

f anteroposterior """*'

Diameters m. i \ nn'iT

I transverse uu.j.j

f anteroposterior 003

Diameters m. iii - (\(\'-.k

I transverse uu.j.)

No. 3.

Length of premolar scries 02G

Depth of ramus at m. i ^^^

' p. m. iv 008

All the specimens of this species are from the Upper Puerco.

ClIIUACUS INVERSUS sp. nov.

The characters of this species are so well marked that I introduce it here,
although the material representing it is slight, a portion of a single mandibular ramus
which supports three premolars, and one true molar. The posterior part of a ramus
accompanies this specimen and may belong to the same species. But of this there is
doubt.

The first premolar has a well-developed interior cusp, and a large heel. The lat-
ter has an internal vertical, and an external oblique side, terminating in a cutting
edge, the internal curving round the posterior border to meet the external. There is
no anterior basal cusp, nor any cingulum. The second and third premolars are com-
pi-essed, and have cutting edges before and behind. The second has the heel slightly
transverse, and a mere trace of anterior basal lobe. Tbe third is larger than the sec-
ond and is more compressed. Externally its face is regularly convex, but internally
its convexity is a vertical median rib, and in front of this the face is concave, thus
maintaining the acuteness of the anterior cutting edge on wear. The heel is small
and comjiressed ; anterior lobe none. The enamel of all the teeth is smooth. No
cingula.

3Icnsurcmenls. M.

Length of posterior tlueo jtrcmolars 015

" " p.m. iv 007

Width " •■ " 0038

Depth of ramus at front of m. i 0115

TRiiSODON Cope.

Amer. Naturalist, 1881, p. 007 ; Tertiary Vertchrata, 1885, p. 270.

But one species of this genus is described, although several have been referred
to it. All but the type species have found a congenial location in the genus Mio-
cla3nus (group Goniacodon). I am now able, however, to add a second species of
ty])ical form.

SYNOPSIS OF THE VERTEBHATE FAUNA OF THE PUERCO SERIES. 3-43

TriSsodon biculminatcs sp. nov.

Parts of the mandibular laini of two individuals indicate this species. One of
these supports the first and second true mohirs, and the other the lirst true molar
only. The character ol" the genus is seen in the great development of the external
cusp of the heel, and the absence of an internal cusp, so that the heel appears to have
one cusp with an oblicpie cutting edge, resembling in this respect Paheonyctis. The
two principal anterior cusps are opposite each other and united for half their eleva-
tion, and the internal is lai-ger relatively than in the T. quivirensis. The lif'th cusp
is low, forming a distinct tubercle in the first molar only, and represented by the in-
ternal anirle of an anterior led<>:e in the second. The heels of both the molars have three
small tubercles on the posterointernal border, which are not so much elevated as the
corresponding ones in the T. quivirensis. A weak external cingulum only. Enamel
obsoletely coarsely rugose with minute tubercles.

Measurements. M.

{anteroposterior Oil
transverse uuo

f anteroposterior 013

Diameters m. ii \ nnnr

( transverse ouuo

Depth of ramus iit m. i 020

From the bottom of the Puerco beds.

DISSACUS Cope.
Amer. Naturalist, 1881, p. 1019 ; Tertiary Vertebrata, 1885, p. 344, 741.

Numerous pieces of the skeleton associated with molar teeth which agree with
those of the D. navqjovius, throw much light on its characters, and on the position of
the genus Dissacus. In the first place the zygapo])hyseal articulations of the lumbar
vertebrte are of the involute type common to all the Creodonta. This is also determinable
lioin the specimen described by uie in the Tertiary Vertebrata, p. 741. In the new
skeleton the fragments of metapodials, i)i-obably metatarsals, furnish evidence of the
existence of five digits, which distinguishes the genus from Mesonyx, where there are
but four. The astragalus is especially interesting in its relation to that of Mesonyx.
It has, as in that genus, a distinct cuboid facet, separated from that of the navicular
by an angle, but the angle is not as well defined as in >[esonyx and Pachyrena.
The trochlea is very .slightly concave, thus differing widely from that of the Meso-
nyehidie, and resembling that of other Creodonta. This bone therefore furnishes a
clear guide to the phylogeny of the Mesonychida). The two facets are well defined
on the proximal extremity of the cuboid. Its distal extremity supports but one, a

A. I*. S. — VOL. XVI. 2k.

344 SYNOPSIS OF THE VERTBBKATE FAUNA OF THE PUERCO SERIES.

coucave facet. The tubercalcis, like the olecranon, is elongate. The coi-acoid is dis-
tinct but short. The peduncle of the ilium is wide. The odontoid process is rather
long and cylindric. There is an entepicondylar foramen, as in Pachysena.

DisSACOs NAVA.iovius Cope, loc. cit.; Tertiary Vertebrata, PI. XXV c, fig. 1.

The paradiapophysis of the atlas has the base extended throughout the vertebra
anteroposteriorly, and the vertebrarterial canal divides it anteroposteriorly, issuing
at about the middle of the superior surface. The atlantal articular surfaces of the
axis are distinct from those of the odontoid. The floor of the neural canal has a
lovr median convex ridge, vv^hich extends to near the end of the superior side of the
odontoid process, from which it is separated, as well as from the articular faces below,
by a shallow groove.

The glenoid cavity of the scapula is an oval, gradually acuminating to the
narrow origin of the coracoid. The spine oi'iginates abru])tly a little posterior to the
middle line. In the humerus the entepicondyle is of moderate dimensions, much
larger than in Mesonyx, but less prominent than in many other Creodonta. Outside
the roller of the condyle there is a distinct border facet, rising to its external border,
which is present in Mesonyx, but is wanting in Oxygena. The roUei' is distinctly
convex. The internal flange is well developed. In the ulna, the olecranon is long,
straight and compressed. Its inferior border is of uniform width from below the
glenoid surface to the extremity, and is but little in excess of that of the shaft fur-
ther forwards. The posterior flange for the humerus is not present on the external
side of the glenoid surface. The i-adial facet is not very oblique, and it is followed
distally on the external side by a deep groove, which runs out on the external side of
the shaft. The latter is marked by a deep and wide longitudinal gutter on its inter-
nal side, which is bounded by a narrow edge above, and a wide border below. It
runs out on the inner side of the shaft pi-oximally, opposite the middle of the external
groove. This gutter is wider and deej^er in this species than in any creodont known
to me, excepting the Pachycena ossifraga. The head of the radius is a depressed oval,
with three articular surfaces ; a median concave, and a narrower reflected bevel at
each end, fitting corresponding faces of the humerus. The inferior ligamentous fossa
is wide and shallow.

The femur is not preserved excepting the head, which shows a rather shallow fossa
ligamenti teris, which is well separated from the border. The astragalar face of the
tibia is quite oblique, especially next to the fibular suture. The internal malleolus
is large, vertical, truncate and furnished Avith a tuberosity in front of the distal
extremity. The articular surface is folded back over the anterior edge for a short

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES. 345

distance on the anterior face, to correspond witli the fossa of the neck of the astra-
gahis. The astragahis is somewhat depressed, and is quite oblique to the vertical
phme. The neck is of moderate length, and is depressed, and the navicular facet
slopes at an angle of forty-five degrees when the astragalus rests bn a plane surface.
The long axis of the trochlea slopes at an angle of thirty degi-ees to the long axis
of the entire bone, outwards and forwards. The posterioi- foi'amen is completely
enclosed. The malleolar face is oblique, the fibular vertical. The articular surface
of the trochlea is continued in an excavation of the neck to its middle, and is ter-
minated by an acute recurved boundary. This indicates a foot frequently held at
right angles to the leg, and a plantigrade walk. The obliquity of the astragalus,
which fits the tibia perfectly, means that the hind legs diverged to the leet in a very
obvious manner. If the calcaneum is placed on a plane surface this divergence is
still greatei-, since the calcaneum is obliquely related to the astragalus, as the latter
is to the tibia. The animal may have, however, walked on the external edge of the
foot. The sustentaculum is not very large as compared with the cuboid facet. The
latter is trapezoid in outline, the four sides all unequal, being related as to length in
the following order, beginning with the longest — inferior, su])enor external, superior
internal, internal. When the astragalar facets are held in a horizontal plane, the long
axis of this face is horizontal. In this position, the inferior sui-face of the calcaneum
is a narrow obtuse ridge, bounded on the external side by a wide deep gutki'. This
is bounded above by an angular border of a nai'row superior-external face, which
widens posteriorly. The superior surface is an obtuse I'idge. The internal sui-face is
undivided by groove or ridge and slopes outward below. The inferior ridge termi-
nates in an acumination at the inferior border of the cuboid facet. In front of the
astragalar facets, the superior face of the calcaneum has three longitudinal fossa)
separated by low ridges. The; internal, which is in front of the sustentaeular, is twice
as large as any of the others. A transverse section through the middle of the cuboid
bone is irregularly triangular. The external ftxce is smooth. On the inferior face the
large tuberosity is flattened, and is subquadrate in outline. It is separated from the
edge of the metatarsal facet by a gi'oove which is, however, closed at both ends, indi-
cating that it has not a pulley-like function for the usual tendon. On the intt lo-
superior face we have j)roximally a smooth band I'nr the navicular, and about the
middle, an oval facet transversely j)laccd f'oi' the ectocuneVfoiin. The cuboid and
navicular do not tluii present in different directions as they do in Oxyjena, but in the
same direction, as in Mesonyx.* The ectocuneiform is deeper than long, and longer

* Scoll, On New anil Little Known Crcodonts, Jour. Phil. Acad., 1886, p. 103.

346 SYNOPSIS OF THE VEKTEBRATB FAUNA OF THE PUERCO SERIES.

than wide. The cuboid facet is continuous, at a i-ight angle with the navicular facet.
The metatarsal facet is simple, and is anteroposteriorly concave. There are two distal
mesocuneiform facets, and one proximal which is turned down for a short distance in
front.

The Dissacus navajovius is a smaller species than any of the other Mesonychidse,
and is especially interesting in view of the ancestral characters which it displays.
It shows that the specialization of the extremities which has occurred in the Car-
nivora has taken place in this line also, in the jirogressive digitigradism ; the reduction
of the digits ; the development of a trochlea of the ankle-joint ; and the loss of
cusps from the molar teeth.

3Ieasurements. M.

Length of base of paradiapophysis of atlas OIG

" " odontoid process of axis 009

" " atlantal facet of axis 013

" " centrum of lumbar vertebriB 018

^ anteroposterior 020

Diameters glenoid cavity of scapula <

^ transverse 014

Transverse diameter condyles humerus in front 017

Anteroposterior diameter of flange of do 015

Length of olecranon 028

Depth olecranon at middle 015

" ulna at glenoid cavity 013

" ulna at middle 013

I vertical 0085

transverse 015

Diameter of head of femur 017

. ( anteroposterior 014

Diameters distal end of tibia {

I transverse 017

Length of astragalus 023

Greatest width of astragalus 018

I anteroposterior 014

' transverse Oil

Transverse diameter navicular facet do 018

cuboid " " 002

Length of calcaneum 043

" " tubercalcis 035

Width calcaneum at sustentaculum 0185

vertical 0085

transverse 013

Dei)th of calcaneum just lichiud cuboid facet 013

Length of cuboid 018

„.,,,. . . , ., ^ anteroposterior 010

Distal diameters of cuboid {

i transverse 010

Diameters of head of radius

Diameters trochlea of astragalus

Diameters of cuboid facet ■

r longitudinal in front 008

Diameters of ectocunr'iform I anteroposterior 010

[ transverse distally OOlj

SYNOPSIS OF THE VEItTEBRATE FAUNA OF THE PUEKf O SERIES. 347

CONDYLARTHRA.

Amon«^ the specimens whicli represent this subordei', some display poitions of
the tenii)oniry dentition. I have aheady described this in Periptychus, but have had
uo light on that of the other genera.

A cranium and a separate set of jaws of Haplocomis cornicalatus display the
last two temporary molars of the superior series. The third superioi- true molar is
just protruding and the two temporary molars in question remain in place considerably
worn. The last temporary molar is scarcely distinguishable in all its details from the
permanent true molars. The penultimate deciduous premolar closely resembles in
form and size the peimanent first premolar, diftering only in the presence of a small
anterior basal cusplct. In a lower jaw of the Anisonclms (jillianus the deciduous last
inferior pi-emolar is in place with the crown of its successor below it, and in front of
the latter is the crown of the penultimate permanent molar. The penultimate
deciduous molar is wanting. The last deciduous molar resembles in every respect the
first true molar. We have now in this kind of dental succession a state of affairs
similar to that which I have described in the creodont genus TriTsodon, and which is
probably common to all Creodonta. The last deciduous molar in both jaws resem-
bles exactly the true molars. "Were this tooth not shed these animals would be like
the Marsupialia in presenting the false appearance of four true molars.

I have shown that in Ectoconus the deciduous premolars iv and iii have the pat-
tern of the permanent true molars (Tert. Vert., PI. XXIX d, fig. 4). The corre-
sponding inferior deciduous teeth are also like the true molars, the penultimate with
the fifth cusp more anterior in position.

In the genus Protogonia I have observed that the last deciduous inferior pre-
molar is rather more complex in its form than the first ti-ue molar. It possesses three
distinct lobes arranged longitudinally, as in the Diplarthra, but the anterior lobe is
not so well developed as in those animals. The specialization of this tooth has not
progressed so far as it has in the Diplarthra, but a little more tlian in the Peripty-
chida^. This character may serve to distinguish the Phenacodontida- lioin tlie Piii|)-
tychida'.

In Periptychus, as I have already shown, tiie last di-ciduous molar is more com-
plex than the premolar which succeeds it, but is not quite so much so as a true inolar.
It has also a rudiment of the third lobe anteriorly, indicating a -step beyond Ilapiu-
conus in antero})osterior enlargement, while it is a step behind in transverse develop-
ment. The three genera, considered together, display such a series of progressive
modification of the last deciduous premolar, as to convince one of its possession of a

348 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

history of its own, and to confirm the Flowerian idea that the deciduous dentition is
an addition to mammalian development, and not a survival of reptilian conditions.
They also show that the proposition of the same author that the Mai'supialia have
four true molars is untenable, as I have previously claimed.

Fragments of skeletons of the smaller Periptychidse are not rare in my collec-
tion. I have been unable so far to fix their species, and as they present no im-
portant diffeiences from the corresponding parts of Periptychus, I do not now
describe them.

The mandibular dentitions of the species of the smaller Periptychidse are very
much alike, and they are with difficulty distinguished one from another. I give the
following table to facilitate their determination.

I. Anterior external cusp of true molars extended forwards, but not incurved.

a. Premolars elongate and compressed.

Molar series about .033 m Haploconus xiphodon.

aa. Premolars shorter, oval in longitudinal section.

Molars .025 B. angustus.

.032 E. lineatm.

" .044 H. cornicidatus.

II. Anterior external cusp of true molars witli anterior ridge directed inwards to a more or less developed fifth

cusp.

a. Premolars robust, with wide section.

Premolars with oval section ; true molars .010 U. entoconns.

Premolars with oval section ; true molars .009 H. cophaier.

Premolars nearly round in section ; molars .019 HejiiithlcBus kowalevskianus.

aa. Premolars narrower, with strong anterior cusp.

True molars .018 Am'sonchus scctorivs.

True molars .014 netnithlavs apiculatus.

True molars .012 Anisonchus gtllianiis.

True molars .011 A. agapetillus.

aaa. Premolars much compressed and elongate.
True molars .040 A. mandibtilaris.

The mandibular dentition of Ayiisonclms coniferus is not sufficiently Avell known
to be introduced into the table.

HAPLOCONUS Cope.
Amer. Naturalist, 1882, p. 417 ; Tertiary Vertebrata, 1885, p. 41.5.

The penultimate and last superior milk molars of this genus have been described
above, together with the inferior milk molars in the genus Anisonchus. It results
from these observations that the i)eculiar form of last and penultimate infei-ior pre-
molars which I have observed in the Haploconus xiphodon are permanent teeth, as I

SYNOPSIS OF THE VEKTEBRATE FAUNA OF THE PUERCO SERIES. 349

have already supposed (Tertiary Vertebrata), and that the inferior dentition repre-
sented in my Plate XXV e, fig. 6, as the milk dentition of that species, is not such, but
is the permanent dentition of the //. Imeatus. The [uvmolars in question are much
worn, so that the absence of the last true molar is jirobably due to accident rather
than to non-protrusion. A second specimen of the //. xijjhodon confirms its char-
acters.

Additional speciniciis of the AnUoiiclms cop/iatcr show lluiL it is i'i>lc'i:ii)lr to this
genus. This, with the new //. corniculatus, increases the species oi" llaploconus
to six.

nAPLOCONrs con^•ICCLATDs Cope, sp. no v.

Five more or less complete crania and a set of jaws repiesent this species. Its
characters are to be seen in its peculiar superior molar teeth, and in its superior size.

The species presents the general chaiacters of the H. iineatus, especially as to
the form of its first premolar, which has the internal cusp an elevated concentric cin-
guhim, and its enamel vertically striated with shallow grooves. The anterior cingu-
lum of the tiue molars however terminates at its interior extremity in an acute erect
cusp which is wanting in the //. Iineatus, and the cusp of the posterior ciiigulum is
isolated by a notch of the lattei-, Avhich develops a second lower cusp immediately
posterior to the first mentioned. This second posterior cusp is seen on the jwsteiior
molar of the H. lineaias, but not on any of the others. The dimensions of the 11.
corniculatus are constantly superior to those of the //. Iineatus, as the following meas-
urements will show. The lengths on the superior molar series to the canine tooth, in
three specimens of each species, are as follows :

n. Mmiculaius 045 .043 .040

H. Iineatus 034 .033 .035

The skull is elongate and is narrow in the cerebral region. The sagittal crest is
low, as is also the iniou. The orbit is small and lateral. This canines are directed ver-
tically downwards.

ifeasurements of Shell. M.

Total lengUi r25

Length from occipital condyle to last molar 058

Width between last molars, inclu.sive 037

" " canines, inclusive 020

The inferior dentition is similar to that of the II. Iineatus, but is more robust.

IIapi.OCOITOS COPHATER Cope. Anisonchus cophattr Cope; Pror. Amer. Pliilos. Sr)C., lRfi3, p. 321.

This sixicies, deseril)ed from a fragment of a mandibular ramus, is now repre-
sented by a large part of a second ramus supporting 'Cwc teeth, ami an anterioi- pari

350 STNOPSIS OF THE VERTEBRATE EAUNA OF THE PUERCO SERIES.

of a cranium with both series of superior molar teeth, lacking the anterior two pre-
molar crowns. The only reason for associating the last specimen with the two others,
is the identity in proportions and dimensions between them, and as the species differ
widely in these respects from the others of the genus, the association is justified by
our knowledge as far as it goes.

The dental characters of the upper series are a diminutive of those of the H.
entoconus. The internal cusp of the first premolar is a well-developed cone. The
fourth or posterior cingular cusp of the true molars is conic, and is almost on the
inner side of the thii-d. The anterior cingulum is distinct and reaches the fourth
tubercle. The external cingulum of the true molars is very strong, but is reduced
to a trace on the first and second premolars. The second external cusp of the last
true molar is very rudimental, so that that tooth is narrower than the others, and than
in the H. lineatas. There is a ledge at the anterior base of the first premolar.

The infraorbital foramen issues above the posterior boi-der of the anterior root of
the second premolar. The zygomatic ridge of the maxillary originates above the
middle of the first true molar. The nasal bones are elongate.

In the mandibular dentition the first premolar is larger than any of the true
molars, and has a well-developed heel. Its section is nearly a half circle in outline,
the external face being convex. The true molars have a cingulum on the anterior
half of the crown ; the jjremolars none at all.

McasiiremeiUs. M.

LengUi of superior dental series (p. lu. iv, estimated) 024

" " true molars 008

f anteroposterior 003

Diameters p. m. i {

y transverse 005

f anteroposterior 0025

Diameters m. ii \

I transverse 006

Width between p. m. ii 010

" " m. iii 009

Length posterior four inferior molars 013

{anteroposterior 00-12
transverse 0025

. ( anteroposterior 0034

Diameters ni. i

transverse 0022

Depth of ramus at m. i Oil

This, with the Anisonchus agapetillus, is the smallest species of the Condylarthra.
It will not be certain until the superior dentition of the latter is discovered, whether
it belongs to this genus or the one in which I have provisionally placed it.

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES. 351

ANISONCHUS Cope.

Proc. Amer. Phil. Soc, 1881, p. 488 ; Tertiary Vcrtehrata, 18S5. p. 408.

Aniso.\ciiu3 mandibularis Cope: Amcr. Naturalist, 1881, p. 831. Mioclwnus jnandibularis Cope; Tertiarj' Vcrtc-
brata, 188.">, p. 339.

The discovery of a second specimen of this species, wliich inchides nearly all of
the dentition of both jaws, i)roves that the position I originally assigned it is the
con-ectone. The second superior premolar is like the first in having an internal cusp,
whose base is, like that of the A. sectorius, concentric with the principal cusp of the
crown.

The great peculiarity of the species, which is displayed by all the specimens, is
the relatively large size of the second inferior premolar. It exceeds the first in ante-
roposterior extent, which is in turn longer than any of the true molars. The third
premolar is also shorter than the second. '^Phe fourth premolar and canine are not
preserved. The fifth cusp is well developed on the first and second true molars, but
is of reduced proportions on the third. The heel supports the two lateral and smaller
median cusps usual in tlris group. On the third molar the laterals are compressed and
the median has a posterior position.

In the superior true molars the second exterior cus]) of the third is well developed.
The fourth cusp is situated well inward.s, furtlici- th;iu in A. sectorius, but not so far
as in A. conifencs. There is a short anterior cingulum, which disappears before
reaching the internal angle of the crown. External cingulum distinct. The second
premolar is as large as the first, and both have a small .inti lior l>asal t iibci-fic. En-
amel surface smooth. Thiid incinolar of irdnced diameters.

ileasurementa. jV.

Length of six superior molars. . '. 0:!li

" " true molar series 010

( anteroposlcrior 00G5

Diameters p. m. ii ■; „„_

> transverse ""'

anteroposterior "OO

Diameters p. m. ii

Diameters m. ii ^ ...

' transverse ""

Length of posterior five inferior molars •''-"

inferior true molars ""'

I anteroposterior ""8

' transverse itOiWTt

Length of p. m . i ^''

^ ( anteroposterior « ^•'

Diameters m. i < noio

I transverse ""'-

A. P. S. — VOL. XVI. 2s.

352

SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUEKCO SERIES.

HEMITHL^EUS Cope.
Amer. Naturalisl, 1883, p. 833.

But two species of this genus care known to me, one of which has not been
described. This is the H. apiculcdus, and it shows such strong tendencies to Ani-
sonchns that it is possible that the two genera will have to be combined. Both
species of Ilemithlfens have an internal cusjJ on the second superior premolar, as in
Anisonehus. The position of the A. haldioini remains uncertain.

Fijr. 11.

Fitr. 12.

Fig. H. nrnillhJinm kowalevs/danun, last, five suiierior niolare, * natural size. Fig. 13. Protor/onia pucrccnsu, supe-

riiir (Icnlitidii, loss foiirtli prciiicilar. i. natural size.

IlF,ivriTin,,mis Anoui.ATrs Cope. Aiu'noneJius iipiriihitiis Cope; Tertiary Vcrtebrala, 1885, PI. XXV c, fig. 7.

The mandibles of this species have been in my possession for a considerable
time, and I have regarded them as indicating a small variety of the Anisonclius sec-
torins. But the discovery of the superior dental series shows that the species is
quite distinct, and belongs between that animal and the Ilemiihhcus koivalevsMmius
in its characters.

As in the t3'])e of Hemithlreus there are an anterior and a posterior cingula of
the superioi- molars. The posterioi* cingulum is produced a little further inwards
than the anterior, but on the first true molar its border does not project further in-
wards than the internal angle of the crown. In the second molar it projects a little
further, while in the third it projects so distinctly beyond the third cusp as to resem-
ble the condition seen in Anisonehus. The anterior edge of the cingulum does not
form a cusp, however, in either tooth, and for this reason I retain the species in the
genus rieniithl.TBus. External cingulum distinct. The internal cusps of the first and
second premolars are intermediate in form between the concentric type of A. sectorins
and the conic form of A. coniferui^. Tiie transverse diameter of the first premolar is
a little greater than tiiat of a true molar, while the crown of the third premolar is
subtriangular in section. The infra{)rl)il;il foramen issues above the second pre-
molar.

In the infci'ior Iruc niol.irs flic lifth cusp is pres(>nt, and is connected by ridge with

synopsis; of the vkktebkatk fauna of the ruEnco seizes. 353

the (.•xtriiial anterior. The opposite cusps of the heel are well developed, but ihe
posterior mediiui is quite rudimental. The first and second premolars are larger than
the molars, but not disjjroportionately so, and they have a characteristic form. Tiie
ci'own is wide at the principal cusp, but the heel is small and is at the inner side of
its posterior face. There is an anterior cusp which is a little to the inner side of the
middle Hue of the principal cusp, and which is distinct at a point much above the
base of the crown, being marked oif from the main cusp below by a shallow groove.
A line passing through the long axis of the crown is an open sigmoid. The last true
molar rises on the base of the coronoid process. Both the exteiiial side ol" the laiuus
and its inferior margin are gently convex. The anterior border of tlie masseteric
fossa is distinct, but there is no inferior border.

Measurements. M.

Leiiglli of superior molars except p. lu. iv 03.')

Length of true molars 0113

f anteroposterior 001

Diameters of p. m. iv ^

j transverse 0005

Dposterior 0035

007

Length inferior molars less p. m. iv and iii 02:5

Length true molars 0125

anteroposterior 0048

{anleroposterui
transverse . . .

1 anteroposterior,
transverse

Diameters of p. ni. i , „„,„

' transverse OO.Ju

.0015

.0038

Depth of ramus at m. i 010

" m. ii (posteriorly) 013

Jaws of eight iiulividiials of this species have come into my ])Osscssion.

IlE.Miriii.Krs KowAi.KvsKiANUS Cope ; Auier. Naturalist, 1883, p.. 833; Tertiary Vertebrata, 1885, p. lO.").

A ciiished cranium of this species shows some points prcviousl}' unknown. The
sagittal crest is low and divides into the temporal crests, which divergi- widely and
rapidly to the postorbital angles. These arc distinct, but not produced. The orbits
are small, and have a partially upward direction. The infraorbital foramen opens
above the antei-ior border of the second premolar. The latter tooth has a well-tlevel-
oped internal cusp. The palate is wide, but as the skull has been crushed this width
is probably exaggerated.

Ill the inferior dentition this species may be readily distinguished by the very
robust first and second premolars which arc shorter and wider than in any other.

354 SYNOPSIS OF THE VERTEBRATE FAUNA OP THE PUERCO SERIES.

PERIPTYCHUS Cope.
Amer. Naluralist, 1881, p. 337 ; Tertiary Vertcbrata, 1885, p. 387.

Peuu'tychus coaectatu.s Cope ; Tertiary Vertebrata, PI. XXIX d, explanation, and flgs. 7-8 ; Amer. Naturalist.
1884, p. 801, fig. 10.

This with the P. brabensis, is characteristic of the lower beds of the Puerco, as
the P. rliaModon is of the upper beds. It is not so abundant as cither of the others,
only five individuals having come into my possession.

This species is of smaller size than the P. rhahdodon and the P. carinidens,
and is especially distinguished from both, in that the cingulum of the inferior
premolars is not continuous on the inner side of the crown, but is confined
to the anterior and posterior bases, sometimes to the posterior base exclu-
sively. The first and second premolars are larger than the true molars, and the latter
diminish in size posteriorly. In the inferior molars the fifth cusp is present, and as
in the premolars there are traces of external cingula. The grooved sti'iation of the
crowns is distinct. The characters which distinguish the species from the P. braben-
sis are the following. The transverse diameter of the superior premolars is relatively
much greater than in the P. rhabdodon and P. brabensis, and the true molars have a
transversely compressed form. There is a faint cingulum on the external base of the
true molars and first premolar in both jaws, at which the grooves of the enamel tei*-
minate abruptly. This is wanting in the P. brabensis. The external faces of the supe-
rior premolars are directed obliquely forwards and inwards, a character not seen in
the P. brabensis.

Measurements of Superior Molars. M.
anteroposterior Oil

Diameters i). ui. i , „,.

transverse 016

f anteroposterior Oil

Diameters p. ni. ii \ ,„„

(^ transverse 016

f anteroposterior 0095

Diameters m. ii ^ „. ,

( transverse 013

PEIUrTYCnUB BRABENSIS Sp. nOV.

Twenty individuals represent this species in my collections, nearly all of them
consisting of jaws only. Of these four only present the dentition of both jaws ; one
exhibits nearly the entire dentition of both maxillary bones, and one the last tempo-
rary molar of the lower jaw.

This species is still smaller than the P. coarctatus, and diifers from it in the
same way, i. e., in the non-continuation of the cingulum across the interior
side of the inferior premolars. But it diflers from the P. coarctatus, as already

SYNOPSIS OF THE VEUTEBltATE FAUNA OF THE PUEIICO SERIES. 355

pointed out, in the smallur tiansversc diameter of the prenioUirs, especially of
the superior series, and in the absence of ein^ula on the exteinal side of the
premolars in both jaws. The inferior true molars also possess but traces of the
cingula. The internal cinguluni of the superior premolars is extensive, contimiing
round to the front and nearly to the external base of the first premolar. In the infe-
rior premolars there is a short posterioi' heel, and on the first, a short anterior cingu-
luiu, which is however nut always present. The infraorbital i'urameu issues above the
front of the first premolar.

Measurements. No. 1 t)/j)e. II.

Length of p. m. i, m. i, and m. ii, superior 029

( iintoroposlerior, superior Oil

Diameters p. m. i

I Inmsverse, " 013

( anteroposterior 012

Diameters p. m. u, inferior <

(^ transverse 008

Superior Molars, No. 2.

Length of molar scries 070

" " true molars 029

{Hutcroposlcrior 010
transverse Oil

^. ..(anteroposterior 098

Diameters m. ii <

[^ transverse 013

Inferior Molars, No. 3.

Length of last five molars 030

^. . .anteroposterior 0115

Diameters p. in. i <

transverse 0095

11. ii <

„. , ... anteroposterior 008

Diameters of m. ii •{

transverse 008

„. . . ...(anteroposterior 0088

Diameters of in. Ill ^

( transverse 007

From the Lower Puerco only. I ). Baldwin.

ECTOCONUS Cope.
Amer. Naturalist, 1884, p. 79.'3 ; Tertiary Vertebrata, 1885, PI. XXXV.

But one species, the £". dilri(/onus Cope, is known. Excepting the briel" notes
contained in the aliovc-mentioned publications the general characters of both genus
and species remain undescribed.

In the superior dentition the first, second and third premolars possess one exter-
nal and one internal cusps. In ili* inHiior series the first and second have an internal
cusp and a heel. The coracoid process is a robust lecurved tuberosity. Tlie astra-
galus is slightly convex anteroposteriorly, and slightly concave transversely. The

356 SYNorsis of the vektebkate fauna of the puerco series.

ti-ochlea is not oblique to the axes of tlie bone. Tlie head presents convex faces an-
teriorly interiorly. The anterior face is convex in every direction, and is continued
to the external side. The internal face is not separated from the anterior along- the
superior border, but is separated below and distally by a strong notch. This interior
facet indicates a large tibiale oi- "internal navicular,'' a bone well known in Rodentia,
and in Bathmodon among the Coryphodontidw. This fiicet is larger than in Peripty-
chus, and in the latter genus it is not cut oft' by a groove as in Ectoconus. The cal-
caneum has a very long tuber. The astragaline facets are in a general horizontal
plane, the internal (sustentaculum) small and a little concave; the external large,
wide, and a little convex. No distinct fibular facet. The cuboid facet is large, and
nearly anterior in presentation. The cuboid has the usual tuberosity, and also a large"
distal facet.

The foot was evidently entirely plantigrade and penladactyle in this genus.

In its dentition Ectoconus presents the most complex known form of the tritu-
bercular type.

EcTooONUs DiTRiGONus Copc ; Aiuer. Naturalist, 1884, p. 790, tig. 4. Periptyclms ditrujoims Cope ; Tertiary Verte-
lirata, p. 404, PI. XXIII g, fig. la. Conoryctes dUrifjouus Cope ; Tertiary Vertebrata, 1885, PI. XXIX d, explana-
tion, and figs. 2-G.

Thirty-two individuals of this species have been sent me, all from the Lower
Puerco beds. The most important of these include the teeth and all the dentigerous
bones, excepting the premaxillaries, with fragments of humerus, scapula and tibia
with calcaneiim, astragalus and cuboid elements entire. Of another, the proximal
parts of the astragalus and calcaneum are preserved with the heads of the first, a
median, and the fifth metatarsals.

The dentition has been described so far as the molars and superior incisors arc
concerned. The first and second true molars usually possess eight cusps, but occa-
sionally there are nine, that is two external cingular cusps. The heels of the first
and second inferior premolai's arc tubercular, and the external cingulum of the infe-
rior true molars is well marked. The enamel is nearly smooth.

The external (fibula) side of the astragalus is a vertical facet. The internal face
is nearly vertical. Beyond it a large depressed tuberosity projects a short distance
inwards but not posteriorly. It bounds tlie sustentacnlar facet of the astragalus
behind. This postsustentacular tuberosity is truncated inwards and downwards {)os-
teriorly. The sustentacnlar pi'ocess of the calcaneum projects outwards and forwards
in a subcirciilar rim beyond the smaller subround facet. The external facet forms
about one-fifth of a circular band, the concavity being next the middle of the proxi-

SYNOPSIS OF THE VEItTEBRATE FAUXA OF THE PUERCO SERIES. 357

mal cud of the bone. Between it and the sustentaeuhir laeet is a ligamentous fossa
In front of it the superior surface descends into a deep ti'ansverse notch which re-
ceives a descending angidar process of the astragahis which is broken from my
specimens. In other words the superior border of the cuboid facet rises ui)wards
abru|)tly in front of this depression. This facet is quite large and narrows to an apex
inwards, where it terminates at a small vertical tuberosity. Inferior to this apex is a
8ul)triangular fossa, which is bounded externally by another tuberosity. A lai'ge
longitudinal tuberosity supports the external border of the cuboid facet on the in-
ferior side.

The skull fragments show that this species differed from all of the known Perip-
tychida; in having a very high and i-obust sagittal crest.

Three vertebra^ of the spccimcu described are preserved, the axis, a lumbar and
two caudals; all without arches. The axis is about as long as the lumbar, omitting
the odontoid process. The latter is rather short and is depressed, the section being a
tiansverse oval. Its articular surface is continuous with the atlantal facets. Body
with a median iufciior keel. The lumbar is short, but exceeds the axis in ti-ansverse
diameter. The articular faces are subplane, while the sides of the ceiitrnin are con-
cave, and not keeled at an}- point. '^Pwo strong foramina perfoi'ate the Hoor of the
neural canal. The bodies of the caudals are depressed and a little longer than wide.
One of them has a median keci-angle below. Other specimens show more distal
caudals of elongate form, showing that this species had a long tail like the Phrna-

o — - — I —

{anicropol
trims vers

Measurements of sp. No. 1. M.

LcngUi of superior true molars willi p. m. i ami ii 044

nnlcroposlorior f07

ise Oil

f nnleroposlerior "Id

Diameters m. i { ,_.,

[ transverse 01..

iaiilcropostcrior 010
tninsverse 01. i

f anteroposterior 000

Diameters m. iii {

(^ transverse 01 J

Lcnglli inferior dental series including canine 073

anteroposterior 008

r • !'
I of canine {

1'

Diameters base (,. .,„ ., ,

transverse OOS,)

Length of true molars 03 1

f anteroposterior 008
transverse <""•>

f anteroposterior OIO.I

Diameters m. ii { ,,.

I transverse '•"'

f anteroposterior 012

Diameters m. iii { __,,

I transverse ""•'

Widths of scapula proximally

358 STKOPSis or the vbktebeate PAirerA of the puerco series.

Measurements of »p. No. 1. M.

Depth of ramus at p. m. iii 0~1

•■m.i 035

( anteroposterior, total 030

Diameter axis \ noo

I transverse, greatest "«>'*

Length of odontoiil process to atlantal facet 009

f anteroposterior 030

Diameters lumbar centrum i . , . f transverse 030

articular face { . ,
I I vertical 031

f anteroposterior 031

Diameters proximal caudal centrum ■ . , f transverse 016

I articular-

[ l vertical 015

anteroposterior, total 041

glenoid cavity 039

neck 033

transverse, glenoid cavity 019

f anteroposterior* 032

Diameters distal end of tiliia \ „„,

( transverse 035

Greatest length of astragalus 035

width " " 036

„. „ ,, (anteroposterior 031

Diameters of trochlea-^

y transverse 018

Length of calcaneum 058

"Width proximally (transversely measured) 038

Transverse extent of sustentaculum 014

Transverse diameter of cuboid facet 018

Length of cuboid 019

Transverse diameter of distal facet do 019

Vertical " Oil

Specimen No. S.

, . , f anteroposterior. ... i 035

Diameters peduncle of ilium <

[ transverse 023

Width of internal side of peduncle of ilium 020

„ , , ( aiiteronosterior 028

Diameters of acetabulum -^

[ vertical 038

Greatest diameter of peduncle of pubis Oil

anteroposterior 027

Diameters of ischium at sjiinc

' transverse 009

Estimated length of femur 140

Width do. at condyles 045

Anteroposterior diameter do. at rotular crests 043

The pelvis has large obturator foi-amina. The ilium is rather short and robust,
and it has a wide anterior face to the crest, but this is not so wide as the interior face,
except at the peduncle, where it is wider. The anterior inferior spine is a vertical
oval tuberosity a centimeter above the acetabulum. The latter has a wide lisjament-

* A small posterior tuberosity estimated for.

SYNOPSIS OF THE VERTEBRATE FAUNA OP THE PUERCO SERIES. 359

ous fossa at the fundus, and the usual ischiadic notch and groove. The pubis is
slender. The ischium is tlat, and the tuberosit}^ is a convexity of the posterior edge
which terminates in a little angular contraction, forming a "spine."

The fossa ligamenti teris of the femur is a fissure which runs out to the margin
of the head. The trochanters are large; the third moderately prominent, and partly
opposite to the lesser tiochanter. The rotular groove is wide and elevated, and its
borders are equal. The external condyle is narrower than the internal. On the
whole the femur is a good deal like that of the Periptychvs rhahdodoa.

PROTOGONIA Cope.

Proc. Amer. Phil. Soc, 1S81, p. 492 ; Tertiary Verlebrata, 1885, p. 424. Phenaeodut pt. Cope ; loc. cit., 1881, p. 493 ;
1885, p. 488.

This genus was originally distinguished from Phenacodus by the presence of
but one external tubercle on the first superior premolar, since there are two in Phen-
acodus. Additional specimens show that the second external tubercle of the P. puer-
censis is normally rudi mental, so that that species may be jii-operly referred to Proto-
gonia. It is further distinguished from Phenacodus by the presence of but one
external cusp on the second superior premolar, in which it agi'ces again with the spe-
cies referred to Protogonia. The latter genus thus defined embraces live species
which difter as follows :

I. Fiflli cusp present on inferior true molars.

Last four inferior molars, .025 ; internal cusp on inferior p. m. i, rudimental P. suniensis.

Last four inferior molars, .0:50 ; internal cusp of inferior p. m. i, large P. plitifera.

II. Fifth cusp wanting from inferior molars, except sometimes m. i.

First superior premolar with rudimental second e.xternal cusp ; inferior true molars, .020 ; last robust premolars with

no. or tubercular heels /'. puerceiisis.

Inferior true molars, .020 ; last small ; premolars with wide, flat, smooth heels /*. calccohita.

As may be seen from the catalogue of species, the material representing this
genus is quite abundant. This is true only of jaws and teeth, for identi liable parts
of skeletons are yet very rare. A very much damaged cranium of the P. puercensis
displays nearly all the dentition, the iuci.sors and the fourth premolar being ab.sent.
The first and second premolars only have internal cu.sps. In the superior canines
the vertical direction is the same as is seen in Ilaploconus sp.

The fifth cusp is wanting from the last inferior true molar in the 7*. y>^/c//*era,
and is in a few specimens absent from the second also. Specific diilerence cannot bo
predicated on the presence or absence of this cusp, although in more specialized types,
its variations are of generic importance. The premolars of the lower series are spaced
in the P. plicifera, while they an- in close succession in the /\ puo'censis.

A. P. S. — VOL. XVL 2t.

360 SYNOPSIS OF THE VERTEBRATE FAUNA OF THE PUERCO SERIES.

AMBLYPODA.
PANTOLAMBDA Cope.
Amer. Naturalist, 1882, p. 418 ; Tertiary Vertebrata, 1885, p. 601.

Additional material belonging to the two known species of this genus, throw
light on points of structure hitherto unknown.

As regards the dentition, the superior premolars are identical with those of
Coryphodou.

In the anterior foot the cuneiform has the general character of that of Corypho-
dou. The pisiform facet is wider, thus approaching the unguiculate types of the same
epoch.

In the posterior foot the cuboid of the P. cavirictus differs from that of
any of the Coryphodontida^ which I have seen in the greater mutual obliquity
of the two proximal facets. That for the astragalus is a wide concave fossa;
that for the calcaneum is a hook-shaped band, the convexity proximad, and
the longer arm, or stem, of the hook anterioi-, and the shorter posterior to
a ligamentous fossa. The anterior band-like facet turns transversely distally. The
position of the cuboid is oblique in the foot, giving the digits which arise from it a
divergent direction externally. The astragalus of this species closely resembles that
of P. latlimodon. The cctocunei'form is much like that of Coryphodou, but is not so
depressed, the anterior face being squai'C. The mesocunet'form has only two-thirds
the longitudinal depth in front. The entocuneiform is narrower transversely than in
Coryphodou, and approaches the form of some of the unguiculates. It indicates a
smaller internal digit than in Coryphodou. The above-described bones all belong to
one individual.

EXPLANATION OF PLATES.

PLATE IV.

Hemiganus otariidens Cope, natural size.
Fig. 1. Part of maxillary bone displaying canine anil two premolar teeth, with ? lachrymal foramen at a; and with

part of premaxillary bone.
Fig. 2. Left ramus of mandible from left side. The ? canine was found separate, and is drawn in; a, same from

above, canine omitted.
Fig. 3. Parts of pariijtal and frontal bones from above ; a, temporal ridge.

Fig. 4. Right frontal and part of i)arietal Ijone. llie sutural surface showing olfactory fossoe on inferior face.
Fig. 5. Left inferior ? canine tootli represented in flg. 3, from front ; ii, from behind.
Fig. 6. Inferior premolar tooth free from alveolus, from front ; a, from inner side ; h, from above.
Fig. 7. First inferior molar of right side, external view.
Fig. 8. Atlas, left lialf, fnun left side ; a, from behind.
Fig. 9. Axis from above ; a, from right side.
Fig. 10. Cervical vertebra without epiphyses ; a, from below.
Figs. 11-12. Cervical vertebra- ; a, from below.

rhil 3oc Vol X\a Pan 11 PlaicIX

.rap.sacl.ions Am rrul sec vol .\ v i h^lT' ii rime

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V 'L.^ ' -fe-

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ifoj^M

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ife'

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.-7 HEMIGANUS OTARIIDENS Vi . 8-9 ONYCHODECTES TISONENSIS

.AGIAULAX MC'

SYNOPSIS OF THE VEltTEBRATE FAUNA OF THE PUERCO SERIES. 361

PLATE V.

Figs. 1-6. nemigailut olariidtiis, parts of specimen figured on preceding plate, natural size.

Fig. 1. Ulna of right side, from within ; <i, from above.

Fig. 3. Proximal extremity of radius from below ; a, proximal view.

Fig. 3. Left femur, proximal half, from behind ; a, external view.

Fig. 4. Distal half of right tibia, from front ; a, from external side ; b, distal extremity.

Fig. 5. Metacarpus of pollex, externoanterior view ; a, internal view ; 6, edge view.

Fig. G. Mctapodial, from side ; a, from front ; b, from behind.

Fig. 7. Ungual phalange, side view ; a, proximal extreniily ; b, from above ; e, from below.

Fig. 8. Onychodectti tisonensis Cope ; maxillary bone of right side, with four molars, from below, § natural size ; tvpi-

ADDENDUM.
On page 311 it is stated that in the genus Hemiganus "there are probably but two true molars." Further
cleaning of°the specimen shows that there are three true molars, as represented in Plate IV, fig. 2 a.

m^^'

vV^

dSiWr^,.

'^

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_ v

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•;iti:iaBiSiSRss:'=

s**xvr^^

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SYNAPSIS OF THE VEUtTEBRATE FAUNA OF THE PUEKCO SERIES. 361

PLATE V.

Figs. 1-6. Iltmiganiit otariideiii, parts of specimen figured on preceding plate, natural size.

Fig. I. Ulna of right side, from within ; a, from above.

Fig. 2. Prosimal extremity of radius from below ; a, proximal view.

Fig. 3. Lefl femur, proximal half, from behind ; a, external view.

Fig. 4. Distal half of right tibia, from front ; «, from external side ; b, distal extremity.

Fig. 5. Metacarpus of poUex, exlernoanterior view ; a, internal view ; b, edge view.

Fig. 0. Mctapodial, froiu side ; fl, from front ; b, from behind.

Fig. 7. Ungual phalange, side view ; a, proximal extremity ; b, from above; c, from below.

Fig. 8. Onyehodectes tisonensis Cope ; maxillary bone of right side, with four molars, from below, } natural size ; typi-
cal specimen ; a, the same from the palatal side.

Fig. 9. Left mandibular ramus of the same species, natural size, broken, external side ; a, crown of second true
molar seen from above, J natural size.

Fig. 10. Ifeoplagiuulax moleitus Cope, fourth inferior premolar, external view, natur.il size ; a, the same from above.

Fig. 11. Neoplagiaulax molesttit, left mandibular ramus, with roots of fourth premolar only, external view ; natural
size.

AKTICLE VI.

ON THE SHOULDEK-GIEDLE AND EXTEEMITIES OF EEYOPS.

BY E. D. COPE.

Read before the Ameiican Philosophical Society, January 20, 1888.

Our knowledge of the genns Eryops, which is one of the most abundant types
of the Rhachitomous Batnichia, extends to the skull in general, the vertebral column,
the pelvis, humerus, femur, and tibia.* The shoulder-girdle, fore foot, and hind foot
are unknown, although the hind foot of a species from the coal measures provisionally
referred to this genus has been described.f In the present paper I can describe the
missing parts from a skeleton found with skull of the Eryops megaceyhalus, which
Avas obtained in Texas, in the Permian beds, by that indefatigable and skillful col-
lector, Jacob Boll.

The shoulder-girdle embraces scapula, coracoid, prsecoracoid, clavicle, and epi-
sternum. The scapula is flat and elongate; its superior portion is rather thin, and gradu-
ally expanded and rounded like the end of a boat's oar. Below it passes directly into the
coracoid and praicoracoid, the suture becoming obliterated early. The prajcoracoid is
larger than the coracoid, is convex downwards, while the coracoid is convex upwards.
This leaves a fossa at their line of junction on both surfaces, and through this the cora-
coid foramen is pierced. The posterior border of the coracoid is regularly convex, and
is not notched as in the Pelycosaurian Theromora.J The internal and anterior bor-
der of the praicoracoid foi'in a continuum of convex outline, the former passing rather
abruptly into the latter. The clavicles are curved bones, forming, as usual, the ante-
rior border of the shoulder-girdle. Their superior portion is directed upwards and

* Proc. Amer. Philos. Soc, 1880, p. 52.

t Trans. Amer. Philos. Soc, 1886, p. 289.

X Theromora is subsliluted for Theromorpha, a name which had been previously used.

ON THE SHOULl>ISR-GIRDLE AND EXTREMITIES OF ERYOPS. 363

backwards, in the foi-m of a band, and it does not extend so far on the scapula as is
usual with Keptilia, and teiminatcs in a rather abrupt flat apex. The inferior
portion becomes horizontal and is directed forwards at an open an<^le, meeting its fel-
low on the middle line. This portion expands a little, presenting a thin edge poste-
riorly, and an oblique truncation inwards and forwards with grooved and finely
digitate edge, at the distal ind. The clavicles underlap the episternum. The latter
is a flat discoidal bone rather wider tiian long, with a regularly convex, thin posterior
edge. The lateral jjortions are overlapped by the prrecoracoids. The anterior border
is coai'sely toothed, as though for the attachment of an omosternum. Of true
sternum I do not observe any trace of sternum.

The ditterences between the shoulder-giidles of Eryops and of Actinodon
(Gaudry) are considerable. The latter has been described and Hgui'cd ])y Professor
Gaudry, and I have had, through his kindness, the opportunity of examining the
typical specimen. An obvious diil'erence is that the latter genus, as in many Stego-
cephali, has the epistei'num and proximal parts of the clavicles sculptured on the
inferior (external) face, with exostosis, which occupies the true skin. The clavicle
articulates externally with a slender bone, which is regarded by Gaudiy as clavicle.
It occupies the position of the anterior thickened portion of the scapula in Eryops.
What its true homology is is not clear to me, but it is in the position of the epicla-
vicle of the fishes. The scapula is co'issified Avith the pra?coracoid. The true coia-
coid is very small, less even than in Eryo])s. An easily observed peculiarity of
Actinodon is that the episternum is longitudinal diamond-shaped, as in Stegoccphali
gem rally.

The shoulder-girdle of Cricotus is in my collection, but is so diflicult to extri-
cate from the matrix that its chai'acters are not all clear. The clavicles are expanded
inwards over the edges of the episternum as in Actinodon, and more widely than in
Eryops, and the expanded surface is sculptured as in that genus. The slender part
of the clavicle is strongly curved upwards, as in Actinodon. The episternum is also
produced posteriorly as in Actinodon.

As compared with the Pelycosaurian reptiles, the shoulder-girdle of Eryops and
its allies shows several im[)ortant points of resemblance, some of which I have already
described. iSuch are the small coracoid and large pra-coracoid, both coissilied with
the scapula in adults. The episternum in Dimetrodon at least, diiVers in its long nar-
row posterior pi-olongation, as in Lacertilia.'' The clavicles have not been described
in Clepsydropidie, although they have been in Diadcctidaj.f In the latter the clavi-

•TraDsactions Amer. Phil. Soc, 188C, p. 292, PI. Ill, fig. D, where I inadvertenlly called it sternum.
tProc. Amer. Phil. Soc, 1883, p. 615.

364 ON THE SHOULDER-OIKDLE AND EXTREMITIES OF ERTOPS.

cles are narrow, and expand but little at their contact with the episternum. They
form a symphysis behind and below the episternum, which has an exclusively anterior
and superior exposure. The latter is wedge-shaped in longitudinal section, thinning
out rapidly posteriorly, where it is not produced on the middle line as in the Clepsy-
dropidse.

In Dimetrodon the form of the clavicle is similar to that of Crieotus. That is
the inferior poi-tion is bent inwards at right angles to the vertical portion, and is
greatly expanded in the anterior direction. Its inner margin is finely digitate by the
production of numerous radiating ridges. This portion of the shoulder-girdle projects
forwards in the manner of the so-called epiplastral bone of the j^lastron of the Testu-
dinata. With the episternum in place this part of the shoulder-girdle of Dimetro-
don resembles the anterior lobe of the plastron of a tortoise. A narroAving of the
prsecoracoid and scapula, and an elongation of the coracoid, would give the Testu-
dinate shoulder-girdle. Such a change in some unknown member of the Theromor-
ous order, together with modifications of the abdominal costoids, has given origin
to the order Testudinata.

The anterior leg of Eryops is short, and very robust. The anterior foot has the
character, entirely exceptional in the Batrachia, of having five digits. The humerus
has been already partially described.* It resembles very nearly that described by
Professor Gaudiy to his Eucliirosaurus rochei.f It is short and very robust, and the
extremities are greatly expanded and almost at right angles to each other. The
proximal extremity with its band-like articular surface, is strongly decurved antero-
exteriorly, ending in a strong protuberance. The distal end has an enormous plate-
like expansion of the internal epicondyle, greater than that exhibited by
Euchirosaurus rochei. The condyle is subglobular and large, and the external
epicondyle is partly posterior to it. This epicondyle forms a process ' flat-
tened on the posterior face, which is continued upwards into the shaft
of the humerus, and is bounded below by a semicircular sharp edge. On the external
border of the shaft above the notch between the condyle and the external epicondjde,
is a short, obtuse, vei'tically compressed process directed at right angles to the shaft.
A similar process exists in EucMrosaurus rochei (Gaudry).

The ulna has little or no olecranon, and has a strong glenoid cavity on the supe-
rior aspect for the condyle of the humerus. It resembles much that of a pelyco-
saurian. The shaft is slender, and the distal extremity is not expanded. At the
latter place the superior face is flat, and the inferior is convex. The radius is a short

*Proc. Amer. Phil. Soc, 1880, p. 54.

t Enchainements du Monde Animal ; Fossiles primaires, 1883, p. 277.

ox THE SHOULDER-GIRDLE AND EXTREMITIES OP ERTOPS. 365

robust element, with both exticmities expanded, the distal much more so than the
proximal. The latter is truncated, with a wide oval outHne, and rests, as in land ver-
tebrates generally, on the coronoid pi'ocess of the ulna. The shaft is suboval in
section, and is much stouter than that of the ulna. The distal end is convex and
is horizontally expanded so as to rest on three carpal elements : radiale, ccntrale 1,
and intermediinn. Its inferior face is Hat distally, and the distal outline is openly
angulate with subequal faces; one principally for the radiale, and the other principally
for the intermedium.

The carpus consists often, possibly of eleven elements. Four of these are in the
proximal row, viz., radiale, centrale, intei-medium and ulnarc. The second row con-
sists of five carpal ia. Between the two rows is centrale 2, which appears to have a
small centrale 3 separated from its external end, but on the palmar face, the distinc-
tion is not clear. The carpals are, in the order of size, as follows : ulnai-e, centrale 1,
centrale 2, i-adialo, carpale 5, intermedium, carpale 2, carpalia 1, 4 and 3. The meta-
carpals are llattcned, with the extremities truncate and expanded ; the proximal the
wider; the distal without indication of condyles. The thiid and fourth are twice as
long as wide distally, and the first is as wide as long. The fourth digit is the only
one in the specimen in wliicli the i>lKilanges are preserved. There are three of them,
the ungual very small and subconic in form, the inferior face flatter than tiie superior.
The two other phalanges are about as wide as long.

MeamretnenU. MM.

Diameters cpistcmum

I. Slioultler-girdlc.
Width at anterior border of pra'corncoid 235

I anteroposterior 110

' transverse 130

Greatest (inferior) width of clavicle 70

Length of coracopr.rcoracoid on upper surface 100

II. Anterior leg.

Humerus, greatest width proxiniallj' '.lO

" " " mcdiaily 40

" distally 103

" " length (part estimated) 130

Ulna, length 112

Ipro.ximally 45

medially 15

distally 30

Radius, length 78

■ proximally 42

Radius, transverse diameter ■ medially 27

. distally CO

Length of carpus from ulna to m. iv 02

Wiillli of carpu."!, proximal row 05

Length of fourth digit 78

366 ON THE SHOULDER-GIRDLE AND EXTREMITIES OF ERTOPS.

This carpus and fore foot is noteworthy from the presence of five digits, an un-
usual character in the class Batrachia. The genus Archegosaurus has but- four,
although according to Baur* there are five carpalia. In the presence of two centralia,
Eryops agrees with occasional specimens of both Cryj)tohranc7tus allegheniensis and
Megalobatrachus japonicus (Wiedersheim). If three centralia are present, the resem-
blance to Archegosaurus decheni is greater, where, according to Baur, there are four.

The posterior foot found at the same time and place as, and having appropriate
proportions to, the parts already described, is not so well preserved. The distal ele-
ments of the leg have subequal widths, but they are folded back to back so that one
of them has lost its tarsal connections. One of them, perhaps the tibia, remains
articulated to two proximal tarsals, probably tibiale and intermedium. The approxi-
mated surfaces of these bones are very thick, as is the internal fiice of the tibiale.
The external face of the intermedium is, on the conti-ary, thinned out. The tibiale is
characterized by the presence of a round, flat, discoid tuberosity on its ? posterior
face, which resembles the disc of a button. Distad from the tibiale is a large trans-
verse centrale, in the position of the Pelycosaurian and Mammalian navicular. It
articulates with the distal extremities of the intermedium as well. Its distal face
articulates with two tarsalia, which are somewhat displaced in the specimen. The
remainder of the foot is in two separate pieces, which represent probably parts of
both posterior feet. In one of these I count five metatarsal bones very much dis-
placed. Two of them are of rather small size. In the other block is a series of four
consecutive phalanges, all wider than long. The distal one is quite small, but the
ungual is not x^resent.

Two small tarsi from opposite sides of perhaps the same individual were found
mixed vsrith the bones of the larger animal. They belong to an allied form. The
tibiale has the same button-like disc on one of its faces, and it articulates distally
with a navicular-centrale. The ? intermedium is a shorter bone and unites with a
single element, which may represent a centrale 2, or a cuboid. It appears to articu-
late distally with two elements, though the matrix does not let it be determined
whether the lines observed are sutures or fractures. And it is not certain whether
the two supposed elements are tarsalia or proximal ends of metatarsalia. There is,
as in the larger tarsus, no indication of an exterior or fibular series of tarsals. The
bone referred to as possibly cuboid, rises on the external side of the level of the
proximal face of the intermedium, and may be therefore a fibulare, which presents
very little proximal facet. This tarsus strongly suggests the presence of but four
digits.

* Beitriige z. Morphogenie d. Carpus u. Tarsus d. Vertebraten, 1887, i, p. 53.

AKTIULE \ II.

'IIR CRETACEOUS AND TERTIARY GEOLOGY OF THE SERGIPE-

ALAGOAS liASIN OF BRAZIJ..

BY JOUX C. nRANNKIl, Ph.D.,

PROFESSOR OK GEOUKiY IN TIIK l^NlVKItslTY OF INDIANA,
STATF. <iEOI.O(il.ST OF AUKANSAS, FORMF.lil.Y A!i<lSTANT GROI.OIilST <1F TIIF. IMPEEIIAI. i:E<>I.OMCAI, SI'IIVIV ol r.lIAZII..

Read before the American Philosophical Society, September 7, 1888.

r.\Ki' I.
Prefatory Notes.

Tn the years 1875 and 187G, while assistant geologist on the Imperial Geological
Survey of Brazil ("A Commissao Geologiea do Imperio do Brazil"), I si)ent several
months in examining the geology of the provinces of Pernambuco, Alagoas and Ser-
gipe. The work done was but little more than a hasty reconnoissance, made without
any maps whatevei-, even of the rudest kind, and without either time, instruments or
appropiiation for making them,

I cannot therefore refrain from expressing my reticence in regard to publishing
what little I know concerning the geology, of the region treated of in the present
paper. Althoiigli tlie work of the Imperial Survey in the I'ogion was all that could
have been accomplished in the time occupied, and though it was much more extensive
than any done prior to my visit, it was far from satisfactory.

Three other reasons have caused this delay in the publication of my observations.
First, I hoped that the Brazilian Government would see lit to have the results of the
Survey's work published more in detail than is possible here. This has not only
never been done, but 'until the pul)lication by that government of Dr. C A. White's
important "Contributions to the Pahuontology of Brazil"' none of the results have
been l^rought out, except as the foiiuer memlicrs have found tiuie to prepare iHicf
papers and [niblish them wherever an opportunity olfcrcd. Evcmi in the case of Dr.

A. r. s. — VOL. XVI. 2u.

370 THE CRETACEOUS AND TERTIARY GEOLOGY

White's jiaper, the delay in its printing was veiy tedious, the report having been
issued four years after the manuscripts and drawings were prepared. Second, I ha\ e
delayed on account of the incompleteness of my observations, and because of my
hesitation lest I should add to that class of writings upon Brazilian geology which is
made up, to so great an extent, of generalizations based upon very limited observa-
tions. A third reason for delay has been the somewhat vague hope that I might yet
have an opportunity of completing my studies. As time goes on, however, the proba-
bility of my being able to revisit the provinces of Sergipe and Alagoas appears to
lessen rather than increase. Since my exploration in those provinces I have returned
to Brazil three times, but as Aracaju and Penedo are small ports off the principal
lines of travel, visited only by sailing vessels and small coasting steamers from Bahia
or Pernambuco, I have never found it possible to make further examination of the
geology of this interesting region. It is hoped, however, that these notes may
be of service to those who may hereafter undertake to add to or finish the work
begun.

Mucli of this i>aper must be of a statistical nature, for the relation of facts to
each other, except in a general way, is not understood, and I do not deem it necessary
or discreet to enter upon the discussion of these relations in detail until more thorough
information is had of the region in question.

The sketch map accompanying this paper makes little pretension to accuracy
and is only introduced for the pui'pose of giving some idea of the geography of the
region under discussion.

The defects of the records are quite evident to myself, but in spite of such de-
fects, I am confident that the facts presented will be useful to those who may here-
after have occasion to do work in this region. As far as I am av/are, no geological
work has been done here since 1876, if I except the trip made up the Eio Sao Fi-an-
cisco by Mr. Derby, the geologic results' of which were published in the Archivos do
Museu N'acional, Vol. IV, p. 87 et seq., but this paper, while it treats somewhat of
the mesozoic geology of the region, does not appear to represent any new work
thereon. What 1 have to say in this paper is upon my own authority, except where
others are quoted and credit given.

I should be remiss in a most pleasant duty did I not refer here to the universal
encouragement which the naturalist, traveler, or stranger moets at the hands of
Brazilians in all parts of that countrj^, and especially in the provinces of Sergipe and
Alaguas. The pooi-est laborer kindly and i)olitely urges you to " make no ceremony "
about sharing his humble iare, while the richest gentleman assures you and makes
you feel that his house and his servants are " yours " indeed. I am under especial

OK Tin: SEKfilPE-AL/VGOAS BASIN OF BRAZIL. 371

obligations to Robert Brown, Esq., II. B. M.'s Consul at Maroim, for assistance in the
prosecution of my woik in the province of Sei-gipc.

OUTLINES OP THE BKAZILIAN CRETACEOUS.

Tlial tliu relations of the Sergipe-Alaguas basin to the otlu-r more or loss similar
deposits of the Brazilian coast may be better understood, I give here l)rielly the
prominent features of the cretaceous geology of the .several basins along the coast as
far as they are known.

I use the term '"basin,'' however, in speaking of the geology of the provinces of
Sergipe and Alagoas partly to separate the locality geographicall}' from others of a
more or less similar character north and south, and partly because it seems to have
been one of the sepai-ate wide-mouthed bajs or indentations — mere curves of the
shore line — which characterized this coast in cretaceous times.

The exposures of mesozoic rocks along and near the Brazilian coast point to the
probability that this portion of the coast line of the Atlantic was, during cretaceous
times, but slightly difi'erent fi-om that of the present day.

The accompanying small map of Brazil will give an idea of the geographical
relations of the known cretaceous exposures to each other, and will suggest, at the
same time, the form of the coast at the time of their deposition.

The most southerly exposures of cretaceous rocks thus far discovei-ed in Brazil
are upon the island of Sao Fi-ancisco in the province of Sta. Catherina. Others
occur along the coast at the Abrolhos, at Bahia, throughout the extension of the
Sei-gipe- Alagoas basin from Estancia to Kio Formozo in the province of Pernambuco,
at Olinda, Maria Farinha and on the island of Itamaricu in the province of Pernam-
buco, at Parabyha in the province of this name, on the Kio ^Slossoro in Pio Grande do
Xorte, and about Crato in Cear;i, while the most noi'tlicMly exposure known along
the coast is IoiiikI on Kio Piabas, province of Paru. More detailed descriptions of
these several basins and localities will be given at the end of the description and dis-
cussion of the Sergipc-Alagoas basin.

TKEVIOUS WOUK.

A glance at the annoted biiiliography of the subject given at the end of this
paper will show that almost no work was done on the geology of the Sergipc-Alagoas
basin prior to 1875.

In IS.38, CJeorge Gardner, the English botanist, during a short voyage on the
Sao Francisco made in pursuit of his botanical .studies, made a ffw notes upon the

372 THE CRETACEOUS AND TERTIARY GEOLOGY

o-eoloo-v of tlie i-co-ion. These notes, however, were so few as not to be worth men-
tioning, except for the fact that lie afterwards made some valuable observations upon
the mesozoic geology of Ceara, and expressed the opinion that the Penedo sandstones
were "identical with those from the upper sandstones of Crato."

Charles Darwin, in his " Observations," makes references to the mesozoic geology
of Pernambuco and Bahia, but he did not visit the coast between those places.

In 1866, Prof Ch. Fred. Hartt touched at Aracaju, Maroim, and Penedo, and
gives the results of his obsci'vations in his " Geology and Physical Geography of
Brazil." He obtained a small collection of fossils at Maroim, but one of sufficient
importance to throw light upon the age of the rocks. He noted exposures at Sapn-
cary (by him called Sapueahy), Maroim, and through. Mr. Lane, who then lived at
Maroim, obtained specimens of Natica. The cephalopods of this collection were
described by Prof Alpheus Hyatt, and the descriptions published in Hartt's " Geol-
ogy and Physical Geography of Brazil," p. 385. "So explorations, whatever, had
been carried on inland, or even along the navigable parts of the streams. The facts
brought out in regard to the Sci'gipc-Alagoas region by Hartt are substantially as
follows :

First. Reporting the following exposures : Estancia red sandstones, and the
limestones of Sao Gongalo, Sapucary, Maroim, Villa Nova, Penedo, Morro de Chaves.

Second. Reference of the Maroim beds to the upper cretaceous, it is presumed,
upon the paleontologic evidence of Prof. Hyatt's descriptions of the cephalopods.
The desci'iptions of these fossils is the only work known to have been done upon the
fossils of the region up to that time. Prof. H^^att in his paper expressed no opinion
concei-ning the age of the beds, but what appear to be Hartt's field labels are given
as " from the cretaceous of Maroim." Such was our knowledge of the geology of
the region here treated of when the writer visited it as a member of the Commissuo
Oeologica do Brazil in 1876.

TFIE GEOLOOIC IMPORTANCE OF THE SERGIPE-ALAGoAS REGION.

Much of the l^razilian coast is very old geologically, no sedimentary beds to in-
dicate the changes through which this part of the continent has passed intervening
between the ocean and the granites and gneisses which are referred to the arehiean,
Along a considerable jjortion of the north-eastern coast, especially from Cape Sto.
Agostinho to Parahyba do Norte, soft sedimentary beds, provisionally referred to the
tertiary, are exposed here and there in abrupt blufls which, are being cut away by the
ocean. These rocks, however, although well exposed, have as yet yielded no fossils,

OF THE SKRGIPE-ALAGOAS BASIX OF BRAZIL. 373

and du not, of themselves, throw much light upon the changes through which the
coast has passed. Xowhere along the Biazilian coast from the frontier of Uruguay
to Cape North are any fossiliferous paleozoic beds exposed, and there are but few
places in which a section from the ocean to the archa\in rocks would pass llirough
anything more than recent deposits and the horizontal tertiary beds referred to above.
Til no place along the wliole coast, however, can a wider and deeper section
be fouml, or one in which the rocks aftbrd a more complete history of the
changes through which this part of the continent has passed from paleozoic times
up to the present than the basin cut by the Kios 8ao Francisco and Sergiiie, and
which lies within the two provinces of Sergipe and AlagOas. The importance of this
region is due to

1. The representation of a geologic range unusual in Brazil.

2. The rich fossiliferous nature of many of the beds.

3. The accessibility of good exposure across the entire section.

Doubtless one of the chief reasons that these two provinces are not better known
geologically, is the fact that they are not on the principal line of travel between Eu-
roi)e and South America, and are, therefore, more or less inaccessible, and if the geo-
logical importance of the region has not been recognized, it must be attributed to the
fact that hitherto nothing, or next to nothing, has been known of it.

With the exception of a canoe voyage by Gardner along the Rio Sao Francisco
to Piranhas in 1837, and a very brief visit to Maroim by Prof. Hartt in the year
I860, none of the writers upon the geology of Brazil, prior to the work done by the
Imperial Geological Sui'vcy, ever visited the Sergipe- Alagoas region, and even those
few earlier writers have but little to say of the geology of this section of the country.

I am of the opinion tlial the kcv to future successful ofcoloofic work in Brazil
lies in the careful study and comprehension of some such typical region as that com-
prised in the [)rovinces of Sergipe and Alagoas. The Bahia basin is an interesting
one, and, as Dr. White has pointed out in his recentl}' published "Contributions to
the Paleontology of Brazil," faunally more interesting perhaps than that of Sergipe-
Alagoas, but the Bahia basin being of lacustrine origin is sui f/eneris, as far as the
raesozoic geology of Brazil is understood.* The Sergipe- Alagoas beds on the other
hand are marine, the rocks above the archrean vary lithologically ami faunally, and
arc more or less exposed across the whole width of the region from the ocean to tlu-ir
inland margins at the base of the Serra d'ltabaiana, while at this lattci- locality, the
lowest of the sedimentary beds are well exposed whcic the seria is cut through

• PIssis, in his memoir publislied l)y llie Frcncli Academy, p. 402, says llial llio tertiary beds of Uic interior of
Bra7.ii, and lying between Uic coast range and llio Serrn de Jfantiqueira, arc lacustrine.

374 THE CRETACEOUS AND TERTIARY GEOLOGY

by the Rio Sergipe, and their relations to each other and to the underlying archaean
rocks are distinctly shown.

Aside from its purely geologic interest and importance, there is, perhaps, no part
of the coast north of Bahia so fertile, the soil being of the black quality known here
as massape and deiived from the decomposition of certain limestones and of organic
matter. Whei-evcr the tertiary beds have been eroded away exposing the underlying
cretaceous rocks, the decaying of these calcareous beds has produced a soil, which, in
spite of indifferent cultivation, yields abundant crops of sugar-cane, cotton, mandioca,
etc. Although the cane-fields upon some of the cretaceous soils have not been re-
planted for thirty years, they still produce abundantly.

TOPOGRAPHIC FEATUKES AND VEGETATION.

The topography of the Sergipe- Alaguas region may be divided into the following
five types :

I. The low, flat lands of quaternary and recent origin, immediately bordering
the coast and streams.

II. The tertiary plateaux.

III. The cretaceous hills. -

lY. The serras, or paleozoic region.
Y. The ti-ans-serra, oi- archseau region.

I. The low, flat lanch. — In some places these flat lands are made up of sand bar-
rens, or they are covered with sand dunes ; in others they are low grounds flooded at
high tide, and usually form what ai-e known as mangues, or mangrove swamps, bor-
dering the estuaries. This type, or rather the mangi'ove part of it, is not confined to
the immediate neighborhood of the ocean, but extends inland as far as the tides are
felt, sometimes for many miles, and in general outline has a dendritic form. On the
Cotinguiba it reaches the town of Larangeiras, on the Sergipe tliat of Riachuello,
and on the Maroim branch it reaches the city of that name.

The vegetation of this belt is a characteristic one. The mangues are in no par-
ticular different from those to be found along the whole Brazilian coast, except, pei-
haps, in the blackness of the mud where it is derived from the decomposition of the
calcareous cretaceous rocks and organic matter. In places these mangues broaden
out until they are from two to five miles wide, and in other places their outer margins
approach the streams where the latter are encroached upon b}^ the hills. Thus they
are at once the characteristic feature of a well-defined topographic type, an important
geologic agent, and an interesting form of vegetation. When the mangrove plants

OF TUE SERGIPE-ALAGOAS BASEST OF BRAZIL. 375

{Ziaguncularia racemosa Gaoit. and Avicenma toinentosa Lin.) have reached matu-
rity, the mangues (swamps) are very nearly, if not quite, impenetrable, and tlic lapid-
ity witli wliicli old vegetation decays and young vegetation seizes uiion new and
reclaimed lands in the tropics, make them a moie powerful agent in the accumulation
of sediment and organic matter than aie the mangrove swamps of Florida.

In its predominant features the belt of sandy lands and dunes does not diil'er in
any marked degree from that of any sandy coast. The sands drift with the winds,
and the conHgui-ation seen at one time is entirely obliterated a few weeks or a month
later.

Kot infrequently these dunes are drivrn into the edges of the thick forests which
grow upon the adjacent topographic l)ell, but beyond this mai'gin tliey cannot pene-
trate far. The effect of the l)lown sand upon the vegetation in sueli cases is ver^^
maiked. The flying grains soon bruise and kill the buds and tender leaves and ulti-
mately a peculiarly rounded, lobed and closely cropped appearance is given to the
windward margins of the forests. Although these dunes do not assume the propor-
tions in this part of the empire that they do in some of the more northern provinces,*
the entire coast of Sergipe and a large portion of that of Alaguas ai-e covei-ed by a
sandy belt which varies in width from five hundred feet to two or three miles.

In some places the dunes predominate, the shifting sands often encroaching upon
and burying mangrove swamps and the xinderbrush of adjacent forests, whilr in
others they form broad, flat glades with a sparse and characteristic flora of cajus,
cacti, etc. From the Barra de Japaratuba to tlie mouth of the Kio Cotinguiba the
unbroken line of dunes thrown up Ijy the prevailing east winds has here am-sted tlie
drainage and turned it westward through the Kio Pomonga which Hows into the Co-
tinguiba just north of Araeaju.f To the south of the Cotinguiba these dunes often
assume extraordinary proportions.:!;

The vegetation of the sandy belt is necessarily sparse at the best. Wherever
the sand is constantly shifting, as it is in the newer portions of the belt, vegetation
has no opportunity to take root. In those portions, however, in which, for any reason,
the sands assume more or less stability, certain drought-resisting forms of vegeta-

* There lias been some speculation regarding the origin of Uiese viisl iicciimnliitions of sand along Uic IJra/.ilian
coast, anil especially about Cape St. Rorjue, M. ile Jonnes holding that they are brought lo these shores by the eqa-
torial current (see Ceaia, by Pompco de Souza, p. Ki). while Dr. Capanenia holds that they come from the Serra do
Araripe. Whatever may be the origin of the sands north of Parahyba, south of that point they are derived dirccllj'
from the rapid cutting away by the ocean of the soft tertiary beds.

fSce also Relatorio da Commissao Hydraulica do Rio Siio Francisco, by W. Milnor Roberts, Rio de .Janeiro, 1880,
p.. I.

t See Geology and Physical Geography of Brazil, by Cliaa. F. llarlt. p. 380.

376 THE CRETACEOUS AND TERTIARY GEOLOGY

tion quickly take root and flourish upon it. The cacti are naturally in this list, and
I have found one species of fern here, while the caju tree, one of the blessings of
the country, seems to reach its highest development in this parched and arid soil.

II. Tlte tevtianj j^lciteaux. — This and the succeeding types can best be under-
stood by anticipating somewhat the succession of geologic events in this region. The
cretaceous formation was here overlain by not less than two hundred feet of hori-
zontal, particolored beds of soft sandstones and clay. Since the emergence of the
region from the ocean, the overlying tertiary beds have been extensively eroded.
Where the tertiary beds have not been entirely washed away, flat-topped, angular-
shouldered hills still cover the cretaceous beds, and form the topographic type
under consideration. These plateaux are not confined to any particular belt, but
may be found anywhere between the ocean and the serras. The tertiary soils are,
as a rule, very sterile, their sterility being due partly and directly to the character
of the soil and partly to its tendency to wash, and its inability to receive and
retain sufficient moisture to support vegetation. The natural growth of vege-
tation upon this soil is strikingly poor, and this region of plateaux is
generally covered with a sparse and stunted growth of forest and sedge, while
the land is extremely barren, and unfit for cultivation. It is also seriously affected
by droughts. Types of this topography occur between Maroim and Sao Christovao.

III. The cretaceous hills. — From what has been said it may be anticipated that
the region of the third type owes its existence to the erosion of the tertiary beds, and
the consequent uncovering of the underlying cretaceous. While this is true in the
main, there are certain elevations in this basin which I am disposed to regard as never
having been covered by the tertiary deposits, certaiiffy not to any considerable thick-
ness. In making a voyage between Aracaju and the mouth of the Rio Sao Francisco,
one may see a range of cretaceous hills lying south of the river and well inland,
though disconnected with the higher lands of the serras. This range of hills stands
at a higher elevation than any of the tertiary hills on either side of it, and for reasons
given further on, it is not believed that the erosion from the summits of some of the
tertiary liills has been very extensive.

The region on the whole is a broken one, the hills being more or less rounded as
a rule, though a lew ol tliem have abrupt faces toward the north and west. Although
the topography of this region is a characteristic one, its value for the purposes of
geologic reconnoissance is somewhat impaired by the presence of the overlying ter-
tiary beds in a great many localities. This type characterizes the region between
Aracajii and Itabaiuna and between Larangeiras and the seri'as. The vegetation is
large and abundant, and within this belt one may often find excellent examples of the

OF THE ^EU(in'E-ALA(;oAS UASI^' OF iji;a/il. 377

idi'al tropical forests having an iimlorgrowtli \\ell-ni<i,h iin])('netrable and trees whose
enurnious trunks reath heights almost incredible.

IV. Tlie semis, or the ixdeo-zoic region. — This type lies to the north and west of
tin- mesozoic region, and forms a well-dt'lincd, natuiai boundary Ixtween it and the
arclwean region which lies farther inland. In the province of Alagoas this border is
formed by the Scrra de Maraba, which, in its continuation to the south-west, is called
the Sena d'ltabaiana, anil still farther to the south Cajahyba. This entire range is
the monoclinal remains of the eroded landward margin of the beds here exposed. Tlic
rocks dip to the east and soutli-east at an angle of from fifteen to twenty degrees,
and underlie the cretaceous and other beds between the serra and the ocean. For
the most part these serras are covered with forests nearly to their summits.

\. The irans-serra (w archcean region. — The topography of this region is an ex-
ceedingly varied one ; now hilly, now mountainous, and now spread out in gently
undulating plains. In some places it is notoriously sterile and is covered, for the most
part, with a sparse growth of stunted timber, while in otheis it is more fertile, and
produces a more vigorous forest. But little time was devoted to the geology of the
archsean region.

THE SERGIl'E-ALAGOAS BASIN.

It will be seen in the resume of what we know of the mesozoic geology of Brazil
that too little is known of the exact limits of the cretaceous basins along the noilli-
eastern coast of South America to permit a single one of them being clearly and
satisfactorily outlined. More work has been done on the Bahia basin than upon any
other one of the nuinl)cr, but of even this the limits arc not known. For the better
understanding of the region, it is deemed best, however, to give such facts as are
availabk' in rouglily outlining the Sergipe-Alagoas basin, before giving the details of
local geology.

The Bahia beds being of lacustrine oi-igin, while those to the north are of marine
origin, it may safely be assumed that these two basins were independent of each
other at the time of the deposition of their beds. The most noith-uasterly point at
which the Bahia cretaceous is known to occur is neai' Cali'i in the province of Bahia,
and the most southerly point at which the marine cretaceous beds of the Sergipe-Ala-
goas basin have been observed, is at and about Estancia, in the province of Sergipe.
The southern margin oi' this basin nin>l, therefore, be to tin; mhiIIi ol' this place. The
rapids in the Rio Heal, known as Passagem das Tedras, suggests the possibility of
the red sandstones, which make the cataracts at Estancia, crossing the liio Heal at
Passagem.

A. r. s. — VOL. .vvi. 2v.

378 THE ( ]lETACEOUS AND TEKTIARY GEOLO(;Y

To the north of Estuucia uo exposures arc known until one approaches the Pedra
da Cajahjba Avbich forms a part of the Itabaiana range of mountains.* From this
part of the piovinec to the Rio Sao Francisco near Propria the mesozoic basin is
bordei-ed on the hindwards side by the eastward extension of this same range of
mountains. In tlie province of Ahigoas, the Serra de Maiaba forms the north-west
boundaiy, and by its ti-cnd leads one to suppose that this north-west margin contin-
ues for some distance in a line parallel to the coast.

Just where this basin ends to the noith-east, and indeed whether it ends at all,
either within the borders of Alagoas, or of Pernambuco even, is not known. The
oreoloo-y along: the coast between Maceio and Rio Formozo has never been examined,
and it is possible that this basin comes to an end between these two points. The
writer feels warranted in the opinion — though it is only an opinion — that Porto Calvo
is in the Sergipe-Alagoas basin, and it is possible that its beds are exposed at Rio
Formozo. An examination of the relations of the arch.'can rocks and what was at the
time supposed to be the tertiary beds at and near the mouth of the Rio Formozo in
the province of Pei-nambuco, does not definitely settle the question as to whether or
not the cretaceous beds continue to the north of this point. The beds seen at Rio
Formozo lie unconformably against the crystalline rocks, but it is not clear whether
the beds supposed, at that time, to belong to the tertiary do really belong to that
period. They are, in general appearance, certainly unlike any tertiary beds seen else-
where in Brazil, being made up of alternate beds of conglomerate, cla}^, chalk and
chalk with flint nodules.

If then the cretaceous beds are continued fiom Alagoas and extend to the north
of Rio Formozo, it is possible that they underlie the tertiary' about Cape Sto. Agos-
tinho, and are continued on the landward side of that cape, underlie Pernambuco,
crop out at Olinda, Maria Fariuha, Iguarassii, the island of Itamaricii, and may even
connect with the mesozoic beds exposed at Paiahyba do J^orte and further north.

LOCAL DETAILS.

The present discussion of the Sergipe-Alagoas basin must be based upon facts
gathered for the most pait in the vicinity of Maroim, the Serra d'ltabaiana, along the
Rio Sao Francisco and the Serra de Maruba.

* The beds of lliis range whicli I have referred lo the paleozoic [irobuljly underlie the erelaceous beds of all this
part of the coast. In ii leiler to lue, iu answer to one making inquiry regarding the nature of the region traversed by
liim from Bahia lo Sergipe, 5Ir. Derby writes: " la the trip across Biiliia and Sergipe, I found a series of shales and
sandstones in the region between the Rio Real and Itabaiana which is most probably the same seen in the mountains.
So far as I could see, there is nothing belweeu thai and the gneiss."

OF THE SEKGIPE-ALAaoAS BASIN OF BRAZIL. 379

These and the other loealities mentioned below are the only ones examined that
are judged to be of sufficient importance, or at which sufticiently valual)le observa-
tions were made, to be worthy of mention here. Hundreds of places were visited in
this region at whieli poor exposures of cretaceous rocks were found, but tliey could
thiuw IK) additional light upon the geologic structure of the region witliout the use
of more trustworthy methods of work than it was possible to employ at the time.

Inasmuch as the tertiary geology of this region is the subject of the second pan
of the pi"esent paper, the beds of this series are not discussed in this place.

In view of its general impoitanee and the light it throws upon the structure of
the entire region the details of the Itabaiana exposures will be taken up first.

Itahaidna. — The best section seen of the rocks underlying the meso/oic beds ai'c
exposed where the Rio Sergipe cuts through the range of mountains of which Itabai-
ana forms a part. Here the crystalline rocks are exposed on the inland side of the
range, and in the gap cut by the I'iver, the unconformable contact between these and
the sedimentary l)eds, which are presumed to be paleozoic, is plainly seen. The
conglomerates, false bedded sandstones, and shales which make up the great body
of the section, have been so metamorphosed that the [)rospect for finding fossils in
them is not very encouraging.

The sandstones are alyiost all changed to exceedingly liaid, glassy quartzites, which
by much jointing have broken into smooth-sided rhomboidal blocks. Following
down the stream one ascends geologically, and finds these quartzites and conglomer-
ates the principal rocks exposed, while those of shale, in which one most hopefully
looks for fossils, are but poorly exposed or entirely covered with talus from the over-
hanging ledges. Th»!se beds have a j)retty uniform dip of 15-20° to the south-cast.
Along the escarpment of the north face of the mountain range the uptiu'ned edges
of the up|)ermost beds are partly exposed, while all the lower ones arc covered with
talus which stands at as high an angle as possible. In this talus large fragments of
shale, such as was not seen in situ were examined for fossils, but without success.

The brief time spent in examining the rocks of this .section lor paleontologic
evidence of their age was very insufficient, and it is altogether probable that careful
detailed examination of these beds will yet furnish the desired evidence.

From the top of Itabaiana the struct me of this range of mountains and its geolo-
gic relations to the regions on either side of it are quite evident. The Itabaiana peak
is at the base of a curve in the ranjre which contimies to the right, as one faces the
ocean, towards the west and south-west, while lo the left it extends to the north and
north-west. About fifteen miles to the north the range turns castwaril and is next

380

THE CEETACEOUS AND TERTIAKY GEOIiOOY

seen promineiuly on the Alagoas side of the Kio Sjio Fiancisco, where it is known as
the Serra de Maiuba. To the south-west diflferent portions of the ranges are known
by diflferent names, the section next to Itabaiana being called the Serra Comprida, and
that next the Cajahyba. It is evident at a glance that this range of hills bears the
same relation throughout to the general Sergipe- Alagoas basin as the Itabaiana beds
bear to those lying immediately between them and the ocean.*

The section across the region given below shows the general relations of the

various beds.

The following sketch looking north-east from the top of Itabaiana, shows the

Looking north-east from Itabaiana.

continuation of the range in a line of detached peaks, the rocks of which all dip
oceanward beneath the mesozoic beds. Fig. B. is a sketch from near the south-ea.st

Pig. B. Serra Comprida from Itabaiana.

end of Itabaiana peak, looking toward the south-east. The beds of the Serra Com-
prida are seen from the inland side, somewhat, and appear to be faulted.

The Itahaiana Section.

The accompanying section across the Sergii)e-Alag6as basin is represented as
being di-awn from the arcluean region north of the Serra, through Itabaiana, Maroim,

*See note by Mr. Derby, on p. 378.

OF THE SER«4IPE-ALAG0AS BASIN OF BRAZIL. 381

and Sto. Ainaro to the sea. No attempt is made to repieseiit ivlative thicknesses
accurately, for no measurements have been made that will render this possible.

Section across the Sergipe-Alagoas basin.

Beginning at the base of the section, M represents the crystalline rocks of the
arclmean, here composed of beautiful greenish gneisses containing amethysts and yel-
low mica. At the point of contact the surface is uneven but hard and rounded as if
water worn, and the conglomerates of the lowest of the paleozoic beds lie uncon-
formably upon it. The gneisses at this place dip to the north-west though much
crumpled in places. The same locks when exposed away from the line of contact ai*e
soft and deeply decayed.

The lowest bed of K is a coarse and very hard conglomerate made up lai-gely of
fragments from the underlying gneiss. The next bed above is of coarse sand and
pebbles, false iiedded, followed by sandstone of varying degrees of coarseness. The
pebbles in this bed are of all sizes np to that of one's fist, and vary in color from the
white of milky quartz to the green of the underlying gneiss from which the latter are
ajjparently derived. Xext follow very hard sandstones with ripple marks, then fine,
fine-grained sandstone. This is succeeded by tine-grained micaceous slaty rock
with ripple marks, then very hard conglomerate followed by solid fine-grained slate,
and coarse sandstone with micaceous bands.

These beds are all exceedingly hard, the sandstones being usually in the form of
jointed and strongly false-bedded glassy quartzites. They have a uniform dip of from
lo"" to 20- to the south-east.

Of the thickness or importance of the rocks between the beds ju.st mentioned and
the next observed, nothing is known. On the sides of the liills immediately east of
Itabaiana a .series of slates, .shales and sandstones were seen, but it was too imperfectly
exposed to admit of a complete section being made, or of connecting such a section
directly with that in the Rio Sergipc gap. On the part of the mountain east of the
river the rocks dip south and south-east. The next overlying beds seen arc limestones
exposed on the stream between Itabaidna and Seira Comprida. Where they were
examined they are considerably distiu'bed, very compact, and traversed by .small veins
of white quartz. In general appearance these rocks resemble the mountain limestones
of England, but no fossils were found in them.

382 THE CEETACEOUS AND TERTIAUY GEOLOGY

It is presumed that these limestones are included in the paleozoic beds which
extend still higher. There is another break in the section here caused by the impos-
sibility of observing the sequence of the overlying beds.

The next rocks seen, and of which the stratigraphic relations are known, are the
mesozoic sandstones found in the hills at Engenhos Araga and Pamonha, and which
appear to dip gently to the north-west, that is, in the direction of the serra.* This
dip necessitates at least one syncline between this place and the mountains. Abov^e
these follow various beds of sandstone and limestone which are described more in
detail elsewhere, all of which aj^pear to belong to the cretaceous, with certain excep-
tional facies referred to in speaking of the age and correlation of the mesozoic Brazil-
ian beds.

The cretaceous beds are somewhat disturbed, and some of them slightly meta-
morphosed. They are richly fossiliferous in places, and have furnished the interesting
series of fossils described by Dr. White in his " Contributions to the Paleontology
of Brazil.''

The horizontal tertiary beds (B) which overlie these do not end along a well-de-
fined line as one might infer from the section, but seem to have covered at one time
almost all the mesozoic rocks, and to have been largely removed by denudation, leav-
ing shoulders here and there, some of them but a few hundred feet and some of them
miles in width. These beds are horizontal as far as the unaided eye can determine,
non-fossiliferous, and are made up of particolored alternating strata of clays and sands,
with here and there calcareous bands and concretions.

The last beds are those of quaternary and recent times which lie against the
tertiary, cretaceous, paleozoic or archsean as the case may be.

Recapitulation. — Explanation of the section :

M. Archasan, (?) schists.

K. Conglomerates, sandstones and shales of the Itabaiana series, partially meta-
morphosed.

II. Not seen.

G, Limestones and shales.

F. Not seen.

E. Sandstones of Pamonha and Araga.

D. Oolitic limestones and sandstones.

C. Chalky beds of Sapucary, Andorinhas, etc.

B. Horizontal tertiary sandstones and clays.

A. Quaternary and recent.

* The fossils from these beds arc referred to in Dr. White's work as coming from "Pamona," which should read
Pamonha.

OF THE SKRfilPE-ALAGoAS BASIN OF BKAZIL. IJSo

EXl'OSUKES ABOUT MAKOIM.

(^luinies have been opened in many plaeos in and about Maioim from which
stone has been obtained lor building walls and houses and iov paving the streets.
One of these quarries on the Avestcrn side of the town is known as the 2)edreira dc
Gambarobe. The rock at this quairy is a brownish giay limestone, more or less
oolitic, and contains some fossils, though the shells are usually broken and dilMcult
of extraction. Lamellibranchs, cephalopods, decapod crustaceans, echinoderms, and
occasionally very lai'ge gasteropods are found b^' the (piurrymcn. Many Ijits of fossil
wood are also found here, but all in a cluured condition.

On the eastern outskirts of the town, and lying more or less in it, at a place
known as Aroeiia are several old, abandoned quarries in calcareous sandstones.
These beds contain many fossils, but the rocks are too hard to allow them to be taken
out in good condition. The dip here is generally toward the east at a low angle.
The fossils described by Dr. White from the Eiacho de Aroeira are all from a small
exposure in the bed of a wet-weather stream to the north-west of the quarries. At
this place the rocks are more sandy iliaii at the quarries rcfen-ed to, and sometimes
are hard as quaitzites. They lie directly beneath the beds exposed at the quarries of
Aroeira and dij) to the S. 80° E., at an angle of six degrees. In the lowest part of
the bed are fragments of the softer yellow lock which lies at a lower geologic level,
forming, with the sand, a kind of conglomerate.

Ziustro.

The richest locality for fossils found in the vicinity of Maroim is one known as
the Laslro, about two miles down thu river from the town, on the east side of the
stream, and just south of the Engenho da Praia. From this engenho the exposures
contiiuie for more than three miles along the hills that border the stream on the east.

The fossils described by Dr. "White from this locality come from the border of
the mangue along the first three-quarters of a mile below the engenho. In this dis-
tance the rocks furnishing the Ibssils are mostly soft, decaying, cream colored to
brown, oolitic limestones, which, by disintegration, have left the fossils free. In
places the small i-ock fiagments aic so full of echinoderms that they can be picked
out like walnuts from their hulls. In some cases the material of the.se fossils is
changed to i)ure calcium carbonate, while in others the more compact beds of lime-
stone have the fossils silicified so that they can be perfectly removed liy the use of
acid to dissolve the limestone.

The weathering out of the fossils at this place is hastened by the tides which

384 THE CRETACEOUS AISTD TERTIARY GEOLOGY

alternately cover and expose them to the air and sunshine. These same beds are
exposed also on the opposite side of the river at a place called Jaque, where the
stone was formerly quarried at low tide. Here, however, the rocks are harder than
those at the Lastro exposure, the fossil fauna being the same.

The rocks at Jaque are oolitic, of a light-brown color near the surface, but where
they have been blasted, or their interior pails arc unaffected by weathering, they have
a bluish-gray color.

Rocks similar to those found at the Lastro and Jaque occur from two to three
miles further down on the west side of the stream. Here too tJie black flints which
seem to have weathered from some limestone beds are widely scattered. The beds
from which the flints have been dei'ived overlie the oolitic rocks exposed at Lastro
and Jaque. On the east side of the stream the rocks overlying the oolitic beds are
soft limestones such as are exposed about Porto da Rede. On the west side the
soft limestone beds are more or less flexed.

The fossils most abundant at the Lastro are conchifers, cephalopods and echi-
noids. There were found here sixteen siiecies of conchifers, of which number ten are
new to science; nine species of cephalopods, six of which are new; ten species of
echinoids, nine of which are new. Gastei'opods are remarkably scarce, but two spe-
cies having been found here, both of which are new.

Pedreira de Jono Pereira.'^'

This quarry is about two miles east of Maroim. The rock is a soft, ci'eam-colored
to yellow, oolitic limestone, affording many imperfect specimens of cephalopods.
The rock is so soft that it is quairied in blocks by cutting trenches in it with picks.
The fossils are bivalves and cephalopods. Three species of the former and three of
the latter have been found here, all of Avhich are new to science. The beds have a
southerly dip.

Oarajdu.

This locality is immediately north of the town of Maroim, and is interesting as
containing the i-ocks from which one of the conchifers described by Dr. White as of
Jurassic aspect (Aucella hraziliensis) was obtained. Most ol' the fossils obtained
here ai-e Ci^phalopods {Ammonites liopTcinsi) and were broken fi-oni compact limestone
blocks whei-e they had l)een separated by disintegration from the main beds, which
crop out here and there about the fields. These beds overlie those of Aroeira, Las-
tro, Ja(pie, Gambarobe, etc., all of which arc oolitic.

*Thi8 locality is ■tttongly given in V>t. AVliilc'.s I'aleoiilology us .]o;lo Forroira in some places, while in others it
ha3 the correct spelling.

OF THE SERGirE-ALACJoAS BASIN OF HI{A/.[L. 385

Jacuruna.

At Jacuruna, a slioit distance iiorth-cast of Maroiin along the estrada real lead-
ing to the village of Rosario, is an exposure of oolitic limestones containing a few
fossils. To the west of the Ponte de Sabao bridge, is a hill having outcrops of simi-
lar rock on either side. An exposui'c of limestone passes east and west through the
village of Kosario. In one place where this stone is quarried, a bed thirty-Hve feet
in thickness is exposed. The rock is oolitic and in lithologic characters strongly
resembles that of the Gambarobe quarry in Maroim. The uppermost ten feet forms
a massive bed. These beds dip at an anglo of twenty degrees S. 45° E. (direction
estimated). Between this point and Campo Redondo the country is low, these cre-
taceons rocks and their hills forming the southern boundary of the valley, while a
range of tertiary hills bounds it on the north. Soft, cream-colored, cretaceous lime-
stones underlie these tertiary hills as was shown in the i)its excavated, north of En-
genho Cami)o Kedondo in a search for coal or gold. From one of these pits, ten feet
deep, a few fragments of fossils were obtained, one of which, Camptonectes idacltus
"White, is desci'ibcd in Dr. White's "Paleontology of Brazil.''

Coqueiro.

Most of the material collected at Coqueiro came from a single locality, a small
exposure in the estrada real within a few hundred yards of the engenho of this name.
The rocks here are sandstones of very uneven texture, being in some places hard, in
others soft, in some places fine grained, in others coarse and porous, while in otheis
still they contain a great many quartz and other pelibles. Among these pebbles arc
some of black quartzite resembling very strongly the dark quartzites found in situ in
the serra d'llabaiana. Owing to local variations in the character of these sandstones,
some portions of the beds contain no fossils, all of them, if any ever existed in such
places, having been lemoved by pei'colating waters. The most abundant fossils are
conchifers, Triyonia suhcremdatu being esjjecially plentiful, and these occni- for the
most part as moulds or casts, the shells having been entirely dissolved out. In some
of the finer-grained portions of the beds many impressions of small and delicate lamel-
libranchs are Inmnl, but llic rock is so {Viable that they can be exti'acted only with
great diflicully. The scarcity of gastero[)ods in tliese beds is somewhat striking.

To the north of the engenho is a small (piarry in which the rocks arc very hard
yellowish sandstones, from which the fossils cannot be successfully removed. East
of the river (Sergifje), at a ])lace called Pocinhos, about seventN'-five feet of sand-
stone is exposed. This exposure also shows a wide range of material in its composi-

A. p. S. — VOL. XVI. 2w.

386

THE CKETACEOUS AND TERTIARY GEOLOGY

tion, being in places very fine, in others containing a few ^Jcbbles, while in others it is
a decided pudding stone containing cobbles half the size of one's head. Fossils are
ver}' scarce here, only a few bivalves being found in the lowest beds. The fact that
the collections made here aflbrd several fossils of Jurassic as^jcct renders the locality
one of sijecial interest.

The beds are identical with those exposed at Porto dos Barcas, Trapiche Maior,
Traj)iche das Pedras Velho and Trapiche das Pedras Novo, though they vary con-
siderably between these various localities in lithologic characters.

EXPOSUKES ALONG THE KIO 8ERGIPE.

Aracaju

Sketch Map

OF fl PORTION
OT THE

PROVINCE""^ SeRGIPE-

■33y ira7in, CjBranner.

Many of the paving blocks and stone steps used in the village of lliachuello
(also called Piutos) ai-e filled with small gasteropod shells and other fossils, and a

OF THE SERGTPE-ALAcAaS HASIN OF BRAZIL. 387

small collection ot" the fossils of the neighborhood was made at one time by J)r. Dio-
nizio Eleutcrio do Menescs, the proprietor of Engenho Moleque near the village. At
this eng-enho is an exposure of soft sandstone, but the fossils in it aie nt)t well pre-
served or readily e.xtractcd. The rocks in the \ icinity of Riaehuello were not exam-
ined thoroughly, and im doiil)! more interesting localities than an}' found remain to
be explored here. Between the village and the engenho Jesus Maria Jose are several
localities which yield good fossils. At the last-named place an exposure in the public
road furnishes an abundance of them. Another locality is the Engenho Sao Jose de
Vieira, and between it and Sao Felix are the largest and best preserved gastero[)ods
found anywhere in the Sei-gipe-Alagoas basin, many of them lying weathered in the
lields. A few of these were collected, but they seem to have been lost, for they were
not included in the collection sent Dr. White with the other mesozoic material from
Brazil to be described. At Sao Felix is the only exposure between the Sito Jose de
Vieira locality and KiachuoUo.

At Trapiehc Santa Maria in the outskirts of this village and on the river l)ank, a
soft, non-fossiliferous sandstone is exposed, having a low dip to the north 15^ east.
This rock, in general appearance, very strongly resembles that exposed at Araga, and
Pamonha. At Trapiche das Pedras many large gastcropods were found in a calcare-
ous sandstone on the east side of the river.

About two hundred yards up the river from Trapiche Maior is a ledge of soft
sandstone exposed at the margin of the stream near a tile flictory, and containing
many fossils, and especially bivalves. At Porto dos Barcos an exposure at the mar-
gin of the river, and which is uncovered at low tide, dips N. 20° E. The rocks at
all the above localities dip beneath the oolitic limestones of the Urubu and Iiubiia
blufls.

I liiid in my field notes, written upon the si)ot, a statement to this effect: That
"the Coqueiro beds are the same as those of Trapiche ^faior, Porto dos Barcos, and
Ti-apiche das Pedras." The importance of these facts will be made apparent in the
discussion of the Jurassic aspect of some of these beds. My notes u])on the detailed
geology of those localities (Porto dos Barcos, Trapiche das Pedras, and Tia|)iche
Maior) are meagre, the collecting at these places having been done principally by
Dr. Freitas.

Urubii and Imhira.

The locality known as Urubu is a bold clilV of oolitic limestone of grayish and
greenish gray color, from one hundred to two hundred feet high, having a X.-S.
trend, and dijiping appi-oximatdy '-M , S. KT E. Along the weatheretl face of this

388 THE CKETACEOUS AND TEltTIAKY GEOLOGY

bluft' many well-preserved specimens of EcJiinohrissus freltasil, AVhite, wei'e fonnd.
Below this point another and similar exposure of cream-colored to gray oolitic rocks,
about seventy-five feet high, is known as Imbira. The rocks here dip about 30°, S.
70° E. These oolitic limestones are Aveathered along the joints, and contain many
large caverns from whose roofs stalactites descend to their floors.

At Canna Brava white to gray oolitic rocks dip S. 45° W. at a low angle. Below
Canna Brava, at a locality on the west side of the river, known as Toque, are good
exposures of limestone. These rocks are somewhat metamorphosed in places, and
only one bed yields many fossils. The rock is firm and compact, and weathers in
jagged, irregular masses which, when struck with the hammer, ring like " clink-
stones," from which peculiarity the place doubtless takes the name " Toque." In a
few places caverns are developed in it. The beds here dip about 20°, S. 65° E.

About a thousand feet below Toque similar beds are exposed at Capoeira on both
sides of the river. On the west side these form a blutf about one hundred feet high.
Being veiy compact and partially metamorphosed, they furnish but few fossils, and
these come from a sing-le stratum and bear a strong resemblance to those found
at the Lastro locality near Maroim. The exposures on the left bank of the river dip
13°, IS". 70° W., while upon the right bank the dip appears to be S. 70° E., angle not
determined. Kocks similar to these are exposed again further down the river at sev-
eral places, the most impoi'tant one being known as the Pedra Branca. Here they
are exceedingly compact and partially metamorphosed, in places resembling marble,
while the fossils have been almost entirely obliterated, the ones found having a general
resemblance to those of the Lastro locality. The exposed upper surfaces of the rocks
weather in sharp, jagged points.

Half a mile from this point on the river, at the margin of the mangue, are found
i'ragments of a bed of limestone cropping out at the base of the hills, which fragments
contain a great many flint nodules.

Tlie only other exposure of importance on this stream, that is, on the Rio Sergipe
proper, and below the last-mentioned place, is at the limestone quarries of Andorinhas.
At several places soft, civam-colored flagstones are quai'ried both for ])aving hitones
and foi- making lime. Fn general appearance these limestones resemble those quai'ri(d
at Sapucary, though they aie not (juite so ])ure or so fine grained. These rocks
contain occasional imperfect im|)ressions of large cephaloi)ods and the remains of
decapod crustaceans. The dip here is generally toward the east, though the rocks
have some appeaiance of being flexed.

The next exposures down stream are those of the Sapucary quarries on the west

OP THE SKRGIPE-ALAGOAS BASIN OF BRAZIL. 'ASd

sidc.'^' The rock liorc is a soft, civam-coloied limestone, in huniuie IVoni one to live
ijiches thick, and coming out as flagstones. These flags are extensively used for
paving-stones in Aracaju. They dip to the east at an angle varying from 2-5'^ to
40°. But few fossils have been found in these rocks, and these have been the scales
of fishes. The locality is described also by Prof. Hartt in his "Geology and Physi-
cal Geography of Bi-azil," p. o83.

These Sapucary beds appeal- to be the highest of cretaceous age exposed in this
region, the next exposures to the east being coarse, ferruginous sandstones, which, in
all probability, belong to the tertiary. If, from Sapucary, one ascends the stream
toward Maroim, many cretaceous exposures may be found in the vicinity of Porto da
Rede. The rocks to the east of this village are limestones, both hard and soft, many
of them containing black flint nodules. They are exposed in many places near the
water's edge, along and in the margin of the mangues. The dip is appi-oxiraately
to the east at an angle varying from 25° to 35°, None of them form prominent fea-
tures in the topography of llu' neighborhood.

EXPOSUKES ABOUT LARANGEIRAS.

The most interesting place in the vicinity of Larangeiras is just outside the town,
along the road leading to ^Nlaroim. Here hundreds of large cephalopods (mostly
Ammonites (JJucJn'ceras) harttii, Hyatt) and echinoderms {Ecldnohrissus freitasii,
"White) lie weathered out in the road. Some of these fossils are badly bruised by the
wear of travel and their being knocked against one another, while others, more re-
cently weathered out, aie fairly well preserved. The material from this locality, and
belonging to the Commissao Gcologica was labeled "Bom Jesus'- after the name of
the engenho to which the lands belong, and this is the locality referred to in Dr.
White's descrii)tions. From this [)oint toward Maroim, at the Engenho I'edra
Branca, is a quarry of iuijiiiic limestone dipping southwest.

A half a mile west of Larangeiras is a place known as ihe Pedra Furada, or
pierced rock. A bed of limestone about twenty feet thick is here exposed in an
isolated bhiif made pi'ominent by lateral weathering. Litliologically this rock
strongly resembles that found at Capoeira and Toque on ilio Sergipe, and of
which bed it seems to be the southward extension. The strata dip about S. 45°
E. at a low angle.

♦ Prof. Ilartt gives the name of this place as Sapiicahy, and in Uic notes sent Dr. While liy Mr. Derby on llic
geology ot the province of Sergipe tlic name is ao given. Sapucaliy being the name of a Brazilian tree, it is qnile nat-
ural that this mistake sliouUl have occurred, and that it should be held as an intelligible word. In spite of this the
people at and about the place called it Sapucary, and Ihis, of course, detcrttilDCs its name.

390 THE CRETACEOUS AND TEUTIAKY GEOLOGY

111 the town of Larangeiras is a qiiari-y on the east side of the stream, the
rocks having- a general resemblance, lioth in lithologic appearance and in fossils,
to the harder portions of the Lastro and Jaqne exposures near Maroim.

Several outcrops of cretaceous rocks occur along the Cotinguiba, the sti'eam
upon which Larangeiras is situated. Above the mouth of the Rio Cajahyba which
flows into the Jacaresica from the west, there is a quarry -of white limestone at the
Caes da liha on the east bank of the river. A section of about twenty feet of this
rock is exposed here. At Oiteiro Galante, on the east side, the same rock shows
again in a less prominent exposure. N'either of these places was carefully examined
by the writer.

ESTAKCIA.

At Estancia the most prominent rock exposures are the red sandstone down
which runs the cataract of the Rio Piauhy at this place. In general appearance
these beds resemble, in a very striking mannei-, the triassic red sandstones of 'New
Jersey, United States, and also those at Penedo on the Rio Sao Francisco. No fos-
sils have been found in them.

Between the city and the port on the river the rocks exposed are dark-brown
sandstones varying to conglomerates containing pebbles of gneiss and clay ironstone
concretions of various sizes up to six inches in diameter. These rocks have a dip of
from two to five degrees to the north-west. I am inclined to think, however, that
this is an exposure of tertiary instead of the harder sandstones so well shown in the
river.

At Ribeira Velha are exposures of the tertiary. The locality of especial interest
in this neighborhood is a place known as Sao Gongalo, where a few cretaceous fossils
were found. The exposure, however, is a very insignificant and unsatisfactory one,
where the soft limestones have been dug out for making lime. In this exposure the
beds are gently arched, the most prominent dip being to the north-west (?). The
uppermost bed is a coarse white sandstone, next below comes a band of line, soft
limestone, then gray limestone of a chalky appearance, and at the base a bed of plas-
tic blue clay. The quarrymen spoke of having found j^^f-nellas (frying-pans) in the
rocks, which, from additional descriptions, were supposed to be large cephalopods.
Ko examples, however, were seen.

THE MARABA SERIES.*

^riiai the jSIai'aba beds bear the same relation to the mesozoic rocks in the i)ro-
vince of Alagoas as do those of Itabaiuna to the mesozoic rocks in Sergipe, there can

* The l-eira tie Mara1)n is said to l)e the liigliest point in tlie i>rovinc(^ of Alagoas.

OP THE SEHGU'K-ALACioAS BASIN 0¥ liUA/AL.

:m

scarcely be a doiiht. Looking norlli-uast lioin llic liio Sao Franci.sco, a shoil tlis-
tance below Piopiia one sees the Maraba as shown in the foUowinii- cut ; the inhmd

• ■"^frF?^)fi^^|s^f««vii*g^|^

-■¥'Si

Serra dc ^laiaba from near Propria, Kio Sao Francisco.

or north-west side of the hills presenting abrupt faces, the south-east slope having a
gentle dip toward the sea, suggesting that, if composed of sedimentary rocks, they
must di]) beneath the horizontal tertiary beds which cover the greater part of the
region in the immediate vicinity of the mountain.

An examination of the country between Penodo and the serra shows that this
impression is tlu' correct one. Leaving out of account the sandstones underlying the
eit}- of Penedo, in going from this place to Maraba, one traverses a wide belt of low,
flat, fertile country which borders the river, and which is more or less flooded by the
enchentes or "freshets" of the Kio Sao Francisco. Prom this low country he rises
gradually and almost imperceptibly upon the taboleiros or plateaux, which are com-
posed of the horizontally bedded clays and sandstones of the tertiary. These
beds continue high up the south-cast face of the Serra de Maruba and end uncon-

Section through the Serra de Maraba.

formably against it. They (the tertiary beds) are cut through by the ordinary pro-
cesses of subaerial erosion, and, if careful seai-ch could be made, it is pi-obablc liiat
exposures of the cretaceous, or at least of rocks higher than those seen on the sum-
mit and face of Maraba and lower than the tertiary could be found. At Igreja Nova,
a hamlet near the south-east base of the mountain, several loose fragments of com-
pact limestone wei'e found .strongly resembling that which occurs near the top of the
Itabaiana series, but the rock was not seen in place.

The serra itself, as seen in the few expo.sures on the top, is composed of li;;ht-colored
sandstones dipping at an angle of about 40° to the S. E.* These roeks are much jointed,
and in niiiny places changed to quartzites. The contact between the paleozoic and

• The alope of the liill where it was seen to correspond with that of the rocks is here given ns the dip. The rocks
are so metamorphosed that it is difficult or impossible, to obtain the di|) dirceUy.

392

THE CRETACEOUS AND TERTIARY GEOLOGY

archsean at the base of this serra was not seen, being covered by soil and the talus from
the abrupt northern face of the ridges. Not far north of this escarpment the gneisses,
such as are scan at the base of Itabaiana, crop out.

From the summit of the Maraba and bearing S. 25° W. is another peak of this
same range known as the Urubii, shown in the following cut, having the abrupt north-

Pico do Uiiibu from Jlaraba.
west face and the south-east dip characteristic of the Maraba. From Maraba, Propria
on the Rio Sao Francisco is visible in the distance, and to the left of that place, though
far beyond the river, appears a broad plateau with its higher face to the inland side and
sloping gradually oceanward, but dying out in the Hat country long before the ocean is
reached. This plateau was judged at the time and on the ground to be the continua-
tion of the cretaceous beds of Sergipe in the Alagoas direction.

Fig. F. Pico da Serra Grande from Maialja.

Bearing N. TO" E. the Pico da Serra Grande (Fig. F) is tlic next prominent peak in
this range, its beds also apparently dipping south-east beneath the tertiary. Farther
away a line of peaks form the north-easterly continuation of this range.

LITHOLOGIC CHARACTERS.

No exposures have ever been found of these SergipcAlagoas cretaceous rocks suffi-
ciently continuous to make it possible to determine with any degree of satisfaction
whether their lithological characters are constant or otherwise, when long distances or
wide areas are taken into' consideration.

Without going into tedious details I may say, however, that all the evidence in my

OF THE SERCaPE-ALAOnAS BASIN OF BUAZIL. 393

possession points in one direction, namely, that the lithologic characters are not only not
constant in these beds, but that they often change very radically withiii a thousand or
two thousand feet.

The oolitic beds, so prominent a feature of the Maroini region, do not appear as such
on the Rio Sao Francisco, where the cMitire Sergipe-Alagoas series is cut through by the
river. The red sandstones of Penedo do not apjjear as such about Maroim, while they
are a conspicuous feature of the geology in the region about Estancia. The shales of
Maceio are not seen elsewhere in the basin at any considerable distance from Maeeio, and
the arenaceous beds of Villa Nova are not known to exist in any other part of the basin.

The rocks vary in texture and character between the following kinds :

1. Coarse, compact sandstone of a bluish-gray color, modified by weathering to
brown, and light gray ; fossils preserved but difiicult of extraction. Example, Riacho
de Aroeira.

2. Coarse to fine friable sandstone of open or porous structure; fossils preserved
mostly as molds or casts. Example, Coqueiro..

3. Oolitic limestone, bluish and greenish, weathering to light gray, cream-colored
and brown. Examples, Urubii, Maroim, at the Garabarobe and Lastro quarries. The
brown and cream colors of the oolitic beds are apparently products of incipient decom-
position, the bluish and green tints predominating where these rocks are quarried to
considerable depths, the brown and cream colors appearing near the surface and where
the rocks are penetrated by joints or otherwise made accessible to meteorologic
agencies.*

4. Cream-colored, lead-colored and white, soft limestone. Examples, the quarries
of Sapucary and Andorinhas.

5. Limestone, more or less compact, without well-preserved fossils, and with flint
nodules. Example, Toque below Canna Brava.

•The following is a brief description of thin sections of specimens of these oolitic limestones examined under the
microscope :

The body of the rock is made up of irregularly rounded patches of calcite, many of which have the characteristic
oolitic structure. There are, besides occasional grains of quartz, a few fragments of grecnisli mica, and here and there
patches of brown liydr().xidc of iron. All these constituents are cemented togelher wilh tine, almost colorless, granular
calcite. The oolites are stained a dull brown or cream color, and stand out prominently in the cement. There is
nothing unusual in the oolites themselves. They are made up of concentric rings of calcite grains one outside the
other, the innermost one usually enveloping a small fragment of organic matter, such as a bit of shell.

Many examples occur among these oolites, however, in which the general mass is net made up entirely of these
concentric ring<i, the formation of these having ceased at a certain point, while beyond this the calcite has been added
principally upon one side of the already formed mass, and without any definite arrangement. In some cases two
partly-formed oolites are cemented by this amorphous calcite, the united mass retaining the usual irregularly-rounded
form In other cases still later formed concentric bands have enveloped the oolites thus united.

A. P. S. — VOL. xvY. 2x.

394 THE CltETACEOUS AND TERTIARY GEOLOGY

THE AGE AND CORRELATION OF THE MESOZOIC BRAZILIAN ROCKS.

Although the mesozoic beds of the region under discussion are here spoken of as
cretaceous, the writer is aware of certain conflicting evidence in regard to their age,
which should not be overlooked.

Prior to the study of the cretaceous fossils of Brazil by Dr. White, the age assigned
to the beds from which these fossils weie derived was more or less unsatisfactory.

Although undoubted paleozoic deposits occur in many places in Brazil, no fossilif-
erous strata have as yet been discovered immediately beneath the cretaceous, while the
overlying beds, referred to the tertiary, have never yielded any fossils, and have been so
referred solely on account of their relation to the strata underlying them. It is plain,
therefore, that the determination of the age of these rocks must be based entirely upon
internal evidence, a determination which, on account, of the incompleteness of the col-
lections or the want of study, has alwaj's been more or less unsatisfactory up to the
publication in 1SS8 of Dr. White's important '" Contributions to the Paleontology of
Brazil."

Prof. Hartt, in his " Geology and Physical Geography of Brazil," p. 385, says that
when, in 1869, the cephalopods collected by him at Maroim were shown to Prof. Alpheus
H}alt, he was at once struck by their Jurassic aspect, although in the description of
these fossils, which is published at length in Prof. Hartt's book (oj) cit., pp. S85-393),
nothing is .said upon this subject, while they are all said to have come from " the creta-
ceous beds of Maroim."

Prof. Hartt says {op. cit., p. 393) that the limestone about Maroim is undoubtedly
upper cretaceous, though he does not tell us upon what evidence this opinion is based.

The Brazilian cephalopods described by Prof. Alpheus Hyatt in Hartt's " Geology "
had been collected by Prof. Hartt from what he considered undoubted cretaceous rocks.
Some of these fossils, however, were of decided Jurassic affinities, and both their strati-
graphic position and peculiar state of preservation warranted Prof. Hyatt in endeavor-
ing to account for their occurrence at such a horizon. Prof Hartt felt confident that the
Sergipe rocks were cretaceous ; the fossils were evidently much worn, and had a decided
Jurassic aspect. Prof. Hyatt therefore suggests* that these cephalopods may have been
eroded from beds at a lower geologic horizon (the Jurassic) and transported to and depos-
ited in the cretaceous. The suggestion is a natural one for a person unacquainted with
the peculiar circumstances under which these fo.ssils occur, but it is only necessary to
say here that a knowledge of these circumstances does away with .the necessity of this
supposition. The fossils were taken from their original beds, and there is no reason for

*Proc. Boston Soc. Nut. Hist., Vol. XVII, p. 370.

OF THE SERGIPE-ALAGUAS BASIN OF BRAZIL. 395

supposing that they have been redeposited, as I shall show hereafter, and the case is
mentioned in this place only to call attention to what is either contlicting testimony
regarding the age of the SeYgipe rocks, or to a mingling in them of Jurassic and creta-
c9om faunas.

M. Emmanuel Liais, in speaking of the secondary deposits of Brazil, in his " Geol-
ogic du Bresil " (p. 197) says: "Ce puissant depot secondaire, tout parait I'indiquer
d'ailleurs, a dii se former pendant au moins une grande partie de Tepoque cretacee, et a
pent etre commence dcs Tcpoque jurassique, au moins dcs la periode oolithique. Pen de
perturbations auront eu lieu dans ces immenses regions pendant cette longue duree, et
par la s'explique comment les especes du commencement de la periode ont pu continuer
d'exister et se meler aux especes posterieures, de sorte que, suivant la tros judicieuse
remarque de Darwin, confirmee, comme nous I'avons vu, par I'union d'especes jurassiques
et cietacees dans les divers depots du Bresil, les deux epoques ne sent pas nettement
separees comme en Europe."

No direct reference, however, is here made to the mesozoic geology of the Sergipe-
Alagoas basin. The writer has not been able to locate this opinion of Mr. Darwin, and
does not know upon what evidence it is based, but from the limits of his (Darwin's) ob-
servations in Northeastern Brazil, it is presumed that such an opinion by Mr. Darwin
must necessarily have been based upon evidence accumulated by him in the Argentine
Republic and Patagonia.

In his "Contributions" Dr. White gives the following reasons for calling the rocks
of the Sergipe-Alagoas region cretaceous (p. 15) : "First, the majority of the types are
such as are generally regarded as characteristic of that period. Second, a portion of the
species are identified with published species of undisputed cretaceous fossils in other
parts of the world. Third, although some of the species have a Jurassic aspect, none of
them arc identifiable with any known Jurassic species. Finally, as all the collections
have been shown to belong to one liiuna, and a part of its species to be certainly of cre-
taceous age, the whole fauna must necefisarily be referred to that period." This decision
is somewhat weakened, however, by a statement (on p. 6) to the effect that " while
such a conclusion would, I think, have been reasonably reached from a study of
the fossils alone, much reliance has been placed in the corroborative testimony of the
geologists of the Brazilian Survey."

If the members of the former Brazilian survey were now asked to give tlit-ir
reasons for calling these rocks cretaceous, they would lay all stress upon Dr. White s
determinations and none at all, or very little, upon any evidence that has been
adduced from stratigraphic relations, simply because, as has already been said, those

39G THE CEETACEOUS AISID TERTIAEY GEOLOGY

relations have, as yet, thrown but little or no light whatever upon the age of these
particular beds.

In the mind of one acquainted with the mesozoic geology of Brazil, this fact
stands forth prominently : that while Dr. White's conclusion that these beds are cre-
taceous, must be accepted, the fauna presents peculiarities of its own in the Jurassic
facies of the Sergipe-Alagdas fossils. Another peculiarity of this cretaceous fauna of
Brazil is that certain of the typical genera, as Dr. White says, if found alone, would
be referred to the tertiary. It may be due to the incompleteness of the collections
made, but as far as we know these tertiary forms, Fusus, Murex and Phorus,* occur
only in the Pernambuco and Para collections, and not a single example is reported
from the Sergij)e-Alag6as basin ; whHe, on the other hand, not one of the Jurassic
forms is reported from the Pernambuco or Para beds.

Thus far the writer has spoken of the mesozoic beds of this region as a whole.
Let us now ascertain, if possible, whether or not both Jurassic and cretaceous beds
exist here, and whether this apparent mingling of faunas may not be due to bad
collecting.

'&•

The StratigrcqiTdc delations of the Mesozoic Beds.

The general relations of the rocks of the separate groups are shown in the sec-
tion through Itabaiaua, and it only remains to indicate the relations of the mesozoic
beds to each other. The details of these relatigns throughout have not been determined
with any degree of satisfaction, the time spent in the province having been taken up
entirely either in hasty reconnoissance or in collecting fossils. In the few more import-
ant instances in which these relations are known they are shown in the accompanying
illustrations. In view of the very interesting departure of the fauna of this group
from what would be regarded as a typical cretaceous fauna, the lack of detailed
information upon this subject is the more to be regretted.

In Prof. Hyatt's paper upon the cephalopods of this region, no definite locality
is mentioned for these fossils than " the cretaceous beds at Maroim." I learned
from Prof. Ilartt subsequently that this collection was taken, almost entirely, from
the paving stones in the streets of Maroim. Now the quarries from which these
paving stones came are so numerous and at such different horizons, that these fossils can-
not be located stratigraphically, and ibr this reason, no valuable paleontological deduc-
tions can be made from them in determining the faunal characteristics and relations
of the separate beds in the vicinity of Maroim.

♦This statement is made solely upon the authority of Dr. White's opinion as expressed in his " Contributions to
the Paleontology of Brazil," p. 17. IJut inasmuch as Fusus, Murex and Xenophora (Phorus) hegiu in the cretaceous
or earlier, it seems to be of doiibtliil value.

OF THE SERGIPK-ALAGDAS BASIN OF BRAZIL.

397

One of these cephalopods, however, Ammonites (Buchiceras) Jiarttii, Hyatt, as it
appeared afterwards, came from the locality known as Bom Jesus. The Jurassic
aspect of this fossil and the eroded condition of the specimens led Prof. Hyatt to
suggest the possibility of its having been transported from some older deposit.

The section criven herewith shows the relation of the Bom Jesus beds to others
of this group.

Iaranqeihas

SECr/ON

Bom Jesus

/?/<? SfHQ/P£r

SECTION

IaV^c

ANQ0RINH;?S

/

:t

InibIra

Porto DOS BflRcos,
TRflP/criE OAS PedR/^'

COQUEIRO, /ARAf A.

r.-'-.i-^--:

T^ff?!^?

■ : • 1 •

1 • 1 - 1 •

■ 1 - 1 -

1 -1 1

• 1 • 1 •

• 1 1

GahajAu

/^RDEIRA-

Lastro
Jhque

"= ITyRBAlANA-

cv..

N

Three sections in the Sergipe-AlagOas basin. (The heavily shaded beds yield fossils of Jurassic aspect.)

The Bom Jesus beds occur in ihe hilltops oast of the town of Larangcira.s. The
rocks are earthy, arenaceous limestones which arc readily attacked by disintpgrating
agencies, and have, therefore, been raiiiilly cotivi rted into residuary earths. The

398 TUE CRETACEOUS AND TERTlAKY GEOLOGY

fossils occur, for the most part, lying loose in the road where they have constantly
been ridden and walked over, or are imbedded in the black residuary earth that
covers these cretaceous hills. In many cases, however, there are irregular blocks of
the matrix preserved along with the fossils. The worn appearance of a great many of
the specimens from this locality is due to two causes : one is the rapid dissolving
action of the rain-water, which when raised to a high temperature by falling upon the
hot rocks, attacks them vigorously and leaves an etched surface such as is produced
by acid acting upon a calcareous sandstone ; the second is the wearing of the rocks
against each other where they are constantly walked and ridden over in the road.
There is nothing in the rocks themselves, in their topographic position, or in their rela-
tions to the underlying strata to lead one to suppose that these fossils have been carried
here from a lower geologic horizon.

Attention is called to the relation of the Bom Jesus beds to those of Larangeiras
and Pedra Furada, and to the fact that the Larangeiras beds are the equivalents of
those of Lastro, etc., at Maroim, and of those of Urubti on the Rio Sergipe.

At Maroim the Lastro beds are overlain by a similar cephalopod-bearing forma-
tion, that of Garajau, which yields other fossils of Jurassic aspect, while those of
Urubii, the equivalents of the Lastro beds, overlie others of decidedly Jurassic facies.

We have then the following arrangement of the strata :

1. At the top are the Garajau and Bom Jesus beds, yielding fossils of both creta-
ceous and Jurassic aspect.

2. Beneath these arc the Lastro, Jaque, LTrubu, Imbira, Toque, Capoeira, Larangei-
ras, and Pedra Furada beds, whose fossils are all referable to the cretaceous.

3. Beneath these are the beds of Porto dos Barcos, Trapiche Maior, Trapiche das
Pedras Novo, Trapiche das Pedras Velho, and Coqueiro, all of which yield fossils of
both cretaceous and Jurassic aspects.

As the Jurassic aspect of both the underlying and the overlying beds might tend
to throw doubt upon the reference of those of Urubii, Lastro, etc., to the cretaceous,
I can only refer to Dr. White's able discussion of this subject in his " Contributions "
(pp. 15-10), where he shows that the weight of evidence favors the cretaceous age,
not only of those particular beds, but of the entire Sergipe- Alagoas series.

It is no longer cause for surprise that beds of decided Jurassic aspects are found
overlying others of just as marked cretaceous aspect. Dr. Blanford, in his presiden-
tial address before the Geological Section of the British Association at Montreal in 1885,
laid timely stress upon these apparently anomalous conditions existing in India and
South Africa, and to tlie grave difficulty or impossibility of an exact and minute

OF THE SERGIPE-ALAGOAS BASIX OF BKA7.IL. 399

correlation of the beds deposited in such widely separated regions as Europe, South
Africa, India, Australia and South America.

It is worthy of note also in connection with the discussion of the mesozoic and
tertiary rocks of Brazil that the fossils of Jurassic aspect are conliuod to the Sergipe-
Alngoas basin and to the Crato district in Cearu,* while those which have a tertiary
aspect are, so far as they have been worked out, entirely from the Pernambuco and
Piabas basins.f

Inasmuch as the divergencies between the faunas of the Sergipe-Alagoas basin and
the Pernambuco basin tend to associate the former with the older and the latter with
the newer geological horizons, the probability is naturally suggested that the Sergipe-
Alagoas beds are more nearly synchronous with those of Crato in Cearu which are
referred by Newberry and Cope to the Jurassic, while the Pernambuco beds are similarly
related to those of Piabas in the province of Para.

THE COXDITIONS ATTENDING THE DEPOSITION OF THE MESOZOIC BEDS.

The rocks represented in the sedimentary beds of the Sergipe-Alagoas sections vary
from the coarse conglomerates at the base of Itabaiana and lying in contact with the
gneiss, through sandstones, quartzites, shales, slates and dolomitic limestones all of possi-
ble paleozoic age. In the mesozoic beds the rocks are sandstones, coarse, fine, and
often calcareous. These are variously colored from dark red through brown to cream
colored. The limestones are arenaceous, partially crystalline with Hint nodules,
oolitic and cherly.

The nature of these mesozoic beds indicates that the changes which took place
during their depo.sition were not very great or ver}' sudden, l^he sandstones lie at or
near the base of this series, the oolitic beds follow, and the soil limestones are the
highest ones seen.

The order and character of these beds appear to indicate a gradual subsidence
of the rejrion from the beefinnini; to the end of the cretaceous. The basal sandstones
often contain pebbles the size of a partridge egg, and sometimes the fossils are rolled
and water-worn, showing that they were deposited in tolerably strong currents and
near the shore. The earthy limestones and oolitic beds which follow indicate an
increasing depth and quieter waters, while the uppermost beds have been deposited

•See opinions of Dr. .1. S. Newberry and Prof. E. D. Cope concerning tlie fossil fislics of Oari in Proceedings of
tlie .Vmerican Pliilosopliical Society, Vol. XXIII, .Ian., 1880.

fSee Dr. C. A. While's "Contributions to the Paleontology of Brazil," p. 17, and Prof. E. I). Cope in tbe Pro-
ceedings of the American Philosophicil Society, ISSfi. In llic latter paper Prof Cope compares llio Pernambuco ver-
tebrate foasils with those of the Fox Ilills (upper cretaceous) of the United States.

400 THE CRETACEOUS AND TERTIARY GEOLOGT

in still deeper and quieter water than any of the lower ones, are more homogeneous,
and contain the remains of fishes and a few cephalopods. No shallow water animal's
remains have thus far been found in them. Then followed the j^ressure that folded
broke, and in places, iiaitially metamorphosed, the cretaceous beds. This ended the
dejjression of the ocean's bottom.

Pressitre. — The cretaceous beds of the Sergipe-Alagoas basin have been sub-
jected to more or less pressure, though this pressui'e has not been uniform.

The following are the evidences of such pressure :

1. Many of the fossils collected in this region are more or less crushed or other-
wise distorted.

2. The beds as seen near Maroim, at the Lastro, and at other places along the
Maroim arm of the Tiio Sei-gipe, and especially along its western bank, are thi-own
into folds.

3. Slaty cleavage is noticeable in the Ilha do Chaves, a mile below Propria.

4. Superinduced cleavage and joint structure are seen in the sandstones of the
Serra d'ltabaiana and in those of the Serra de Maraba.*

In some localities the limestones are metamorphosed, and their fossils completely
obliterated, while in many other places the obliteration of the fossils is but partial.

The folds in these rocks indicate that the pressure by which the region was dis-
turbed came from the south-east. Although this pressure was sufficient to flex the
cretaceous beds, and, in places, to metamoi'phose them more or less, there is nothing
to show that it was great. The flexures are usually gentle and the rocks seldom dip
at a high angle, as will be seen from the accompanying table of dips.

ECONOMIC.

It is not to be wondered at that the popular notion that precious metals must
occur in all mountainous regions, has led to the supposition that the serras of Itabai-
aiia and Maraba arc auriferous. These mountains have been examined repeatedly in
the search for gold, silver and what not, but without success. As long ago as the
seventeenth century they were explored and found barren of such riches.f At the

* Certain oUier saadstones of tliis region are partially metamorphosed. In some places this metamorphism is com-
plete, while in others it is but slight. Tlicre seems to be some doubt as to the exact place of these metamorphosed
siindstone beds in the scries, but the weight of testimony seems to sliow that they are tertiary. The metamorphism of
lliL'se particular beds cannot, I think, be adduced in support of the tlieory of pressure, for it is entirely due to
weathering.

f "Within the jurisdiction of this captainship ('Sergipe del Rey') is the mountain of Tabayana ; from whence
several valuable ores were presented to the council of 19 ; but, upon proof, were found not worth farther looking
after." 1<)41. Voyages and Travels into Brazil by John Nieuliotl". Pinkerton's Voyages, Vol. XIV, p. 706.

OF THE SEUt.'irE-ALAtti^AS IJASIX OF BIIA/.IL. 401

time when the gold and diamond diggings of Brazil were at the height of their suc-
cessful operation, extensive explorations were carried on in these mountains. Even
during the last twenty years such exi)lorations have not entirely ceased, though the}"
have never given any promise of success.

There is nothing in the geologic structuic, as ftir as it is known, to lead one to
expect deposits of economic value in the mountains of Sergipe and Alagoas.

It is possible that from the soft chalky beds at Andorinhas and Sapucary com-
mercial chalk could be manufactured, though I have nothing beyond the general
appearance of the rock to lead me to this supposition. It may be worth testing for
this purpose. The tertiaiy beds would yield mineral paints of vaiious red, yellow
and brown colors, but owing to the absence of demand for such paints in Bi-azil, they
would have no value at piesent.

The last member of the tertiary beds, the conglomerate which is scattered far
and wide over the region, yields some limonite iron ore, but this ore has not been
found in any considerable quantity as yet, and in view of the vast deposits of excel-
lent iron ores in Sao Paulo and Minas Geraes, it is scarcely possible that a limonite
ore could be profitably handled in this part of the Empire.

There were the usual reports through the province, at the time of my stay, of
the discovery of coal, silver, copper, lead, etc., but in no case did I find such reports
well founded or worthy of serious attention. The coal said to have been found at
Campo Redondo was the charred I'cmains of a tree buried in the tertiary clays, while
the copper discovered in the same neighborhood was a piece of the manufactured
metal.

Building Stone. — The time will doubtless come when the beautiful oolitic lime-
stones of the Kio Sergipe will be utilized for architectural purposes. This stone
resembles the famous oolitic limestones of Indiana, so highly ])rized and extensively
quarried and used in the United States for building jiurposes.

The ease with which the Sergipe rocks can be quarried and sawed and cut into
blocks of any size and shape, its hardening upon exposure, its rare beauty, its pleas-
ing tints of green, gray, brown, and cream color, its convenience to transportation by
water to Aracaju and thence by sail and steamer to all parts of the Brazilian coast,
may yet make it a successful rival of the beautiful crystalline rocks of Kio de
Janeiro.

A. p. s. — vor.

402 THK CUF,TA( EOUS AND TEIITIAUY GKOLOGY

Pakt II.

THE BRAZILIAN MESOZOIC BASINS OTHER THAN THAT OF SERGIPE-ALAGOAS.

In order that the relations of the Sergipe-Alagoas basin to the mesozoic geology
of other portions of Brazil may be the better iinderstood, brief descriptions are here
given of all the other known mesozoic localities in Brazil. The locality in the bay of
Sao Francisco, Province of Santa Catherina is here described for the first time.

SAO FRANCISCO DO SUL, PROVINCE OF STA. CATHERINA.

Of the cretaceous rocks at this place but little is known at present, and that lit-
tle is due to a small collection of fossils made here in 1876, by Mr. Luther Wagoner,
then topogiapher on the Imperial Geological Survey of Brazil. As the specimens
brought away by Mr. Wagoner were collected incidentally, no observations were
made by him of the extent of the deposit, and no one seems to have given the matter
attention since that time. The writer visited the place in 1881, but as he was there
one night only, he was unable to gain any knowledge of it. The position of the out-
crop suggests that this rock may underlie the country about the bay of Sao Francisco.
It certainly does not appear in the surrounding hilly country.

Lithologically it is a brown, earthy, calcareous sandstone resembling that found
at Olinda in the province of Pernambuco. It was observed only upon a small island
about six miles from the port and covered only about one acre.

The outcrop, Mr. Wagoner tells me, is very inconspicuous, near the water's
edge,* and suggests the geology of Bahia and Maria Farinha, except that the expo-
sure is very limited. The specimens collected contained among others a number of
fossil gasteropods. It should be stated here that this reference of these beds to the
cretaceous is not based upon a detailed study of the fossils, but mainly upon the
striking general resemblance of the fossils to those found in the cretaceous beds fur-
ther north. It is to be regretted that this small collection of material was not sub-
mitted to Dr. White in connection with other mesozoic material collected by the Bra-
zilian Survey. Of the relation of the cretaceous at this place to the older rocks noth-
ing is known.

* It was suggested at the time that these specimens might be fragments carried to the place as ballast, but Mr.
Wagoner was toiilidenl that this could not be the case, for the rocks do not occur in fragments, but in a solid bed.

OF THE SEKCilPE-ALAGUAS BASIN OF BUAZIL. 403

THE ABi:oLIlOS.

This locality is described by Prof. Hartt in the American Naturalist, Vol. If,
March, 18G8, and in his Geology and Physical Geography of Bi-azil, pp. 174-214. It
was not visited by the writer, and the following facts aie taken from Ilartt's writings
and from the seiies of photogi-aphs made there in 1876 by j\Ir. Feri-ez, the photog-
rapher of the Brazilian Survey. For further details reference should be made to
Prof. Hartt's book.

The sedimentary i-ocks bear a close resemblance lithologically to those of Penedo
in the Sergipe-Alagoas basin, and they seem to be referred to the cretaceous largely
on account of this resemblance. They are made up of arenaceous limestone (?) be-
neath, blue shale above, followed by a yellowish shaly sandstone containing obscure
plant impressions. These sedimentary beds are covered by igneous rock, and they
all have a north-westerly dip at an angle of from ten to fifteen degrees. Neither
the thickness nor the superficial distribution of these rocks is given, but both the size
and the height of the islands preclude the possibility of either great thickness or ex-
tensive distribution.

THE BAIIIA BASIN.

The geology of this basin has been written upon by Pissis, Ilartt, Allj^ort, Derby,
Kathbun, Mai-sh, Cope, and White. The description and discussion by Mr. Derby,
published in the Archivos do Museu Nacional, Vol. Ill, pp. 135-158, written after
the most thorough exploration of the region that has yet been made and with all the
facts in hand that had been accumulated by previous observations, is the most nearly
complete that has ever been published.

The cretaceous rocks of the Bahia basin were deposited in a fresh-water lagoon,
and are therefore quite distinct from those of the cretaceous beds along other portions
of the coast. The limits of the basin have never been determined except in a very
general way. The following outlines are taken mainly from Mr. Derby's jiaper above
referred to.

The eastern margin of the basin extends from the city of Bahia to the station of
Cat 11 on the Bahia and Siio Francisco railway, a distance of fifty-four miles, '^rhe
western limits are buried beneath the tertiary hills near Cachoeira and end somewhere
east of Nazareth, while the erosion of these tertiary beds gives the margin of the creta-
ceous exposures a ragged edge, outliers or buttcs of the tertiary being scattered here
and there throughout the central ])ortion of the area. The northern margin is binied
beneath the tirtiary i)lateau which extends from near Cachoeira north of Sto. Amaro,

404 THE CKETACEOUS AND TEKTIAUY GEOLOC.'T

and crosses the Bahia and Sao Francisco railway at Pqjuca. The south end of the
basin has been cut away by the ocean, the island of Itaparica having been included
in it.

Near Inhambupe, about thirty miles north-east of Alagoinhas, Mr. Derby found
disturbed beds of schists, sandstones and limestones, which he is disposed to refer to
the paleozoic. It is unfortunate that he does not report the dip of these supposed
paleoz'oic beds. While on the Rio Joannes on the east side of the basin he saw the
cretaceous lying inconformably against the gneiss.

The cretaceous rocks of the basin are conglomerates, sandstones, shales and lime-
stones. The invertebrate fossils are described by Di*. White in his Contributions to
the Paleontology of Bi'azil.

Of these fossils Dr. White remarks, that " all the types which the fauna em-
braces, so far as they are determinable, are represented among moUusks now living "
and that " a part of these types are at present known to exist or have existed only
upon the western hemisphei'e."

THE PERNAMBUCO BEDS.

The most southerly exposure of the mesozoic beds seen in the province of Per-
nambuco, by the writer, was near the mouth of Rio Formozo. This exposure, how-
ever, seems to be the northern extension, and near the end, of the Sergipe-Alagoas
beds, as has already been suggested. But whether it is or is not the continuation of
the former, these Rio Foi'mozo mesozoic beds do not pass to the northwai'd of Cabo
Sto. Agostinho, along the oceanward side. This cape is of quartz-porphyry,* and on
three sides is built against unconforraably by particolored tertiary beds of sands and
clays, a characteristic view of which is given in the part of this paper treating of the
tertiary dcposits.f Whether the mesozoic beds underlie the tertiary on the landwai-d

"Specimens of llie crystiilline rocks from the vicinity of Cape Sto. Agostinlio, one from Estagao da Illia, collccled
by Professor Ilarlt, and two from the island of Sto. Aleiso, collected by the writer, were prepared for microscopic
examination and submitted to Dr. George H. Williams, of Johns Hopkins University, Baltimore. Concerning these
specimens Dr. Williams writes : " * * * They must be described as quartz porphyries or rhyolites, according as
they are of pre-tertiary or tertiary age. They arc quartz orthoclase aggregates with almost no bisilicate constituents,
but their structures are very varied. One of the specimens from Sto. Alci.\o has a granular, holocrystalline ground-
mass of quartz and feldspar, some chlorite, possibly representing original hornblende or mica, and considerable blue
tourmaline. Tlie other Sto. Alcixo specimen is a fine granophyre. Porphyritic quartz and feldspar lie in a holocrys-
talline groundmass whicli is filled with beautiful splierulitcs showing the black cross between crossed nicols. The
specimen from EstaQao da Illia bus large porphyritic sanadincs with chlorite inclusions which were once glass. The
groundmass is mostly composed of rectangular sanadincs with much finely disseminated chlorite and iron hydro.xide.
Magnetite octahedrons and beautiful zircons are abundant. There is no nephelene in any of these rocks.

t Sec page plate of "The Uuinas de I'almyra."

OF THE SKRGll'E-ALAGOAS BASIN OF BRAZIL. 405

side of the cape is not known, and it lias been assumed tluit tiiey do not, altliougii
there is nothing in the topography of the region to make such connection between the
Sergipe-Aiagoas beds and the Pernambuco beds impossible.

■ At Olinda the cretaceous rocks are exposed in the southern face of the hill upon
which the church stands. The rocks are soft, earthy, somewhat calcareous, fossilifer-
ous sandstones. This and an exposure of white limestone a short distance west of it,
and known as the Forno de Cal, are the only exposures of cretaceous rocks now
known in the immediate vicinity of the city of Recife, the next one to the north being
in the vicinity of Kio Maria Farinha.

The region, rocks, and fossils of the Maria Farinha beds have been described
by Kathbun in his paper upon the cretaceous lamellibranchs,* and by Derby in Dr.
White's contributions,! and only such of their characters as may be useful for future
comparison with the other cretaceous localities of the coast need be mentioned here.
Lithologically the beds vary from cream-colored and white pure limestones to earthy
sandstones, all of which dip at a very low angle to the east. They are undisturbed,
and none of the beds are oolitic in character. It is to be remarked that, of the several
fossils of Jurassic aspect in the Brazilian collection described by Dr. AVhite, none
have been found at the various Maria Farinha localities. It is also noticeable that
all of the genera in this collection suggestive of the tertiary age are represented in
the Maria Farinha locality, but none of them have been found at the Sergipe-Aiagoas
localities. These facts seem to suggest, at least, that the Pernambuco beds are not
so old as those of the Sergipe-Aiagoas region from which the collections were made.

It is the opinion of the writer, though it is expressed with that hesitation which
should come from a lack of sufficient facts to suppoi-t it pioperly, that the INIaiia
Farinha and Olinda beds belong to the same horizon as those exposed three miles
south-east of Pencdo at Villa Nova, and that the latter overlie all the cretaceous beds
from which the collections were made in the province of Sergipe.

The extent of the Pernambuco deposits has not been determined, even approxi-
HKilely, and it is not known, perhaps cannot be from surface exi)osure« alone, whether
the Olinda beds are, or ever were, connected with those of Maria Farinha, and whether
the latter are continued by those found at Iguarassu and Goyanna, or whether those
at Parahyba are also a continuation of the Pernambuco beds.

The coast between Olinda and Maria Farinha is low, and there arc no lock ex-
posures along it to aid one in connecting the two localities. From near Maria Farinha

• Preliminary Report on llic Cretaceous r.amellibrnnclis collected in llie vicinity of Pornainhiico, Hrn/.il. Hy Kidi-
ard Kathbun. Proc. Bost. Soc. Nat. Hist., XVII. 1874, pp. 311-256.
t Contributions to the Paleontology of Brazil. By V. A. Wliitc.

406 THE CRETACEOUS AND TERTIARY GEOLOGY

to near the mouth of the Rio Parahyba do !N^orte the coast is formed of a series of
soft, horizontally bedded, particolored tertiary sands and clays which are readily cut
and washed away by the ocean, the talus of which obscures any beds which might
otherwise appear at their bases.

Cretaceous beds are exposed on the low gi'ounds of the island of Itamarica,
while the tertiary beds form the higher lands.* The relative positions of the creta-
ceous and tertiary upon this island favor the opinion that these same i-elations con-
tinue northwaid, and that the cretaceous of Parahyba is simply the northern exten-
sion of the Pernambuco beds. It is possible that a careful examination at low tides
of the beds along the coast and streams from Itamarica to Parahyba would settle this
question.

No rocks referable to the paleozoic have thus far been found in the vicinity of
the Pernambuco basin.

THE PARAHYBA EXPOSURES.

N^o examination of the mesozoic exposures at Parahyba do Norte was made by
any member of the Commissao Geologica, and the only information that we have of
them is from a few brief notes by Williamson, Agassiz, and Capanema. The rocks
are soft, impure, gray limestones, dipping gently to the east, and containing but few
fossils. They occur near the city of Parahyba and at Minas da Cachoeira. Their oc-
currence at the latter place seems to add to the probability that the Parahyba beds
are simply a northerly continuation of the Pernambuco beds.f I have been unable to
obtain information regarding the age of the beds immediately underlying the creta-
ceous rocks of Parahyba.

THE MESOZOIC BEDS OF CEARA AND PIAUHY.

Our knowledge of the cretaceous beds of the interior of the province of Ceara
is derived almost exclusively from the observations and collections of Gardner made
just fifty years ago. With the exception of Capanema, the many writers who refer
to these beds have derived their information from him.

The facts of most importance brought out by Gardner are the geological section

* Relatorio dos Traballios da Coiiiinissm) Geologica do Brazil, por Cli. F. Hartt, Rio dc Janeiro, 1875, p. 10.

t Among Uie fossils sent Dr. Wliiio from the Miiseu Nacional at Rio were some broken specimens of Toxaster altim-
cuius White, taken from the walls of ihe fort at the mouth of the river. As these specimens were collected by myself, I
quote from my fiehl notes written at the time, November the 25tli, 1875 : "I found some echinoderms and shells in the
walls of the old Dutch fort, at the nuiutli of the Uio Parahyba, but upon inquiring for the quarry from which the
stone came, I was told that it was brought from Europe ; at least that there are no quarries in the neighborhood yield-
ing such slonc."

OF TIIK SEKfaPE-ALAOOAS BASIN OF BRAZIL. 407

of the Scrra do Araripe, which, bogiiHiin<ij at the base, consists of horizontal beds of
lignite, limestone, sandstone, with chnlk and flints, the sandstone stratum forming the
mass of the plateau. The total thickness of the section is from I'lOO to 1500 feet.
The underlying rock appears to be slate, presumably paleozoic. The fossil fishes
found occur in concretions scattered about the surfjxce, which he supposed to have
been water-worn. The exact position of the fossils in the section is not known.
Gaidner believed them to have been weathered from the sandstone bed, but TIartt,
who, however, never visited the locality, expresses the opinion that they come from a
bed beneath the sandstone, which he thinks, together with the overlying beds, belongs
to the tertiary. The serra here spoken of trends N. N. E., and is an elevated plateau
about thii'ty miles wide, cut down rather abruptly on both sides, the valley on the
Ceara side being lower than that on the western or Piauhy side. The geographic dis-
tribution attributed to these beds by Gardner, that is, from this point to Maranhao,
is not warranted either by the facts given by him, or by any that have subsequently
come to light. The fossil fishes collected at and about Crato, by Gardner, were de-
scribed by Agassiz, who believed them to be of cretaceous age.* It should be men-
tioned in this connection, however, that Dr. J. S. Newberry and Dr. E. D. Cope
regard these fishes as of Jurassic age.f

OTHER CRETACEOUS EXPOSURES,

Jiio Mbasoro, Province of Rio Grande do Norte. — The exposure of the creta-
ceous on the Rio Mossoro in Rio Grande do Norte was reported by Major Coutinho.
Nothing is known of the character of the beds or their geographical distribution.
V. Dr. White's Contributions, p. 10, foot-note.

Bio Pidbas, Province of Para. — Concerning the Piubas locality I personally
obtained from Snr. Ferreira Penna substantially the same information as that fur-
nished Professor Hartt, and given in Dr. AV'hite's Contributions, p. 9. Snr. Penna
assured me, however, that he had never seen these rocks exposed elsewhere than at
the mouth of the Rio Piabas where they aw. uncovered at low tide. lie found the
exposure wliiji' making a trip along the coast in a .small boat, and spent but a lew
minutes in making the collection from which Dr. White describes so many species.

Pio S'lo Francisco above the Falls of Paulo Affonso. — The secondary nature of
some of the rocks of the Silo Francisco valley above the falls was first made known

•Edinburgh New Pliilosophical Jour., .Jan., 1841.

tA Conlribulion to llie Vertebrate Paleontology of Brazil. By E. D. Cope. Proc. Am. Pliil. Soc. January,
1886, pp. 1-21. The opinion of Dr. Newberry is quoted as here cited. It should be staled, however, that there is
possibly a mistake in crediting him with this opinion, for in a recent conversation with the writer. Dr. Newberry re-
ferred to tliese fossils as of cretaceous age.

4f)8 THE CRETACEOUS AND TERTIARY GEOLOGY

by Mr. Derby in his article published in the Archives do Miiseu l^acional, Yol. IV,
for 1879 (published in 1881). These are horizontal beds exposed in the isolated hills
which extend along the left bank of the river from near Piranhas to Itaparica, but
they form more continuous chains from the serra de Tacaratu toward the interior,
while what ajjpear to be plateaux of the same rocks stretch away to the south of the
river also. In the immediate neighborhood of the river they are continuous from near
the mouth of Pajeu to Itajiarica. Above Pajeu, hills, apparently of these same rocks,
stand back fi-om the river and overlie the gneiss. These beds contain silicified wood
in abundance, besides a few other fossils. At a place called Atalho, fossil cyprids,
bones, teeth, and scales of fishes and of i-ej^tiles have been found. At Caissara several
leagues further up the river these beds occur again.

The only determination of anj- of these fossils made by Mr. Derby was that of
scales of Lepidotus. He ventures the opinion that they are secondary and prob-
ably cretaceous, resembling somewhat the fi'csh-water cretaceous of Bahia.

I regard the discovery of these dejiosits by Mr. Derby as one of great import-
ance in the study of the mesozoic geology of Bi-azil, for this locality seems to furnish
the facts necessary for the determination of the relations between the coastal beds of
the Sci'gipe-Alagoas region and the interior, and somewhat more elevated, beds of
Ceara.

JRio Purus, Valley of the Amazonas. — The only unquestionable cretaceous de-
posits thus far known in the Amazon valley are on the Rio Aquiry, an afHuent of
the Rio Purus. These deposits were discovered by Chandless and are very briefly
described by him in an ai'ticle published by the Royal Geographical Society, Yol.
XXXYI, p. 119 et seq. The Aquiry enters the Puriis on the right in latitude 8° 45'
South, longitude G7° 23' "West. From the mouth of this stream to 11° South latitude
the water, when low, uncovers rocks in the middle of the stream, in which fossils are
Jbund. Yertebraj from these beds were seen by Agassiz at Manaus, and he pro-
nounced them to be those of Mosasaurus. Nothing is known of the general charac-
ter or extent of these beds. The theory that the cretaceous outcrops around the rim
of the Amazonian basin is not supported by any known facts.

CORRELATION OF THE MESOZOIC OF THE COAST AND THE INTERIOR.

Owing to the lack of knowledge of details a very brief discussion of the rela-
tions of the mesozoic beds along the coast with those of the interior is undertaken
with much hesitation. Indeed, were it not that this subject has already been touched
upon by Mr. Derby in such a manner as to lead geologists to suspect that we have
here unusual and very peculiar conditions, it would not be attempted at all.

OFTIIK SEIiCarK-AI.AfJitAS RASrx OF RIIA7.IL.

40^

It will be observed uijon tbe small map on this page that, besides the small basins
along the Biazilian coast, there are, leaving the Purus exposures out of account, two
mesozoic localities at some distance from the coast, one at and about Crato in Ceani,
another about Taearatu, a short distance above Paulo Alfonso, in the valley of the
Kio Sao Francisco,

The cretaceous beds of the Sergipc-Alaguas basin overlie paleozoic rocks, those
of the Sao Francisco basin appear to ovei'lie metamorphic rocks, which may be either
paleozoic or archiean, while the Ceara beds overlie slates which ai'e pi'obably paleozoic.
The region separating the Sergipe-Alagoas ex-
posures from those of the Sao Francisco l)asin
are granitic and metamorphic, while metamor-
])hic rocks lie between the Ceaia exposures and
the sea coast. If we take into consideration the
cretaceous basin of Bahia we find that also sep-
arated from the exposures of the Sr>o Francisco
basin by crystalline and metamorphic rocks.

Gardner reports fossil fishes in the vicinity
of Jaidim. a town south-east of Crato. The
serra de Tacarati'i on the northern side of the
Kio Sao Francisco above the falls of Paulo
Affonso is saitl by Mi-. Derby to be composed of
rocks of secondary age. This serra is about one
hundred miles south-east of Jardim, but Mr.
Derby says* that " these secondary dej^osits be-
come more continuous and form a long line of
hills, which beginning with the sena de Taca-
ratii stretches away into the interior of the province of Pernambuco." The iiUer-
vening country has not been examined, but it seems highly probable then that the
Tucarutii beds are the southern extension of those of Crato and Jardim.

As nothing is known of the details of the mesozoic geology of the Mossoro
l)a>in, nothing can be ventured regarding it fnither than that its proximity to the
Crato exposures natiu-ally suggests that the beds of the two localities are identical,
the intervening beds having, in all probability, been removed by erosion.

Mr. Derby intimates that there is something pueuliar about the relations of
the coastal mesozoic beds to those of the interior. The ohjeetion olTcMed by him to
the correlation of the Ceara and the Sao Francisco valley beds with those along the

* Arcliivos do Miiseu Xiirioniil, Vol. IV, p. 92.
A. T'. S — vol.. XVT. 2z.

EA&TiitN Brazil

THE 016T«I8UTION
or THL

410 THE CRETACEOUS AND TERTIARY GEOLOGT

coast, namely, that they stand at different elevations, reduces itself to the suggestion
of a classification of geological formations by comparative elevations.* If all the beds
of the cretaceous both npon the coast and the interior Avere found to be perfectly
horizontal, with no evidences of faulting between, we might possibly be justified in
such an assumption. The position of the Sergii)e-Alag6as beds, however, shows that
this is not only not the case, but that in this basin, at least, the beds have a veiy
decided dip toward the ocean, while their inland margins rise to a height of from 400
to 500 metres or more where they overlie the paleozoic beds of the Itabaiaua. At
their inland margins they rise to the same elevation above a large number of expo-
sures of the same beds at tide level near the coast. If no connection had been ti'aced
out between these Sergipe localities we should be obliged, according to this hypsome-
tric classification, to separate those beds geologically. This suggestion must be
regarded, therefore, as a geological non seqiiitur. Mr. Dei-by assumes also that the
coastal cretaceous beds do not reach an elevation greater than 100 metres above tide.
This is erroneous. The serra of Itabaiaua is, according to Mouchez, from 700 to 800
metres above tide, and the cretaceous beds overlie the paleozoic beds of Itabaiana to
within 150 metres of its summit, which would make the elevation of the cretaceous
at this place above tide about GOO metres.

Accoi'ding to the elevations along the Rio Slo Francisco, determined by Col. W.
Milnor Roberts, the river near Tacaratii and in the vicinity of the secondary beds
reported by Mr. Derby, is 320 metres above tide, and, according to him, the hills to
the north are about 303 metres above their bases,f or, say 620 metres above tide.

The Ceara beds, according to Gardner, rise to an elevation of 600 metres above
tide, which would also indicate that, as far as elevations are concerned, there is no
impoitant difierenee between the interior and the highest of the coastal d(!posits. But
I lay no stiess upon this elevation of the coastal beds, for, to my mind, it is a matter
of but little importance, and I return to my original statement, that even if there
were very marked difterences of elevation, nothing could be deduced therefrom
iu regard to the relative geologic ages of the beds of the various localities. The ele-
vation of the Brazilian coast has not been an even one, and thei'e is certainly no rea-
son for supposing that it would be. The elevation or depression of coast lines, and
especially of lines as long as that from Rio de Janeiro to the mouth of the Amazon,
has seldom or never been known to be even or regular. It is true that the tertiary
beds along the Brazilian coast have a generally horizontal position, but this appear-

* "It is (linicuU li> iulinit the contemponuieous ilepositioa of beds at sucli different elevations," Ai-chivos do
Museu Nacional, Vol. IV, p. 94.

f Arc'hivos do Mus. Nac, 1879, Vol. IV, p. 91.

OF THE SERGIPE-ALAGUAS BASIN OF BKAZIL. 411

ance does not conflict in any respect with the idea of unequal elevation, for this eleva-
tion would need to be very unequal indeed to make itself perceptible to the unaided
eye*

All of the little evidence we have seems to suggest the identity of the coastal
with the inland beds — certainly it does not suggest any obstacle to their correlation.

Agassiz pronounced the fossil fishes, collected by Gardner, cretaceous, while
Xewberry and Cope believe them to be of Jurassic age. Dr. White pronouncLS the
majority of the Sergipe-Alagoas fossils to be cretaceous, though he and Prof. Hyatt
declare many of them to have a Jurassic aspect. If these facts suggest anything in
regard to the relations of the rocks to these localities, it is that the probabilities favor
theii-, at least, ai)proximate identity. While I cannot, of course, maintain this corre-
lation, I must, at least, dissent from a separation without better evidence than has
thus far been adduced.

*S«e G£ulogie du BresU, par EmniADuel Liais, p. 249.

412 THE CKETACEOUS AND TERTIARY GEOLOGY

Part III.

THE TERTIARY.

The thorough understanding of the tertiaiy geology of Brazil, not to say of
South America, would, in my opinion, be of more interest and value to geology than
that of any other geologic horizon represented on that continent. A full discussion
of it cannot be undertaken here, is not intended, and indeed would be impossible with-
out a vast amount of field work. The formation is too widespread and the questitms ■
presented by it too complicated to admit of discussion in this place, further than it
relates to the geology of the Sergipe-Alagoas basin and to the cretaceous formations
of the eastern coast.

In Brazil this formation is represented by horizontal beds of sands and clays form-
ing a system of marine terraces from 100 to 250 feet thick, extending from some-
where between Eio de Janeiro and Bahia more or less continuously along the coast
to the north, being especially prominent fi'om about Maeeio nearly to Cape St. Roque.
In the Amazon valley they form the table-topped hills and are widespread from the
Rio Araguay westwai-d. The beds of the Amazon valley were thought by Agassiz
to have been deposited in an inland lake during the glacial epoch. Pissis says that
the marine tertiary beds of the coast are represented in the interior by beds of la-
custrine origin. The origin attributed to the tertiary by Prof. Agassiz need not be
discussed, as in all i^robability no one now admits its correctness, while a lack of
knowledge of the lacustrine deposits of the interior i-enders the discussion of the
theory of Pissis impossible. It must be admitted, however, that this theory of Pissis
is a very plausible one, and such observations as the writer has made upon the geol-
ogy of the interior of Brazil leads him to accept it as a favorite working hypothesis.
In none of the supposed tertiary beds of the interior, at least in the provinces of Rio
de Janeiro, Minas Geraes and Matto Grosso, has the writer found unquestionable
evidence of the tertiary age of such deposits.*

Character of the Beds. — The general features of the Brazilian tertiarj' have been
several times described by such good observers as Agassiz and Ilartt. The uniformity

* Since the above was written a letter, making inquiry concerning tlic tertiary, has been directed to Mr. James E.
Mills, of San Francisco, Cal., who has traveled in many parts of the interior of Brazil. In reply Mr. Mills says :
"I cannot give you any definite information from my notes about the tertiary deposits in Brazil, because in Rio Grande
do Sul, Minas Geraes, Rio de Janeiro, and other parts of the country where I had opportunities for observations and
study, there are no deposits which I know certainly to be of tertiary age."

OF THE SEUGIPE-ALAGUAS BASIN OF HltAZIL. 413

in its general charactei's between such widely separated regions as the Amazonas and
Sergipe or Bahia is worthy of note. The description b}' Hartt of the tertiary hills
of Parauaqiiara''' might have been written of some of the Sei'gipe exposures with the
exception of the sequence of the various beds. The following is his section, begin-
ning at the top :

1. A few feet of fino, light, brick-red earth, consisting of a mixture of clay and
fine sand.

2. Red sandy clay with iron nodules.

3. Tauatinga clay, grajish-white. Heavy bed, not laminated.

4. White clay, partly pure tauatinga, partly sand, resembling brick of two im-
perfectly mixed clays. Bakes hard in the sun and resists denudation.

5. White or cream colored, soft, fine-grained sandstones.

6. Variegated sandy clay.

7. Argillaceous sandstone vai'iegated with bands and mottlings of delicate shades
of white, red, purple, brown and yellow.

The following section was observed near Maroim, and may serve as a type of this
foi-mation along this part of the coast :

1. Surface soil, sandy, 4 feet.

2. ^Mottled red and white clays, 3 "

3. Blown sandstone, 1 foot.

4. Chalky above, red beneath, 6 feet.
6. Terra-cotta colored sandy clays, 7 "

6. Dark red, hard clays, 2 "

7. Soft, light-brown sandstone, 3 "

8. Soft white sandstone, 10 "

9. Soft earthy sand, 1 foot.
10. Talus, princiiJally from Xo. 7, 15 feet.

Not enough attention was paid to the tertiary to enable the writer to say
whether the secpience of the beds is at all uniform even wilhin the limits of the Ser-
gipe-Alagoas basin.

Evidences of the Age of this Formation. — The series of rocks here referred to
the tertiary are so called upon stratigraphic evidence alone, for it is a vciy remarkable
fact in regard to these beds that, widespread as they an;, no fo-^sih liave cvci- b 'cn
found in them, though diligent search was made for them by the writer, and doubtless
also by others. But their relations to the cretaceous and their general resemblance to
the tertiary beds of other parts of the world scarcely leave any doiibl alidut the

* Kill, liiif. Hoc. Nal. Scl., Jim , 1H71,

414 THE CKETACEOUS AND TERTIARY GEOLOGY

correctness of this reference. What portion or portions of the European or North
American tertiary they re])rescnt it is quite impossible to say. In general appearance
the deposits bear a strong resemblance to some of the later tertiary formations of the
Southern United States, though the materials of the two are derived from very differ-
ent geologic sources.

Absence of Fossih. — It is a peculiar feature of the Biazilian tertiary that with
the exception of a few plants found in the Amazon region, fossils have never been
discovered in it. That fossils should not be preserved in beds of such thickness and
vi^de distribution, made uj) of strata of sands and clays variously commingled, is cer-
tainly to be wondered at. The statement of the fact has often been questioned, and
explained by the assumption that careful search has not been made for them. That
they may yet be found is of course highly probable, but the writer has searched miles
of exposures in vain for any recognizable ti'ace of organic remains.

Several hypotheses suggest themselves in explanation of the non fossiliferous
character of these rocks. After examining the geology of the Amazon region,
Agassiz concluded that an enormous glacier once moved down that valley, and built
up across its eastern end a gigantic moraine ; that as the ice melted the formations
which ai-e now believed to be tertiary were deposited from the cold, muddy waters
flowing from the glacier into a great lake. This, to his mind, explained also the
absence from these beds of organic remains other than leaves. But even if Prof.
Agassiz's theory of the origin of these Amazonian beds wei"e accepted* it would not
explain the absence of fossils from the same rocks along the south-eastern coast of
Brazil.

Two hypotheses are offered : The first is, that these rocks were deposited so
rapidly, and from water so overloaded with mechanically suspended matter, that ani-
mal life in them was impossible. It may be said in regard to this theory that while
most of the teitiary beds are of a nature which would support it, there are many
beds of clays and very fine sands which would hardly be deposited rapidly or from
strong currents (see section on p. 410).

The second hypothesis is that these beds once held organic remains, but that
they have been dissolved out by infiltrating waters.

Present Extent of the Tertiary. — The present distribution of the tertiary beds in
the Sergipe-Alagoas basin cannot be given except in general terms, but from the
explanation of its relations to the cretaceous roclvs of the region (y. p. 37G) one will

* Agassiz's theory has never been accepted by the geologists who are acquainted with the geology of the Amazons,
and he is said to have abandoned the theory of the glaciation of Brazil before his death. See Glaciers, by Slialer and
Davis, p. 47.

OF THE SKRGIPE-ALAGOAS HASIN OF BRAZIL. 415

have no difficulty in knowiiioj what is to be expected. It is much more widespread
through the western than the eastern portion of the province of Sergipe, while in
Alagoas it is i-epresented by a succession of mutilated terraces. In the western part
of Sergipe the dry, thirsty soil of this formation prevents the growth of important
forests, and gives to it the name of ngreste in distinction from the wooded cretaceous
soil of the eastern portion.

Xear the coast, along the Siio Francisco river and the other larger streams of the
region, there are scarcely any i-emnants of the tertiary to be seen, but as one recedes
from the valleys, outliers of the tertiai'}' become more and more common until it forms
continuous taboleiros, deeply gashed at their margins and sometimes abutting uncon-
formably against the paleozoic or cretaceous rocks of the serras. On the sketch map
on page 'ASH the outliers of tertiary in the vicinity of Aracaju and Maroim are located
approximately. The tertiary hills are, as a rule, characterized by having Hat tops. It
not infrequently happens, however, that small outliers have been so extensively eroded
that tlie upper and most resisting bed of clay has been washed away and the lower
beds have broken down rapidly, leaving a more rounded and less characteristic topog-
raphy.

Toward the northern end of the basin the tertiary beds approach nearer and
nearer the coast, until, north of Maceib, they form a chain of hills and bluH's which
continue with occasional interruptions to near Kio Grande do N^orte.

T/ie Oiirjiiial Extent of the Tertiary. — The original extent of the tertiary beds
is indicated partly by the horizontality and the approximate uniformity in thickness
and character of the strata, and, secondly, by the distribution of the existing remnants.

A satisfactory statement of the original limitations is not possible with our
present knowledge, because we are ignorant of and have no means of determining
the amount of erosion that has taken place from the up[)er surface of these beds.
It is evident, however, fiom the continuity of the lithological characters of preserved
beds where exposed over wide areas, ihat the strata were originally continuous over
the whole of the lower parts of the Sergipe- Alagoas basin, and over many of its con-
siderable elevations as well.

It is easy thus to calculate a part of the erosion that has taken place, but in
view of the impossibility of knowing the oiiginal thickness of these beds, we cannot
make a trustworthy estimate of the whole. It is a noteworthy fact, however, that
in the Sergij)e-AIaguas basin the remnants of the tertiary beds do not extend
beyond the limits of the cretaceous basin into that ul the crystalline rocks. They
end uni'onforniahly against the beds of the Serras d'ltaliaiana and M n;i').i, and only

416 THE CRETACEOUS AND TEKTIARY GEOLOGY

where the cretaceous beds are not limited on their inner margins liy such ranges do
they rest upon or against the cr3\staliine rocks.

Denudation. — The present distribution of the tertiary, the horizontality of its
beds, and the high angles of their exposed mai-gins, all go to show that the denuda-
tion of these rocks has gone on and is still going on at a rapid rate, whatever the
causes. Erosion and denudation are still in progress, and some of the causes, at
least, may be studied upon the ground. Tlie tertiary beds frequently form broad,
dry, and sterile plateaux or table lands known in Brazil as " chapadas " and " tabo-
leiros." In places these plateaux are almost barren of vegetation, and are deeply
scored with gullies, ravines, and gorges as much as two hundred feet in depth. Such
spots are quite common about the mai'gins of these plateaux. It is not an uncommon
thing, however, for hills having more or less vegetation over them to yield to erosion
and to develop such eases as that shown in the accompanying illustration of the
" Ruinas de Palmyra," near Cabo Santo Agostinho. This case is a typical one (y.
Plate III). One is often struck, on looking at these rapidly cutting exposures of
tertiary, with their resemblance in miniature to the topography of parts of the Grand
Canon region of the United States.

The great denudation of the tertiary beds is due to the following five principal
causes :

I. The impenetrability of the surface clays.

II. The sterility of the soil resulting in barrenness of vegetation, and the absence
of the protection arising therefrom {v. p. 370).

III. The unresisting nature of the rocks {v. p. 410).

IV. The great precipitation concentrated within a few months of the year in-
creasing in geometric ratio the abrading and transporting powers of the Avater.

V. The temperature of the water considerably elevated by falling upon the bare
hot surface clays.

I shall si)eak of these causes in their order. (I) The impenetrability of the soil
prevents the pi-ecipitation from soaking into the ground, obliges it to seek its level
along the surface at once and is therefore conducive to Hoods and to corrasion. The
tertiaiy clay through this region being more or less impenetrable, sheds its water
almost perfectly, and thus hastens erosion by floods and freshets whose abrading and
transporting powers are enormous.

It will be observed that in almost all the descriptions of Brazilian tertiary sec-
tions there are alternate sandy and clayey beds. When erosion acts readily upon
the sandy ones they are soon removed until a bed containing sufficient clay to turn
the water readily is reached. This bed then becomes the surface clay of the country

OF THE SEUGIPE-ALAOoAS B^VSIN OF UIIAZIL. 417

shedding the great precipitation of the region and sending it in powerful torrents down
the soft margins of the formation whence enormous quantities of earth are swept
away.

(II) The sterility of the soils derived fi-om the tertiary is notorious. This steril-
ity is due largely to the tendency of the surface clays to turn the rainfall instead of
allowing the water to soak into the ground. But little water is able to penetrate the
ground, and for this reason dry weather is invariably fatal to crops in tertiary soils.
The natural vegetation is remarkably scant, except in the deep ravines. The wiiv
grass, stunted trees, and cacti that grow over these sterile plateaux, or "chapadas"
as they are called, offer but little resistance to the water flowing from the surface.

(III) In the description of sections in the tertiary it may be seen that there are
no beds in the whole column capable of long resisting the ordinary processes of sub-
aerial erosion in the tropics. The beds are usually clays and sands variously commin-
gled. I have seen but one indurated rock in the series — a xcvy hard, glassy quail zitc,
metamorphosed in weathering, but this rock does not appear to form a continuous bed
at any one place, or to occupy a definite position in the series. In such soft and easily
eroded material the heavy rainfalls of Brazil do the greatest possible amount of work.

(IV) Predjiitation. — That })reeipitation in the tropics is not as evenly distrib-
uted through the year as in temperate regions is a well-known fact.

In Brazil the year is divided as naturally into rainj' weather, "tempo de chnvn,^''
and sunny weather, "teinjyo de sol,^'' as in temperate regions it is divided into winter
and summer; but it is important to note also that the rainy season is the hot season,
the " tempo de sol," referring to continuous sunshine.

The rainy season does not set in at the same time in all parts of the country,
but may begin one, two or three months later or earlier in one place than in anolluM-.
This, however, does not all'ect the general results. One of the best records of rain-
fall in Brazil of which I have knowledge is that kept by the St. John del Key Gold
Mining Company at Morro Velho, in the province of Minas. This record covers the
l)eiiod from 1855 to the i)resent. A resume of these observations published in 1880
shows* the average annual lainfall up to that time to be 1G37 millimetres, and that
89 per cent of this, or 1457 millimetres, fell during the six months from October to
March inclusive, while from April until September only 180 millimetres fell.

A series of observations kept at the city of Sao Paulo during the year 187.)t

* See " Relatorio do W. Milnor Roberts sobro o Uio Sao Francisco," Itio ilc Janciri), ISSn. .""1 /.'. r/«^l ,1,- EiK/m.
karid for May, 1880.

t The Uio Newt, February 5, 1880.

A. r. S. — VOL. XVT. .'Ja.

418 THE dlETACEOUS AKD TERTIAllY GEOLO(ir

shows that out of the total rainfall of 1287 millimetres during the year, ll.:2 milli-
metres fell from November to Apiil inclusive.

Observations made during 18S1 at Sao Bento das Lages in the province of Bahia
give a total precipitation of 3984 millimetres for the year, of which 1439 millimetres
fell from March to August inclusive.*

Observations made at Uberaba, province of Minas, from 1880 to 1882 inclusive,
show an average annual precipitation of 15G1 millimetres, of which 12G4 millimetres
fell from October to March inclusive.f

Records of the rainfall in Ceara during the twenty-eight years from 1840 to 187G
show an average precipitation of l-'^4(i millimetres during one half the year, against
143 millimetres during the other half J

These examples are sufficient to give a correct idea of the distribution in
time of the precipitation in Brazil. A rainfall so unevenly distributed cannot fail to
do an unusually large amount of erosion.

On the whole, however, the erosion of these tertiary beds does not appear to be
going on so rapidly as formerly, and indeed I am not disposed to believe that the
extensive denudation of the tertiary beds which has taken place in Brazil is to be
attributed to subaerial erosion alone. It seems probable that the greater part of this
work was done at the time of and immediately after the emergence of these beds
from beneath the ocean, while the beds were even softer than at present, and before
the surface was taken possession of by vegetation. In many places over the tertiary
region where the country is thickly clad with vegetation, erosion is practically nil.
The great width of many of the valleys and the precipitous faces of the tertiary
i-anges that border them suggest that these hills faced the ocean, or were the shores
of bays during the time of the land's elevation from beneath the tert'ary seas.

At many places along the Brazilian coast the ocean is also attacking these terti-
ary beds upon the edges, undermining them and spreading the material of which they
are composed over the ocean's bottom. The bed of the ocean north of Rio de Jan-
eiro is, in most places, covered, not with the glauconite ooze i;sually found along the
shores of continents, but for the most part with I'ed mud carried down by the
streams, or washed away directly by the waters and undertow from the soft red tei-
tiary rocks of the coast.§ The red cliffs, so noticeable to one sailing along the coast
in sight of the shore, are all tertiary, and wherever the water breaks against the bases
of these cliffs, they arc being rapidly cut away.

* V. ItevUla dc Kii<jcnhitri<i for Jlay, 1883.

t V. Hcvisla de Eixjenhiiria, Vol. V. (1883), p. 2.")1.

J F. E isaio Estalislico dc Pomiieu Oo fiouza IJiazil, p. 10.") (in part).

t- V, Challenger Reports ; Narrative, Vol. I, Pari I, pp. 315-217.

OF THE SERGIPE-ALAGuAS BASIN OF BUAZIL. 419

The greater portion of this red mud is derived directlj' from the cliffs along the
shores where the ocean is a more direct agent of erosion, transportation and distribu-
tion of this material than the streams themselves. I make this statement advisedly.
The streams of the region do not now, as a rule, have sufficient current to be such
powerful agents of transportation as they formerly were. In the lower portions of
their courses they have generally cut down to or near their base levels, and are
therefoi-e comparatively powei'less. During the season of I'ains the finer sediments are
washed from high grounds into all the streams, but the Rio Sao Francisco is the only
stream flowing into the ocean in Brazil south of the Amazon, which brings down any
considerable ((uantity of sediment. Such streams as the Scrgipe seldom carry muddy
waters into the ocean, the material brought in suspension from the higher country
being precipitated near the head of tide water, which is usually miles inland.

It happens that nearly all these tertiary beds have ahead}' been worn away from
the immediate neighborhood of the ocean in the Sergipe-Alagoas basin. They arc
well exposed, however, along the coast between Maria Farinha and Parahyba iu
cliffs, some of which are more than four hundred feet in height.

In a few places tertiary beds are exposed along the ocean's beach, having no
cousideiable thickness of their own formation overlying them. In such places they
offer more than the usual resistance to the encroachment of the sea, and though the
beds aie by no means hard, they are worn into fantastic forms by the surf. An ex-
ample of this kind is shown in the plate illustrating the erosion of a tertiary beach
near Rio Formozo in the province of Pernambuco (y. Plate IV).

Horizontality of the Beds. — These beds are, everywhere I have seen them, so
nearly horizontal that no dip is apparent. Exposures have been seen where the beds
appeared, at first sight, to have dips as high as twenty degrees, but upon closer in-
spection they have turned out to be delusive. In other places the writer has seen steep
dips caused by the caving and tumbling down from l)anks of fragments so large that
they might easily deceive the observer into believing them true dips. Pissis gives
three examples of dips iu the tertiary, one of which is as high as 15°.''' The cases
cited are possibly delusive ones, such as are referred to above,

Metamorphism in the Ih-tiarrj. — The fiict that there are compact, glassy quartz-
itcs among these tertiary beds might, if taken alone, lead one to surmise that this
formation had undergone dynamic disturbances. 8uch, however, is not the case.
These quartzites seldom or never occur forming beds continuous over considerable
distances, but arc derived from soft beds of sandstone which become indurated here

• .Mcmoirc sur la position g<;,)logiquc, etc., p. 107.

420 THE CKETACEOUS AND TERTIARY GEOLOGY

and there upon exposure, forming quartzite blocks. As the surrounding portions of
the beds weather away, these blocks are usually left lying loose on the surface where
they exfoliate somewhat under the same influences that hardened them. That this
metamorphism is produced by weathering scarcely admits of doubt. The beds
which are metamorphosed are high ;ip in the tertiary series, the underlying beds being
made up of soft horizontally stratified beds of clays and sands. It is quite evident
that these rocks have not been subjected to any unusual lateral pressure, such as that
which so often produces metamorphism. The jiressure from above is and always has
been less than that upon the underlying soft beds of sand, so that its metamorphism
cannot be attributed to pressure from above. The strongest evidence that this meta-
morphism is a process of weathering is found in the condition of some of the par-
tially metamorphosed masses of sandstone. The writer has seen at three separate
exposures large masses of this glassy qaartzite protruding from banks of soft sand-
stone, which, upon being broken ofi' two or three feet beneath the face of the expo-
sure, showed the uncovered end of the block to be nothing more than sand in piocess
of hardening, the mass becoming harder and harder toward the exposed surface.
When these blocks are entirely separated from their surroundings and lie fully exposed
to sunshine and rain, they become as hard as glass and have a similar conchoidal
fracture. Heated as they usually are by the direct I'ays of the sun during the day
and cooled by radiation at night or by cool rains, their angles exfoliate until they are
almost })erfectl3^ lound. Prof, Hartt at one time thought that these boulders were
brought to this region by glacial action. Although these quartzites arc tertiary,
there are occasional quartzitic beds in the cretaceous also, yet there is but little
danger of confounding them ; those of the tertiary being barren of fossils, while those
from the cretaceous usually contain a few fossils. The hard cretaceous sandstones in
the Riacho de Aroeira approximate in general appearance to the tertiary quartzites.
It would not be safe, however, to assign all the non-fossiliferous quartzites of this
region to the tertiary, for some of the cretaceous sandstones have been sufiiciently
porous to allow percolating waters to dissolve out the fossils, and subsequently they
have been changed to quartzites. I find in my notes the opinion expressed that a
certain bed of quartzite more or less buried beneath black cretaceous limestone soils,
is cretaceous, and uncovered by the decay of overlying beds of limestone.

Tertiary quartzites were observed in the Sergipe basin at the following localities :

]. Sitio da Ribeira, on the Rio Sergipe as one ascends the stream from Aracajii

towards Piutos, on the north-west side, opposite Andorinhas quarries. These blocks

have evidently been derived fioni beds at a higher elevation and have been left here

by denudation.

OF THE SERGIPE-ALAGOAS BASIN OF BKAZFL. 421

2. Oil the hill-tops -west of Maroim, and beyond (from the town) the church
Maroini de Cirau.

3. Fazenda de Sto. Antonio, three-quarters of a mile west of the church. These
(iiKut/iti's are also on the hill-tops.

4. East of Sto. Antonio, and visible from the church of Maroim de Cima is a
hill covered with quartzite.

5. Immediately north-west of Maroim on the hill-top, exfoliating in blocks.
Along- the Estrada Real also north-west of Maroim, a bed one to two feet thick is
exposed. The Santa Cruz chuich north-west of Maroim is on a hill of ferruginous
conglomerate, bare of vegetation.

SUKFACE GEOLOGY.

Post- tertiary. — On the hills west of the Cotinguiba at a sitio near Maroim known
as Sitio de Bclcmges, and again along the hills about the Santa Cruz church north-
west of Maroim, are types of a formation overlying the teiliary. This formation ia
spread over the hills and valleys of the Scrgipe-Alagoas basin and over the adjacent
counti-y in the form of a thin coating of cobblestones, pebbles and sand, sometimes
loose and sometimes cemented into a pudding-stone as much as ten feet in thickness,
and, when exposed, stained black by manganese. It caps the summits of the tertiary
plateaux or their outliers, and it is frequently strewn along down the sides of hills
and accumulated in the valleys. It is not confined to the geographic limits of the
cretaceous or tertiary, but is found further inland and far beyond the present limits of
these formations. It is everywhere more or less irregular in thickness, and nowhere
can it be said to be univcisal or continuous. The writer has seen this material
throughout Sergipe and Alagoas, in Parahyba, and as far inland as the headwaters of
the Kio Ipancma in the interior of the province of Pernambuco, where there is no
remnant of stratified tertiary beds. Between the lower Rio Sao Francisco and the
frontier of the province of Alagoas, and indeed in many parts of the province of Per-
nambuco, this water-worn material is found minfjled in boirs with the remains of ex-
linct, gigantic mammals.

One of the marked characteristics of this post-tertiary formation is that it is
much coarser inland, and grows finer as the coast is approached. The explanation
of this water-woin material seems to be that the tertiary i)eriod was closed by a
dci)ression along the present coast, which carried the beach liue far inland, or that
it was already there. Then followed a gradual emergence, during which the whole
aica MOW covered by this widely distributed water-worn material was passed gradu-

422 THE CERTACEOUS ANB TERTIARY GEOLOGY

ally through the condition of a beach upon which the then loose, angular, surface
rocks of the country were rounded and worn into the boulders, cobbles and peb-
bles which we now find scattered over this region. While the surf was beating upon
and wearing the hard crystalline and metamorphic rocks of the interior it was unable
to produce any very marked effect upon the topography of the country, but when,
in the course of the land's emei-gence, the soft, sandy and clayey beds of the
tertiary were brought within its reach, the work of land sculpture it was able to do
was enormously increased. During the emergence of these tertiary beds they were
deeply eroded, and the mud which originally made part of them was washed seaward,
and their coarser materials were concentrated upon the slowly receding beach. In some
places these accumulations assume unusual proportions, as if they had been brought
together by the gradual beating of waves along a beach, or had been reconcentrated
by later streams. An example of this kind, outside the Sergipe-Alagoas basin, occurs
at the Camassari diamond washings in the province of Bahia (v. Plate V). It would
seem that during this period some of the wider valleys excavated in the tertiary were
formed. The concentration of coarse material left sjiread over the surface of the
remaining portions of the tertiary, and indeed over the whole country rising from the
bottom of the ocean, the thin covering of cobblestones, and pebbles such as we now
have. Later, by the oxidation of the iron in these beds they have been changed here
and there to a ferruginous conglomerate.

The post-tertiary deposit is veiy widespread in Brazil, and has been referred by
both Agassiz and Hartt to glacial action.

o

3
C

o

C3

P

OF THE SERGIPE-ALAGUAS IJASIX OF BRAZIL. 1-3

Part IY.

bibliography.

BiIiliof/rci])Jn/ of the Geology of Sen/ipe-AIar/uas Basin.

AsoNTMOCs Ahticles : A Cominissuo Geologica do Brazil, published iii "O Vulgarisador" newspaper of Rio de
Janeiro, Brazil, November 3, 1877.

The same article was reprinted in " O Novo Mundo," an illustrated periodical published in New York, Vol.
VIII, January, 1878, pp. 18-19.

The article gives a brief sketch of the region explored and the collections made
by the Imperial Survey.

COMSTOCK. T. B.— American Journal of Science, X[, June, 1877, pp. 4GG-473, contains an abridged report by Prof.
Charles F. Flartt upon the work of the Commissao Geologica do Brazil, translated by T. B. Comstock from
the report printed in Portuguese. See Ilarlt, ]{elalorio Pruliuiiuar do Traballios da Commissao Geologica na
Provincia de Pernambuco.

Dehby, Orville A. — O Brazil Gfiographico e Ilistorico ; A Terra e o Homem, por J. E. Wappaeus, Rio de Janeiro,

18S4.

This book iii)on the pliy.sical geoj:frapliy, natural history, etc., of Bi-azil, is the
work of several collaborators, the f^eology (Chap. V, p. 44 et seq.) having been writ-
ten by Mr. Derby. He mentions the provinces in which cretaceous and tertiary
foi-mations arc found, and gives a small map showing their general distribution, and
remarks that the relation between the cretaceous along the coast and that at highei-
elevations is not known.

Deubt, O. a. — Contributions to the Paleontology of Brazil, by Charles A. While.

The descriptive part of this important work, pp. 7-14, is by Mr. Derby, and is
the most important paper yet published upon the stratigraphy of the mesozoic and
tertiary geology of Brazil. The leading facts known to liiin legardiug all the Bra-
zilian cretaceous localities, except those of S.io Francisco do Sid, are given. The
theory advanced by Mr. Derby, however, to the effect that the Ceai'a beds '"appear to
be a distinct series*' because they ri.se to a greater elevation than other kuowu locali
ties, is criticised on pages 408-410 of the present paper.

DEtiny. O. .\. — Contril>ii:(;>s para o Eatndo da Geologia do ViiUe do Rio SAo Francisco, pel") Dr. O. \. jtrihy, piib-
liOicd in llie Ardiivo.s do Muf^eu Nncioiial, Vol. IV. 1879, ]t. 87 et xq.

In counccticin with the "iio\ocr\ of the lower Sao Francisco, Mr. Deiby takes
occasion to sketcli and discuss the cretaceous and tertiary geology of the euliri-

424 THE CRETACEOUS AXD TERTIAKY GEOLOGY

Brazilian coast. He announces the imiwrtant diseoveiy of secondarv beds at Atalho
and Caissara on the S.lo Francisco, above the falls of Paulo Affonso, and expresses
the opinion that they extend to the south along that side of the valley.

These beds are said to resemble those of Bahia, but doubt is expressed regarding
their being the same, because the author finds it " difficult to admit the contempora-
neous deposition of beds at such ditferent elevations."

Gardner, George. — Travels in the Interior of Brazil, principally through the Northern Provinces and the Gold and
Diamond Districts during the years 1830-1841, by George Gardner, F.L.S., London, 1846, 113-147. There
are but few important notes on the geology of Sergipe-Alagoas. On p. 119, he says the rocks at Penedo dip
west, which should read east. On p. 198 et scq., he describes the chalk formation of Ceara. This is substan-
tially a repetition, with less detail, of the article from the Proceedings of the Glasgow Philosophical Society,
of April, 1843, referred to below.

IIartt, Cii. Fued. — Geology and Physical Geography of Brazil, by Ch. Fred. Ilartt, Boston, 1870.

This book contains all sorts of valuable scientific information regarding all
parts of Brazil. The observations of the wi'itcr, who visited Minas Geraes and the
provinces along the coast north of Ixio, are supplemented by a study of almost all the
best authorities regarding the gcogi-aphy, geology and natural history of the Empire,
so that the work represents, better than any other, the state of knowledge of Brazilian
geology at the time of its publication. Following Agassiz, he refers the surface geol-
ogy to the glacial drift, gives excellent descriptions of the tertiary at many localities,
refers the sedimentary beds of the Abrolhos, and the fresh- water deposits of the Bahia
basin to the cretaceous. He touched at Aracaju, Maroim and Penedo, and from
Maroiin took away a small collection of fossils, which lead him to refer the rocks of the
region to the cretaceous. On pages 555 et seq. he gives a resnm6 of the mesozoic
geology of Brazil. The name " Sergipian group " is proposed for the limestones of
Maroiin, and the " Cotinguiban group" for the cream-coloi'ed limestones of Sapucaiy.
The work is illustrated with many carefully executed wood-cuts, among which are
numerous valuable geological sections.

lIiiNDJ'.usoN, James. — A History of Brazil, by Jiuues Henderson, London, 1831.

He mentions the occurrence of dints and limestones in the province of Sergipe.

IIvATT, Ai.i'HF.iia. — Report on the Crelaceous Fossils from Maroim, Province of Sergipe, Brazil, by Alplicus llyiiU.
In Ilartt's Geology and Physical Geography of Brazil, p. 38") ct neq.

Five species of fossils, mostly cephalopods, are described fi-om a collection made
at Maroim by Prof. Ilartt.

Hyatt, I'uOc. Alpheus. — The Jurassic and Cretaceous Ammonites collected in South America by Prof James Orton,
with an Appendi.v upon the Cretaceous Ammonites of Prof. Hartl's collection, by Alpheus Hyatt. Proceed-
ings of the Boston Society of Natural History, Vol. XVII (May, 1875), pp. 365-373, Boston, 1S75.

This paper has a brief note upon BucMceras hartlii Hyatt, described by him in
Hartt's Geology of Brazil, p. 38G, as Ccrat/ies hartlii.

OF THE SERGIPE-ALAGUAS BASIN OF BUAZIL. 425

MoDCHEz, Capt. Ernest.— Les Cdtes du BriSsil, Descriptions et Instructions Nautiques, par M. Ernest Mouchez, Cap-
tain de Vaisscau. Preniifiro Section. Du Cap San Roque a Baliia, Paris, 1874.

Although this work has nothing directly upon geology, it contains many facts of
great value in studying the geology of the northeastern coast, and especially the terti-
ary of Brazil, which is so intimately connected with the present aspect of the coast in
many places. On p. 16 are observations upon the continental shelf; on pp. 20-21,
the currents of the coast and tides and reefs are discussed, while the elevations of a
great many points along the coast are given. On p. 145, he says that gold and dia-
monds are found in Itabaiana, and speaks of a great quantity of "toMrfte combustible"
and the probability of coal being fountl, and adds that *' Cette province est egale-
ment celebre par les richesses fossiles qu'on y rencontre, principalement sur les bords
du Sao Francisco."

WuiTE, CiiAULES A. — Contribui(;oes a Palcontologia do Brazil (com o original cm inglcz), iwr Charles A. Wliitc,
M.D. Archives do Museu Naclonal, Vol. VII, Rio de Janeiro, 1887.

An extract from the Archivos was made of this paper and issued with an an-
nouncement and "errata," by Dr. White, from Washington, D.C., dated January 2,
1888, under the title, "Contributions to the Paleontology of Brazil ; Comprising De-
scriptions of Cretaceous Invertebrate Fossils, Mainly from the Provinces of Sergipe,
Pernambuco, Parii and Bahia."

This work is by far the most important one ever published upon the paleontology
of Brazil. The bibliography of the invertebrate mesozoic paleontology of South
America is given, followed by a brief but comprehensive sketch of the mesozoic geol-
ogy of Brazil, by Mr. Derby.

315 species of mesozoic fossils are described, 170 of which are new. These in-
clude the lauK'Uibranchs, gasteropods, cei>haIo])ods and cchinoderms, collected by the
Imperial Geological Survey in the provinces of Sergipe, Bahia, Pernambuco and Para.
All are accompanied by excellent plates of 445 figures, drawn by McConnell and litho-
graphed by Sinclair, of Philadelphia. In a discussion of the geologic age of the Ser-
gipe beds. Dr. White concludes that, in spite of the Jurassic facies of sev'cral of the
fossil forms, the burden of testimony favors the reference of these beds to the cieta-
ccous.

Bibliojraphy of the Cretaceous and T'.rtiari/ Qeolorpj of Brazil hearing upon that of the

Sei-(/ipe-Alaf/oas Basin,

The mesozoic and tertiary geology of Brazil is so intimately connected with that
of the other portions of South America that the bibliography of one portion must of
necessity include much of that of the other. For thi.s reason some titles are given

A. p. a. — VOL. xvr. ^m.

426 THE CRETACEOUS AND TERTIARY GEOLOGY

here which do not relate directly to the mesozoic or tertiary geology of Brazil. There
are besides many valuable works upon the geology of the River Plate basin, the west
coast and north-eastern South America, by such writers as Darwin, D'Orbigny, Bur-
meister, Castelnau, Humboldt and Spix and Martins, to which one naturally turns for
trustworthy information in studying the geology of the continent. With the excep-
tion of the publications of Spix and Martins, which the writer has been unable to
consult carefully, the most pertinent of these works are referred to, the others are
omitted. The titles given under the preceding head must, of course, be added to
this list.

Agassiz, Louis.^Edioburgh New Philosophical Journal for January, 1841. Description of the Fossil Fishes Collected
by George Gardner in the Province of Ceaiii, by Louis Agassiz.

Prof. Agassiz referred the beds from which they were taken to the cretaceous.

Agassiz. Louis. — Tlie Atlantic Monthly (Boston, Mass.) for July and August, 1866. Physical History of the "Valley
of the Amazon, by Louis Agassiz.

In this article the author expresses the opinion that Ceara and the region north
of that province belong geologically to the Amazon Valley region, describes briefly its
tertiary deposits, and refers them " to the ice period in its earlier or later phases."

To explain its aqueous origin, he postulates a gigantic terminal moraine closing
the valley at its eastern end, behind which these beds are supposed to have been de-
posited in cold fi'esh water.

Agassiz, Prof. Lodis. — Geological Sketches, by Louis Agassiz. Boston, 1886, Vol. II, p. 153 et seq.

The chapter on the "Physical History of the Valley of the Amazons" is the
same as that published under this title in the Atlantic Montlily for July and August,
18(50. .

Agassiz, Louis and Ma.ior Joao Martins da Silva Cootinho. — Sur la Gdologie de I'Amazone, par MM. Agassiz
et Coutinbo, Paris, E. Biot, 1867, 8. E.\trait du Bulletin de la Socieie Geographique de France.

Sub.stantially the same views are here given as are published in the Atlantic
Monthly article rcfeiTcd to above.

Agassiz, Phof., and Mrs. Louis. — A Journey in Brazil, by Prof, and Mrs. Louis Agassiz, Boston, 1868.

On pp. 146-7, are remarks by Prof Agassiz upon the so-called glacial drift at
Bahia, Pernambuco, Maceio, Parahyba and Para. He refers to the fossils found at
Parahyba do Korte, and he discusses the so-called drift phenomena of the Amazon
Valley region. The clays and "sands i-eferred to here as glacial drift are tertiary, and
the watei"-worn material found at the places referred to is the " wash," or concentrated
coarse material left scattered over the region, as the laud rose from beneath the ocean
at the close of the tertiary.

OF THE SERGIPE-ALAGUAS BASIN OP BItAZIL. '127

AoASsiz, Prop. Louis.— Com'es Rentlus de VAcadamie Francsiis, Vol. XVIII, p. 1007.

Letter from Louis Af^assiz to Elie de Beaumont describing the fossil fishes
from Cearu. Seven species are mentioned by him, and he aflirms his beHef in the
cretaceous age of the Ceara rocks.

Allen, J. A. — Notes on the Geological Character of the Country between ChiqueChique, on the Uio de Sao Francisco
and Bahia, BraEil, by J. A. Allen. In Hartfs Geology and Physical Geography of Brazil, pp. 309-318.

A brief but very important contribution to the geology of Eastern Brazil. The
author does not attempt to give the horizons of the geological formations found upon
his ti-ip, but his descriptions enable one acquainted with the geology of the country to
assign them to their various equivalents along the coast.

Allpout, S. — On the Discovery of Some Fossil Remains near Bahia in South America. Quar. Jour. Geol. Soc, Lon-
don, Vol. XVI, Pt. Ill, pp. 2G3-208.

The article is illustrated, and, besides a brief description of the eastern portion of
the Bahia basin about Montserrate and Plataforma, is accompanied by notes on the
fossils by John Morris and Prof. T. Kupert Jones. The vertebrate remains described
by Mr. Allport are figured in four plates.

Anontmocs.— Annual of Scientific Discovery for 1866-7, pp. 270-3.

Agassiz's Lowell Institute lectures are quoted as to the glacial origin of the ter-
tiary of the Amazon.

AsosT.Mons.— Annual of Scientific Discovery for 1871, pp. 24fr-7.

These notes appear to have been taken from Prof Hartt's writings. It is stated
that the cretaceous beds probably underlie the tertiary of the whole Amazon Valley.

Bates, Henuy Walter.— Tlie Naturalist on the Amazons, by Ilcnry Walter Bates. 4th ed., London, 1875.

A few notes are given in this work concerning the table-toi)ped hills of Almey-
rim and the serras north of that point, and upon the parti-colored cliffs about Obidos.

Brown, C. BAnRiNOTON. — Tertiary Deposit of the Soliniuos and Javary Rivers in Brazil, by C. Uiirringlon Brown, with
an Appendix by R, Etheridge. Quarterly Journal of the Geological Society, February, 1879.

Reference is made to the old loess-like river deposits, and several sections of the
tertiary are given. N^oting that the tertiary had already been traced from Loreto,
Peru, to Tabatinga, the author says that he has not seen this terrane further east than
Sao Paulo, 150 miles below Tabatinga and 1.'350 miles from the mouth of the Ama-
zon. It is suggested that the tertiary beds have been disturbed. They are said to
occupy here an area of 300 miles in length by ^O in breadth, and contain both fresh
and brackish water shells.

428 THE CRETACEOUS AKD TERTIARY GEOLOGY

Brown, C. Babkington.— On the Ancient Ri%-er-(leposit of llie Amazony, by C. Barrington Brown. Quart. Jonr.
Geol. Soc, Vol. XXXV, 1879, p. 763 ct seq., with illustralions.

This paper treats principally of quaternary and recent deposits, but some refer-
ences are made to the tertiary, while the sections given indicate its lelations to the
later formations.

BoRMEiSTER. H.— Description Physique de la RepuWique Argentine d'Apres des Observations personelles et i-trangers,
par le Dr. H. Burmeister, Paris, 1876, 3 vols.

The third volume is dated Buenos Ayres, 1870. This work is a translation from
the German. The second volume is upon the geology of the Eepublic, and the third
upon the living and extinct vertebrates. It contains no references to the mesozoic
geology of Brazil, but it is useful in connection with its study.

Caldcleugh, Alexander.— Travels in South America, during the years 1819-21, etc., by Alexander Caldcleugh, 2
vols., London, 1825.

In Yol. I, p. 48, he refers to the discovery of vertebrate remains, apparently
quaternary, near Rio das Contas in the province of Bahia.

Capanbma, GuilHERME S. DE.— Trabalhos da Commissao Scienlifica de ExploraQao, Part 1, Rio de Janeiro, 1862.
SecQao Geologica, pp. 120-143, by Guilherme S. de Capanema.

This commission was made up exclusively of Brazilians, and extensive explora-
tions were undertaken. Dr. Capanema was chief of the geologic section. He
visited ]S"azarcth and the island of Itaparica in Bahia, Parahyba and Ceaia. He
speaks of the cretaceous rocks at Parahyba, hitherto unknown, and says that the cliffs
at Crato, said by Gardner to be chalk, are silicate of alumina.

Chandless, W.— Notes on the River Aqniry, the Principal Affluent of the River Puius, by W. Chandless. Jour.
Roy. Geog. Soc., Vol. XXXVI, 1866, p. 1H> et »eq.

The geology of the region treated of is only incidentally referred to in this
article. The localities are mentioned from which were taken silicified woods, the
remains of Mosasaurus, and of extinct turtles.

COBISTOCK, T. B.— American Journal of Science, 1875, pp. 464-6. Note by T. B. Comstock upon the Work of the Com-
missao Geologica do Brazil.

This note refers principally to the work done by the Brazilian Survey in the
Amazon Valley.

Cope, E. D.— A Contribution to the Vertebrate Paleontology of Brazil, by E. D. Cope. Proceedings of the American
Philosophiciil Society, Vol. I, No. 131, January, 1886, pp. 1-31.

Descriptions of mesozoic fossils from Bahia, Sergipe and Pernambuco. In this
paper Prof, Coi>e attempts to correlate the Brazilian mesozoic beds with those of the
United States. The occurrence of the teeth of one species of fishes {Apocopodon

OF THE SERGIPE-ALAGOAS BASIN OF BKAZIL. 420

se)'icciis Cope) induces him to believe the Maiia Farinha beds of Pernambuco cno
probable equivalents of the Fox Hills of the United States, or to the iMjvstriclitian
cretaceous age. Another species from Sapucary leads him to say (p. 7) that these
rocks " probably belong to the cretaceous." One species from the north-eastern part
of the province of Bahia leads him to refer those rocks to the pliocene-pampiean.

CouTiNHO, JOAO Maktins DA SiLVA. — L'Emboucljure de L'Amazone, par Don Joao Martins da Silva Coutinho. Bui.
leiin de la Societr de G<;ograi)liie, Paris, Octobre, 1807.

The statement is made that the same geology (the tertiary) prevails from the
mouth of the Iluallagua in I'ei-u to Marajo and to Piauhy.

Dauwin, Chaules.— Geological Observations on the Volcanic Islands and Paris of South America Visited During the
Voyage of II. M. S. "Beagle," by Charles Darwin, M.A., F.R.S., etc., London, second edition, 1870.

Though there is but little in this book which deals directly with the mesozoic
and tertiary geology of Brazil, Chapter YIII, upon the elevations of the eastern coast
of South America ; Chapter XI, on the formation of the Pampas ; Chapter XII, upon
the older tertiary of Patagonia and Chile, are rich in suggestions which must be kept
constantly before the mind in studying the cretaceous and tertiary of Brazil. Appen-
dices to the second part of the volume contain descriptions of secondary and tertiary
fossils from South America.

Derby, O. A.— A Bacia Cretacea do Bahia de Todos os Santos, por Orville A. Derby, M.S. Archives do Museu
Nacional, Vol. Ill, third and fourth Trimesters, Rio de Janeiro, 1878, pp. 135-1.58.

In this memoir Mr. Derby brings together all the observations made, up to the
time of publication, upon the fresh water cretaceous basin of Bahia, and the accom-
panying tertiary and crystalline beds. Inasmuch as Mr. Derby has done more work
upon this particular region than anyone else, his paper is the most comprehensive yet
published. Its value has now been greatly increased by the publication of Dr.
White's Invertebrate Paleontology of that basin. The article is in Portuguese, and
has never been published in any other language.

Derbt, O. a.— Revista de Engenharia (Rio de Janeiro), Vol. Ill (1881), Nos. 0, 8, 9, 11, 13. Rcconhecimento
Geologica do Valle do Sao Francisco, por Orville A. Derby, M.S.

The article published in the Archivos do Museu Nacional, Vol. IV, 1879, p. 87
et seq., has all the geological facts more fully discussed.

Derby, O. A. — A Contribution to the Geology of the Lower Amazonas, by Orville A. Derby, M.S. Proceedings of
the American Pliilosophical Society, Vol. XVIII, 1879, pp. 1.55-178.

The known cretaceous localities of the Amazon region arc mentioned and brioHy
described on pp.174-"); the tertiary on pp. 17G-7. A Portuguese version of this
pa{)cr is also published in the Archives do Mu.scu Xacional, Vol. IT, 1877, jip. 77-104,

430 THE CRETACEOUS AND TEUTIARY GEOLOGY

under the title: Contribuisoes para a Geologia da Regiao do Baixo Amazonas, pclo
Prof. Orville A. Derby, M.S.

D'OjiBiGNT, Alcide.— Voyage dans les deux Ameriques, public sous la direction de M. Alcide D'Orbigny, Paris,
1859.

Several references to Brazilian geology as made for pp. 120-185, but as this book
was intended for a popular work of travels, it contains nothing of importance upon
geology. On p. 185, it is remarked, however, that " le calcaire s'y trouve en bouconp
d'endroits."

D'Orbigny, Alcide. — Voyage dans TAmerique Aleiidionale, Geologic et Cartes, Paris, 1843.

The only maps referring to Brazilian cretaceous and tertiary geology are included
in Plate X, Figs. 1, 4 and 5. The text in which these general charts are explained
relates mainly to the geology of the River Plate basin, Patagonia and the "West Coast.
Though this publication represents a great deal of valuable geological details, it con-
tains a vast amount of speculation which, as far as Brazil is concerned, is of doubtful
value. There is nothing directly upon the Brazilian cretaceous and tertiary, but the
latter portion of the work, pp. 209 et seq., " considerations generals sur la Geologic de
TAmerique Meridionale," discusses the dynamic movements and building up of the
continent. Of the cretaceous terranes, he says (p. 233) : " Qu'ils sont tout a foit
inconnus aux regions orientales et centrales de I'Amerique Meridionale." The
mesozoic and tertiary geology of Brazil can best be understoood by a study of that
of the central and western parts of the continent, which he treats at some length, in
the Geologic and Paleontologie of his voyage. On p. 237, he discusses the existence
of the Jurassic in South America.

D'Obbigny, Alcide.— Coinptes Rendus de I'Academie des Sciences, 1843, Vol. XV, p. 771. Considerations generales
et coup d'ceil d'ensemble sur les grands faits geologiques dans I'Amerique Meridionale a ete le theatre, par
Alcide D'Orbigny.

The conclusions of this brief paper are highly imaginary.

Duu.\ND, L'Anmi. — Considerations gijnerales sur rAmazone, par I'Abbo Durand. Bui. de la Soc. de Geograpliie, Paris,
Novembre, 1871.

It is stated that the tertiary seems to be entirely wanting in the Amazon basin ;

that the bottom of the basin is cretaceous, overlain by from seven to forty metres of

clays, above which is a formation of sands, clays and gravels, which are referred to

the "trias ou terrain de vieux gres rouge." A very wide distribution is attributed to

this formation.

Ethekidgk, R.— Notes on the Mollusca collected by C. Barriiigton Brown from the Tertiary Deposits of Solimoes
and Javary Rivers, Brazil, by R. Etheridgc. Quart. Jour. Geol. Soc, XXXV, 1879, p. 83.

An appendix to Mr. Brown's paper, q. v., consisting of specific descriptions of
tertiary fossils.

OT TTIE SERGIPE-ALAGOAS BASIN OF BKAZIL. 431

Gardseu, GEOncE. — Geological Notes made during a Journey from the Coast into tlie Interior of the Province of
Ceard, in the North of limzil, embracing an Account of a Deposit of Fossil Fishes, by George Gardner, Esq.
Edinburgh New Philosophical Journal, April, 1841, p. 75 el »<q.

Hautt, C'h. Fued.— The American Naturalist, Vol.11, March, 18G8, No. 1. A Naturalist in Brazil, by Ch. Fred.
Harlt.

Brief description and illustrations of the geology of the Abrolhos. The same is
given more fully in the author's larger work upon the Geolog}' and Physical Geog-
raphy of Brazil.

Hartt, Cu. Fred.— On the Growth of the South American Continent, by Ch. Fred. Hartt. The Cornell Era (Ithaca,
N. Y.), December 12, 1808.

This paper deals with the dynamic movements of the continent, and gives the
general order and distribution of the various formations,

Hautt, Prof. Cb. F.— Amazonian Drift, by Prof. Ch. F. Hartt. American Journal of Science, April, 1871, pp.
294-296.

The writer expresses the opinion that the horizontally bedded sandstones and
clays of the Amazon region referred by Agassiz to glacial drift are tertiary.

H.^uTT, Prop. Charles Fred.— On the Tertiary Basin of the Maranon, by Charles Fred Hartt. American Journal of
Science, Vol. IV, July, 1872, pp. 53-58.

This article deals with the teiliary beds of the Upper Amazon region.

Hartt, Chas. Fred.— Recent Explorations in the Valley of the Amonzonas, by Chas. Fred. Hartt. Journal of the
American Geographical Society, New York, Vol. HI, 1872, pp. 231-252.

The then known extension of the cretaceous in the Amazon Valley is mentioned,
and there are a few notes upon the tertiary. The author reiterates his belief in the
tertiary age of the Serras of Pai I'l.

Hartt, Chas. Fred.— Preliminary Report of the Morgan E.xpedilion, 1870-71. Report of a Reconnoissance of the
Lower Tapajos, by Chas. Fred. Hartt. Bulletin of the Cornell University (Science), Vol. I, No. 1, Ithaca,
N. Y., 1874, pp. 1-37.

This paper sketches the work done by the Morgan Expedition, and contains a

few unimportant references to the tertiary beds in the Lower Amazon region.

Hartt, Cn. Fred.— Contributions to the Geology and Physical Geography of the Lower Amazonas, by Chas. Ficd.
Hartt. Bulletin of the Buffalo Society of Natural Science, January, 1875, pp. 201-235.

Contains good descriptions and sketches of the tertiary geology of the Lower
Amazona.s.

Hartt, Charles Fred.— Pelalorlo Prclimlnar dos Trabalhos da CommissAo Geologica na Provlnciii de Pcrnnmlmro,
por C'h. Fred. Harlt, Chcfe da mesina Coinmissilo, Rio dc Janeiro, 187.';.

This paper contains a few brief notes upon the cretaceous geology of the prov-
ince of Pernambuco.

432 THE CRETACEOUS AND TEKTIARY GEOLOGY

HoMBOLDT, Alexander von. — Personal Narrative of Travels to the Equinoctial Regions of the New Continent dur-
ing the years 1799-1804, by Alexander Von Humboldt and Aime Bonpland (English translation from the
original French), third edition, London, 1823.

Humboldt and Bonpland did not enter Brazil, and their observations upon the
geology of adjacent territory are referred to as valuable supplementary material in the
study of the geology of Brazil.

Jones, T. Rdpert. — Note on the Fossils Entomostraca from Monserrate (Bahia), by T. Rupert Jones. Proc. Geol.
Soc, Vol. XVI, p. 266.

Descriptions of five species of Cypridse are given, and Prof. Jones says that
they appear to be allied to the recent and tei-tiary species. This note is a part of an
article by S. AUport, on the cretaceous fossils of Bahia. See under All port.

LiAis, Emmanuel. — Climats, Geologic, Faune et Geographic Botanique du Bresil, par Emmanuel Liais. Paris, 1873.

The author of this book traveled extensively in Brazil, and his remarks upon the
geology of that country represent much personal observation. For many of his
statements, however, he gives no authorit3\ Hei-e and there discussions of the
geology of the entire South Amei'iean continent and correlations of terranes of widely
sepai'ated regions are attempted. Chapter V is devoted to the discussion of secondary
geology, and Chapter VI to that of the tertiary and quatei-nary. On p. 186, he says
that fossils collected by M. Meyen from a locality on the vrest coast of South America
and studied by Von Bach contained both cretaceous and Jurassic species. Much
doubt as to the value of the book is caused by a considerable number of serious
errors. What appear to be facts of great value are so interwoven with figments of
the imagination that, to those best acquainted with the geology of Brazil, it does not
merit the confidence one would like to give it.

Maush, Prof. O. C. — Notice of Some New Reptilian Remains from the Cretaceous of Brazil, by Prof O. C. Marsh.
American Journal of Science, May, 1809, pp. 390-393.

These fossils are from the B.ihia basin, and include Crocodilus, Thoracosaurus,
Megalosaurus. Lepitodus scales are reported.

Morris, John. — Note on the Molluscan Remain from Monserrate (Bahia). Quart. Jour. Geol. Soc, London, Vol.
XVI, p. 266.

This note, containing the description of one species of Melania, is part of an
article by AUport. See under Allport.

Peueirv, Fi:r,ipi-n Frvncisco.— U )teiro da Costa do Norte do Brazil dosdc Maceio ate Paiii, por Felippe Francisco
Pereira, Pernambuco, 1878.

This work contains geographical notes of value in this connection. The hills de-
scribed and figured along the immediate coast between Maceio and Para are nearly-
all tertiary, except that of Cape Santo Agostinho, which is of igneous origin.

OF THE SEKGIPE-ALAGOAS BASIN OF BRAZIL. 4.']3

Pissis, M. A. — M^inoirc sur la position Gcologique des Terraine de la Parlic Australe du Bresil, ct sur les Soultve-
mcnts qui, k Diverses Kpoqucs. out Cliangt' le Relief dc cette Contrue, par M. A. Pissis (priiscute a rAcade-
mie das Sciences le 27 Juiii, 18-12).

This i)aper relates mostly to the geology of the crystall'me rocks of Rio de
Janeiro, Sao Paiilo and Minas Geraes. On pp. 3i»7-8, he speaks of the tertiary along
the coast between Rio and Bahia, and on pp. 398-9, on the Bahia basin.

In the plates accompanying this memoir, he gives a section at Monscrrate, Bahia,
while his geological map represents the Bahia fresh water cretaceous basin as tertiary,
with a fresh water division covered by marine beds. On this point, see Mr. Rathbun's
paper upon the geology of Ithaparica. On p. 403 et seq., he discusses the "grands
movements du sol "' and gives a list of dips. Of the tertiary, he says that the marine
tertiary of the- coast is represented by lacustrine beds in the interior.

PoMPEO DE SoczA Brazil, Tuomaz.— Ensaio Estatislica da Provincia do Ceara, por Tliomaz Pompco dc Souza Bra-
zil, 180:i.

This work mentions, besides a list of minerals found in the province, the occur-
rence of cretaceous and quaternary fossils at a number of places (pp. 144 -IGO).

FOBTO Seouuo, Visconde de. — Illsloiia Geniil do Brazil antes de sua Scpara(;,io e Independcnciu do Porlugiil, pelo
Visconde Porto Sogiiro, Hio de Janeiro, without date. Two vols.

In Yol. I, p. 353, it is stated that the location of the city of Parahyba was fixed
by the existence there of a calcareous sandstone, Avhich is in places a true limestone
and in others marble.

Rathbcn, RicnARD.— Sketch of llic Life and Scientific Work of Prof. Cbarles Fred, llartt, by Richard Rallibun^
Proceedings of the Boston Society of Natural History, Vol. XIX, pp. 238-304, 1878.

A list is given of tlu; localities at which cretaceous, tertiary and post-tertiary
rocks occur. A brief sketch is given of the work accomplished by the Commissao
Geologica do Brazil, under Prof. ILutt, and of the geological structuie of the various
regions explored.

Ratudun, Richard. — ObscrvaQocs sobre a Geologia da Ilha de Ilaparica na Bahia deTodosos Santos, por Mr. Richard
Ralhbun. Archivos do Museu Nacional.

This paper deals with the geology of the island mentioned, but comparisons arc
made with the geology of the mainland. Most of the i)aper is occui)ied with a de-
8cri])tioii of the reefs.

Ratiihun, UiniARD. — Preliminary Report on the Cretaceous Lamcllibranchs Collected in the Vicinity of Pcrnaniburo,
Hrazil, t)y Richard Rathbun. Proceedings of the Boston Society of Natural History, XVII, lH7t, pp. 2I1~2.")(>.

This paper describes twelve new species of Lamellil^ranchs of the collection made
in 1870 by the Morgan expedition, under Prof. Hartt. Tlic jm latory j)ortioM of the

A. P. S. — VOL. XVI. 3c'.

434 THE CRETACEOUS AND TERTIARY GEOLOGY, ETC.

paper contains brief notes upon the stratigraphic geology, which are credited to the
notes of Mr. Derby.

Reade, T. Mellard. — Denudation of the two Americas, by T. Mellard Reade, CE., F.G.S. American Journal of
Science, Vol. XXIX, No. 172, April, 1885, pp. 290-300. Substance of Presidential Address to the Liverpool
Geological Society, Session 1884-5.

A part of this paper is devoted to the rate of denudation of the Amazon basin,
and references are made to the nature and distribution of the rocks.

Sampaio, Theodoro Fernandes. — Revista de Ertyenliaria, Vol. VI (1884), pp. 52-54. InformaQSes a respeito dos
caracteres geologicos do territorio comprehendido entre a cidade de Alagoinhas e a do Joazeiro, por Theodoro
Fernandes Sampaio.

Short but valuable notes upon the geology of the region along the railway line
from Alagoinhas to the Rio Sao Francisco. The second chapter treats of the tertiary
region, which is said to extend from Alagoinhas to Agua Fria, a distance of fifty-six
kilometres.

Smith, Herbert II.— Do Rio de Janeiro a Ciiyaba. Kotas de um Naturalista, por Herbert II. Sniilh, Rio tic Janeiro,

1887.

Reference is made, i)p. 10-11, to the evidences of the elevation of the eastern
coast of Brazil (Rio) during the quaternary.

Williamson, B. — On the Geology of the Parahyba and Pernambuco Gold Regions, by E. Williamson. Transaction
of the Manchester Geological Society, Part Vll, Vol. VI.

This paper is devoted to the occurrence of gold and to the geology of the crys-
talline and metamorphic rocks of the region. A valuable note is given on the lime-
stones and the tertiary beds which cover them.

Woodward, Henrt. — The Tertiary Shells of the Amazon Valley, by Henry Woodward. From the Annals and
Magazine of Natural History, for January and February, 1871.

—Challenger Reports. Narrative, Vol. I, Part I, p. 215-217.

The soundings and dredging along the Brazilian coast indicate that the ocean's
bottom is here remarkable for the absence of animal remains and glauconite, and that
it is covered by fine red mud (pj). 215-217).

ARTICLE YIII.

DESCRIPTIONS OF NEW SPECIES OF FOSSILS

FnOM TUE

CLINTON, LOWER IIELDERBERG, CHEMUNG, AND WAVERLY GROUPS, FOUND IN THE COLLEC-
TIONS OF THE GEOLOGICAL SURVEY OF PENNSYLVANIA.

BY GEORGE B. SIMPSON.

Read before the American Fhiloaopbical Society, December 21, 1888.

In determining the species in the collections of Messrs. Hall, Sherwood, Fellows,
and Others, I fonnd a number of species which I could not reconcile with any known
to me, or with any published in the books to which I had access.

Prof. James Hall, of Albany, N. Y., was kind enough to review these specimens
with me, and decided that many of thera were new. Of these new species nineteen
are described in this paper. The eleven other species I had no opportunity to submit
to him, but I have no doubt that they also are species that have not hitherto been
described.

George B. Simtson.

43G

DESCRIPTIONS OF NEW SPECIES OF FOSSILS FEOM THE

INDEX.

Name. Figure. Page.

Ortliis pennsylvanica 1 437

Orthis subcircula 2 437

Cboneles punctata 3 438

Cyrliaa triplicata 4 439

Syringothyris angulata 5 440

Syringothyris randalli 6 441

Meristcllaincerta 7 442

Rliyn. (Stenochisma) loevia 8 443

Rhynclionella medialis 0 444

Rhynchonolla striata 10 444

Aviculopecten a-qiialata 11 44.5

Lyriopecten alternatiis 12 446

Leptodesma leiopteroiilcs 13 447

Leptodesma parallelum 14 448

Ply(,'liopteria obsoleta 1.5 448

ModiomorpUa rigid ula 16 449

Name. Figure. Page.

Modiolopsis subrhomboidca 17 4.50

Goniopbora curvata 18 450

Nucula sinuosa 19 451

Nuciila subtrigona 20 452

Tellinomyd (Pala'oneilo) ciincata 21 453

Tellinomya (Palreoneilo) dimhiuens.... 33 453

Platyceras breve 33 454

Platyceras dorsale 34 454

Platyceras intequalc 25 456

Platyceras mitelliformc 26 456

Platyceras striatum 37 457

Platyceras varians 28 458

Acervularia communis 39 459

Cladopora rectilineata 30 459

Ilomalonotus trentonensis 31 460

ALPHABETICAL LIST.

Pnge.

Acervularia communis 459

Aviculopccten requalata 445

Cbonetes punctata 438

Cladopora rectilineata 459

Cyrtina triplicata 439

Goniopbora curvata 450

Homalonotus trentonensis 400

Leptodesma leiopteroidas 447

Leptodesma parallelum 448

Lyriopecten alternatus 446

Merislella incerta 4 12

Modiolopsis subrhomboidea 450

Modiomorpha rigidula 449

Nucula sinuosa 451

Nucula subtrigona 4,52

Orthis pennsylvanica 437

Page.

Orthis subcircula 437

Platj'ceras breve 454

Platyceras dorsale 454

Platyceras insequale 456

Platyceras uiitelliforine 456

Platyceras striatum 457

Platyceras varians 458

Ptycbopteria obsoleta 448

Rbyn. (Stenochisma) Uevis 443

Rhynclionella medialis 444

Rhynclionella striatii 444

Syringothyris angulata 440

Syringothyris randalli 441

Tellinom3'a (Pala;oneilo) cuneata 453

Tellinomya (Pahconoilo) iliminiieiis 453

( LINTOX, LOWEU IIELUEUIJEUG, ClIEMUXG, AND WAVKKLV GUOUI'S. -A-')!

OKTIIIS PENNSYLVANICA Simpson, n. sp., Fig. 1.

Shell essentially circular, in nearly all the specimens observed, the height and
width being equal ; hinge line short, length equal to half the width of the shell ; car-
dinal extremities rounded ; lateral and basal mar- yms
gins regularly rounded, except in the middle basal
margin of the ventral valve, where there is a slight

/>^V<.

constriction.

Dorsal valw .somewhat gil)l)()us, greatest con- 'jsT ^^^y^,^

vexity a little above the middle; ra[)idly curv- a^s- ttis-3''

ing to the cai'dina! and lateral margins; a little more gradually to the basal margins.
Along the middle of the valve is a flattened or slightly depressed area, narrow at the
beak, gradually growing wider, and compai-atively broad at the base.

Ventral valve. A perfect specimen has not been observed, but gutta-peicha casts
have been taken from impressions of fragments in the rock which probably belonged
to this species. The valve is flattened, or of much less convexity than the dorsal
valve, with a slight elevation along the middle, corresponding to the depression of the
opposite valve.

Siuface marked by piomincnt, subangular, radiating stria?, increasing by bifur-
cation, of uniform size at the margins, where there arc twelve in the space of 5 mm. ;
a .short distance below the beak there are twenty in the same space. The radii are
crossed by fine indistinct concentric strife, which on many specimens are ob.solete ;
also, at irregular intervals, by lines or varices of growth.

On the cast of the dor.sal valve the distance from the beak to the lower margin
of the muscular impression is about one-half the length of the valve. Impression ovate
in outline ; width equal to or very slightly less than the length, with a deep depies-
sion along the middle. On the ventral valve the impression is ovate, bilobed. The
margins are distinctly lobed in four divisions. The greater portion of the speciuiuns
observed have a diameter of Irom 20 to 25 mm.

This species may be distinguished from Orthis impressa, of this formation, by its
smaller size and its more circular outline; from Ortliis tior/n, by its more circular out-
line, less conspicuous sinus and elevation, and ovate muscular impression.

Formation and localities. Chemung gi"oup, Sullivan township, Tioga county ; P.
and E. Railroad, between Ludlow and Wetmore, and Kin/ua creek near west line nf
McKcan county; and at various other localities in Pennsylvania.

ORTIILS SUBCIRCaLA Simpson, n..'=p., Fig. 2.
Shill liioadly oval, nearly circulai- ; height usually about three-fourths the width :
gi'catest width just below the middle; cardinal line shoil, length less than lialf the

4:58

DESCRIPTIONS OF NEW SPECIES OF FOSSILS FROM THE

A.P.S,

iTr7l889i

■width of the shell ; extremities curving outward, the lateral margins abruptly rounded,
and the basal margins broadly rounded ; on the ventral valve slightly constricted at

the middle. Dorsal valve flattened, greatest convexity a little
below the beak, gently sloping to the front and lower lateral
margins, more abruptly curving to the cardinal extremities;
at the middle of the base a shallow depression, which con-
tinues about one-half of the distance to the beak. Ventral
valve more convex, greatest convexity about one-third the length from the beak ;
beak incurved and projecting beyond the area line.

Sui-face marked by conspicnous i-adiating strife, which are sometimes of unifoi-m
size, but usually near the base there are very fine alternating striiB. The large striae
are marked at infrequent intervals by elongate pits or openings. The radiating striae
are crossed by fine, indistinct concentric stria?, and at greater intervals by distinct
lines or varices of growth.

On the casts of the interior of the ventral valve the muscular impression is ovate
in outline, the length equal to more than two-thirds that of the shell; width two-
thirds the length ; bilobed by the callosity of the adductor muscle. The sides are
somewhat distinctly lobcd in three divisions.

This species most closely resembles OrfJiis circulas of this foi'mation, but the
valves are incqual, the outline less circular, the greatest width being below the base,
and is constricted at the base. The muscular impression is larger.

Formation and locality. Clinton group, above fossil ore, from McKee's ore bank,
seven miles north-west of Lewistown, Miftlin count}' ; also, ore mine north of Black-
log creek, Orbisonia, Huntingdon county, Pennsylvania.

CriONETES PUNCTATA Simpsou, n. sp., Fig. 8.

Shell semicircular, plano-convex or concavo-convex ; greatest width at about the
middle of the shell ; height from two-thirds to three-fourths the width ; length of

hinge line usually less than the width of
the shell, but sometimes equal to it and
occasionally greater. The cardinal ex-

it-'

tremities are rectangular, or sometimes
produced in slight acute extensions, rarely rounded ; lateral and dorsal margins reg-
ularly rounded.

Ventral valve, varying fiom moderately convex to gibbous; greatest convexity
at the middle, gently curving to the beak, more rapidly to the latero-basal margins.

CLIXTOX, LOWER IIELDEUUERG, CllEMUNC;, AND WAVEIJLY GUOUPS. 4."]!)

and abruptly towards the hinge line. At the cardinal extremities there is a flattened
area 2 mm. or more in width, gradually growing nari-ower to the beak.

Dorsal valve, variably concave, sometimes nearly flat, at other times following the
contour of the ventral v:ilvo; always with a flat area corresponding to that of the
opposite valve.

Surface marked by rounded or subangular striiv, which increase by bifurcation ;
at the base there are four in tlxe s])ace of 1 mm. The radiating stria' ai-e crossed by
extremely fine concentiic striie, about twelve in the space of 1 mm., and at irregular
intervals by more distinct lines of growth.

On ihe hinge line are evidences of eight spines, four on each side of the beak,
short, slightly oblique.

The inteiior of the ventral valve shows a narrow median ridge, with narrow oval
occulsor muscular impressions. Corresponding to the radiating striae of the exterior
are rows of small conical nodes, the bases of which are nearly in contact. They con-
tinue nearly half way to the beak.

The impression of the interior of the ventral valve has numerous punctai or
holes, caused by the mucronate nodes of the interior ; it presents a peculiar clathrate
appearance, which is sutticient to distinguish it from any other species.

Fonnalion and locality. Lower Heldcrberg gronp, Mansing's quariy, near Ilaz-
ardville, Carbon county, Pennsylvania.

CYRTINA TRIPLICATA Simpson, n.sp., Fig. 4.

Shell more or less triangular, subpyramidal ; hinge line equal to the greatest
width of the shell ; proportional length and width somewhat variable ; length of the

4

r

iC.PS ' Yr.l083.

dorsal valve from one-half to two-thirds the width ; length of ventral valve about
equal to the width ; height of the area greater than the length of the dorsal valve.

Ventral valve quadrilateral, subpyramidal ; most i)romiuent at the beak, which
is variable in elevation, straight, not arching over the area, usually attenuate ; lateral
margins regularly rounded. Mesial sinus narrow at the beak, rapidly widening as it
extends forward, and becoming deep ; where it reaches the margin the shell is pro-

440

DESCnil'TIONS OF NEW SPECIES OP FOSSILS FKOM THE

diiced in a conspicuous sublinguiform extension. Sides of the sinus flat, abruptly
sloping, making the bottom angular. Area large, triangulai", varying in height ;
sometimes nearly equal to the width of the shell, at other times not more than half
the width ; slightly oblique ; striated in both dii*ections ; the longitudinal striae dis-
tinct ; the vertical striae much fainter and frequently obsolete ; fissure moderately
narrow and closed by a convex pseudo-deltidium.

There are three or four plications on each side of the sinus, the two bounding the
sinus being much more prominent than the others.

Dorsal valve semicircular or subtrigonal in outline; length from one-half to
two-thirds the width. Mesial fold broad, prominent, extremely elevated in front,
bounded by broader furrows than those between the plications. There are two plica-
tions on each side of the mesial fold, the two plications adjacent to the fold being
much more i^rominent than the others ; on some of the sj^ecimens nearly as promi-
nent as the fold itself, the other two plications being obscure.

The surface of both valves is ornamented by very fine and delicate concentric
striae, which are obsolete on all but the perfectly j^reserved specimens. There are also
more conspicuous lines of growth which are most prominent on the anterior portions
of the shell.

This species most closely resembles Cyrtina JiamiUonensis, but may be distin-
guished by the two prominent plications adjacent to the medial fold; this feature will
serve to distinguish it from other species of the genus at present known.

Formation and locality. Chemung grouj), three miles north-west of Warren,
in Warren county, Pennsylvania.

SYRINGOTirf RIS ANGULATA Simpson, n. sp., Fig. 5.

This species closely resembles 8. randalli ; but the specimens are usually of
smaller size; the mesial sinus and fold proportionately narrower; the cardinal ex-
tremities angular and frequently attenuate.

J.

APS, Tr. IS39.

Formation and locality. AVaverly group, near Warren, Warren county, Penn-
sylvania.

CLINTOX, LOWER IIKLDEKBKUG, CllEMUJS'G, AND NVAVEULY CJliOUPS. 441

SYRINGOTIIYRIS RANDALLI Simpson, n. sp., Fig. 6.

SliL'U transversely somi-elli])tical or semi-circular; vcntricosc, becoming gibbous
with age; k'ugth usually about one-half the width, but sometimes three-fourths;

VIII f.

APS. Tr. 1889.

hinge line straight, length equal to the greatest width of the shell. Cardinal angles
more or less rounded, not attenuate.

Ventral valve with a high vortical or slightly sloping cardinal area, Iroin the
apex of which the sides of the shell curve outward to the antcro-basal margins.

A median sinus begins at the beak, rapidly widening as it extends forward, and
becoming deep, with abruptly sloping sides ; where it reaches the margin the shell is
produced in a conspicuous linguiform extension.

The deltidial aperture is covered foi- about one-half its length from the beak, by
an arched transverse callosity or pseudo-dcltidium. The edges of this callosity unite
with the strong dental lamelhe, which divide the rostral portion of the shell into three
chambers; and from the inner posterior surface of the callosity extends the syringo-
thyi-al tube, which is unusually broad near its posterior extremity, but tapers rapidly
to an open termination, sloping into the internal cavity. This tube is split for its
entire length along its outer surface, and appears to have been thickened and fdlcd in
its posterior portion with the increasing age of the animal.

Dorsal valve convex, greatest convexity at about one-third the length of the
shell from the beak ; convex to the cardinal line, becoming somewhat (lattoned at the
cardinal extremities; gradually curving to the lateral and basal margins. Mesial
fold narrow at the beak, raj/idly widening and becoming pi'ominent below; pioduccd
at the margin corresponding to the linguiform extension of the ventral valve.

Surface of the valve ornamented by from forty to sixty costaj, which occur lioth
on the sides and the siiuis. liadiating .striae crossed by concentric stri.i', wliicii, in
the specimens observed, are most conspicuous on the mesial fold and sinus. There
arc also lines of growth, which are usiuilly the .strongest on the anterior portion of

A. r. s. — vol,. \vi. .'}i>.

442 DESCRIPTIONS OF NEW SPECIES OF FOSSILS FKOM THE

the shell. Horizontal lines of growth, without vertical cross lines, are conspicuously
developed on the cardinal area.

On the ventral valve the muscular scars are strongly developed, and extend over
nearly two-thirds the length of the shell ; ovate in outline, the width being equal to
three-fourths of the length ; the area being largely occupied by the cardinals, between
which lie the narrow linear adductors. The casts of the dorsal valve show the marks
of the deeply striated cardinal process and elongate tooth-sockets. In casts of the
ventral valve the whole ujDiier portion and the area are marked by numerous irregu-
larly disposed prominent pustules.

From the external characters alone it would be impossible to separate this species
from Spirifera disjuncta, but the intei"nal differences are generic.

Formation and locality. Chemung group, near "Warren, Warren county ; and at
Union City, Erie county, Pennsylvania.

MERISTELLA INCERTA Simpson, n. sp., Fig. 7.

Shell subrhomboidal, greatest width at or a little below the middle ; length of
the ventral valve equal to the width; of the dorsal valve, slightly less. Yalves con-

'fV^ 2' vex, the ventral valve being the most gibbous. An-
4 '>. tero-basal margins gently curving outward, at the
* , middle abruptly rounding and the front produced in

■ '. ■ \ / a broad extension.

APS. Trisas.^ Ventral valve the more convex; gi-eatest con-

vexity a little above the middle, abruptly curving to the cardinal mnrgins, and more
gradually to the front. A comparatively deep, broad sinus extends from the beak to
the base, forming one of the most conspicuous features of the species. Umbo promi-
nent. The beak is broken away on the specimens observed, but enough remains to
show that it was rounded, closely incurved, nearly at right angles to the plane of the
axis.

Dorsal valve less convex than the opposite, greatest convexity above the middle,
regularly curving to the antero-basal margins, elevated towards the base into a mesial
fold, which is much less conspicuous than the corresponding depression of the ventral
valve; beak small, incurved, lying below that of the opposite vah^e.

The general aspect of the surface is tb.at of a smooth shell with a few strong
lines or varices of growth. There are indications of radiating stria?, and it is possi-
ble that specimens in a better condition of preservation would show both radiating
and concentric striie.

CLFNTON, LOWER HELDEKBERG, CHEMUNG, AND WAVERLY GROUPS. 4-43

The form of this species is very simihir to that oi Meristella hella, of the Lower
Helderberg group, but that species has a depression on both the ventral and dorsal
valves, while this species has a fold on the dorsal valve. The subrhomboidal form
distinguishes it from any species of the Upper Helderberg gi-oups.

formation and locality/. Chemung group, near Warren, Warren county, Penn-
sylvania.

RnYNCHONELLA (STENOCIIISMA) L.EVIS Simpson, n. sp., Fig. 8.

Shell ovate or subtrigonal ; valves subequally convex in young shells ; in
older shells the dorsal valve usually the most gibbous. The width is slightly less
than the height ; greatest width about two-thirds the length from the
beak ; raai'gins from the apex to this point slightly curved outward,
nearly straight, then somewhat abruptly rounding, and at the base pi'o-
duced in a slight extension.

Ventral valve depressed convex, slightly gibbous at about one-third ^■''■^- ''''■'^^■
the length from the apex, curving abruptly to the cardinal and antero-basal margins,
more gradually to the lower lateral margins, becoming depressed in a shallow sinus,
which commences at about one-third to one-half the length from the beiik; beak
slightly inciu'ved, strongly projecting beyond the beak of the opposite valve.

Dorsal valve convex, greatest convexity at or a little below the middle, gradually
curving to the apex and cardinal margins, more abruptly to the lateral margins ;
mesial fold commencing at a little above the middle and often becoming prominent at
the front; beak small and nearly straight.

Surface marked by from ten to twelve rounded or subangular plications, wliicii
become obsolete on the upper portions of the valves. The mesial fold is composed of
three plications much more piominent than the others. In the sinus there are three
plications smaller tliiiii the others; the two plications bordering the sinus larger ilum
those on other portimis of the shell.

The specimens observed have a length of 9 or 10 mm., and a widlli of IVoni 7 to
9 mm; a transverse section broadly oval, having a width of 5 or G mm.

This species is easily distinguished from any other of this formation at present
known. The outline is very similar to some of the forms of Ji. (Slenocliisma) exhnin,
of the Chemung group, but the shell is smaller, the plications of the foUl and siiuis
are fewer and the plications become obsolete on the upper half of the valve.

Formation and locnUty. Clinton group, limestone, two miles south-west of litll's
mills, Blair county, Pennsylvania.

444 DESCRIPTIONS OF NEW SPECIES OF FOSSILS FROM THE

RHYNCHONELLA MEDIALTS Simpson, n. Pp , Fig. 9.

Shell broadly oval or subti-igonal in outline ; height a little less than the Avidth ;
greatest width about two-thirds the length of the shell from the beak. Margins from
the apex to the widest portion of the shell nearly straight, then abruptly rounding
and slightly produced in front in a broad extension.

Ventral valve convex at the sides, depressed in the middle ; mesial sinus com-
mencing at the apex and growing wider to the base, where it occupies fully one-half
of the width of the valve.

Dorsal valve unknown.

Sui'face marked by about twenty i^lications, of which eleven occupy the mesial
siiins, niid are smaller than those on other portions of the valve, there being five iu
the same sjDace occupied by three of the others ; plications
rounded or subangular. There are also very fine radiating striae
covering the plications, four or five in the space of 1 mm. The
plications are ci'ossed by lines oi* varices of growth. ]S^o con-
^P5 ""^-iAikiiii-^jjg^ centric strije have been observed, though they may occur on more
perfectly preserved specimens.

The specimen from which the above description was taken has a height of
25 mm., and a width of 30 mm.

This species is associated with JRliyncIionella , but may be reodily dis-
tinguished by its size and the much laiger number of plications in the mesial fold.
The only species approaching it in that respect is li. (S.) venubtula, of the Hamilton
group, but they are so unlike in other respects that there will be no difficulty in dis-
tinguishino: them.

Formation and locality. Waverly group, near Warren, Warren county, Penn-
sylvania.

ELIYNCnONELLA STRIATA Simpson, n. sp., Fig. 10.
Only the ventral valves of this species have been observed, but they differ so
much from known forms that it is necessary to consider them
as belonging to a new species.

Shell subtriangular ovate, apex pointed ; length and width
about equal. Margins from the apex to below the middle
nearly straight, broadly rounding below, and at the base pro-
duced in a broad extension.
^■^■^' Tr.issa Ventral valve convex at the sides, depressed at the middle.

Mesial sinus beginning near the apex and continuing to the base, becoming very
broad as it ajiproaches the front.

CLFNTON, LOWER HELDERBERG, CHEMUNG, AND WAVEHLY GROUPS. 445

Surface marked by thirteen plications, of which five occupy the mesial sinus, the
thi-ee central ones being larger than the outer ones, but all smaller than those on
tlie other portions of the shell ; plications subangular or angular. There are also
very fine radiating striie on the plications, three in the space of 1 mm. The plica-
tions are ci'ossed by fine concentric sti'iae, which are most prominent on the fi-ont of
the valve, becoming obsolete above, and also by stronger imln'icating lines or vari-
ces of growth.

The specimens observed have a height of about 35 mm.; width just below the
middle equal to the height.

This species may be distinguished from any other of this formation by its large
size, deep sinus and radiating striations. This species closely resembles lihyncho-
nella missoitriensis of the Kinderhook group, but the shell is larger, the apex more
pointed, giving an angular appearance to the upper portion of the shell, and the
greatest width is below the middle.

Formation and locality. Waverly group, four miles south-west of "Warren,
AVairen county, Pennsylvania.

AVICULOPECTEN .-EQUALATA Simpson, n. sp., Kig. 11.

Shell of medium size, equilateral, not oblique, nearly circular ; length a little
more than the width ; basal and lateral margins below the ears regularly and broadly
rounded.

Left valve flattened; greatest convexity a little more than one-fourth the length
of the shell from the beak ; very gradually sloping to the \ ~ "~~~' "^ i

lateral and basal margins. '"'^ ^^

Kight valve unknown, /^r;

Hinge line straight; length a little more than two- ,
thirds the width of the shell, central, not extending as
far as either lateral margin.

Beak obtuse, I'ounded, straight ; umbo ]>rominent,
well defined by its slightly gibbous form and its rapidly
sloi)ing aides. \, . „c ^r >«„

' o "v.' A ns. Tr. 1809

Ears subequal, narrow, triangular, and each separated from the body by a nar-
row sinus; lateral margins convex.

Surface ornamented by narrow but distinct rays of essentially unil'onu size ; at
the base there are six or seven in the space of 5 mm. ; a little above the middle of
the shell there ai'c twelve in the same space ; interspaces wider than the rays. The

/

446 DESCRIPTIONS OF NEW SPECIES OF FOSSILS FROM THE

rays are crossed by fine, concentric, ci'ennlating stripe, which become fasciculated on
the anterior ear. IS^ear the base there are frequent concentric undulations.

The specimen from which the description was made measures as follows : height
50 mm. ; width 50 mm. ; length of hinge line 35 mm.

Formation and localitij. Chemung group, one mile north of Warren, Wai-ren
county, Pennsylvania.

LYEIOPECTEN ALTERNATUS Simpson, n. sp., Fig. 12.

Shell large, broadly ovate, nearly circular ; height about equal to the longitudinal
diameter ; basal and lateral margins i-egularly rounded.

Left valve moderately convex ; greatest convexity above and a little anterior to
the middle, gradually sloping to the base, more abruptly curving to the cardinal and
"' '" " ■ - ' , antero-basal margins.

/ Right valve unknown.

irTT^^ i^jf Hinge line straight, length a little more

; 1 than one-half the transverse diameter of the

; \ \ valve.

1^'//// Anterior ear short, triangular, not well de-

• fined; posterior ear triangular, flat; limits not
/< v>l well defined except near the beak; margin
</, v./ slightly concave, nearly straight; extremity.

v., V 1

V • , ./ angular.

X^: , ■ / °

X ; ', . •^'' Surface marked by more than thirty com-

■^-, Va>^

APS Tr i?89 "*-*^ paratively strong, sharply rounded rays ; between

adjacent prominent rays ai-e usually three smaller rays, the central one of which is

larger than the others, occasionally nearly as prominent as the principal ones ; the

other two are much finer. On and ne ir the posterior ear, and also near the anterior,

this disposition is not conspicuous, the radii being more nearly of the same size. The

radii are crossed by occasional lines of growth. .No concentric striae have been

observed.

This species somcvvliat resembles Lyrlopeden tricostatas of this formation, but
may be distinguished by its more nearly circular form, by the greater number of and
less prominent radii, antl the sti'oiiger central one of the three intermediate stri:e ;
from L. magnificns and L. mricrodontus by its smaller size and the decided difference
in the ornamentation of the surface.

Formation and locality. Chemung group, one mile north of Warren, Warren
county, Pennsylvania.

CLINTON, LOWER IIELDERBEIUJ. CIIEMUNO, AND MAVEKLY CUOUPS, 447

LEPTODESMA LEIOPTEEOIDES Simpson, n. sp., Fig. 13.

Shell above the medium size, subrhomboidal ; body elongate ovate, oblique at an
angle of about fifty degrees to the hinge line; height a little less than two-thirds the

Vlllg..

\

\

'->

1

^--

A'r.S. trl'iis.

length ; antebyssal margin rounded, much constricted at the b3'ssal sinu.«, then broadly
rounding, somewhat abruptly recurving at the postbasal extremity.

Left valve gibbous above the middle, becoming flattened on the jjostero- basal
portion. Kight valve unknown.

Hinge line straight, length unknown, as a perfect specimen has not been observed.

Beak subanterior, prominent, oblicjue, directed forward; uiiihonal region gib-
bous, descending more abruptly on tin; posterior than on the anterior side.

Anterior end short, angular at the extremity, rounded below, separated from the
body by a distinct, nearly vertical byssal depression. Wing narrow-, triangular,
joining the body at a little more than one-third the length of the valve from the
postei-ior extremity ; margin oblique below, becoming deeply concave and abruptly
curving backward, and probably produced in a spiuiform extension.

Siufaee marked by concentric stria' which are most distinct on the wing near the
body, being often fine, sharp, and lamellose in appearance; frequently on the body
becoming fasciculated and producing an undulated surface. The wing is sepa-
rated from the body by a well-delined, naiTow groove, which regularly curves inward
from the beak to the junction of the wing with the body.

The specimens vary in pro|)ortional height and length ; two left valves having re-
spectively a length of GO and 50 mm. and a height of 40 and 45 mm. The width
of the body of the longer specimen at the junction of the wing with the body is 30
mm., of the shorter one 35 mm.

The specimens usually have somewhat the appearance of the genus Leiopteria.

Some of the specimens resemble Leptodesma billinf/si of this foiniation, l)Ut may
be distinguished from that sjjecies by the greater width of the lower ])ortion of the
body, the larger wing and the very distinct separation of the body and wing.

448 • DESCRIPTIONS OF NEW SPECIES OF FOSSILS FROM THE

Formation and locality. Chemung group, near Warren, Warren county, PL-nn-
sylvania.

LEPTODESMA PARALLELUM Simpson, n. sp., Fig. 14.

Shell small, slightly oblique; body ovate; the basal margin frequently nearly
parallel with the hinge line; length usually a little more than twice the height;
t""^i_K'-~:s=n.-"-- anterior margin rounded; basal margin nearly straight, a little con-

^^^. ' :j stricted anterior to the middle; abruptly recurving at the postei'O-

fAlRs.'Tr.isas. basal extremity ; the posterior margin being obliquely truncate.

Left valve convex, greatest convexity at the nmbonal region, becoming flattened
at the postero-basal portion.

Right valve a little moi-e convex than the other but not gibbous.

Hinge line straight, length about three-fourths that of the shell.

Beak about one-fourth the length of the shell from the anterior end, prominent,
directed forward ; umbonal I'egion convex, descending much more abruptly on the
posterior than on the anterior side.

Anterior end short, angular at the extremity, rounded below. A depression ex-
tends from near the beak to the basal margin a little anterior to the middle. Wing
very narrow, triangular, joining the body nearly at the posterior extremity ; margin
of wing straight, oblique ; extremity obtusely angular. The wing is distinctly sepa-
rated from the bodj^

Surface marked by concentric striae, which fiequently fasciculate on the anterior
portion of the shell. There are also concentric undulations.

This species most closely resembles Leptodesma jyropinqiim, but the wing is more
distinctly separated fi-om the body, and the body is much less oblique, the basal
margin being sometimes nearly parallel with the hinge line; this latter feature will
serve to distinguish it from any other species at present known. When a little
covered by the rock it might easily be mistaken for some form of Sphenotus.

Formation and locality. Chemung group, hill north of Warren, Warren county,
Pennsylvania.

PTYCIIOPTERIA OBSOLETA Simpson, n. sp.. Fig. lo.

Shell of medium size, subrhomboidal in outline; body ovate oblique, at an angle
of about forty degrees to the hinge line; height about three-fourths the length.
Anterior margin abruptly rounded ; basal margin curved outward at the extremities,
concave at the middle ; posterior margin somewhat abruptly' recurved.

CLINTON, LOWER UELDERBERG, CHEJIUNG, AND WAVERLY GROUPS. 449

Right valve unknown. Left valve moderately convex; greatest convexity at
the unibonal retrions.

Hinge line essentially straight; length a little less than that of
the body.

Beak situated at about the anterior fourth of the shell, small, (\
extending beyond the hinge line. A shallow broad sinus extends
from the beak to the basal margin a little anterior to the middle. *ps t-

Anterior extremity acuminate, margin rounded.

"Wing triangular, joining the body at the posterior extremity ; margin for a short
distance straight, then abruptly curving forward, and just before reaching the cai-
dinal line curving upward. Wing convex ; flattened immediately below the cardinal
line; separated fi'om the body .by a narrow but conspicuous depression.

Surface ornamented by faint radiating strite, which arc obsolete, except on i)ei--
fectly i)reserved specimens, and also by fine concentric striie, which frequently become
fasciculate on the anterior portion of the shell.

This species may be distinguished from any other at present known by the slight
obliqueness of the body of the shell, the basal margin being nearly parallel with the
hinge line; the margin is decidedly convex; in nearly all other species it is straight
or concave ; the wing extends beyond the body of the shell. The outline is very
similar to that of some forms of the genus Cypricardinia.

Formation and locality. Chemung group, hill north of Warren, Warren county,
Pennsylvania.

MODIOMORPHA RIGIDULA Simpson, n. sp., Fig. 16.

Shell of medium size or smaller, subquadrangular in outline ; height a little more
than three-fifths the length of the shell ; basal margin regularly and gently curving
from the anterior to the postbasal extremity ; posterior margin gently ..i.y ^r-"^ -
curved, slightly o1)lique, sometimes neaily at right angles to the basal
margin; cardinal line essentially straight ; anterior rounded al)rui)tly, '/

extended, without limitation by a sinus. ""' -'

Beaks a little more than one-fourth the length of the shell from the anterior enil;
umbonal ridge prominent, extending from the beaks to the postbasal extremity.
Valves convex towards the basal margin, becoming gibbous above the middle and in
the umbonal region ; po.sterior slope convex near the beaks, becoming fiattened as it
approaches the posterior margin.

Surface marked by concentric stritc which frequently become obsolete on poriion.s
of the shell.

A. p. s. — VOL. xvr. yK.

450 DESCRIPTIONS OF NEW SPECIES OF FOSSILS FROM THE

On casts of this species the palllal line is sometimes so strong as to give a
distorted appearance to the specimen.

This species may be distinguished from Modiomorpha rigida, of this formation,
by its greater gibbosity, the less oblique posterior margin, less clearly defined umbonal
ridge, and the more prominent beaks.

Formation and locality. Chemung group, Tioga village, Tioga county, Penn-
sylvania.

MODIOLOPSIS SUBRHOMBOIDEA Simpson, n. sp., Fig. 17.

Shell of medium size, rhomboid ovate in outline ; length twice the height ; basal
margin slightly convex along the middle, curving to the extremities ; posterior mar-
la.. ^- gin abruptly rounded below, somewhat more gradually recurving

\--^''-" :^ to the cardinal line ; cardinal margin slightly arcuate ; anterior

Kj -^^^TtlS^ . ii/ margin sharply rounded.
/'." ""^ Tri£89 Valves flattened, greatest convexity at the umbonal ridge.

Hinge line slightly oblique, extending a little more than two-thirds the length of
the shell.

Beaks appressed, situated about one-fourth the length of the shell from the
anterior end ; umbonal ridge not distinctly defined ; posterior slope rounded, becom-
ing flattened just before reaching the cardinal line.

Surface marked by fine concentric linee, and at irregular distances apart by
varices of growth.

The anterior muscular impression is moderately large, well marked, and situated
just within the anterior margin below the beak. The best preserved specimen has a
length of 24 mm., and a height of 13 mm.

This species may be distinguished from M. suhcarlnatus by the less clearly
defined umbonal ridge, the somewhat arcuate hinge line, and the absence of a constric-
tion in the basal margin.

Formation and locality. Clinton shale, above fossil ore, at McKee's ore bank,
north-east of McKee's house, Ferguson valley, seven miles north-west of Lewistown,
Milllin county, Pennsylvania.

GONIOPIIOllA CURVATA Simpson, n. sp.. Fig. IS.

Shell trapezoidal, medium size or smaller ; length usually about twice the height,
sometimes a little less; anterior end i-apidly declining from the beaks, abruptly
rounded below; basal margin gently rounding, sinuate a little anterior to the middle;

CLINTON, LOAVER HELDERBERG, CHEMUNG, AND WAVEKLV GROUPS. 4.")1

posterior margin obliquely truncate; cardinal line very slightly oblique, straight;
hinge line about two-thirds the length of the shell.

Valves convex below the umbonal ridge, gibbous in the umbonal region; um-
bonal sIoj)e flat or a little concave. vmt.^ /-^W'^-^^^^^i:-^

Beaks at the anterior end, acute, incurved; umbonal '' ,k

ridge very prominent, angular, extending from the beak to

the basal extremity, curved. Along the middle of the pos- Atj i..^}

teiior slope is a low rounded or subangular ridge, moderately conspicuous.

Test marked by concentric stria?, which become fasciculate on that portion of the
shell below the umbonal ridge. The test is raised in a crest along the umbonal
lidgc.

Anterior muscular impression situated at the anterior edge, ovate width two-
thirds the length ; pallial line near to and parallel with the basal margin, appearing
as a shallow groove on the casts ; posterior muscular impression not observed,

A specimen of medium size has a length of 40 mm., and a height of 18 mm.
Other specimens preserve about the same pioportion.

This species closely resembles Goniophora truncata of the Hamilton gi'oup, but
■ the hinge line is much longer, and the posterior margin correspondingly less oblique.
From GoniopJioi-a chemungensis it is distinguished by its smaller size, more distinctly
curved umbonal ridge, and the ridge along the middle of the umbonal slope; from O.
jyerangtihi by the less angular form of the shell, the much less oblique posterior mar-
gin, and the ridge on the posterior slope ; from O. suhrecta by its much smaller size
and the ridge on the posterior slope,

Fm'malion and locality. Chemung group, near Warren, "Warren county, Penn-
sylvania.

NUCULA SINUOSA Simpson, n. sp., Fig. IS).

Shell small, ovate cuneatc in outline, subnasute behind; height varying from
one-half to two-thirds the length, usually slightly more than one-half; anterior and
middle portions of the basal margins rounded, somewhat ab- va
ruptly constricted towards the posterior end ; posterior margin ' i

obliquely truncate; anterior end abruptly rounded; cardinal \
line, anterior to the beak, sharply declining ; more gradually
sloping to the iwsterior. A

Valves slightly convex, greatest thickness of the shell a
slioit distance below tlie iitnlio.

452 DESCKIPTIONS OF NEW SPECIES OP FOSSILS FKOM THE

Beaks from one-third to three-fifths of the leno^th of the shell from the anterior
end, extending above the hinge line, compressed, sharp, not prominent,

Umbonal ridge distinctly defined, subangnlar, with a shallow depression below
it, which is most conspicuous at the postbasal margin, becoming obsolete on the
upi^er half of the shell ; posterior slope marked by fine, sharp, slightly divergent
strife. There are also fine concentric striations, which on the specimens observed are
obscure, the surface appearing smooth.

Hinge line marked by frequent ci-enulations.

Three specimens measured have each a length of 10 mm. ; height varying from
5 to 7 mm.

In the striation of the posterior slope this species resembles Nucula poststrtata,
of the Trenton and Hudson River groups, but may be distinguished from that species
by its less gibbous form, the constriction of the postbasal margin, and the conspicu-
ous depression below the umbonal ridge.

Information mid locality. Clinton group, McKee's ore bank, seven miles north-
west of Lewistown, Mifflin county, Pennsylvania.

KUCULA SUBTRIGONA Simpson, n. sp., Fig. 20.

Shell somewhat variable in form, usually subtrigonal ; length and height about
equal ; basal margin i-egiilarly rounded, not constricted toward the posterior end ;
viii| ,os^'^2 -rt<^^ 2 posterior margin rounded or obscurely truncate;

i^ *^ ■ anterior margin abruptly rounded ; cardinal line

very abruptly declining anterior to the beak, more
A.p.s. Trio;;a. gi'ailually declining to the posterior.

Valves slightly convex, somewhat flattened as they approach the basal margin.

Beaks about one-third the length of the shell from the anteiior end, not promi-
nent, compressed, extending above the hinge line ; umbonal I'idge obscure, very
slightly arching upward ; posterior slope very narrow, rounded.

Surface marked by fine concentric stria? and occasional varices of growth. The
concentric 8tri:e are often very obscure, the shell appearing essentially smooth.

Hinge line, postei-ior to the beak, marked by a row of fine transverse teeth.

Three specimens representing the extremes in form measure respectively 10, 4
and 4 mm. in length, and 10, 3 and 2 mm. in height. Larger specimens occur.

This species may easily be distinj^mishcd from iV. sinuosa by the absence of a
constriction in the basal margin, furrow below the umbonal ridge and striie on the pos-
terior slope.

CLINTON, LOWER nKLDERBEHG, CHEMUNG, AND WAVERLY GROUPS. 45^^

Fcyrmation and localilij. Chemung gi'oiip, north of Blacklog creek, Orbisonia,
Huntingdon county, Pennsylvania.

TELLINOMYA (PAL.EONEILO) CUNEATA Simpson, n. sp., Fi^L'. 21.

Shell small, ovate cuneate in outline; length twice the height; basal margin
broadly rounding, becoming constricted or arcuate at about one-third the length of
the shell from the antei'ior end; posterior margin short, obliquely t*-.
truncated; cardinal line essentially straight, sloping at nearly the ^j ^ ' '''w
same angle anteriorly and posteriorly to the beaks ; anterior end
large and regularly rounded.

Greatest convexity of the valves above the middle nnd in the umbonal rcirion.
Beaks about central, slightly incurved, extending a little above the hinge line ; um-
bonal ridge clearly defined, subangular ; posterior slope declining regularly aiid
abruptly to the cardinal line, lielow the umbonal ridge there is a broad shallow
depression, extending from near the beaks to the base and constricting the basal
margin.

Surface marked by strong, lamellose, concentric stria? at regular distances apart,
and by very fine concentric lines between the lamellose striic.

On the specimens observed there are eight or nine transverse teeth on each side
of the beak.

The specimens measured have a length of 12 mm., and a height of G mm.

This siiecies may be distinguished from T. (P.) dimimtens of this formation by
its smaller size, more distinct lamellose striations and the less abrupt constriction of
the posterior portion.

Formation and localitij. Clinton group, seven miles north-west of Lewistown,
Mifflin county ; and north of Blacklog creek, Orbisonia, Huntingdon county, Penn-
sylvania.

TKLIJNOMVA (PAL/EOXEILOi DIMINUENS Simpson, n. sp., Fi-. 22.

This species is very similar to the preceding, but is usually larger, the specimens
obsei-ved having a length of from 23 to 2.j mm., and a .
height of 11 or 12 mm. The posterior portion is more
abruptly constricted, the lamellose striations are not so
])rominent and are more closely arranged.

Fomnation and locnlUij. Clinton gnmp, McKee's ore .
bank, north-east of McKee's house, Ferguson valley, seven miles norlh-west of Lew-
istown, MitUin county, Pennsylvania.

454

DESCRIPTIOlSrS OF NEW SPECIES OF FOSSILS FROM THE
PLATYCEEAS BREVE Simpson, n. sp., Fig. 23.

Shell of medium size, apex not incurved ; body straight ; rapidly increasing in
size ; -width at the base equal to the length of the anterior side ; length of the poste-

A.P.S.

Tr.l889.

rior side from two-thirds to four-fifths that of the anterior ; both sides convex ; on
the anterior side there is a prominent subangular elevation, commencing at the apex
and continuing to the base ; on each side of the carina a slight depression ; other
portions of the shell without plications or elevations.

Aperture circular; peristome, as far as can be ascertained, not sinuous.

Surface marked by elongate pustules, subregularly arranged, giving to the sur-
face the appearance of being coarsely striated, three in the space of 5 mm. Thei'c
are also faint indications of concentric striae.

A specimen of average size measures as follows : Diameter of aperture 23 mm.;
length of the anterior side 20 mm.; of posterior 15 mm.; height 15 mm.

The characteristics of this species are the short conical form, the rapid enlai-ge-
ment from the apex to the base, and the elongate pustules of the surface.

From those species having a carina it is distinguished as follows: From Platij-
ceras carinatum by its conical form, the straight apex, the absence of plications and
the non-sinuosity of the peristome ; from Platyceras mitelliforme by its larger size,
conical form and the ornamentation of the surfjice ; from Platt/ceras conicum by its
shorter form and the absence of conspicuous plications and the consequent non-sinu-
osity of the peristome. It most closely resembles Platyceras dorsale of this forma-
tion, but it is shorter, much less oblique ; posterior side straight or convex ; the apex
more central and does not project beyond the base, while in that species the apex
projects beyond the base a distance nearly equal to half the diameter of the aperture.

Formation and locality. Chemung group, near Warren, Warren county, Penn-
sylvania.

^ PLATYCERAS DORSALE Simpson, n. sp., ig. 24.

Shell obliquely subconical ; anterior side curved ; posterior straight or concave ;
apex not incurved, laterally compressed ; body of the shell regularly increasing in

CLINTON, LOWER UELDERBERG, CHEMUNG, AND WAVERLT GROUre. 455

size, rounded; width at the base equal to three-fourths of the length of the ante-
lioi- side; Icnf^th of the posterior side one-half that of the anterior; ri<^ht and
left side usually equally developed, but on one speci-
men the right side has a little the greater develop-
ment. On the anterior side there is a conspicuous
rounded or subangular elevation extending from the
apex to the base ; on the posterior side is an elevation
beginning at about one-half the length of the side
from the apex, and continuing to the margin ; on
each side of this elevation a comparatively broad shal-
low depression, of the same extent as the elevation.

Apeiture circular; peristome slightl}' sinuous at the elevations.

Surface marked by fine radiating lines ; at the base there are three in the space
of 1 mm., above they are much more closely disj^osed ; the radii are crossed by fine
concentric strife ; the surface presenting a cancellated ai)pearance when the specimen
is well preserved.

One specimen measures as follows : Length of the anterior side 32 mm., of pos-
terior 18 mm., height of shell 18 mm. Another specimen, which probably belongs
to the same species, is flattened, but that may be due to pressure ; it measures as
follows : Length of the anterior side 45 mm., of the posterior 22 mm., width at the
base 35 mm.; height of shell 18 mm.

This species most closely resembles Plati/ceras hreve of the same locality, but
the anterior side is much longer and the posterior side proportionally shorter, the shell
more oblique and not so regularly conical ; it has an elevation and two depressions on
the posterior side, and the surface has fine radiating stritc, the surface of P. hreve
being marked by elongate pustules ; from Platijceras striatum it may be distinguished
as follows: It is more oblique, the posterior and anterior sides are more unequal, and
it is without the conspicuous ])lication8 characteristic of that species. It resembles
Platycfras mitelliforme in having a prominent carina, but it is a larger form and the
apex is not bent or incurved. From Platyceras conicum it may be distinguished by
the absence of conspicuous plications ; and from P. carinatum by the straight apex
and the absence of conspicuous plications.

Formation and localitij. Chemung group, three miles north-west of Warren,
Warren county, Pennsylvania.

456 DESCRIPTIONS OF NEW SPECIES OF FOSSILS FROM THE

PLATYCERAS IN.EQUALE Simpson, n. sp., Fig. 25.

Shell small, subangularlj ovate ; apex minute, incurved, making less than one
turn, very thin and angular; inclined or twisted to the right. The body of the shell

expands rapidly ; anterior side curved, sometimes
forming nearly half a circle ; posterior side also
curved, but in a lesser degree; width of base from
one-half to three-fourths the length of the anterior
side ; posterior side one-third the length of the
anterior.

The left side is flattened or only very slightly convex, and is often nearly or
quite at right angles to the base, making a sharp angle or ridge between the side and
the back of the shell. The right side is convex and much more developed than the
left. In front the shell becomes flattened near the margin. On the left side, about
half way between the base and angular carina, there is a low rounded ridge, com-
mencing near the apex and continuing the length of the shell, though this feature in
many of the specimens is obscure.

Aperture circular or broadly oval.

Most of the specimens observed are casts or macerated so that the surface char-
acters are obsolete. On some of the specimens there are evidences of strong radiating
striiE or elongate pustules, and when well preserved there are numerous concentric
stria\

The characteristics of this species are the pinched appearance of the apical
portion, and its inclination to the right, and the much greater development of the
right side; in the latter feature it resembles Platyceras cymbeum, but it is a much
smaller shell, and the plications are very much less prominent ; from Platyceras mitel-
llforme it may be distinguished by the inequality of its sides, in that species the
prominent ridge being in the middle of the shell, both sides being equally developed.
Formation and locality. Chemung group, four miles north-west of Warren,
Warren county, Pennsylvania.

PLATYCERAS MITELLIFORME Simpson, n. sp., Fig. 26.

Shell small, obliquely arcuate from the base ; apex incurved, making part of one
volution; below which the body volution rapidl}^ increases in size. Width at the
base equal to three-fourths the length of the anterior side ; length of the posterior side
less than one-half of the anterior ; right and left sides equally developed.

Anterior side convex, with a prominent, broad, rounded carina along the middle;

CLINTON, LOWER IIELDKRBEUG. C.UEMUXG, AND AVAVEHLY GROUPS. 457

on each side of the carina, towards the base, a shallow depression. On some of the
specimens there are indications of very slight plications on the posterior side.

Aperture oblique, broadly elliptical; peristome a lit-
tle sinuous at the carina, and sometimes slightly sinuous, ^ ' ^ f\\
corresponding witli tlu' faint folds of the posterior side. /

Surface marked l)v concentric lines and by broad uu- ""*"' ^ -*

dulations, which sometimes give to portions of the shell s.?^. Trissj. *-^-i.__>i_>i

a lobed appearance.

A specimen of about the average size measures as follows : Height 12 mm. ;
length of the anterior side 25 mm. ; of the posterior 8 mm. ; width at base 22 mm. ;
thickness 18 mm.

The characteristic feature of this species is the prominent carina or ele\ ation
along the dorsum.

It may be distinguished from J'latijceras iuiequale of this formation by its some-
what larger size, the prominent carina, and the equal de^■eIopment of the right and
left side ; from Platijceras ihrsale by its smaller size, its Hattened form (a transverse
section being oval), and by the partial volution of the apex ; from Platijceras {Or-
thonychia) striatum by its arcuate form, the prominent carina and t!ie absence of con-
spicuous plications; from Platijceras carinatum by the equal development of the right
and left sides, and the absence of conspicuous plications ; from other species at
present known, by its small size, decidedly curved form, and prominent carina.

Formation and locality. Chemung group, near Warren, "Warren county, Penn-
sylvania.

PLATYCERAS STRIATUM! Simp-son, n. sp., Fig. 27.

Shell subconical ; ape.v nol incurved or bent ; body essentially straight ; sides a
little curved from the base to the apex ; right and left sides equally developed ; width
at the base, length of anterior and posterior sides equal. Ou the anterior .><ide is a
narrow, angular, conspicuous carina, extending from viii^ /^Ik. ' >-^x z.

the apex to the base ; on the posterior side are three

oblique, broad, prominent, rounded ridges, with de-
pressions between them, which are wider than the
ridges. j

Ajjcrture oval. The bases of all the specimens ^ , -•*

observed arc attached to the rock so that the form of the peristome can not be defi-
nitely ascertained.

A. 1'. >. — VOL. \vi. .3f.

458 DESCRIPTIONS Or NEW SPECIES OF FOSSILS FROM THE

Surface mai-ked by comi3aratively strong, radiating stria^, which are sometimes
continuous, at other times intei-ruiited, then having the appearance of very elongate
pustules ; at the base about six in the space of 5 mm. 'No concentric striie have
been observed, though it is possible that they exist on more perfectly preserved
specimens.

A typical specimen measures as follows : "Width at the base 30 mm. ; thickness
20 mm. ; height 30 mm. ; width of sides equal ; width of i-idges <mi posterior side
3 mm. ; of depressions slightly more ; of carina 3 mm.

This form resembles the two specimens which are figured in the Pal. of New
York, Vol. V, Pt. II, PI. I, Figs. 20-23, which differ materially from the other figured
specimens placed under that species. The ether specimens, though perfectly pre-
served and showing concentric striae, have no indication of radiating strife, while in
the specimens described they are one of the most characteristic features ; they are
also shown in the figures mentioned above. The plications are also stronger and the
form more regularly conical.

This species resembles P. striatum from this formation, but may be easily distin-
guished by its straight form, the equal length of the sides, the coarser striations, and
the strong plications of the posterior sides ; from P. hreve it is distinguished by its
more elongate form, oval aperture, more continuous radiating strije, and the conspic-
uous plications of the posterior side.

Formation and locality. Chemung group, four miles north-west of Warren,
Warren county, Pennsylvania.

PLATYCERAS VAEIANS Simpson, n. sp., Fig. 28.

Shell small, subconical, curved, slightly oblique; apex not incurved or bent;
width at the base two-thirds the length of the anterior side, and about equal to the
posterior.

Plications frequent, variable in number. On the dorsum there is sometimes a
faint indication of a carina; on the left side is a deep conspicuous furrow, which ex-
viiig. tends about thi'ee-fourths the length of the shell from the base; beyond
this there is a slighter furi'ow ; the area between them elevated,
I'ounded and very prominent. On the posterior side of the shell are
usually two or three slight furrows, but occasionally a stronger one, and
""iBBm on the left side there is one nearly as strong as on the opposite side.
Sometimes on the posterior side the ai'ea between two of the smaller furrows is ele-
vated, having the ajipearance of a ridge or carina, which is occasionally prominent.

CLIXTOX. L(JWEK IlELDEHBEHC;. CIIEMUXG. AND WAVEULV GROUPS. 459

The deep tuirows on the ri^ht and left sides of the shell ap[)ear to be constant fea-
tures, the others are somewhat variable.

Aperture broadly oval, nearly circular ; peristome concealed on all the speci-
mens observed, so that its true form can not be ascertained.

All the specimens occurrinj:,- in the form of casts, the surface markings are
obsolete.

This species may be disting-uished from Plalyceras mitelliformis by its straighter
form, absence of a prominent carina, and by its conspicuous plications; from P. (0.)
stn'atuin by its smaller si/e, more frequent i)lications, and the absence of radiating
stria', the convexity of the anterior side and the concavit}' of the posterior ; from
Platijceras incqnale by its equally developed right and left sides and its conspicuous
plications; from P. (0.) breve by its curved, more slender form, and plications;
from Phttyceras dwsale by its smaller size and conspicuous plications; from P. (0.)
attenuntum by the straight apex and the strong lateral furrows ; from Plalyceras
carinutnm by the absence of a prominent carina and by the conspicuous lateral furi-ows ;
from P. (0.) com'cum by its smaller size and curved form.

Forviotion unci locality. Chemung grouj). four miles north-west of Warren. "War-
ren county, I'cnn'iylvania.

ACERVULARIA COMMUNIS Simpson, n. sp., Fig. 20.

Growing in large convex or globular masses ; under side covered by a wrinkled
epithecal crust. Stems multiplying by gemmation from the calyces; usually in con-
tact, but sometimes free and circular. vi.

Calyces in contact, polygonal, generally pentagonal or he.\-ago-
nal ; surrounded by acute, linear, crested edges ; diameter a little
variable, but usually about 5 mm. Sides of calyces al)ruptl\' curved
downward to the central cavity. ^ps Tr.i889

Jiays thin, sharp; about thirty in nuinljcr; alternately long and short, but of
uniform size at the margins of the calyces. Diai^hragms thin.

Formation and locality. Lower Helderberg group, from .\. 15. Miller's farm.
Warrior lidge, Barree township, Huntingdon county, Pennsylvania.

CLADOPORA RF:CTILTNKATA Simpson, n. sp., Fig. .30.

Zoai'ium I'ainose, l)il'urcating, rarely anastomosing ; diamctci- of Itranches fi-om
l.JU to 3 mm.

460

DESCRIPTIONS OF NEW SPECIES OF FOSSILS, ETC.

;-<vI;"S 2; »:- jyS; J,-- 3': Vi, li i; j-

• :y :p ■>•■?- r- s * Jt a » g s-

* ■»■ 3- y > fr I jg s J-- r »■ •"*

■ -a •». * *- :^ > $■ T: «- I « >• .

Cells tubulai-, nearly at right angles to the axis ; rapidly increasing- in size to the
aperture ; apertures arranged in regular longitudinal i-o\vs, which are sometimes sepa-
rated by a slight ridge ; from five to nine rows on a branch ;
from eight to ten apertures in the space of 5 mm. longitu-
dinally; diameter of an aperture from .33 to .40 mm. Lips
riitiimt,tn^Kr. ^ PS Tr IS39. promineut, in well-preserved specimens, projecting over tlie
aperture, and giving to the cell-tubes the appearance of opening very obliquely.

This species somewhat resembles C. mulUpora, but the branches are smaller and
straighter, and the cell apertures are much more regularly arranged in parallel, longi-
tudinal rows ; in its size and manner of growth it is very similar to 6'. niacrojjhora,
but the surface of that species has not been observed, so no comparisons are possible.
Formation and localities. Lower Helderberg group, one and a-half miles south
of Rock Hill furnace, Orbisonia, Huntingdon county ; also north of Tyrone Cit}-,
Blair county, Pennsylvania.

HOMALONOTUS TRENTONENSIS, Simpson, n. sp., Fig. 31.

r '

"*\ /•.

:^mi::i i

APS. Tt. !2°9.

'Homa'ofoNs ^ren^onen3(3

SirTipson.

The figures represent a few of many si)ecimens in the State Collection, found l)v
Mr. C. E. Hall in strata of the Trenton group, cropping out just aljove the milldam
at Reedsville, in MiiUin county, Pennsylvania.

ARTICLE IX.

TOE MAMMALIA OF THE UINTA FORMATION.

PAUT I. THE GEOLOGICAL AXD FAUXAL HELATIOXS OF THE UINTA FORMATION.
PAHT H. THE CREODOXTA, UODEKTIA AND ARTIODACTYLA.

BY WILLIAM B. SCOTT,

PROFESSOR OF GEOLOGV IN PIUN'CETON COLL

PAUT III. THE PEHISSODACTYLA.

PART IV. THE EVOLUTION OF THE UNGULATE FOOT.

BY HENRY K.VIRFIKLD OSBORN,

PKOFSgSOR OF CO.\IP,VRATlVE ANATOMY IN PKIKCETON COLLGGE.

Il.I.USTUATED BY FiVE LlTHOORAPIlrC PlATES ASD TiIIRTEES DiAOKAMS.

Kead before the American Philosophical Society, May 17, 1889.

This memoir i.s principally descriptive of a collection made by the Princeton Sci-
entific Expedition of 1886 in the Bad Lands of the Wiiitc Kivor, Northea.stern Utah.
The party was composed of Messrs. Keynolds, Harlan, Ilervey, Baucus, Paton,
Kane, all Princeton students, under the able leadership of Mr. Francis Speir, Jr.,
whose name is connected with so many important pahcontological discoveries. The
expedition exi)erienced great difficulties in the field and some risk from the hostile
threats of the White River Utes.

The collection has been skillfully prepared and mounted by Dr. Franklin C.
Tlill. The drawings are by Mr. Rudolph Weber.

A. p. s. — VOL. xvr. 3<;.

462 THE MAMMALIA OF THE UINTA FORMATION.

Part I.

BY AYILLTAM B. SCOTT
GEOLOGICAL AND FAUNAL RELATIONS OF THE UINTA FORMATION.

While the palaeontological relations of the various series of Eocene deposits in
the upper Green Eiver valley are, for the most part, quite clear, the stratigraphical
relations are as yet hut imperfectly known, and leave many questions still open. Dr.
White (iJ^o. 18, p. 35) believes that the entire series is conformably deposited : " In
the great region now drained by the Green river, there are three well-marked groups
of strata, that come in their order above the Laramie group, and which all agree in
referring to the Tertiary period. These are the Wasatch, Green River and Bridger
groups, named in ascending order. The Wasatch group is the lowest of a series
of three fi-esh-water Tertiary groups, all of which are intimately connected, not
only by an evident continuity of sedimentation throughout, but also by the pass-
age of a portion of the molluscan si^ecies from one group up into the next above.
Not only were the three groups, aggregating more than a mile in thickness, evidently
produc(!d by a continuous sedimentation, but it seems equally evident that it was
likewise uninterrupted between the Laramie and Wasatch epochs, although there
was then a change from brackish to fresh waters and a consequent change of all the
species of invertebrates inhabiting these waters."

King, on the other hand (N^o. G, p. 353 et seq.), adduces evidence to show, not
only that the Wasatch and Laramie are very clearly separated by unconformities, but
that the three Eocene series are likewise divided by lack of conformity with each
other.

Pala^ontologically the arrangement of the series is less obscure, though if we
accept White's view of a continuous sedimentation from the Laramie to the Bridger
we shall meet with very formidable dillicuUies. Thus no place is left for the very
peculiar and primitive fauna of the Pucrco group, which is not at all represented in
the noi'thern basin ; the same reasoning will apply to the transition fauna of the
Wind River group, a formation estimated as being 1000 feet or more in thickness
(St. John, No. 12, p. 200), bui so far as is yet known confined to the Wind River
valley. To assume that the I*uerco is contemporaneous with the Wasatch, the Wind
River with the Bridger, and the beds of the Bridger basin with those of the Washakie

THE MAMMALIA OF THE UINTA FORMATION. 463

basin, Avould involve very complicated hypotheses of barriers and mij^rations for
wliich there is no present evidence. The fauna of the Wind River deposits has been
shown by Cope (No. 1) to be intermediate betAveen the "Wasatch and the Bridger,
mingling types which elsewhere are found only in one or other of these formations
with some forms peculiar to itself Thus such typically Wasatch genera as Coryph-
odon, Phenacodus, Didymictis, Calamodon and E$thonyx are associated with equally
typical Biidger genera, Palceosyops, Lamhdotheriwn, Mkicis and Microsyop^, while the
earliest and smallest known member of the Dinocerata, Bathyopsis, is confined to
these deposits. Thus the Wind River formation occupies the same intermediate
position between the Wasatch and Bridger palreontologically as does the Green River
stratigraphically. One is therefore led to infer with Cope (No. 2, p. 453) that the
Green River and Wind liiver are parts of the same formation, though the entire
absence of mammalian fossils from the former renders this determination somewhat
uncertain. It may perhaps be objected to tTiis view that in South westei-n Wyoming
the Bridger beds lie conformably upon the Wasatch. This conformity is, however,
not improbably deceptive, for the following reasons : (1) King states (No. 6, p. 380)
that he has observed unconformities between the Wasatch and Green River beds.
"In the region east of the Wasatch a large amount of the Vermilion Ci'eek [Wa-
satch] series was left in a nearly horizontal position and the sediments there sank
quietly through deep water upon an appi'oximately level bottom, accumulating in
strata nearly conformable with the underlying Vermilion Creek rocks. From the
manner in which tlie rocks of the Green River group abut westward against the Ver-
milion beds, it is evident that there was in the region included between the Wasatch
and Uinta a highland lifted above the lake of the Green River period." (2) The
direct contact between the Wasatch and Bridger strata at the west of the Bridger
basin takes place by the latter bed's overlappiiir/ the Green River and thus reaching
the Wasatch. If we assume that the Green River shales represent a distinct series,
this oveilap can be explained only by supposing either that these beds had been
swept away from the western part of the basin before the deposition of the liridger
series, oi', what is much more probable, that a series of disturbances first contracted
and then expanded the waters of the lake. (3) It is im[)03sible to account for the
faunal differences between the Wind River and tlie Wasatch on the one hand and the
Bri<lger on the other on geographical grounds alone, as this area is encompassed
]>()tli on the north and the south by typical Wasatch beds.

The balance of evidence would thus seem to point to the conclusion that the
Wind River beds are to be regarded as forming the base of tiic Bridger scries, and
this view has the further advantage, thai the Bridger, thus deliued, is characterized

464 THE MAMMALIA OF THE UINTA FORMATION.

throughout by the presence of some form or other of the Dinocerata, and that this
extraordniary group of animals is entirely confined to this formation.

A still more intricate and difficult problem is that with reference to the relations
of the various areas of Bridger deposits (apart from the Wind River series) to each
other. Of these areas there are three : (1) The Bridger basin west of the Green river ;
(2) the "Washakie basin east of the same river, and (3) a small area to the south-
east of the latter. Supposing the Bridger beds of the Washakie and Bridger basins
to have been deposited conformably in the same lake which laid down the Green
River series and to have been uplifted together with the Green River in a post-Bridger
upheaval, it is not a little remarkable that erosion should have removed the Bridger
from all parts save the middle of these two basins. The few observations which bear
upon this point in the way of the dips of the two formations combine to indicate that
the movement took place at the end of the Green River period, that the western lake
[^". e., the supposed extension of the Green River lake west of the Wasatch moun-
tains] was extinguished by this upheaval, and that the waters of the period formed a
lake of restricted area altogether within the basin of Green river. Even with this
supposition, which I conclude to be the most probable until it may be varied by
future evidence, there is left the shadow of a doubt, whether the three Bridger bodies
which appear upon our map — that of the Bridger basin, the Washakie basin, and the
region east of Vermilion creek — were parts of a continuous sheet, or whether they
themselves were areas of special lakes in the same general basin, but characterized
by great fauna resemblances." (King, TsTo. 6, p. 391).)

The facts of stratigraphy, so far as at present known, leave this question an
open one, but the asseml>Iage of fossils seems to point to the conclusion that there
were at least two distinct lakes, and these not contemporaneous, but successive, a
conclusion which we have already indicated elsewhere (Osborn, No. 10, p. 13 ; Seott,
Xo. 15). The faunal lists of the two basins seem to show that the Bridger contains
slightly older deposits than the Washakie, though the two may be in part contem-
poraneous. Some of the differences to be noted are, no doubt, due in part to the
fact that the Bridger basin has been much more frequently and thoronghl}' explored
than the Washakie, and others probably to conditions of preservation, for within
the limits of the Bridger basin there are localities and strata which are especially
rich in certain soils of mammals which elsewhere arc much rai-er. As an example
of this may be mentioned the abundance of th(! lemuroids and other small mammals
at Twin Biittes. Future explorations may perhaps therefore diminish the number of
faunal differences between the two areas.

A striking fact is the greater richness and variety of the forms found in the

THK MAMMALIA OF THE UINTA FOUMATION. 405

western basin, especially of the smaller mammals, the crciodonts, lemiiroids and insec-
tivores, which are comparatively rare in the AVashakie and very abnndant in the
Bridgcr. No tillodonts have yet been reported from the former. The perissodac-
tyls are quite diifcrent. l*al<eos;/ops is somewhat less abundant in individuals in the
eastern basin, but /*. major and palndosus are both present, and there are in the
Princeton Museum two undescribed species of this genus from the AVashakie which
differ markedly from any yet found in the Bridger, and one of them, l)y its great size,
seems to approximate closely Diplacodon. Hyrachyus is represented by the same or
similar species and in about equal numbers in both areas. On the other hand the
Uinta rhinoceros Aovjnodon (== Orthocynodon nobis) is found only in the eastern
i-egion, and the same is almdst certainly true of Triplopus, while Colonoceras occurs
only in the Bridger basin. Among the artiodactyls the chief diiterence is to be
Ibund in the presence and frequency in the eastern basin of the large species of
Achvnodon, A. insolens and ^1. rohistiis. JTeloIiyus and Ilomacodon have not yet
been reported from this region, but as representatives of these series occur in the
Uinta, we may confidently expect that they will be found in the AVashakie.

Another strikiny; difference between the two basins is to be found in the fades
of the Dinocerata so common everywhere. The great variability of these animals
precludes the making of any very sharp distinctions between those of the two ai-eas,
but in a general way and with some exceptions they are in rather strong contrast
with each other. The prevailing type of skull in the Bridger basin is comi)aratively
short and broad, the nasal tuberosities are of moderate size, the parietal protuberances
are placed well forward, and the parieto-occipital crest is of moderate height. The
Washakie forms have generally a long narrow skull, with very large nasal tuberosi-
ties, the j)arietal protubei-ances are placed far back, the parieto-occii)ital crest is very
high ; the cervical vertebraj in these species are also as a rule shorter than in those
of the Bridger area, and the animals tend to become of a larger size.

Making all due allowance for sources of error, it is difficult to believe that these
differences are to be explained by geographical considerations, while on the other
hand the strati graphical relations of the AVashakie beds to the underlying Green
Kiver scries present objections to the assumption that the latter could have been a
land surface for any great length of time. But these relations ai'c by no means clear
and very much need to l»e re-examined.

AV'e have no information relative to the small area east of Vcrtniliou cn-ok. but
in all j)robability it is simply an outlier of the AVashakic.

From pi-esent knowledge, we are therefore justilied in making a tlirciliiM sub-
division (>r liu- Bi idger epoch, willi ilic Wind liiver licds at llic l>;t<c and the

466 THE MAMMALIA OF THE UINTA FOKMATION,

Washakie at the top, and in regarding the Dinocerata as the group characteristic of
the entire epoch. It is interesting to note that where the fauna of the "Washakie
basin departs from that of the Bridger basin, it does so in the way of resemblance to
the Uinta.

The geology of the Uinta formation is even more impeifectly known than that
of the earlier formations. Dr. White says of it (ISTo. 18, p. 37): " Resting directly,
but by unconformity of sequence, upon all the Tertiary or Cretaceous groups in the
region surrounding the eastern end of the Uinta Mountain range, is another Tertiary
group that has received the name of 'Uinta Gi'oup' from Mr. King, and 'Brown's
Park Group' from Prof Powell. It is jwssible that this group was deposited
continuously, at least in part, with the Bridger group, but at the places where
the junction between the two groups has been seen in this region, there is an evident
unconformity, Ijoth l)y displacement and erosion." Dr. White has also informed us
(orally) that the moliusca of these beds are strikingly different from those of the
earlier Eocene. The Uinta deposits are composed of sandstones and clays much
like those of the Bridger in appearance. They are of no great thickness and are best
developed in the valley of the White Rivci-, a tributary of the Gieen, in eastern
Utah and Western Colorado. The mammalian fauna of the Uinta formation, while
very distinctly of more modern character than that of the Bridger, is nevertheless
quite closely allied with it, more especially with that of the Washakie subdivision,
and it should he further remembered that this fauna has as yet been only very par-
tially explored, and we may therefore expect that future discoveries will fill up many
of the gaps still existing.

The more extended study of this fauna confirms the generally accepted view
that the Uinta group should be placed at the summit of the Eocene, forming the
transition to the White River Miocene, with which it has many affinities. The most
striking and obvious difference fi'om the Bridger fauna consists in the absence of
the great Dinocerata so characteristic of this series, of which no trace has yet been
found, nor is it likely that they will be discovered in the future. If present at all, this
group must have formed but a rare and unimportant factor in the assemblage of
Uinta mammals. Another Bridger type of which no member has been discovered in
the Uinta (with a possible exception hereafter to be mentioned) is that of the Tillo-
dontia, which seem to have finally disappeared, unless, as some have supposed, we
are to look for their successors in the Edentata. The smaller mammals are very
much more scantily repi-esented than in the Bridger, though some of this difterence is
perhaps due to conditions of preservation, as indeed the Uinta fossils are scantici- and
less well preserved than in the immediately preceding or succeeding epochs. A simi-

THE MAMMALIA OF THE UINTA FORMATION. 4G7

lar, though less striking, ditiereiice exists between the Bridger and Washakie basins
with regard to the number and variety of the smaller forms. The great assemblage
of Icmuroids and pseudo-Icmuroids which the Wasatch and Bridger show so abun-
dantly is thus far represented in the Uinta only by the single species IJijoyi^odus
(iracilts Maish ; the Insectivora, Chiroptera, and Marsupialia have not been found at
all, though there is hardly room for doubt that they existed, inasmuch as they occur
both in the Bridger and the White River. Rodentia are also rare and only one form
can be certainly named, a species of Plesiarctomys, which genus is abundant in the
Wasatch, Wind River, and Biidger beds. A second species of rodent is apparently
implied by a lower incisor with a fragment of the mandible; it is, however, much
larger than any known rodent of the Bridger or White Kiver, and may possibly be-
long to some tillodont ; but this is not at all probable, as the tooth is much more
like that of a rodent, besides being smaller than that of any Bridger tillodont. We
have still therefore to search for the forerunners of the beavers, mice, and hares of
the White River.

So far but a single creodont is certainly known to occur in the Uinta fauna, a
very large species of Mesonyx. This family ranges therefoi-e throughout the Eocene
of Nortii America, being represented by Dissacus in the Puerco, PacJujama in the
Wasatch and Mesonyx in the Bi'idgcr and Uinta. The absence of the upper molars
from the specimen before us precludes the possibility of a final reference of it to
Mesonyx ; but there can be no doubt that it is a member of the famil3^ A second
creodont is jjcrhaps indicated by a small lower sectorial molar and a premolar ; if so,
the animal must have belonged to the Miacidiv, or it may be a true carnivore. The
creodont nature of the specimen is the more probable on account of the greater like-
ness of the Uinta fauna to the Bridger. In any case this animal very probably rep-
resents the foreninner of the dog-like forms of the White River, the so-called AmpJii-
cyon and Galecynus. and possibly of Dinictis as well. Nothing is as yet known of
any Uinta forms which would serve to connect the hya^nodonts of the AVhite River
with the Oxycena and Protopsalis of the Bridger.

The Perissodactyla of the Uinta are of great interest, as they are plainly inter-
mediate between those of the Bridger and those of the White River. The curious
family of Titanotheiidn' has its Uinta member in Diphicodon, which thus comiecls the
Bi idgei- genera I'ahi'osyops and Leurocrplndus with the I'itanollieriam (Moiodns) of
the White River. IIyr(idiynf>,Ho frequent in the Bridger, has apparently disajjpeared,
its place being taken by Tri]d(qms. which already occurs in the Washakie bed."?,
though sparsely. In the Uinta it is one of the nK)st abundant of all the lossils, and
is highly characteristic of the formation. Another genus, rare in tlir Washakie.

468 THE MAMMALIA OF THE UINTA FOllMATION.

abundant in the Uinta, is the rhinoceros-like Amynodon, a hornless form which is fol-
lowed in the White River by Metamynodon and Aceratherium. The TTinta tapir is
Iscotolo'phus, which is as yet represented by a single abundant species of small size.
This genus appeals to be represented also in the Bridger, at least there is a well-pre-
served specimen in the Princeton Museum which we cannot distinguish from it, and
seems to have its representative in the White River beds, which Dr. Leidy has
doubtfully referred to Lojiliiodon, though only for the reason that the specimen, an
isolated last lower molar, could not well be distinguished from the corresponding
tooth of that genus. As Leidy has suggested, this specimen most probably belongs
to a very different genus (No. 7, p. 239). The equine series is represented in the
Uinta fauna by two species of Orotherium, (JSjnhijjpus) which stands just interme-
diate between PUoloj^Ims ( Oroldpims) of the Bridger and Mesoliip)pus of the White
River, though it also is found in the Bridger.

Perhaps the most striking change in the facies of the Uinta fauna as compared
with that of the Bridger lies in the importance suddenly assumed, by the selenodont
artiodactyls. The Bridger is not known to contain any forms which can be strictly
called selenodonts, though Homacodon and Helolryus are obviously commencing to
develop this type of dentition. The Uinta contains at least two clearly mai-ked gen-
era of selenodonts, if not more, which are very rich in individuals and thus give a
very diflei-ent aspect to the fauna as a whole. Of these the most abundant is Pro-
toreodo7i, the unmistakable ancestor of the Oreodontidce, the peculiar family which is
so eminently charactei'istic of the American Miocene. By the five-lobed construction
of the upper true molars, Protoreodon indicates the line of connection with the buno-
selenodonts of the earlier Eocene, especially, if we may judge from the molar teeth
alone, with Ilelohyus. The other clearly defined Uinta selenodont is Leptotragulus,
the most ancient unequivocal member of the Tylopoda and closely related to the
White River genus Poehroiherium, though in many respects more primitive. The
absence of the upper molars from our specimens prevents comparison of this type
with Homacodon, from which it was perhaps derived. The Uinta has thus yielded
forerunners of the two most important and characteristic groups of White River sel-
enodonts ; it remains for future investigation to bring to light the ancestors of Lepto-
riieryx and Hyopotarims, though the latter is not improbably an immigrant fi'om the
Old Woi'ld. No Bunodonta have as yet been found, and the origin of the White
River genera, Perchocrus, Entelodon, etc., is still obscure. Our knowledge of the
Bridger bunodonts is still very imi^erfect.

It will be evident from the above statements that the aillnities of the Uinta
fauna are most closely with the Bridger, as at least five genera are common to the

THE MAMMALIA OF THE UfNTA FORMATION.

460

two horizons, Plesiarctomys, Mesonyx, Triplopus, Aviynodon and Orotherium, while
on the other hand no genus passes through from the Uinta into the White River,
unless one agrees with Cope (!N'o. 2, p. 455) in regarding Metamynodon as identical
with Amynodon. While this is true, the Uinta fauna is no less clearly the forerunner
of the White River fauna ; what there are strong reasons for believing to be the an-
cestors of the hyracodonts, the titanotheria, the rhinocei'oses, horses, tapirs, camels
and creodonts, the dog-like carnivores, and the sciuromorph rodents of the White
River, all ai'e to be found. We cannot therefore agree with Schlosser (No. 13, p. GO)
in regarding this formation as Oligocene, which in fact seems to be an unnecessary
term in the classification of our Tertiary lacustrine deposits, the division between the
uppermost Eocene, lepresented by the Uinta, and the lowermost Miocene, represented
by the White River, being quite sufficiently clear.

The following partial list of genera will indicate the relations between these
three formations. The Wasatch genera, which penetrate only into the Wind River
beds, are omitted from the list :

Bkidger.

Dinocerata

Tillodonlia

Slvpoloplius

Jli'sonyx

Itliacis

Oxyjpna

Protopsalis

Antiac-odon
Anisacodnn
Passalacodon

Ictops
Plesiarctomys

C'olonymys
Mysops

Omoinys

Hyop^odiis

Tomitlieriiim

Liiiinollieriiim

Microsyops

Xipliacodon

lliloliyus

Hoinarodon
Arliiunudnn

Uinta.

Mesonyx
? Miacis
?

?
Plesiarctomys

Uyopsodus

t

Protoreodon

1
Lcptolragiiliis

?

Larobdotlipriiiiii

A. r. 9. — VOL. XV r. 3h.

WlUTIC RiVKK.

Hyfcnodon

DapliiPnus (Ainpliicyou)

Cyiiodiclis

Diniclis

Iloplophoneiis

Lcptictiii

Ictops

Sciurus

Gymnoptyclius

Ileliscoinys

Iscliyromys

Eiimya

Palicocastor

Paltuolagus

Laopilliecns
Mfiiotherium

Didelpliys

Oreodon

AiirinchnTU'*

I'di'liroilicritiin

Enlclndon

Pi-rcluiTiiM

Li'plocliD'riis

IlyopotaiiHis

Li'l)loiii<'ryx

IIypi^*odu8

470

THE MAMMALIA OF THE UINTA FORMATION.

Bridgek

Uinta.

White River,

Palscosyops
Telmatotherium
Isectoloplms
Hyrachyus
'? Triiilcipiis
OrollRnium
Pliolopluis
Colonoccras
Amyuodon

Nyclilestes
Vesperngo
Nyclithcirium

Diiilacodon
Isectoloplius

Triplopus
Orotherium

Amyuodon

TitanotUcrium
Mesotapirus

Hyracodon

Mesoliippus

Motamynodon
Acerathfi'ium
Domniua

THE MAMMALIA OF THE UINTA FORMATION. 471

Part 1 1.

nv wrr,LTA:\r r, scott.

THE CREODONTA, RODENTIA, AND ARTIODACTYLA.

PRIMATES.

HYOPSODUS GRACILIS Uar^h.

This species is the only one of the pseudo-lemuroids whieh has as yet heen
reported from the Uinta deposit s. This group is exceedingly abundant in tlie Bridger,
especially in the Bridger liasin ; it has become moi'c rare in the Washakie, and
although the Uinta has yielded but one species as yet, others Avill no doiilit be found.
The change from the Bridger will, however, almost certainly remain a vei-y striking
one in the reduction of these animals. They are not certainly known to occur in the
White River, though some specimens have been I'efcrred to the group {Laopithecus,
Menotlierhan).

CREODONTA.

MESON YX Cope.

M. uintexsis Scott (No. 10, pj). 1G8, KiO); PI. X, Fig. 0. The specimen upon
which this species is founded consists of five isolated lower molar and premolar teeth,
an incisor, a canine, one upper premolar and fragments of the mandible, including the
condyle. The reference to Mesonyx is somewhat uncertain on account of the absence
of the upper molars ; there can be no doubt, however, that the species belongs to the
family MesonyrJifdup, and it is of interest as being the largest and latest known mem-
bei- of that long-lived and most curious group of creodonts. This species may be
distinguished from those of the earlier Eocene formations by very much greater size
and by the fact that the anterior basal cusps of the lower molars arc much reduced.

The incisor is peculiar and consists of a l)road and rounded sim]>le crown; it is
of large size and contrasts strongly with the incisors of the Bridger species 3L obfn-
s ideas f the canine and upper premolar dilRi' from the corrcHponding teeth of the

472 THE MAMMALIA OF THE UINTA FORMATION.

Bridger species only in size. Since the first account of this sjDecies was written, we
have found that the order of the teeth was there incoi-rectly given ; what was there
described as the second premolar being probably the third molar. The most anterior
premolar of the lower series pi-eserved in the specimen is probably the second (see
PI. X, Fig. 9). This is a relatively small tooth, implanted by two fangs and with
a low, rather elongated crown, which consists of a median cone and faintly separated
anterioi' basal tubercles ; this tooth is therefore somewhat differently constituted from
the second lower premolar of M. obtusidens, in which the anterior tubercle is not in-
dicated at all, and the posterior one scarcely so. The fourth premolar has nearly the
shape and size of the true molars, consisting of a high and massive, backwardly
directed principal cusp, which is separated by a deep and narrow cleft from the lai-ge
trenchant heel. The latter is as long from before backwards as the main cusp and is
strongly convex on the outer side and concave on the inner. This tooth diftei-s from
the fourth premolar of 3f. oltusidens only in the absence of the anterior basal cusp.
The first true molar has the same construction as the tooth just described, but is
larger, and the main cusp is remarkably massive; there is an indication of the anterior
basal cusp, but it is much less distinctly marked than in the Bridger species. This
tooth is the largest of the series and considerably exceeds in size the lower sectorial
of a full-grown lion. The second molar difters from the first only in being a little
smaller and in the still greater reduction of the anterior basal cusjjs. The develop-
ment of these cusps varies greatly in the ditferent genera of the family ; in Dissacus
of the Puerco they are fairly well developed; in the Wasatch genus Pacliycuna they ai-e
greatly increased in size, becoming as large as the heel, which gives the inferior
molai's a trifid appearance when seen from the side. In the Bridger species of Meso-
nyx these cusps become much smaller, being greatly exceeded in size by the talon,
while in M. uiniensis they have almost disappeared, entirely so in fact, from the
fourth premolar. The third molar is very small, and is even more reduced in size
than in M. obtusidens, which reduction chiefly affects the talon ; in the Bridger spe-
cies the talon is of good size, but in the specimen before us the heel is a mere rudi-
ment. The main cusp is also more compiessed and pointed. The third molar is very
differently developed in the various genera of the family ; in Disaacus and Pacliyieaa
iiT^B is nearly or quite as large as m7^ ; in Mesonyx obtusidens it is distinctly smaller,
the proportions being 7 : 9, and in M. tiinfensis it is still smaller, as 3: 5. This reduc-
tion in 3Iesonyx is, of course, correlated with the loss of the last upper molar, and the
pi'ojwrtions in the Uinta species would seem to indicate that the second upper molar
was undergoing a similar reduction.

It has been geneially assumed that the molar teeth of the Mesonychidoe are of

THE MAMMALIA OF THE UIXTA FOUMATION". 473

an exceedingly pi-imitive pattci-n. As far as the upper molars are concerned this is
no doubt tiiio, thcise teeth never vaiying from a simple tritubercular paltein. T>is-
sacus, however, shows that the lower molars were originally of the tuberculo-sectorial
pattern, as the postero-internal cusp is distinctly present, from which it follows that
the simplicity of the Mesonyx molars has been attained by the suppression of parts,
somewhat in the same fashion as has been done in the lower sectorial of ITi/<i',aodon
and the Felidce. These lower molars of Mesonyx can, however, hardly be called sec-
toiial, as the shearing action is limited by the small height of the outer tubercles of
the upper molars. The cusps of the lower series are, therefore, worn upon the sum-
mits and very rapidly become blunted and worn down.

A second specimen, consisting of parts of two metatarsals and a phalanx, should
probably be referied to this species, as their shape and character are very similar to
those of M. ohtusidens. The latter species is remarkable for the small size and weak
development of the feet, but if these bones from the Uinta are correctly referred, M.
uintensis must have had very different proportions, for they are considerably shorter
and much more massive than in the lion. It is obvious that the Uinta Mesonyx must
have been a very formidable animal, much more so than the more highly organized
carnivores and creodonts (except Uein/pscdodon) which succeeded it in the White
Kiver epoch.

Measurements. S/.

Diameter of incisor (fore anil aft) Oil

" " (transverse) Oil

Dianu'tcr of caiilnp 033

LoDglli sfcond lowpr premolar 014

" fourth 028

Tliifkncss " " " 013

Length lower molar series f'TS

" first molar 031

Thickness first molar OI.T

Length second " 02U

Thickness second molar 015

Length third " 018

Transverse diameter of mandilxilar condyle 0:17

MI ACTS Tope.

This genus of creodonts is the one which most closely approximates the true
Carnivora and is characterized by the sectorial pm J and m i. This is also true of
Didymiclis, but here the last upper molar is wanting and the lower sectorial is of
very diil'erent character.

474 THE MAMMALIA OF THE UINTA FORMATION.

?MiAcis vuLPiNus S. & O. (Syn. ? AmjjMcyon vulpinum S. & O., No. 17, p.
255). Since our preliminary acconnt was written we have had the opportunity of
examining various specimens of Amphicyon, which show that our tentative reference
of the Uinta species to that genus must have been incorrect. It is equall}' clear that
the species cannot belong to Cynodictis, or to any of the flesh-eaters known to occur
in the White River beds, whereas it may very well be a species of Miacis, and in
view of the similarity of the Uinta fauna to that of the Bridger, this will probabl}'
turn out to be the case. The specimen consists of an isolated inferior premolar and
first molar.

The premolar, probably the third, consists of a high, acute and compressed cone,
with rudimentary posterior heel ; a cingulum runs entirely around the crown and is
most conspicuous on the anterior surface. The principal cusp is not complicated by
the development of accessory tubercles upon it. The sectorial is canine in appear-
ance; the anterior portion consists of three cusps arranged in the ordinary triangle,
of which the external is the larger and flattened on the outer side ; the anterior cusp
is still very low and the sectorial blade is therefore much less completely developed
than in Cynodictis and hardly more so than in the Bridger species of Miacis. The
heel of the sectorial is low and small and not very distinctly basin-shaped.

RODENTIA.

PLESTARCTOMYS Bravard.

Of all the rodents of the American Eocene, this genus is by fai' the most abund-
ant and characteristic, being found in all of the formations above the Puerco, which
indeed has hitherto yielded no I'odents at all.

This genus has been defined by Cope (No. '3, p. 175-G) as follows: "The crowns
of the inferior molars suppoit four rather small anil strictly marginal tubercles, which
inclose a median valley. The anterior inner tubercle is more elevated than the others,
and the posterior two tubercles are connected by a low ridge on the posterior border,
which may be more or less tubercular on the last molar. In some of the species the
marginal tubercles are merely elevations of the margin, while, in others, the adjacent
tubercles of a pair approximate, so as to form a pair of interrupted cross-crests.

" There are five superioi- molars, of which the anterior is of small size. They
resemble those of Sciurus, but the transverse crests are obsolete or wanting. The
l)ositions corresponding to their exU/rual extremities are maiked by more or less dis-

Till-; MAMMALIA OK THE UINTA FOKMATIOX. 475

tinet cusps. There is a single internal tubercle of the crown. In the thiid and
fourth molars of P. delicatissimus, I observe rudiments of a second internal tubercle.

" The incisor teeth are compressed, with narrow anterior face. T!ie enamel is
not grooved and is little or not at all inllected on the inner side of the shaft, while
it is extensively so on the external face. There is a large vowndi foramen infnwrhitale
like that of Ischyromys and Fiber, and entirely unlike that of Gi/mno2)tychus and Sci-
unts, conforming in this respect to the forms of the extinct group of the Proloinyidui
of Pomel."

PLESiARCTOiAn'S sciUROiDES S. & O. (PI. XT, Figs. 1-2). The specimen upon
which this species is founded consists of a skull, lower jaw and several liaib-bones.
It dilfers from the Wasatch and Bridger species principally in the character of the
superior molar teeth, in Avhich the transverse crests, wanting in the earlier members
of the genus, are clearly though not very prominently developed. There are also
several ditlerences of minor importance in the structure of the skull. These charac-
ters may possibly be regarded as of generic value, but the changes are so slight and
the agreement with the typical species of Plesiarctomys so close, that we do not feel
justified in forming a new genus. In size the Uinta species is very small, much in-
ferior to P. delicatissimus.

The skull is very much like that of the last-named species, but with some not
unimportant differences. The muzzle is rather shorter, though both longer and
broader than in Sciurus ; the nasal bones are considerably broader behind than in
front, thus reversing the proportions found in Arctomys ; their anterior ends are some-
what thickened and slightly divergent, and posteriorly they terminate on the same
transverse line as the premaxillaries ; the latter are quite lai-ge and exhibit a consid-
erable surface upon the dorsal side of the skull. The frontals have a broad and nearly
straigbt anterior edge, but nari-ow very rapidly, showing a very strong postorbital
constriction of the ci-anium and no trace of postorbital processes, as is the very gen-
eral rule among the more ancient rodents ; they also seem to have a smaller antero-
posterior extent than in Sciurus, though this cannot be definitely decided. This poi*-
tion of the skull has quite a diffei-ent aspect from that seen in the two other species,
of which the structure of the cranium is known, P. delicatissimus and P. hians. It
differs from the former in that the postorbital constriction is less decided and less
elongated from before backwards, while the frontals are much more widely exi)anded
in advance of the constriction and over the orbits ; the superciliary ridges seem also
to be somewhat more prominent. So far as one can judge from Cope's ligure (J^o. 3,
PI. XXIV, Fig. 5) the postorbital constriction is deeper and the cranium more
tapering in front in /'. scinroides and P. hians.

476 THE MAMMALIA OF THE UINTA FORMATION.

As in the other species the infraorbital foramen is in the species before
us large and situated more immediately beneath the orbit than in Sciurus. The
shape of the lachrymal cannot ver}'^ well be made out. The parietals are apparently
very long ; they are broader than in Arctomys, narrower than in Sciurus, and the sagit-
tal crest is barely indicated. The squamosal has, of course, a corresponding de-
velopment ; the zygomatic arches are very slender and compressed ; they do not
arch out very strong!}' from tlie sides of the cranium and are somewhat shorter than
in P. delicaUssimus. The anterior part of the arch is shaped much as in Sciurus, but
the jugal has a somewhat greater vertical depth.

The occiput is very broad and low ; in the median line it forms a slight convexity
over the vermis of the cerebellum, and on each side of this there is a rather deep
depression. The paroccipital processes are very short, and the foramen viagmim
veiy large. The base of the skull is much injured, but enough is preserved to show
that the basi-occipital is rather broad and flat, and that the tympanic bullae are mode-
rately inflated. The posterior nares are rather long and their anterior edge extends
to the front of the last molar ; in Sciurus and Arctomys it is altogether behind the
molar series. The mandible is relatively more robust than m P . delicaUssimus ; the
masseteric fossa is, however, not so deep, and extends forward to beneath the second
molar ; the coronoid process is shorter, more slender and pointed, and the notch be-
tween it and the condyle is not so deep.

The train is in general like that of P. Mans, but the cerebellum is proportion-
ately larger, the hemispheres shorter, broader behind and tapering much more rapidly
forwards.; the olfactory lobes are also narrower.

The Dentition. The molar formula of this species, like that of the other mem-
bers of the genus is pra. ?, m. |. In the upper series the penultimate premolar (pm.

3) is implanted by a single fang and has a very small and sim-
ple ci-own, thus dittering strongly from the large pm. 3 of
Arctomys. Pm. 4 is nearly as large as the molars. The
crown is of triangular shape and consists of three tubercles at
/£x:J^ the anales of the crown, with a fourth one intercalated be-
DIAc..l.-UppeI•andlo^yermo. tween the internal and postero- external cusps, thus forming
\iu-& oi p. sHuroides. xf an interrupted posterior crest; the antero-external cusp is the
largest and most prominent of the three. This tooth resembles much the correspond-
ing one of Arctomys, but with the difference that the posterior line of tubercles have
not coalesced to form a crest and that the cingulum is much less strongly developed
on the front of the crown. The molars are very closely like those of Sciurus, but of
more distinctly triangular shape. In m. 1 and 2 there are two external cusps from which

THE MAMMALIA OF THE UIXTA FORMATION. 477

run two crests, meeting at the internal cusp and thus forming a V. In Sciurus the crown
is somewhat expanded behind the posterior crest, thus making the inner side of the
tooth nearly as broad as the outer ; in 1*. sciuroides this expansion is much less
marked. M. 3 has but a single transverse crest with a broad basin-like expansion
behind it. The upper molars of Plesiarctomys are obviously of the same fundamental
type as those of Sciurus and Arctomys ; their chief diflerence from the latter consists in
the fact that the cingulum on the anterior side of the crown is not strongly developed
and is scarcely visible from the side, whereas in Arctomys, this portion of the cingu-
lum is very prominent and gives the side view of the tooth an appearance of being
composed of three diistinct transverse crests.

The lower teeth are much worn, and it is therefore rather difficult to analyze
their construction. The single premolar is composed of a pair of anterior cusps, of
which the inner one is high and pointed, the outer one very small and but faintly
distinguished from its neighbor, and a low heel. The molars have four cusps sur-
rounding a median depression, of which the antero-internal one is much the most
prominent. The presence or absence of connecting crests could not be clearly made
out. The resemblance of these teeth to the lower molars of Sciurus is very close.

The humerus is short and very slender with large head and low, inconspicuous
tuberosities ; the bicipital groove is distinctly deeper than in /*. delicatissimus, and
the deltoid ridge is quite long and prominent. The distal end of the specimen is
broken, but enough remains to show that the epicondylar foramen is present and that
the supinator ridge is somewhat less prominent than in the Bridger species so fre-
quently referred. Whether the Uinta species had such a prominent internal cpicon-
dyle is not certain.

The tibia is quite long, slender, and strongly arched forwards ; the distal end is
quite deeply grooved for the asti-agalus and the internal malleolus is well developed.
The fibula, although not preserved in the specimen, was evidently entirely distinct, as
the outer face of the distal end of the tibia is uninjured and exhibits no trace of co-
ossification with the fibula.

Aleat'irements. M.

Length of skull OKI

Brcadlh of skull in front of orbits 025

" behind ■' 010

Length of lower jaw 028

Lcnglli upper molar scries 013

lower " " Oli

Anteroposterior diameter of lower incisor OOH

Transverse " " •• W2

Length of humerus (esliniiilid) 04ii

•' of libia 0«0

A. V. S. — VOL. XVI. 'h.

478 THE MAMMALIA OF THE UINTA FORMATION-.

The chief interest wliieh attaches to Plesiarctomys lies in the light which it thi'ows
npon the difficult problem of the origin of the rodent dentition. As to this question
Schlosser says : " Die Ziihne dieser altesten Formen erinnern einigermassen an das
Omnivoren-Geh\&s. Doch lasst sich die urspriingliche Beschaffenheit nicht mehr
erraitteln. "Wahrscheinlich bestand der einzelne Backzahn aus sechs Hiigeln
('Denticnles '), deren je drei in einer Zahnhalfte vorhanden waren. Der Typiis eines
solchen PriV/i/iM'-Omj/it'oreji-Zahnes ist noch am besten in den oberen Molaren von
Diclwhune erhalton. Hier tiitt nicht selten noch der sechste Hiigel auf, worauf schon
Kowalevsky aufmerksam gemacht hat. Bereits zu Anfang der Eocaenzeit erscheint
eine grosse Anzahl verschiedener Niiger, die indess nur insofei-ne unter einander ver-
wandt sind, als sie mit Ausnahme von ganz wenigcn Arten — Mijomoiyha — als Mittel-
formen zvvischen den in der Jetztvvelt ziemlich scharf getrennten Uystncomorphen
und Sciumorphen angesehen werden miissen. Auch fehlen keineswegs Mittelformen
zwischen diesen and den Myomoiyhen'''' {'Eo. 14, pp. 98-99). After a further discus-
sion of the subject Schlosser concludes that the rodents are to be derived from the
miirsupials.

Plesiarctomys lends but little snppoit to these views, and this genus is entitled
to special consideration as being one of the oldest if not the very oldest known form
of rodent. Here the superior molars are plainly of the tritubercular pattern, forming
distinct crests only in the later species. The inferior molars show the anterior trian-
gle of three cusps with a talon behind, or what Cope has termed the tuberculo-secto-
rial molar. This is the type of dentition which is all but universal in the Puerco
fauna, ungulates, creodonts and lemuroids all exhibit it, and Plesiarctomys seems to
show that the rodents are to be derived from the same generalized group of primitive
placental mammals, the Bunotheria, to which Ave refer the origin of the types just
mentioned. It is, to say the least, very suggestive that the most ancient known
types of rodents should exhibit the tritubercular plan of molar structure so unmis-
tably, though it must he admitted that this of itself is not conclusive against their
derivation from the marsupials. It removes, however, the necessity of such a deriva-
tion, and the other reasons assigned by Schlosser for his view do not appear to be
very cogent.

Plesiarctomys is, on the whole, to be regarded as a member of the Sciumorpha,
but cei'tain features, such as the lai'ge infraorbital foramen and the absence of i)0st-
orliital processes, are jjoints of resemblance to the Hystricomorpha. This association
of characters, as is shown in the passage already quoted from Schlosser, is very gen-
erally found to occur in the Eocene rodents.

TIIK MAMMALIA OF THK UINTA FORMATION. 419

AKTIODACTYLA.

LEPTOTRAGULUS S. & 0.

Our pieliminaiy account of this genus was founded upon the mtindible and infe-
i-ior dentition. Since then a considerable amount of new matcnal has been brought
to light in the collection which mateiially modifies some of our conclusions. The
genus may be thus defined: Selenodont artiodact^'ls with probabl}"^ unreduced den-
tition; lower premolars simple and ti'cnchant ; molars with short crowns; the cres-
cents of the lower molars are more or less trihedral, and the internal ones not flat-
tened into thin lamina?. The ulna and radius closely applied, but not coalesced; the
number of functional digits in the manus probably four, in the pes two.

Leptotkagulus pkoavus S. & O. (PI. VII, Figs. 9-1(5). This species, the only
one as yet certainly indicated fi-om the Uinta deposits, is represented in the collection
by sevei-al specimens, including the mandible, inferior molar series and last premolar,
some fragments of vertebra3 and portions of the humerus, ulna and radius, metacai-
pus, phalanges, scapula, tibia, tai'siis and metatarsus. Imperfect as these remains
are, they suffice to show the principal characters and systematic position of the
genus with sufficient clearness. Unfortunately no part of the skull or of the upper
teeth has as yet been identified.

The Mandible. Only the horizontal ramus is pi-eservcd in any of the specimens;
this is elongated, compressed and slender, becoming especially thin beneath the ante-
rior premolars, and thus offering a strong contrast to the short deep mandible of /*ro-
toreodon with its abrupt chin. The symphysis extends back nearly as far as the
front of the second premolar; two minute venous foramina are placed, one above the
other, beneath the third premolar.

Dentition. The fraguientary state of the specimens prevents the determination of
several important points with refeience to the lower teeth of this genus. Thus, the
number and character of the incisors and canines are altogether uncertain, though [nob-
ably they were present in lull. In front of the third ()remoIar is a short diastema, and
in advance of this occurs a pair of oblirpiely placed alveoli, which may have contained
the first premolar, or the canine and lateral incisor, which latter view seems the more
probable on account of the procnmbency indicated, though this would necessitate an
exceedingly short symphysis. In this case, the number of lower premolars would be
but three. The premolars ai-e very simple and trenchant, and in general appearance
quite like those of IViif/ulus. Pm. 2 has a very compressed and sh'.irjjly trenchant
crown; it has a sharper posterior edge than in Traf/ul us ixnd is ap|)arontly not divided,
as in that genus, by a cleft. Of pm. 3 only the fangs are left. Pm. 4 is slightly

480 THE MAMMALIA OF THE UIISTTA FORMATION.

more .complicated than its predecessoi's ; it consists of a high, acute, and compressed
cone with a small and sharp antero-internal tubercle ; from the inner side of the apex
of the principal cusp there runs downwards and backwards a thin ridge, which en-
closes a very narrow valley between itself and the outer posterior edge of the crown.
The corresponding tooth in Tracjulus has quite a ditferent appearance.

The molars are very brachyodont and are rather short in the antcro-posterior
direction ; the inner cusps are conical and thick, not flattened out into thin plates ;
the outer crescents are somewhat trihedral in section and rather angular than curved.
The valleys are widely open at the top, but rapidly narrow below, becoming
mere slits after a moderate degree of attrition. The two valleys of the same tooth
are not quite connected, being separated by the deep constriction between the two
outer crescents and by the coalescence of the posterior turn of the anterior crescent
with the inner wall of the tooth. They are, however, closely adjacent and even in an
advanced stage of wear do not become widely separated. The cingulum is quite
sti-ongly marked upon the anterior and posterior faces of the crown, as well as be-
tween the outer crescents, where it foi'ms a distinct accessory column, which is largest
on the first molar. The heel of the last molar is relatively large and encloses a broad
valley ; a narrow cleft appears upon its inner wall.

The teeth associated with the skeletal fragments about to be described are so
much damaged that their reference to this genus is not quite certain, though very
probable ; they are somewhat larger than those of the type specimen, and may, per-
haps, represent a second species.

Vertehrce. An isolated atlas is probably to be referred to this genus. It just
reverses the proportions of the atlas of Protoreodon in being long and narrow, and
with but slightly extended transverse processes ; between the anterior and posterior
articular surfaces the body of the vertebra is deeply constricted. The anterior co-
tylus for the occipital condyle is very deep and transversely extended, narrow from
above downwards, and is slightly notched on the outer edge. The transverse p)"0-
cesses are thin plates and do not appear to be perforated at the base by the verte-
brarterial canal. The surfaces for the centrum of the axis are flat, oval and placed
almost transversely ; there is no facet upon the posterior edge of the inferior arch of
the atlas, for the centrum of the axis beneath the odontoid process, such as occurs in
the camels and true ruminants. Fragments of other vertebrai occur in the collec-
tion, which undoubtedly belong to Leptutragidu^, but they arc not sufficiently well
preserved to warrant desci-ii)tion.

Of the scapula, the distal portion is preserved. The glenoid cavity is oval in
shape, the antero-postcrior diameter exceeding the transverse ; the coracoid is promi-

THE MAMMALIA OF THE UIXTA KOIIMATIOX, 481

nent and but little recurved. The position of the spine is somewhat different from
that seen in the recent selenodonts, in which the spine is phiced in advance of the
median line, thus making the prescapidar fossa much smaller than the postscapular.
This inequality is also shown in a less degree in Zieptotroyulns, in which respect this
genus agrees with the earlier selenodonts, such as Anojjloihermm and Oreodon.

Of the humerus we have only the distal end, which, however, shows some inter-
esting characters. The internal epicondyle is not pi-orainent and is pushed to the pos-
terior side ; the inner condyle of the trochlea is much extended transveiscly, while
the outer condyle is narrow; the intercondylar ridge is not prominent, and is above
rounded and rather broad, but rapidly becomes narrow and sharp. This portion of
the humerus forms a sti'ong contrast to that of the oreodonts, v/hich finds a parallel
onlj' in AnopIoVierium. Tlie anconeal fossa is deep and nariow and appears to per-
forate the shaft.

The ulna and radius are very closely applied together, but they do not coalesce,
at least in the proximal portion, whatever may be true of the distal end. The ulna
has a heavy and prominent olecranon and a deeply incised sigmoid notch, with a re-
curved superior margin. The contact with the humeral condyles is limited to very
small inner and outer facets, which are but continuations backwards of the corre-
sponding radial facets. The shaft is quite broad at the level of the head of the radius,
but rapidly tapers below this point and becomes very slender and reduced. The
head of the radius is both broad and deep ; it is only slightly concave, and the division
into facets only obscurely marked ; the anterior margin rises somewhat towards the
inner side ; the groove for the intercondylar ridge is very shallow, and the front margin
is not notched. Below the head, the shaft tapers and becomes more rounded. Un-
fortunately neither the distal ends of the forearm bones, nor any of the carpal ele-
ments, have been found.

The femur is represented by a much damaged pioximal end, which, however,
presents some interesting features. The head is small, hemispherical, and set upon
a long and very distinct neck; the pit for the lie/amentum teres is but faintly marked.
The great trochanter is broken away, but it was obviously large and, with
the strongly projecting neck, makes this region of the bone unusually broad. The
second trochanter is large, rugose and prominent; what appears to be the remains of
a ridge, connects this trochanter with the greater one, as in Ili/ojtuUnnus, to the femur
of which this fragment bears considerable resemblance. The shaft is broad and flat-
tened alx)ve, but appears to taper rapidly and to become more rounded below.

Of the tibia we have only the distal end, with fragments of the proximal con-
dyles, which, however, are too much injured for description. The lower portion of

482 THE MAMMALIA OF THE UIN^TA FORMATION.

the shaft is transvcrsel}' extended and quite massive, expanding but little into the
distal end. The articular surflices for the astragalus are quite deeply incised and
separated by a prominent intercondylar ridge. The outer edge of the distal surface
shows a narrow facet for the fibula, indicating that the latter had commenced to shift
beneath the tibia; though the narrowness of this facet and the large rugose surface
upon the external side of the tibia, make it probable that the redaction of the fibula
was by no means complete. It has, however, clearly begun.

The foot structure of Leptotragulus is very curious, though we arc unfortunately
not in a position to give a complete account of it. The manus seems to have pos-
sessed four functional digits, and is represented in the specimen before us by the
proximal ends of what appear to be the second and fourth metacarpals. Mc. ii (?)
is slender, but hardly more reduced than in Oreodon. Its proximal end shows three
facets, of which the largest is a concave one for the trapezoid, and behind this a
separate one for the trapezium ; a small oblique sui-face towards the ulnar side seems
to be for the magnum, thus indicating an " unreduced '' type of manus. Mc. iv is
much stouter and has a trihedral shaft and a simple, saddle-shaped surface for the
unciform. On the ulnar side is a deep depression, apparently for the fifth metacarpal.
This account is given with reserve, as the fragments described are too imperfect for
certain identification, and the differences indicated between the fore and hind foot are
vei'y remarkable.

The pes is fortunately much better preserved than the manus. The calcaneum
is rather long, compressed and deep, with approximately parallel edges. The fibular
facet is prominent, strongly arched from befoi-e backwards and descending abrujjtly
in front. An astragalar surface is on the inner side of this facet and forms a shai-p
angle with it. The distal end of the calcaneum is very narrow, but of considerable
antero-2:)Osterior extent, and bears upon its inner side an unusually large surface for
the astragalus. The latter is strikingly high and narrow ; the proximal condyles are
small and quite widely separated ; the outer one is divided by a considerable interval
from the cuboidal facet, while the innei' extends further down and reaches the navicu-
lar surface. The fibular facet upon the outer condyle is very long. The distal trochlea
is unevenly divided between the facets for the navicular and cuboid, the latter being
very narrow. The inferior surface foi" the susteatnctdicm tali is long and narrow and
separated by a deep pit from the overhanging external proximal condyle. The cuboid
is high, narrow, and deep (antero-posteriorly) ; the calcaneal facet is arched from
before backwards and in front descends much below that for the astragalus; it is also
slightly broader than the latter; upon the tibial side of the cuboid are two small
and widely separated facets for the navicular, the position of which indicates that the

THE MAMMALIA OF THE UIl^TA FORMATION. 483

latter hone was comparatively low, and the cuneiforms high. The distal end is en-
tirely occupied by the large facet for metacarpal iv, though what seems to be a small
facet for nit. v occuis upon the external side. The po.sterior hook-like process of the
cuboid is long, heavy and lecurved.

The metatarsus apparently consists of two functional mombors, though grooves
and facets upon the sides of these indicate the presence of at least the proximal
jjortions of the lateral metatarsals. Mt. ii appears to have been less reduced than
mt. T. The median pair are more slender than the corresponding metacarpals, and
were i^robably more elongate; their shafts are flattened and the approximate surfaces
very closely applied, but show no tendency to coossify ; both have strong posterior
projections from the proximal ends. The head of nit. iii would .seem to indicate that
the nieso and ectocuneiforms were not coossified. Mt. iv covers the entire distal
surface of the cuboid, allowing to nit. v only a lateral connection with that bone.
The distal ends of the median metatarsals at once suggest the eharacteiistic cannon-
bone of the Tylopoda, and are i-emarkably like those of the AVhite Kiver representa-
tive of that group, Poehrotheriwn. The articular surface is but slightly marked off
from the shaft, and the keel, though piominent, is confined entirely to the palmar
side. When the ends of the two metatarsals are placed together they exhibit a slight
divergence, as is the case in a much more marked degree in the recent Tylopoda.

The i)Jialan(/es of the proximal row are long and slender and curved outwards ;
the proximal surfaces are somewhat symmetrical, causing a slight divergence of
the toes. Tiie ungual phalanges are high, compressed and pointed, and somewhat
flattened on the internal (approximate) sides; they resemble the unguals of the ante-
lopes much more tlian those of recent Tylopoda.

TJie Sydemalic Position of LeptulnKjulin^.

In our preliminary account, drawn up IVoni the mandibular dentition only, we
inclined to the view that this genus was allied to J'rodreniotherium, and was, perhaps,
the forerunner of the White Kiver 2ycp<on<eri/a;. The new material iilcntilicd .since
that description was written, as well as direct comparison with original .speciniL'U.s of
J'ro(Iremothfriu7i), shows that our former conjecture is entirely untenable-. The teeth,
and still more strongly the skeletal fragments, prove that Lrp/otriiynliis i.s very clo.sely
allied to J'oehrdlhcriiaii, and is, theiefore, the earliest known iei)resentative of the
Tylopoda. If more perfect specimens should confirm the douhl I'lil indieatioii in tiie'
type of the loss of the first lower premolar, this species, at least, could not be jjlaced

484 THE MAMMALIA OP THE UINTA FORMATION.

in the direct line of descent, though in any event it is but slightly removed from
that line.

The inferior premolars are composed of the same elements as in Poebrotherium,
but have quite a different appearance, owing to their moderate antero-posterior extent,
while in the latter genus they are very much elongated ; the difference between the
two being least marked in the ease of the last premolar. The most important dis-
tinction in the molars of Leptotragulus from those of the White River genus consists
in their completely brachyodont chaiacter; they are also not so far removed from the
bunodont pattern ; the inner cusps are stout and conical, and the outer distinctly tri-
hedral, while in Poehr other turn the crescents, and especially the internal ones, are
compressed into thin laminae. The crescents are not so sharply separated by deep
constrictions, and the valleys are more nearly confluent in the Uinta genus ; the heel
of the last molar is also pro])oitionally larger and contains a deeper valley. Another
difference consists in the absence fi-ora the molars of Poebrotherium of the small col-
umns developed from the cinguUim between the outer crescents. The elongated
slender shape of the mandible recalls that seen in Poebrotherium.

The atlas is almost a reduced copy of that of Poehrotherhim, but the notch at
the side of the anterior cotylus is not nearly so deep, and there is no articular surface
developed on the hinder edge of the inferior arch to connect with the centrum of the
axis below the odontoid process.

The scapula has a more oval glenoid cavity, a less prominent coracoid, and the
spine is placed more nearly in the median line ; the intercondylar ridge of the distal
end of the humerus is narrower and less prominent.

If we may trust the account given above, the manus is very different in the two
genera in that the manus of Leptotragidus is but slightly reduced, and that the sec-
ond metacarpal retains its connection with the magnum. The fibula seems also to be
less reduced, though the process has obviously begun. The tarsus, metatarsus and
phalanges are much like those of Poebrotherium, with only slight difierences, such as
the lai'ger size of the cuboid hook and the relative narrowness of that bone. The
unguals are very closely alike in the two genera, and indicate that the peculiar foot
structure which characterizes the camel and llama could not have occurred in the
Uinta and White Kiver representatives of the group, which probably had feet with
the general appeai'ance of those of the ordinary ruminants. The pad or cushion
developed, no doubt, 'i)ari-2KiM^a with the formation of the peculiar cannon-bone with
its diverging distal ends. The ungual phalanges of Protolabis and Procamelus have
not as yet been identified, but there is every reason to suppose that they exhibit suc-
cessive stages of reduction.

THE MAMMALIA OF THE UINTA FORMATION. 485

Leptotmgulus is, therefore, to be regarded either as the direct ancestor of Poehro-
therium, or as very closely connected with that ancestor. Unless, however, the doubtful
indications of tetradactyl feet in the Uinta genus should be confirmed, it throws no
groat amount of light upon the vexed question of the origin of the camels and their
relations to other selenodonts. Among the imperfectly known and sparsely represented
artiodactyls of the Bridger, the one which most clearly suggests itself as a forei-unner
of Leptotragulus is Ilomacodon. This animal is described by Marsh (!N'o. 9, p. 364)
as being " very neai'ly allied to Heloliyns, and but a single step away from this genus
towards the selenodonts. * * * This piimitive selenodont had forty-four teeth,
which formed a nearly continuous scries. The molar teeth are very similar to those
of IL'lohijus, but the cones on the crowns have become paitially triangular in outline,
so that when worn the selenodont pattern is clearly recognizable. The first and sec-
ond upper molai-s, moreover, have three distinct posterior cusps, and two iji front ; a
peculiar feature, which is seen also in the European genera, Dicliohune and Caiao-
therium. There were four toes on each foot, and the metapodial bones were distinct.
The type species of this genus was about as large as a cat. With Helohyus this
genus forms a well-marked family, the Ileloh tjidLe."

The association of these two genera into a single family is (piite inadmissible,
as it ignores a fundamental ditference in the construction of the upper molars ; in
Hdolnjus the fifth cusp is in the anterior half of the tooth, while in Ilomacodon it is
in the posterioi- half This latter genus belongs clearly to the DichobanidiK, and the
puljlished description shows no reason for separating it fi'om Dichohune itself. Before
identifying it with the European form, however, it seems best to wait for fuller infor
mation with regard to the American species. Similarly, in the absence of the upjjcr
molars, it would be premature to state that Leptotrar/nlus is to be derived from IIo-
inacodon, though there is nothing in what is known of the two genera to forljid such
a derivation, which may thci-efore be fairly assumed. If lliis connection should be
confirmed, it would necessitate revision of the current views upon the relationships of
the Tylopoda. Schlosser (No. 13, p. 42) regards this group as a branch of the same
general stem which developed the Ano])lothen'dce, the Anthracolheridit', etc., and espe-
cially as being connected with the oreodonts, a suggestion which has also been made
by Uutimeyer (Xo. 11, p. 98). On the other hand, Schlosser derives the tragulines
and true ruminants from the JJichobunidti', thus making a very radical distinction
between the Tylopoda and the other existing selenodonts. If, however, the sugges-
tiiui hcie made, that Ilomacoflan and Leptolraqxhis are genetically connected, be con-
firmed, it follows that the Tylopoda are also to be derived from the Dicholmiiiddi, and
therefore from the same origin as the other recent selenodonts. Such a result would

A. P. S. — VOL. XVT. .'5.1.

486 THE MAMMALIA OF THE IJTNTA FOEMATION,

further conclusively demonstrate that the oi-eodonts are but remotely connected with
the camels, as is indeed made exceedingly probable by what we already know of the
earlier members of the two groups. Protoreodua, which will be fully described here-
after, shows the unpaired cusp in the anterior half of the upper molars, just as in
Heloliyus, from which, indeed, it seems to have been derived, and its i-esemblance to
such types as Anoplutherimn and the (so-called) Eocene species of ILjopotaruus is
very marked, and Schlosser is fully justified in associating the oreodonts with
these phyla. On the other hand, the resemblances between the camels and the
oreodonts are only such as are common to all the more primitive selenodonts,
and indeed this peculiar family has decidedly more similarity to the tragulines than
to the camels. Lejitotragvlvs is almost as different from its contemporary Protoreo-
don, as Oreodon is fi-om Poeirothermm, or Merycliyus from Procamelus. The Bi'idger
representatives of these two phyla are apj^arently Ilelohyus and Ilomacodoji, which
not imjirobably take their rise from a common starting point in Pantolestes of the
Wasatch, which would thus be the progenitor of the two chai'acteristically Amei*-
ican groups of selenodonts, as well as through the DicJwhwiidfi} of the European
series also. Of course, in the present state of knowledge, these suggestions can
only be made tentatively ; they are not yet capable of demonstration.

Whether we adopt the scheme of genetic relationships propounded by Sclilosser,
or the one here suggested, it necessarily follows that the word, selenodont can only be
used as a term of description and not of classification ; for it seems clear that this
pattern of dentition has been independently assumed by many groups of artiodactyls.
Nearly all the types of selenodonts may be traced back step by step to the general-
ized buno-selenodonts of the Eocene, and the resemblance between the molar teeth
of such forms as Trayulus and Cainotherium, Oreodon and Cervus is much more
probably due to mechanical conditions than to genetic relationship.

Measurements. ^f. M. M.

* Jlomitrodoii •carjiins. \ Lcjitvtriigiiliis proavus. Poihrothcrium wilsoni.

Lonsilli lower premolar , 'erics (last three) .021 .030

Lasl lower premolar antorcpost. diumeter .007 .011

" " transverse " .003

First lower molar anlcropost. diameter .008 .011

transverse " .005 .000

Length lower molar series 017..') .020f .042

Last lower molar aiitero-post. diameter 007.3 .012t .01!)

" transverse " ,001 .OOOf -007

Height of astragalus 014 .020

* These measurements from Marsh, Am. .Jour. Sti. and Arts (3). IV. p. 120.
f Second specimen.

THE MAMMALIA OF THE UINTA FORMATION. 487

rilOTOREODDX S. & O. (PI. VII, Figs. 1-s).

This genus is the most perfectly known and after TriplopuK the most abundant
of all the Uinta mammals. The Princeton collection contains portions of a large
number of individuals representing nearly all the important parts of the skeleton and
pertaining apparently to two species. The genus is to be defined as follows : " Sel-
enodont Artiodactyla with unreduced dentition ; inferior canines with the form and
function of incisors, first lower premolar cuniniform ; superior molars of five crescents,
the unpaired one nuicli reduced and situated in the anterior half of the crown, outer
wall erect and flattened ; crowns very low and broad, valleys widely open ; inferior
molars much like those of AgriocJia'rtis with conical inlernal crescents. ^N'o diaste-
mata in lower dental scries and a very short one behind the u])per canine. Cranium
narrow and very much elongated, orbit open behind, apparently no lachrymal pit.
Lunar resting more directly upon the magnum than in Oreodon ; inanus pentadactyl ;
pes tetradactyl and inadaptively reduced.

Before proceeding to describe the osteology of this remarkable genus it may
be well to glance at previous notices of the Uinta selenodonts. The first one de-
scribed is the Arfriochmrus yumiluii Marsh (No. 8, p. 250), which is founded upon the
inferior dentition. It seems highly imi)robable that this White River genus should
occui- in the Uinta, and the lower molars of Protoreodon being, as we have already
seen, much like those of Agriochoerus, one is led to infer that this supposed A'jrio-
chuinis belongs to Protoreodon, 1)ut in the absence of the upper molars this cannot l)e
certainly determined. Prof. Marsh (No. 9, pp. 361-5) subsei[uently named three
genera fi'om this formation, of which he says : " In the Diplacodoii horizon of the
Upper Eocene, the selcnodont dentition is no longer doubtful, as it is seen in most of
the Artiodactyla yet found in the^c beds. These animals ai'e all small, and belong to
at least three distinct gencia. Oni' of these, IiJomeryx, closely resembles Ilomacodon
in most of its skeleton, and lias four toes, but its teeth show well-marked crescents
and a |)artial transition to iliu tfftli of Ilijopotanuis from the Eocene of Europe.
With this genus is another (Parameryx), also closely allied to Homacodoii, iuit appa-
rently a straggler from the true line, as it has Init three toes behind. Tlu^ most pro-
nounced selcnodont in the Upper Eocene is the Oromeryx, which genus apjjcars to
be allied to the existing deer family, or Cproidr,, and if so is the oldest known repre-
sentative of the group." "The least specialized, and apparently' the oldest, geiuis of
this group [the oreodonts] is Ayriochrrrn-i, which so nearly resembles the older Ily-
opolatiiits, and the still more ancient Eomeryx, that we can hardly doubt that they
all belonged to the same ancestral line." ''A most interesting line, that leading to

488 THE MAMMALIA OF THE UINTA FORMATIOlSr.

the camels and llamas, separates from the primitive selenndont line in the Eocene,
probably through the genus Paranisryx." Whether any of these genera includes the
previously described Agriochoerus ■pumiliis, we are not informed, but the reference to
the i-esemblance between the dentition of Eomeryx and that of ffyojwtcnmcs and
AgriocJmrus may perhaps be taken to imply that the former is founded upoH the
superior dentition of A. pumilas, and may therefore be the same as Protoreodon.
That Parameryx may be identical with Leptotragulus is suggested by the reference
of the former to the camel line. However, these meagre accounts, with no species
descriptions, and no accompanying figures, are insufficient for identification.

The osteology of Protoreodon is in most res[)eets vei-y similar to that of its
White River successor, Oreodon, though in several respects it is more primitive and
presents many interesting indications which serve to point out the line of descent
thi'ough which this remarkable family has passed, as well as its relationships with
other groups of selenodonts, both American and Eui-opean.

The sJcull, which fortunately is in a fairly good state of preservation, presents
some features of great interest. It resembles in general construction that of the
Oreodontidce, being in some respects like that of Oreodon, in others more like the
skull of Agrioclicerus, while, in several ways, it is more primitive than either. As a
whole the skvdl is longer and narrower than in the Miocene forms, a statement which
applies especially to the cranium and more particularly to the posterior region of it
behind the post glenoid processes, the length of which region is to a remarkable
degree greater than in the later genera of the group. The face, on the other hand, is
rather short, the anterior edge of the orbit being above the first molar. The cranial
and facial axes are in the same straight line, and the upper contour of the skull is
also nearly straight, as it is not complicated b^^ the development of large air sinuses
in the frontal region. The sagittal crest is very long and prominent, the occiput low
and narrow.

As is general among the Oreodontidx the parietals have a very great antero-
posterior extent and form with the squamosals almost the entire cranium ; posteriorly
they are very narrow, but become much broader in front of the squamosals, where
they send down great processes to meet the alisphenoids. The high sagittal crest
extends the entire length of the pai'ietals. The occiput is rather strongly convex,
but is low and narrow, as are also the exoceipitals, which do not extend upon the
side walls of the cranium. The paroccipital processes are long and slender, and are
much like those in Agrioclicerm, but are furthei- in advance of the occipital condjdes
and even more widely separated from tlic postgleuoid processes than in that genus,
the elongation of this region of the cranium being especially characteristic of Pro-

THE MAJrMALIA OF THE UINTA FOKMATION. 489

toreodon. From the crushed condition of this part of the skull it cannot ver}- well
be determined whether or not the peiiotic was exposed upon the surface, but in all
pi-obability it was. The frontals form but very little of the roof of the cranium, as
they terminate just behind the orbits and are chiefly confined to the covering of the
orbits and nasal cavity. The supraciliai-y ridges are prominent and diverge from the
sagittal crest, as in Oreodon, terminating in long, overhanging postorbilal processes,
but. as in A(jriochcerus, they do not leach the jugal and the orbits are widely open
behind.

The squamosals ai-e very large and form most of the side walls of the cranium ;
they articulate with the parietals both superiorly and anteriorly. The zygomatic
processes are very long ; the free poition is compressed laterally and quite slender,
but the bases have a considerable antero-posterior extent, as in Oreodon ; the post-
glenoid processes are less massive, and are less produced transversely and more ver-
tically than in that genus. The glenoid cavity is also somew^hat different ; Leidy
thus describes it in Oreodon (No. 7, p. 75): "The glenoid articulation exhibits a broad
surface extending outwardly on the under pait of the posterior root of the zygoma.
Its fore part is nearly straight and hoiizontal transversely, and is modei-ately convex
antero-posteriorly, inclining forward internally and backward externally. Postero-
internally it descends upon a remarkably large and strong postglenoid tubercle,
which is antero-posteriorly compressed, mammillary in shape." This description
applies also to the glenoid cavity of P/oforeofZo/i, except that the surface has a smaller
extent from before backwards, it is rather more strongly convex in front and is not pro-
duced into an elevation internally; the postglenoid process is also much less massive.

In correspondence with the great elongation of the cranium the zygomatic arches
are very long; the jugal is long and quite slender and has but a rudimentary post-
orbital process, which is widely separated fioin that of the frontal ; it is not notched
to receive the anterior end of the zygomatic process, as in Oreodon, and has a some-
what greater extension upon the face than in that genus. The lachrymal, on the
contrary, would seem to be smaller, and apparently had no depression or pit, though
this statement cannot be made positively.

The base of the skull is unfortunately much injured mikI allows but little to be;
made out with regard to its structure. The alisphenoids aie very large and have
large pterygoid plates, which bound the much elongated posterior nares ; the latter
exteiids far forward between the molars and ends anteriorly in a pointed arch, so that
both in shape and position it agrees closely with that f)f . (r/;-/ot'//"'//<s, and is (piile
dillerent from that of (freodon. It cannot be ascerLaiued, whelhcr or not the tym-
panic bulUe were intluted, but we may infer that they were from the iact that such

400 THE MAMMALIA OF THE UINTA FOKMATION.

inflation occurs botli in Agriochoerns and in the oldest species of Oreodon which are
found in the lowest horizon of the White River formation, the Tttanotherium beds.

The maxillary resembles that of Oreodon, being very low beneath the orbit and
extending up to the nasals in advance of the lachrymal ; the alveolar portion is low
throughout in correspondence with the exceedingly low crowns of the teeth, and even
the portion of the maxillary which forms the side wall of the nasal cavity is propor-
tionately lower than in the Miocene genus. The palatine processes are rather narrow,
and a narrow incision occurs between the alveolus of the last molar and the palatine
l^late; this incision is also present in Oreodon, but is broader. The infraorbital fora-
men is situated above the penultimate (pm. 3), which is the position it occupies in
Agriochcerus, while in Oreodon it is placed somewhat more anteriorly. I^^either
nasals nor premaxillaries ai-e preseiwed in any of the specimens.

The mandible is short and deep, with an abruptly rounded chin and long, very
steeply placed symphysis. The condyle is shaped very much as in Oreodon, but the
coronoid process is more like that of Agriochoerus in being higher and more deeply
sepai'ated from the condyle, and consequently less like that of the peccar3^ So far
as can be determined the angle of the mandible was not so much thickened as in
Oreodon; the two rami are not closely interlocked and even in old individuals show
no tendency to coossification.

The dentition is of especial intei;est as it serves to connect the isolated and pecu-
liar family of the Oreodontidce with the other selenodonts of the American and Euro-
pean Eocene.

Upjjer jaw. The superior incisors are not repi-esented in the collection, and only
the fang of the superior canine; the latter, however, shows the characteristic D-
shaped section seen in Oreodon, and doubtless the crown presented no important
ditlerences. The same statement applies to the most anterior pi-emolar (pm. 1).
The second premolar, seen from the outer side, resembles much the coi'respond-
ing tooth in Oreodon,, having a compi'essed conical crown with trenchant edges.
Seen from below, however, it is more simply constructed, as the small internal access-
oi-y valleys are barely indicated, and the cingulum is but feebly developed. The
third premolar is moi-e transversely extended than the preceding one and has the ex-
ternal wall slightly concave, but is constructed essentially like it, and is consequently
simpler than the same tooth in Oreodon. Loidy describes these premolars in Oreodon
as follows (]S[o. 7, p. 81) : "The anterior thi-ee upper premolars are constructed after
the same plan. They decrease successively in size and in the degree of develop-
ment of theii- details of form, fi'om the third to the ftrst of the series. Theii-
crown is a trilateral [)yramid, with a pointed apex and a broad external cordiform

THE MA.'\1MAI.1A OF THE CINTA FOK.M ATIOX. -JOl

surface. The iiaiiowcr internal sui-faces appear as triangular inclined planes, sepa-
rated by a median acute ridge extending from the point to the base of the crown.
The anterior of the internal surfaces forms at the base a pair of shallow pouches, de-
fined by a double festoon. The posterior of the same surfaces forms a single and
larger pouch at the base of the crown, included by a single and thicker festoon. This
latter in the third premolai- almost assumes the dignity of an additional lobe to the
crown, resembling the inteiuial lobes of the true molars." In I'rotoreodon these fes-
toons are not more distinctly marked on pm. 3 than on pm. 2, l)«ing only very fuiiit
elevations of the cingulum and single, not double, on the anterior part of the crown.
Pms. 3 and 4 are each implanted by three fangs, pms. 1 and 2 by two. The last pre-
molar has the form seen in Oreuduii and the ruminants generally ; it consists of two
crescents, an internal and an external and an internal, with a strong internal cingulum.
The outer crescent still retains the cordate shape and the trenchant edges seen in the
anterior premolars, and is considerably larger than the external crescents of the
true molars. The cingulum is not so strongly developed on the outer side, nor is the
external wall of the tooth so deeply concave as in Oreudon.

The anterior premolars of Protoreodon, with their great simplicity of construc-
tion, are decidedly moi-e like those of Ar/riochcerus than those of Oreodon, Init the
fourth premolar resembles that of the latter genus, while in Ar/riochcerics it has
I'cached an extraordinary' degree of complication, having almost completely assumed
the molar pattern, and differing only in the rudiiTKJUtary state of the postero-inlernal
lobe.

The molars of Protoreodon increase in size successively from the lirst to the
third; the crowns are very low and broad in proportion to their antero-postcrior ex-
tent ; the valleys are very shallow and widely' open ; the internal crescents ai'C massive
pyi'amids, and the horns are very faintly mai-ked ridges. The external crescents are
intermediate in character between those of Oreodoa and of Ar/riochoerus, in being less
concave and overhanging and less deeply separated by an outward extension of the
median valley than in the latter, more so than in the former, with an additional
resemblance to Oreodon in the compressed shape of the buttress between the outer
lobes and in the faint convex ridge which rnns up from the apex of these crescents.
The balance of resemblance, as far as the outer wall of the tooth is concerned, in-
clines therefore towards Oreodon, while the construction of the inner half of the crown
and the general appeai'ance of the whole is more like that of Af/riorlKma*. A very
imi)0rtant difference from the molars of all the oi-eoilouts liillu-rto known consists in
the presence of a lilili 1((1)l' situated in the anterif>r ii.ill' of ilic crown; it i-; but
slightly separated from the antcio-iiitciiial crescent, with wliicli it is appairnlly

492 THE MAMMALIA OF THE UINTA FORMATION.

beginning to coalesce. These five-lolx-d upper molars are found among nearly all the
selenoJonts of the American and European Eocene, and the discovery of Frotoreo-
don confii-ms Schlosser's conjecture (No. 13, p. ^2) that the Oreodontidce have been
derived from animals vf\t\\ molars of this pattern, though this ancestral type is clearly
not what he supposed it would be, namely, the common form from which both oreo-
donts and Tylopoda have descended.

Lower jaw. The incisors increase in size fi-om the median to the lateral one ; their
crowns are proportionately higher than in Oreodon, and of somewhat different shape,
in that they are more flattened and quadrate in outline and the cutting edges are
straighter. In the median one the cutting edge is placed nearly at right angles to
the lateral edges, while in Oreodon the incisors are more acute and pointed. As in
all the later members of this family, except Plthecistes, the lower canine has assumed
the form and functions of an incisor, and forms the largest of that series, while the
fii-st premolar has become caniniform, a transformation which is entirely peculiar to
this line of selenodonts. Analogous changes occur in XijjJiodontherium, where pm. 2
is caniniform, and in Hyplsodus whei'e the canine and first two premolars have gone
over to the incisor series, ])ut only the oreodonts show the enlargement of the first
premolar and such a change of shape, that it almost deserves the name of a tusk. In
Protoreodon the caniniform pi-emolar is directed upwards and outwards, scarcely at
all forwards ; the outer side is quite strongly convex, the inner side is divided by
a ridge, running down from the acutely pointed apex, into two somewhat concave
surfaces, of which the anterior is somewhat the larger ; the anterior and posterior
edges of the crown are sharp and trenchant. In Oreodon both the outer and inner
sides of the crown are more flattened; that is to say, in Protoreodon the transforma-
tion is less complete, the tooth retaining very clear marks of the fact that the func-
tional canine is in reality a premolar. The other premolars have considerable resem-
blance to those of Oreodon, but are of less complicated construction and are more
compressed and trenchant. Pm. 2 is a simple compressed cone, with very small
transverse and considerable antero-posterior diameter ; on the hinder edge is a very
narrow and shallow depression, Avhich is much more marked in Oreodon, but not in
Ayrioclmrus, where this tooth is entirely simple and reduced in size. Pm. 3 is an
enlarged copy of pm. 2, except that the internal cingulum is more pronounced and
the posterior valley somewhat larger. In Oreodon this tooth is proportionately much
larger, the valley is enlarged, curved inwaidly and completely enclosed ; the anterior
half is also much more strongly concave on the inner side. Pm. 4 is more or less
bi-oken in all the specimens, but enough remains to show that it was more simply
constructed than in Oreodon, which Leidy thus describes : " From the median point

THE MAMMALIA OF THE UIXTA FORMATION". 49.'}

an oblique ridge desoends internally and terminates in a huge trilateral pointed
tubercle which springs from the middle of the base of the crown and rises nearly as
high as the principal point." In Protoreodon this tooth has a very much smaller
transverse diameter than in the Miocene genus ; the anterior half of the crown is
much less concave on the inner side, and it would seem that the internal tubercle was
I'udimentary and the posterior valley small. At all events, the greater simplicity of
the tooth can be seen from the anterior half of the crown, which is uninjured.

The true molars differ in several important respects from those of Oreodon and
are more like, but not identical with, those of Agrioclicerus. The inner crescents are
more conical and less compressed than in the former genus, and the outer cusps are
less distinctly crescent-shaped ; they are also more widely separated from the inner
crescents, so that the valleys are much broader and shallower. The anterior pair of
crescents is much more completely sepai'ated from the posterior than in Oreodon, by
a depression which runs across the crown. All this is equivalent to saying that in
Protoreodon the selenodont pattei-n of the lower molai's is much less completely de-
veloped than in Oreodon. Compared with the molars of Agrioclicerus, those of the
Uinta genus present the following dilt'erences : The internal crescents are convex on
the inner side, instead of being concave with a median ridge, and the basal tubercles
on these crescents are smaller ; the outei- crescents are less flattened and the bones
slightly more prominent, though leaving the valleys open in front. As in this genus,
the heel of the last molar is proportionately larger than in Oreodon.

The hrain of Protoreodon (PI. Ill, Fig. 1''), as indicated by the shape and size
of the brain ease, is very narrow and considerably elongated ; the hemisphei'cs aie
especially small and more simply convoluted than in Oreodon ; the convolutions are,
as in this gonus. latiiL'r broad and have a nearly straight fore and aft direction, con-
verging in fi'ont ; the niedihiteral gyrus is not very distinctly mai'kcd. The posterior
region of the brain including the cerebellum and medulla is very long in proportion
to the hemispheres, and apparently the corpora qnadrn/eimna were partially uncov-
ered V)y the interspace separating the hemispheres from the cerebellum. As a whole,
the brain is distinctly smaller in jjroportion than in the Miocene genera of the group.

The vertebral column is rejjresented in the collection by many s|)ecimcns fiom all
the regions; but as they are not in an especially good state of preservation, and as
they do not show any very striking dilVerenccs from the vcrlcbriv of Orrodmi. they
will not rer[uii-c an extended description.

The atlas is very similar to that of Or'odon and lescnibles tlKrrrur.' that of the
tragulines rather than that of the Pecora ; it is short in the antero-posterior direction,

A. I'. S.— Vf)I,. \\[. 3k.

494 THE MAMMALIA OF THE UINTA FORMATIOjISr.

but with widely extonded transverse processes, which are perforated by the vertebrar-
terial canal. The anterioi- cotylus for the occipital condyles is cpiite shallow, the
posterior faces foi- the axis are quite flat and with j^reater vertical than transverse
diameter. The axis is likewise similar to that of Oreodon, though with differences.
The atlanteal faces are narrower than in that genus, but higher, and form a consider-
able part of the side walls of the neural canal; the odontoid process is narrower and
more peg-shaped, and the articular surface on its lower side is not continuous with
the facets for the atlas, but separated by a faint ridge; the upper surface of the pro-
cess is marked by a quite high and strong i-idge, which loses itself posteriorly in the
floor of the neural canal. In Oreodon the upper surface of the odontoid is either flat
or, as is the case in \mmj specimens, it shows an approximation to the spout-like
form of the ruminants in the elevation of the edges, so that it becomes somewhat
concave from side to side. The shape of the process is thus seen to be quite different
in Protoreodoit. The centrum of the axis is keeled and quite strongly opisthoccelus ;
neither neural spine nor ti-ansverse process is pi-eserved in any of the specimens.

The other cervical vertebra^ ai'e i-ather longer in proportion than those of Oreodon
and have somewhat more markedly opisthocoelous centra ; the anterior ones at least
have either obsolete or very low spines, and apparently all except the seventh exhibit
the vertebrarterial canal.

Except for their light and slender construction, neither the dorsal nor the lumbar
vertebrae present any special peculiarities; the posterior dorsals are long and com-
pressed, the lumbai-s become broad and depressed in the hinder part of the region.
The artiodactyl characteristic of cylindrical and interlocking zygapophyses in the
posterior dorsal and lumbal* vertebr;e is quite as well developed as in the White
Kiver genera of the group. Several caudal vertebrae indicate that the animal pos-
sessed an unusually long and stout tail, even more so than in Oreodon, as is made
probable by the size of the transverse process upon the more anterior vertebrje.

Of the scapula only the distal portion is preserved, which is, however, sufficient
to show its more essential characters. The glenoid cavity is subcircular in outline,
much as in Oreodon and Hyopotamus ; the coracoid process is recurved and promi-
nent, and the spine rises abruptly from the neck, as in Oreodon and the ruminants,
and having a very different shape from that which occurs in the pigs. As far as can
be judged from the fractured condition of the specimens, the spine may be said to
divide the blade into nearly equal pre- and ])ostscapulai- fossae. Thjs position of the
spine is rendered the more probable from the iact that it occurs in Oreodon and
Hi/ojJotamtis, with the scapula of which the jrortions of the shoulder blade of 7Vo-
toreodon which are preserved in the collection closely agiee.

THE ArA^iAr.VLTA of the uin^ta formatiox. 495

Tlie hinnfras agrees closely with that of Oreodon, which, as Cope has pointed
out (No. 4. p. 508), is very peculiai- and difteiing from that of all recent artiodactyls,
finds its nearest analogue in Anoplotlierium. "'The greater tuherosity is large, rising
ahove the head; and is incurved, terminating inwards in an acuminate apex. Its
border at the base is thrown into an obtuse angle. The lesser tuberosity is small, and
is well separated from the gieater by a deep and wide bicipital gi-oove. The deltoid
ridge is distinct. The condylar extremity is more transversely extended than in any
recent artiodactyl, owing to the fact tiiat the posterior internal distal tube-
rosity is placed interior to the tiochlea instead of partially behind it, and that there
is, in addition, an iiiti'inal (.'picondyle not seen in the recent suilline or ruminant mem-
bers of the order. The intei'condylar I'idge is strong, and wider than in most recent
ruminants; in the suillines it has nothing like such a development. Another pecu-
liarity is the rtange-like free border of the exteinal trochlea, which is especially re-
curved at its superior part." With a few modifications this description will apply
equally well to the humerus of Protoreodon, The shaft is rather more slender and
the deltoid ridge much less massive and prominent; the anconeal fossa is narrow and
very deep, perforating the shaft, as is also genei-ally the case in the Miocene genus.

The ulna, as would naluially be expected, shows no tendency to coossify with
the radius; the olecranon is unusually long and stout and deeply grooved at its upper
end; the sigmoid notch forms somewhat less than a semicircle and passes below into
two small facets for the head of the radius. The shaft is stout and trihedral in the
upper portion, but becomes very much compressed below and presents a deep groove
on the internal side. The distal end is not preserved in any of the specimens. This
bone shows more difterences from the corresponding element of Oreodon than does
the humei'us; the most important of these are the proportionately greater size of the
olecranon and the sinjjular llatteniny; and ":roovin<>: of the shaft.

The radius differs but slightly from that of Oreodon, as far as tlie fragments pre-
served allow of comparison. The surface for articulation with tlic humerus is a
rather broad head, which is much compressed from before backwards; the articular
facets form three [)ortions, a median concavity for the intercondylar ridge, or tuber-
osity it might be called, and a more flattened sui-face on each side of this. The
external beveled surface for the curious flange on the humeral condyle above men-
tioned, is more concave and descends lower on the anterior far-e of the bone than in
Oreodon, and the anterior edge is somewhat more sinuous than in that genus, though
there is no emaigination for the intercondylar ridge, such as occurs in the anterior
proximal edge of the radius in the recent ruminants, and, in a much less marked
degree, in the suillines. This structure of tlu' head of tiu' radius is highly chaiac-

496 THE MAMMALIA OF THE UIXTA FORMATION.

tei'istic of the Oreodontidoi and is cori-elated with the equally characteristic shape of
the trochlea of the humerus. The ulnar facet on the posterior side of the head of
the radius is much less conspicuous than in Oreodon. The shaft has a somewhat flat-
tened, ti-ansversely oval shape, which does not tend to become cylindrical, as is the
case in Oreodon. Unfortunately we have not been able to identify the distal end of
the I'adius.

The carpus. As one Avould naturally expect, the structure of the carpus of
Protm-eodon is very much like that of the Miocene members of the family, though
some variations of importance may be observed, esi3ecially that the various elements
are less cuboidal in outline and of lighter construction.

The scaphoid is more extended transversely and less antero-posteriorly than in
Oreodon; the proximal surflice is more deeply concave, the rising in front more
abrupt and the radial facet descends further on the anterior side; on the distal sur-
face the facet for the trapezium is distinctly larger than in the Miocene t3'pe, where
it is exceedingly minute ; the trapezoid facet is also larger in proportion, while that
for the magnum is correspondingly smallei', nor are there any such anterior ridge and
posterior concavity as are to be seen in the later representatives of the family. In
Oreodon the lunai', as Cope has pointed out, has the remarkable peculiai'ity of resting
almost entirely upon the unciform, while the magnum has moved almost completely
under the scaphoid, a tendency which I'caches its maximum in 3ferycoc7iosrus and
Merychyus, when there is only a lateral contact between the lunar and the magnum.
Among recent artiodactyls, the only group where such a displacement of the carpal
elements is to be found is that of the Tragulina.

In Protoreodoii the same peculiar consti'uction of the carpus appears, but it has
not been carried to quite the same extent. The lunar has a gi-eater antero-posterior
diameter, the radial surRice is more extended in the same direction and is not nearly
so strongly convex. The distal beak-shaped prolongation, which passes between the
magnum and the unciform, has not moved quite so far toward the radial side, and
even in front the lunar rests somewhat, though but slightly, upon the magnum, while,
behind, the magnum is proportionately much larger and presents more directly down-
wards, instead of being rather more lateral than distal, as is found to be the case in
Oreodon. The unciform facet of the lunar is of course smaller than in the last-named
Ibrm. The Uinta genus thus presents a transitional stage between the carpus of the
more typical artiodactyls and that characteristic of the Orcodontida;, though the
tendency towards the latter is already very distinct.

The cuneiform dilfers more from the cori-espouding bone in Oreodon than does
any of the other carpal elements. Indeed, the specimen which we have regarded as

TllK MAMMALIA OF TIIF. UINTA FOK.MATIOX. 4'.)7

tlie ciiiK'ifonn of Protorefdoii may possibly belong to some other genus, though this
is not at all probable from the association in wliich it was found. It is much more
extended transversely and less antero-posteriorly than in Oreodon, as is also the
scaphoid; the ulnar facet is a simple groove which does not descend up(m the outer
side of the bone, thus agreeing with OrevJua and differing very markedly from
Dicuhjles: the pisifoi-m facet is very large, nearly tlat, and occupies the entire pos-
terior surface; the unciform facet is likewise quite diQerent from that of Oreodon in
being shallow, of less antero posterior and greater transverse extent. In general the
cuneiform of Protoveodon is quite low and broad, with an /-shaped upper contour,
highest on the internal or radial side and sloping down towards the external side.
Neither trajje/.ium nor trapezoid is represented in any of the specimens, but judging
from the facets on the scaphoid, both of these bones were better developed than in
Oreodon, implying a larger relative size of the lateral digits.

The magnum, on the contrary, is smaller and of a somewhat different shape ; the
proximal surface is divided nearly evenly between the facets for the scaphoid and lunar
which meet at a high angle so as to form a sharp ridge along the superior median
line. Seen from the side the upper contour of the magnum forms a ilattcned arch,
whereas in Oreodon the magnum is quite low in front and rises abruptly l)ehind, and
nearly the whole of the proximal surface is taken up by the facet for the scaphoid.
Another difference consists in the long hook-like process which in Protoreodon is
given off from the posterior surface of the bone, and which in Oreodon is represented
by a mere rudiment. When the l)one is in its natural position the distal surface for
mc. Ill presents obliquely downwards and outwards, even more obliquely than in On-
odoii. As usual in unreduced artiodactyls there is a small facet on the radial side of
the magnum for nic. ii.

The unciform likewise presents some imijortant dillerences from that of Orrodan.
In this genus the lunar and cuneiform facets are of very nearly equal size, while in
Protoreodon the latter considerably exceeds the former, as naturally Ibllows from the
already-mentioned fact that the lunar rests less completely on tiic unciform than in
Oreodon. The surface for the attachment of the cuneiform is less distinctly convex
and extends more closely to the external side of the bone than in the Miocene forms.
On the distal surface the facet for mc. iv is smaller and that for mc. v larger than in
Oreodon, and the latter is more distal in ])ositinn aTul less crowded to the external
side V)y the increased development of nic. iv than in Oreodon. This dillerence l)rings
about a further one in the shape of the inferior contoui-. which is more i-egularly
curved from side to side ami imt so distinctly angulated in /'roOn-f'0(/<>ji. Tiie lat-
eral facet for nic. iii i- ui" aliont the same proportions.

408 THE MAMMALIA OF THE UINTA FORMATION.

The metacarjmls are but scantily represented in the collection, and only one of
these exhibits the proximal end. Fortunately, however, this is the first metacarpal
or pollex, the existence of which would naturally be infeired from its presence in
Oreodon. There can no longer be any question as to the pentadaetyl character of the
fore foot in Oreodojt, as two specimens of O. Culbertsoni, Colorado and Dakota, with
the pollex in position, are preserved in the Princeton Museum, and in the Museum of
Comparative Zoology there is a beautifully preserved manus of 0. gracilis in
which all five digits lie in their natui-al position in the matrix. In Protoreodon the
first metacarpal is proportionately better developed, being both stouter and longer,
than in Oreodon. Its proximal end has a nearly flat head for articulation with the
trapezium. In none of the specimens of Oreodon mentioned have any phalanges
been found in connection with mc. i, though the distal end is rounded and fiiintly
keeled on the palmar side, indicating the probable existence of phalanges. Such
were certainly present in Protoreodon, as is shown by a small proximal phalanx be-
longing to the pollex. Its proximal facet is very oblique, sloping strongly outwards,
and is but slightly concave. Its distal surface plainly shows the articulation for the
ungual. The pollex, though of small size, was thus present in all its parts, and the
importance of this fact as connecting the aitiodact} Is with the always pentadaetyl
Condylarthra is obvious. It seems almost certain that artiodactyls with unreduced
anterior feet existed through the Bridger and Wasatch periods, though it is some-
what surprising to find them persisting so late as the White River Miocene.

The ])elvis is not well preserved in any of the specimens. Several fragments
seem to show that in construction it is essentially like that of Oreodon, and therefore
rather suilline in character. The ilium has a long compressed peduncle which expands
rather abruptly into a large terminal plate for articulation with the sacrum. There
is no supra-acetabular fossa. The length of the ischium could not be ascertained.

The femur would seem to be longer and stouter proportionately than in Oreo-
don. The rotular trochlea is very prominent and quite narrow, and has the inner edge
higher than the outer. A difference from Oreodon is shown in the presence of a fossa
for the plantaris muscle instead of a rugose sui'face, though, as Kowalevsky has
shown, no great importance can be attached to this character.

The tibia is only imperfectly represented ; some points may, however, be made
out with regard to it. It is entirely unankylosed with any portion of the fibula, and
has a stout shaft of transversely oval section below, above it is more trihedi-al. The
distal end is very like that of Oreodon, with deeply incised grooves for the astragalus
and a very well-developed malleolai- process.

The ^.bula is entire but very slender. Its distal end is closely applied to the

THE MAMMALIA OF THE UINTA FORMATION. 409

astrugaliis, foi-mini^ a large external malleolus, and as in all aitiodaetyls rests ui)()n
the lil)ular laeet of the caleaneum.

The hind foot is very eloseij' siniilai- to that of Oreodon, even more so than is
the foi-e foot. The astragalus is somewhat narrower in proportion to its height and
the ditTerence between the external and internal condyles in size is somewhat more
marked. The internal condyle passes directly into the navicular facet, though inter-
rupted by a slight prominence. The distal end is very unequally divided between
the navicular and cuboidal facets, the former being much the larger; the difference is
perhaps even more marked than in Oreodon. These two genera agree thus with
Ano])lot}ierium and the Suina and differ from most ruminants in which the two facets
are of more nearly equal extent. The calcaneal facets and those for the malleolai'
processes of the tibia and fibula resemble the corresponding parts in Oreodon, ex-
cept that the distal calcaneal surface is largei*. The caleaneum is much more slender
than in Oreodon and is especially less expanded at the distal end. The fibular facet
lises to a greater height and distally is more abruptly marked, and the interval
between this fixcet and the distal end is considerably greater than in Oreodon. The
sustentaculum projects further internally, though decidedly small, as in all oreodonts.
The cuboidal facet is more arched and concave from before backwards, and narrower
from side to side, while the distal astragalar surface is Hatter and larger. In fact,
the caleaneum diffei's from that of Oreodon moi-e than does any other tarsal bone.

The cuboid is higher and narrower than in Oreodon and the asti-agalar and cal-
caneal facets more equal, though the latter is somewhat broader. In correspondence
with the greater length of this part of the caleaneum, the cuboid is much more deeply
incised by the calcaneal facet, and the anterior edge of the navicular surface rises
high above it. The distal surface of the cu1)oid displays the usual facets for the
fourth and fifth metatarsals ; the latter is lai'ger and the former is smaller than in
Oreodon and of somewhat different shape. Another difference from this genus con-
sists in the greater extension dI' the hook-like projection from th ■ rear of the cuboid.
In lact the differences are so striking as to raise the suspicion that the si)ecimen
described belonged to some other genus, as unfortunately it was not associated with
teeth. lint from its correspondence with the caleaneum it very probably should be
referred to I'rotoreodon, and its dilferei'ces from the cuboid of ruminants and pigs
are even more marked than from Oreodon.

The navicular is likewise higher and nariower than that of ( hmdon. l)ut other-
wise resembles it very closely ; its proximal surface is somewhat more deeply con-
cave and its anterior edge rather more sinuous. Cope's statements (No. 4, p. 510)
with regard to the cuneiforms of Oreodon, are in some respects inacctn-ate, and

500 THE MAMMALIA OP THE UINTA FORMATIOlSr.

require correction before we proceed to the comparison with Protoreodon. Cope's
description is as follows : " The ectocuneifoi-m is distinct, and much wider than lon^.
The mesocuneiform is extero-posterior in position, and the transverse diameters are
small. It is produced distally overlapping the head of the second metatarsus.
Entocuneiform wanting." In reality the ecto- and mesbcuneiforras are coossified,
the line of junction between them being marked by a slight step, or difference of
level on the distal surface, which indicates the two facets for the second and third
metatarsals i-espeetively, and what Prof. Cope has called the mesocuneiform is really
the entocuneiform. If Pi-of. Cope's statements were correct, Oreodon would present
the remarkable anomaly of having the ectocuneiform support two digits, while the
mesocuneiform supports none at all, or in other words, having the second metatarsal
shifted outwards from its ordinary attachment. Besides this, the ectoeiineiforra per-
sists, with remarkable constancy in tlie ungulate series, and its absence in such an
unreduced pes as that of Oreodoi/, would be very extraordinary. But as we have
seen, these anomalies do not exist. In Protoreodon almost exactly the same conditions
are found as in the Miocene genus, only here the difference in height between the
meso- and ectocuneiforms and consequently the distal step is more pronounced. The
entocuneiform is not preserved in any of the specimens, but the facets on the navicu-
lar and second metatarsal show that it was shaped very much as in Oreodon.

The metatarsals are four in number and entirely free ; no trace of the hallux has
been found, and in all probability none existed. The metatarsals are of more equal
development than in Oreodon, the lateral ones being somewhat largei* in propoi-tion to
the median, as would be inferred from the structure of the distal row of tarsals ; in
other respects they are closely alike. The second metatarsal has a somewhat latei'al
bearing on the ectocuneiform, and metatarsals iii and iv are closely interlocking.
The ridge on the distal end of the raetapodials is confined to the palmar surface.

T^he, 2^cda7iges also resemble those of Oreodon, those of the first row being long
and depressed, those of the second i-o\v shorter and with the distal trochlea even less
asymmeti'ical than that of Oreodon., showing a less degree of convergence of the
hoofs than in that genus, and hence very much less than in any of the recent artio-
dactyls except the Tylopoda. The ungual phalanges are, as would be inferi'ed from
this, but slightly asymmetrical ; they are higher, narrower, more pointed, and alto-
gether more claw-like than in the Miocene members of the group, with the exception
perhaps of Merycliyus.

'Wc only cleai'ly characterized species of Protoreodon is 1*. parvus S. & O., of
which the type specimens are the skull and lower jaws figured on PI. I, Figs. 1 and
2. This was a very small animal, inferior in size even to Oreodon gracilis. The

THE MAMMALIA OF THE UINTA FOKMATIOV. 501

measurements agree fairly well with those of the so-called AgriocJioirus pumUiis
Marsh, with which this species may prove to be identical.

Jleasuremetiis. }f.

LoDglh upper mohir scries (entire) •; .05:5

" " preiiioliir series ? .020

" " true molar •' 027

Length lower moliir series (entire) 054

" " premolar series 027

" " true molar " 027

Third upper molar, anlero-posterior diameter 010

" " " transverse " (lOS

" lower " anteroposterior " 012

" " " transverse " 000

Although the teeth preserved in the Princeton collection all belong to P.jiarvus,
there are several limb and foot bones which exceed those of the type species so
greatly in size, that they very probably belong to a larger species. They differ,
however, only in size and must unquestionably be referred to the same genus. Size
alone is not a very satisfactory criterion, but it seems unlikely that such differences
can be within the limits of mere sexual or individual variation. The Ibllowing
measurements will show the size of these larger specimens of I'rotoreodou as com-
pared with that of Oreodon Culbertsoni.

JUeaiurermnU.

Protoreodon. 0. Culbertsoni.
Af. M.

Breadth of humeral trochlea 016 .021

Height " " •• (Ill .(ii;t

Length of calcaneiim OoO .0)2

Sysietnatic I^o.^ition and lielalionsliips of Protoreodon.

Tt will be obvious from the foregoing description that Protorpodon is very closely
allied to the White Kiver genus Oreodon, and may safely be regarded as the ancestor of
that genus. This relationship is made clear b}' a comi)arison of the skull, the teeth and
the feet, where the differences which Protoreodon exhibits from its Miocene successor
are just these tendencies towards the simplification that we should naturally expect to
find in the ancestral type. More particularly the i)resencc of the liflh cusp in the
upi)er molai-s is a welcome indication of the connection between the oreodonfs and
the buno-selenodonts of the earlier Eocene. On the other liaiid. I'rotoreodon has
many points of resemblance to Aiiriodnvrus, which are somewhat as follows : (1) IMie
shape of the cranium and the remarkable clougalion of its posterior portion: (i!) the

A. v. S. — VOL. XV'I. '6\..

502 THE MAMMALIA OF THE UINTA FORMATIOX.

open orbit and probable absence of the lachrymal depression ; (3) the position and
shape of the posterior nares ; (4) the position of the infraorbital foramen; (5) the
character of the anterior premolars ; (G) the character of the lower molars and of the
internal part of the upper. These difterences of im[)ortance separate Agrioclicerus
from Protoreodon, namely, that in the former there are considerable diastemata in both
upper and lower jaws, that the last pi-emolar has assumed more or less completely the
pattern of the molars, and that the outer crescents of the upper molars are over-
hanging.

Can Protoreodon then be regarded as the common ancestor of both sections of
the Miocene family, the Oreodontmce and the Agriochoerinoe f As to the derivation of
the former subfomily from this genus there seems to be no I'easonable doubt; its rela-
tions to the second are more obscure. If we accept Schlosser's view that a closed
dental series is always secondary, and a sign " dass der betrefiende Stamm am End-
ziel seiner Entwicklung angelangt ist, wenigstens finden wir diesen Zustand, nur bei
solchen Formenreihen, welche gerade im Aussterben begriffen sind " (No. 13, p. 40),
it is clear that Agn'ocJmrus, with its diastemata, cannot be derived from Protoreodon.
But even if this principle be generally true, it is not without exceptions, as the long
persistence of the oreodonts themselves demonstrates, and the diastemata are, there-
fore, of themselves insufficient to decide the question. A more important difficulty
is the constitution of the outer crescents of the upper molars, Avhich in the known
species of Protoreodon have commenced to assume the flattened shape characteristic
of Oreodon. The Uinta genus therefore stands very near to the common ancestor of
Oreodon and Agriochoerus, but seems itself not to be that ancestor, at least so far as
we may judge from the teeth of P. parmis ; it is quite possible that another species
of the same genus may stand in the same relation to Agrioclicerus.

The forerunner in the Bridger fauna of Protoreodon would seem to be Helohyus
Marsh, known as yet only from the teeth. In this genus the upper molars have five
cusps, the unpaired one in the anterior half of the tooth, which are of a pyramidal
shape, and refer the genus to the generalized group of buno-selenodonts, which is
represented by the Eocene iiyopotamids of Europe. If more perfect specimens shall
confirm this supposed connection between Helohyus and Protoreodon, it will demon-
strate the connection of the Oreodunti'd'n with the Anthracotheridce, which has been
already often surmised. With regard to the connection of the former group with the
Tylopoda, we have seen in considering Leptotragulus that the relationship is proba-
bly a very remote one, and that the Tylopoda seem to pass backwards into the Dich-
ohunidtp, from which Schlosser derives the typical ruminants and the Tragulina.

In his later papers (No. 5, p. 384) I*rof. Cope has included Protoreodon among

THE MAMMALIA OF THE UnSTTA FOHINrATION. 503

the Xiphodoiitnla; on account of the presence of the fifth cusp in the anterior half of
the upper molars. This airangement is, however, quite untenable, as this unpaired
cusp is common to the great majority of the selenodonts and even the bunodonts of
the Eocene. On the other hand, the peculiar structure of the premolars in Xipho-
(lon, the caniniform premolar of Protoreodon, and the entirely divergent structure
of the feet in the two genera, show that their relationship to each other can only be
a I'emote one, and that any association of thoui in a single family must be arbitrary.
This is especially the case when we consider the obviously close relationship between
the Oreodont idee ami Protoreodon, which is recognized in Cope's scheme {I. c, p. 387).
K" tiie presence of the fifth cusp be looked upon as a family character, then it will be
necessary to form a new ftimily, the Protoreodontidce, for the reception of this genus.
But this seems to be unnecessary, and a more natural method would be to regard the
Uinta type as forming a subfamily of the Oreodontulce, characterized by the five-lobed
up2)er molars, and in other respects combining the features of the Miocene subfami-
lies, the Oreodonthue and the Agr/ochain'ni£. This last fact is of particular import-
ance, as it proves the connection between AfjriocJioerus and the true oreodonts, and
shows that Cope's reference {I. c, p. 388) of this genus to the Diclwdotdidiv, on account
of the complication of the last premolars, does not represent the natural arrangement.
"Whether we adopt Leidy's view, that Agriochcei-us is the type of a distinct family, or
with Gill regard it as representing a subfamily of the Oreodontulce, is a matter of
comparatively little importance ; the essential fact being the close genetic connection
of the two.

List of Papers Quoted.

E. D. Cope.— I. On the Vertebrata of the Wind River Eocene Beds of Wyoming. BuUutin U. S. Geological and Geo-
graphical Survi-y of the Territories, Vol. VI. No. 1, pp. 183-202.

2. The Meso/oic and Ctenozoic Realms of the Interior of North America. Amcriciiii NatunUi^l. Vol. XXI.

pp. 445-462.

3. The Verlehmla of the Tertiary Formations of the West. Report of the U. 9. Geological and Geographical

Survey ol' the Territories, Vol. HI, Pt. I, Washington, 1884.

4. Synopais of the Species of Oreodontida;. Proceedings of the American Philosophical Society, Vol. .XXI.

pp. 5()3-.'"j72.

5. The Classification and Phylogcny of the Artiodaclyla. Ibid., IS.ST, pp. 377-JOO.

Clarence King.—fi. U. 8. Geological Exploration of the Fortieth Parallel. Vol. I, Systematic Geology, Wash-
ington, 1878.

Joteph Leidy. — 7. The Extinct Mammalian Fauna nf Dakota and Nebraska. .Iourn;il of the ,\c.iilemy of Natural
Sciences, Philadelphia. 1800.

0. C. U'lrth. — H. Notice of New Tertiary Mammals. American Journal of Science and Arts, 3d Ser., Vol. IX, pp.
230-250.
!i. Introduction and Succession of Verlchrate Life in America. Ibid., Vol. XIV, p. 337.

504 THE MAMMALIA OF THE UINTA FORMATION".

E. F. Osborn. — 10. A Memoir upon Loxolophodon and Uintatherium. Contributions from the E. M. Museum of Geol-
ogy of the College of New Jersey, Princeton, 1881.

L. Rutimeyer. — 11. Beitriige zu einer natiirliehen Geschichte der Ilirsche. Abliandlungen der Schweizerschen Palse-
ontologischen Gesellschaft, Vols. VII, VIII and X (separatim).

Orestes St. JoJm. — 12. Report on the Geology of the Wind River District. Twelfth Annual Report of the U. S. Geolog-
ical and Geographical Survey of tlie Territories for the year 1878, pp. 175-269.

M. Seldosser. — 13. Beitriige zur Kenntnis der Stammesgeschichte der Hufthiere, etc. Morphologisches Jahrbuch, Bd.
XII, pp. 1-336.
14. Die Nager des Europiii.schcu Tertiiirs, etc. Palieontographica, Bd. XXI, pp. 21-164.

W. B. Scott. — 15. The Eocene Lacustrine Formations of the West. Proceedings of the American Association for the
Advancement of Science, 1887.
16. On some New and Little Known Creodonts. Journal of the Academy of Natural Sciences of Philadelphia,
Vol. IX, pp. 155-185.

W. B. Scott and H. F. Osborn. — 17. Preliminary Report on the Vertebrate Fossils of the Uinta Formation. Proceedings
of the American Philosophical Society, 1887, pp. 355-364.

C. A. W7ate.—18. Report on the Geology of a Portion of Northwestern Colorado. Tenth Annual Report of the U. S.
Geological and Geographical Survey of the Territories, 1876, pp. 3-60.

THE MAMMALIA OK THE UINTA FORMATION. 505

Part III.

BY HENRY FAIRFIELD OSBORN.

THE PERISSODACTYLA.

It is necessary to open this section with some observations upon the synonymy
of the Eocene Perissoclactjda. With the assistance of Prof Marsh, the writer
recently examined the types in the Yale College collection, in comparison with those
at Princeton. The result shows that the entire nomenclature of these genera is in
utter confusion, arising from the attempts which have been made by others to work
from the brief descriptions given by Prof. Marsh and without the aid of figures.
This confusion extends through all the American and foreign literature which relates
to the American Eocene fauna, and invalidates a great deal of otherwise very useful
work.

The synonymy of Helaletes (Marsh) is DUophodon (Scott) and Desmatotherium
(Scott). Lophiodon nanus (Marsh) also belongs to this genus. //. latidetis (nobis)
must be separated from it. It was originally distinguished from H)jraelujus by the
presence of a third lobe on the last lower molar ; this lobe is very small and forms
the only distinction between the type and the mandibular dentition of Diloplwdon
minusculus. The maxillary dentition is precisely like that of Hyrachyiis, except in
point of size and in the presence of two internal lobes upon the thiid and fourth pre-
molars. The latter feature was given by us as the generic distinction of Desmato-
therium. The dental structure (excepting only the rudimental third lobe) and the
tarsal characters remove it entirely from the tapir series, and relate it to Tr/plopus,
from which, however, it is probably generically distinct. Colonoceras has precisely
the dentition of Hyrachyics agrarini^ Leidy. The rudimental horn cores upon the
nasals may serve as a generic distinction, although, in the very limited knowledge
we have of the nasals of Ilyrachyus, this character is not altogether satisfactory.
Limnoliyus is a nomen nudum, having been applied to a type already preoccupied by
PaliP^osyops. Prof. Marsh will apply a new generic name to his L. laticeps.

The primitive horses arc in much'confusion. Eohippus* (Marsh) is a synonym

* Am. .lour. Sri. mid Art.s. Nov., 1870, p. 401.

506 THE MAMJIALIA OF THE UINTA FOKMATION.

of Hyra colli erium (Owen). Oroniermm* (= Lophiotherium) was based uiDon a good
type lower jaw ; it has the hist lower premolar like the molars ; it is identical with
and pi-eeedes OroJirp]ms,f which was founded upon an uncharacteristic type {0. im-
milus, a numbei- of loose upper molars), and was not fully defined until later upon
the discovery of O. agilis. Both genera are, however, synonymous with Pliolophus
Owen, which is defined by the last premolar being like the molars. The species, 0.
agilis, is however founded upon a type in which the third and fourth upper premolars
have two internal lobes ; it thus falls within the definition of Upihipjms.X Epihip2ms
is thus in turn close to Ancliilop)lms, but may be distinguished from it by the more
dejjressed crests and less complete union of the para- and metaconules into transverse
crests. Then follows Mesolrippus with three premolars like the molars, and crests
like those of Amchiloplms.

There is a remarkable parallelism in the assumption of the molar pattern by the
premolai-s in all the perissodactyl series. "We can almost predicate of any Wasatch
genus, fourth premolar unlike the molars ; of any Bridger genus, fourth premolar
like the molai's ; of any Uinta genus, third and fourth premolars like the molars ; of
any "White River genus, second to fourth preraolai's like the molars.

PERISSOD ACT YL A.

AMYNODON.

Numerous errors have found their way into the descriptions of the skull and
dentition of this genus which require correction before its affinities can be discussed.
There are now three skulls known which may be referred to three species. Fii'st, the
type Amynodon advenus Mai-sh, fi-om the Uinta beds ; second, the type of A. ( Orilio-
cynodon) antlquus, nobis, from the Washakie beds ; third, the type of A. intermedius,
spec, nov., from the Uinta.

Prof Marsh's description of the type is as follows :§ "The skull is intermediate
in I'orni between that of a tapir and a rhinoceros, but the molar teeth are entirely of
the latter type. The premolars are all unlike the molai's, and the canines above and
below are very lai-ge. The incisors are small and the inner one in each jaw is lost in
the present adult animal. The lower canines are placed nearly horizontal, and, taken
in connection with the I'cst of the anterior dentition, they prove conclusively that the
large lower teeth usually regarded as incisors in Aceratherium * * * are really
canines." The number of digits is stated to be 4-3. This description is erroneous

*Am. Jour. Sci. and Ails, Sciil., 1872, p. 217.
t Op. ciL, proposi'cl Sept., 1872 ; ilclined May, 187:!.
Ilnlvort. and Succession of Vcrl., etc., Ann;., 1877, ii. 2X.
§ Am. .lour. Sci and Arts, Third Scr., Vol. XIV, p. 251.

THE MAMMALIA OF THE UINTA FORMATION. 507

in most of the particulars, partly owing to the fact that it was diawii up before tlie
type specimen had been removed from the matrix and put too^ether, and in this way,
even after examining the type ourselves, as Prof. Mai'sh kindly allowed us to do, we
were formerly led to consider the Bridger species as a distinct genus {Orthocynothm).

A second examination of the type specimen recently made brings out several
important diagnostic characters in addition to those already noted in oin- i)reliminary
bulletin, p. 262. There are three lower and probably three upper incisors ; the first
lower premolar is wanting. The canines are erect. There are rugose postorbital,
antorbital and infraorbital processes. The nasals are very short and slightly over-
hang the anterior narcs. The type of A. antiquus is still found to resemble that of
A. adi'enus closely, with the important exception that there are four lower premolai-s
instead of three, the first lower premolar being fully functional and bifanged ; the
first upper premolar is missing. It may subsequently be found to represent a distinct
genus as we at first supposed, for which the original name Ortliocynodon would stand.
The -.i. intermedius, besides its much larger size, may be clearly distinguished from
the two foregoing by the jirocumbent position of the canines and by the retention of
a single fanged first upper jjremolar.

These and other characteristics throw a new light upon the phylogenetic jjosition
of the Amy nodonl idee, which will be discussed later.

AMYNODONTID.E S. & 0.*

Rhinoceros-like animals extending from the Middle Eocene to Lower Miocene ;
skull deeply excavated in front of orbit ; incisor border very broad; nasals short and
hornless ; canines and incisors present and functional in both jaws ; pattern of the
molars like that of the rhinoceros except for the non-reduction of the external crest
of the third upper molar; probably four t(jcs in fi-ont and ihree behind; proliably
astragalus rests upon the cuboid as in the ihinoceros.

This defmition is considerably modified from that previously given by the writers,
in order to embrace the Miocene genus.

The genera embraced in this family are:

Lower Miocene ( White Hivei-).

Metamvnodon,! nobis. Dentition, i. 1., c. I, pm. 3, m. '. ' Ui)pcr and lower
canines obrKjuely placed, the latter fitting sonu'wlial inti'rnal to the former when the

• E. M. Museum Bullniii, No. :i. 1H8:(. p. 4.

\ lliillcliii ni' thf iMiiNfiiiii 111' C<>iii|>nmlivi- Znology, Sf|il., \>->sT, l>. 105.

508 THE MAMMALIA OF THE UINTA FORMATION.

jaw is closed. The second, third and fourth upper premolars of the molar pattern.
Postglenoid and posttympanie processes widely united. Periotic not exposed.

Upper Eocene ( Uinta).

Amtnodon* Marsh. Dentition, i. % c. I, pm. |:f, m. f. Upper and lower
canines erect or obliquely placed. The third and fourth premolars of the molar pat-
tern. Postglenoid and posttympanie processes separated. Digits : 4-3 (Marsh).

A. ADVENUS Marsh. Premolars f. JSTamber of upper incisors uncertain. Up-
per canines obliquely placed, lower canines erect.

A. iNTERMEDius, spcc. nov. Premolars *; first upper premolar rudimentary and
single fanged. Canines in both jaws very large, semi-procumbent, oval in section.
Three functional upper incisors.

Middle Eocene ( Washakie).

A. (Orthocynodon) antiquits, nobis. Premolars I ; first upper premolar bi-
fanged. Canines in both jaws vertical, triangular in section. Number of upper and
lower incisors uncertain. Mastoid portion of periotic exposed.

AMYNODOlSr INTERMEDIUS.

Plate X.

This species is represented by the base of the anterior portion of a skull (No.
10,309) in beautiful preservation, showing the complete characters of the teeth, the
palate, the position of the orbit and the lowei- portion of the premaxillaries. The four
canines and an upper molar of another individual are preserved. Part of the lower
jaw and a premolar of a third, and the mandibular symphysis of a fourth. There are
also numerous skeletal fragments, which however cannot be referred to Amynodon
with any certainty.

Dentition (Figs. 10, 10a). The three incisors arc equidistant ; the median one
is slightly larger than the lateral and is placed about as far from the premaxillary
suture as the lateral incisor is from the canine. The crowns are pointed, convex on
the anterior and posterior faces, and slightly compressed laterally. The canines are
very powerful, with lance-shaped crowns and a decidedly flattened oval, quite unlike
the trihedral cro>yns in A. antiquns ; they project forward at angle of forty-five de-
grees, and slightly outwards ; the fangs are powerful and deeply rooted beneath the
first ])remoUir. There is a considerable diastema.

» Aim. Jimr. S<i. iiiul Art-, Scpi., 1877, p. 2")1.

THE MAMMALIA OF TIIK UIXTA FORMATION. 509

The first preniohir is represented merely by the fang, which is riidiinenlary
on one side. The second preniohir has a slight swelling on the outer face which
anticipates the posteroexternal lobe; there is a single prominent internal cusp united
by two low subcqual ridges witli the external crest; surrounding the inner lobe is
the prominent cingulum which also characterizes all the succeeding molars. The
third and fourth premolars are interesting as showing a mode of development of the
2)OSterioi- ci'ests quite unlike that in the tapir or Equus series ; there is as yet no redupli-
cation of the internal lobe as in the latter series, but the posterior crest grows directly
out from the bilobate external crest in the valley formed by the prominent cingulum
and the much broader and more elevated anterior crest ; these two teeth are practi-
cally similar and submolariforra, except that the third premolar has a less distinct
posterior lobe and crest.

The first true molar is very similar to that in Aceralheriuin ; the anterior crest
has a slightly marked "anticrochet'" which is entirely wanting in ni.- and m.'; the
external crest is perfectly flat and decidedly inclined inwards. The second molar has
the same characters with a very decided extension of the external crest beyond the
posterior transverse crest. In the third molar the external crest arches out widely
beyond the posterior crest; the base of the crown is subquadrate in outline.

ilensurements.

A. antiquus. A. inUrmedius. Metamynodon planifrons.

M. M. M.

Antero-postcrior (liami'ter of nicdiiin lower incisor OH (ii|)per do. ) .012

Fore and aft (linmcter of crown of lower canine al base. .015 (upptrdo.) .031 .035
Upper jaw.

Fore and aft diameter of third premolnr 01!) .021

Length ot entire molar series .187 .225

" true molar 101 100

Fore and aft diameter first molar 037 .044 .017

Transverse " " " 0:n .043 .0(W

Fore and aft diameter, second molar 04."i .058

Transverse " " " 037 .052

Fore and aft diomcler. third molar 028 .046 .OfiO

Transverse " " " 030 .040 .Olil

Measurement of canines, outside .115 .130

The sluU. The most striking features of the skull are at once seen to l)e the
great breadth of the ni .i x i I lari es opposite the canines and their deep excavation
between the canines and the antorbital ridge; the snout is thus broader than the

A. r. s. — VOL. xvr. 3.m.

510

THE MAMMALIA OF THE UINTA FORMATION.

DiAG. 2. — Amynodon iniermeiHiis. Side view of the anterior por
tion of tlie skull. X \.

anterior premolar region. The anterior nai'es were apparently widely open. The

p r e m a X i 1 1 a r i e s are rather nar-
row above and spread inferiorly,
rounding and arching forwards into
the broad incisive border ; the rela-
tions of these bones in Metamynoclon
indicate that they had a deep area of
union in front, contrasting strongly
with the separate slender premaxillfe
of Aceralherium. The m a 1 a r pro-
jects widely below the orbit which is immediately above the second molar and the
maxillaries are deeply depressed and incurved in front of the antorbital ridge.

The upper and posterior regions of the skull are entirely wanting in this speci-
men, and have been fully described in the E. M. Museum Bulletin upon the skull of
A. (OrtJiocynodou) antiqims.

The lielations of the Amynodontidce.

The Washakie sjiecics, both in structure and time, is the most primitive form, as
indicated by the complete dentition and the perfectly erect position of the canines.
Then follows the A. advenns from the Uinta with small erect canines and the reduc-
tion of one premolar, otherwise very similar in size and dentition to the foregoing.
The A. intermed.ius, while retaining the rudiments of the first premolar, which were
probably lost in fully adult specimens, shows a vei-y decided transition towai'ds Me-
tamynodon in the enlargement, shape and procumbent position of the canines, also in
the broadening of the snout and compression of the antorbital region. It is sing-
ular to note how little i)rogress the premolars of Melamynodon show ; the second pre-
molar is like the third, but in none are the posterior crests very prominent.

Pliyloyenetic Position of Aynynodon.

The discovery of Amynodon.^ with its full complement of teeth and markedly
rhinocerotic molar pattern, naturally led Pi'of Marsh to suppose he had found the
long-sought Eocene representative of the true rhinoceroses.* This opinion has been
adopted by every one. But the discovery of Metamynodon, which is certainly far from
the I'hinoceros line and is an undoubted successor to these Eocene forms, directs atten-
tion to a number of developmental features which suggest an entirely distinct supposi-

*IntroiUiclion and Siiecossiun of Vcrlelirale Life in Anuiica, n. ;iO.

TIIK M.V.MMALIA OF THK UINTA FORMATION. Gil

tion, viz., that this is anothiM' line of pseudo-rhinoceroses parallel to the JIi/iaco(h)ii-
tiike. For while the molar dentition and [icrliaps also tlir lower canines point
directly to Acevatherium, the development of the other teeth and man\- cliaracters of
the skull are very diverse. Prof. Cope* i)resents a third alternative in deriving
I[ijrac(Ml(>ii from Amynodon, placing these genera in one family, and not placing
the latter in the Aceratlierinni {^Cwiiopu-i) series at all.

The features which unite the known AinynodoididtH with the aceratherine line
are: (I) The structure of the lower molars and of the superior molars to the
second molar inclusive ; (2) the structure of the astragalus, in extending upon the
cuboid. The dietiugnishing features of the Amynodontidx are: (1) The retention
of fully functional incisors in both jaws, and the great enlargement of the canines;
(2) the retention of the lophiodont character of the last ui)per molar in its sulxpiad-
i"ate form and complete cvternal crest ; (3) the retarded development of the postei'ior
crests of the superior premolars ; (4) the deep excavation of the maxillaries in front
of the orbit and flattenin"r of the cranium, tofjether with the shorteninof of the nasals.

In Aceratherium, on the other hand, from the lowest American Miocene, we fuid
the upper canines entirely wanting, also one of the incisors; the premaxillaries are
narrow and weak, and support two subequal incisors. In the lowTr jaw^ are the large
lateral teeth and small median pair, of very doubtful homology. Upon the discovery
of ^Imynodon, Prof Marsh suggested that the outermost lower tooth of ^li'cra-
therium was a canine, instead of an incisor as previously supposed, but there is little
additional basis for this opinion. A. occidentale has, moreover, long nasals and con-
vex maxillaries. In fact, excepting in the molar series we lind no support for the
supposition that Aceratherinin is a descendant of any known species of Aiinjnodon.
The resemblance of Hyracodon to llyrarhyinA, of Mcxohiyims to Jfymrolherhim, of
Titiinotherlian to I'ali'()!<ynps is altogether of a different character, and in each ca.se
is relatively far more signilicant of direct descent.

So far as Prof Cope's suggestion of the descent of Ifyracodmi from Aunjiioduii,
is concerned, the following ol))ections may be made: (1) The dentition and o.stcology
of llyrnchyii.-i fill in every respect the conditions which we should expect to find in
an ancestral foi-m of /////v/ro/A^//, boili in the displacement and reduction of the foot
bones and characters of the teeth ; (2) a wide dillerence between the aceratherinc
and hyracodon series is found in the structure of the carpus and tarsus. The posi-
tion of the Atnyiiodiiididin it appears turns ui)on these parts. An astragalus found
near the skull of .1. rndif/itn.i (see Diag. 12) resembles closely that of Acerathnnun
in its broad cuboidal facet. If, in addition, as we think highly prol)ablc, Ainynodon

•Tlif IVrissodnctyli. Am. Niil., 1S87, \<. 9!)U.

512 THE MAMMALIA OF THE UINTA FOKMATION.

is found to possess four toes in the manus, tlie series can be readily placed interme-
diate between ILjracodon and the rhinoceroses ; (3) these j^enera have in common
a complete incisor series, but in contrast with Amyaodon, IJyracodon shows a decided
degeneration of the canines ; (4) the molars of Ilijracodon, while imitating those of
the rhinoceros, entirely lack the pecnliar proportions and inclinations of the extei-nal
and transverse crests, which are so distinctly rhinocerotic in Amyaodon.

The difficulties which have been indicated in any attempt to derive the rhinoceros
from any known species of Amyaodon do not preclude this derivation from some
member of the Amyaodoatidcii ; the point insisted upon here is, that such a member
has not yet been discovered ; secondly, that the later Amynodontidfe represent a dis-
tinct line of pseudo-rhinoceroses, probably intermediate between the true line and the
Hyracodon series. They approach the true rhinoceros in part of their tooth structure
and probal^ly in the foot structure, and the latter series in the remainder of the den-
tal structure.

DIPLACODON Marsh.

As observed by Marsh at the time this genus was established,* Diplacodon is
intermediate between the Bridger genus Pcdoeosyops and the Lower Miocene Tltano-
tJieriidie. Nothing is known of the skull, the type consisting merely of the palate
and complete maxillary dental series. Unlike the Bridger genera the fourth premo-
lar is like the molars and the second and third premolars are developing the second
internal cusp, and in process of assuming the molar pattern. We may anticipate
that the skull will show the initial development of the great horns of Titaaotherlum.
The numerous remains of the skeleton in the Princeton collection enable us to fully
confirm the relationship suggested by Marsh. Tlie skeleton is in every detail inter-
mediate between that of Pala'osyops and Titaaothermm, and presents a remarkable
intermingling of characters persisting from its smaller Bridger progenitors and antici-
pating its great Miocene successor. Allowing for a moderate elongation of the dor-
sal spines, IHplacodmi stood about five feet seven inches at the shoulder, while Pakii-
osyops stood a little less than four feet, and Titaaotherlaia considerably ovci" seven
feet. The metapodials i-etained the stilted spreading character seen in Palceo^yops ;
much longer than in this genus and less bulky than in Titaaoiherinni. On the other
hand the cervical vertcbrte in some species arc much flattened, indicating a short neck
similar to that in the Miocene genus. The detailed comparative measurements given
below are taken from a Pala'osyops species of the middle size, probably P. pala-
dosus ; the measurements of Diplacodon are mostly from a skeleton belonging to a

»Aiu. .Iiuir. Sci. aiul Ails, March, 1875, \\ S-IG.

TIIK MAM>rAT.TA OF THE UINTA FOiniATIdX. 513

single individual ; the Ttlanothen'ain measureraents aio aiiproximatc from specimuns
which we have referred to T. yrontll.

It is remai-kable that the comparatively small genus Palwosf/ops anticipates in
so many features the gigantic Miocene genus. The foot structure in almost every
detail is persistent, this being the only line of Perissodactyla known in which there
is no reduction of the fifth digit. Then we find in Falwosi/ops, as shown in tiie dia-
grams, every tarsal articular facet reproduced in the later genus. The neck, in some
species at least,* undergoes considerable change in i)roportion, but the doisal >iiiues
of the Eocene genus are decidedly elongate and anticipate the great liuinp of Tltaa-
otherium. The skull undergoes an entire remodeling, consisting principally in the
elevation of the occiput, this being correlated with the elongation of the dorsal ver-
tebral spines and development of a powerful ligametdum nuchce to support the nasal
horns. The earlici- species of Titanotherium, T. (Megaceratops) coloradense, for exam-
ple, retain the long nasals overhanging the premaxillaries which are so characteristic
o? I^aheosijops. A feature common to the three genera is the prominence of the lesser
trochanter.

In adtlition to the features ali'cady mentioned as distinguishing the Miocene
and Uinta genera from their Eocene ally, we notice the prominent and reeuived del-
toid hook of the humerus; the cversion of the major axes of the iiniominate bones,
with the marked expansion of the suprailiac border. In the manus and pes, while
the facets remain the same, the proportions of the different elements are much altered,
as pointed out in a later section. In the pelvis and tarsus, however, as already stated,
J)/])hicodon stands nearer the oldL'r than the more recent form.

In the Washakie beds is found a large sjiecies, about the same size as /^ valli-
(huis Cope, which is i)i-ovisionally referred to PaJceosijojis {P. Jnjixjnatlnis, spec, nov.,
Princeton collection, No. 10,27.'3). This is represented by a lower jaw seven-eighths
as large as the type mandil)le of i>/"/>/«cofZo/i. As in the latter, llu' incisors forma
close procumbent series; the tips forming a gently aiched line when seen from above.
The sym[)hysis is extremely long (11 cm.) and shallow ; the canines are rather small
and semi-procumbent. The molar-premolar series measures 24.") cm., the last molar
measures 6.5 cm., the transverse measurement outside of the canines is 9.(3 em. ; in
Diplacodon ehitns the same measurement is K) em. Unfortunately the premolar
crowns are broken; it is probable that one or two of the premolars will be fotmd to
be like the molars. The characters of the ehiu and symphysis are significant ofdose
relalionshii) to Jjiplacodon elalas.

• In Prnl. Miirsli's collcrlion IlifTi' on- some cen'kal vcrlcUrii; rcrerruU lo Dii'lucoilon wlii' i' l''v. .il i ili. .^nn.'

)>ri>|ionicin!il Irnglti iis in Pitlito»yi>ii.

514 THE MAJNIMALIA OF THE UINTA FORMATION.

It is not possible to determine tlie species to wliieh our skeletal remains of Dip-
lacodon belong, as we have but a portion of a single upper molar. They may be
referred to D. elaius. Palceosyops has hitherto been referred to the Chalicotherudce,
but the discovery of the foot bones of Chalicotherium by Filhol shows that the genera
are Avidely separated. The discovery of the skeleton of Diplacodon, however, links
Palmosyops very closely to Titanotherium, and the differences between these three
genera are principally in three characters, viz., the assumption of the molar pattern
by the premolars, the development of frontal horns, and the loss of the incisor teeth.
If these charactei's are given a family I'ank we cannot decide where to place Dipltico-
don. It seems best to group the three genera in the single family TitanotJieriidce.*

DiPLACODON ELATUS.

Generic characters. Dentition, fourth and third upper premolars like the
molars. Last upper molar with single internal cone. Digits 4-3.

8p(cijiG characters. Second upper premolar with a rudimentary postero-internal
cusp, like the molars.

As above stated, the specific reference of these specimens is uncertain. The cer-
vical vertebras are much shorter than those associated with the Yale College speci-
mens. It is probable that they represent a distinct species.

The Skeleton.

Plate VIU.

Cervicals. The axis (Xo. 10,396a), Fig. 15, has a broad spine overhanging
the postzygapophysis. The laminae are very slightly notched. The postzygapophy-
ses are an elongate oval. The transverse processes are hooked, turning inwards at
the tip and perforated at the base. The centrum presents a sharp inferior keel. The
remaining cervical'^ and dorsals belong to a single individual (N^o. 10,39(5). The
cervicals preserved are probably the 3d, 4th, 5th and Gth. The 5th is the most com-
plete (Fig. 1) ; the spine is pointed, vertically placed and grooved posterioi'ly ; the
zygapophyses are very stout with the facets placed at angles of -15° ; the vertical
diameter of the centra is much gi-eater than the transverse; they are quite strongly
opisthocoilous 3 the transverse processes do not extend below the level of the
centrum.

The centra of seven dorsals arc preserved. The one figured is between the

* Tliis generic luid lamily iiiiiuu has lici'ii ailoplcil liy llu; writcT because Menodiis Pomul is louiul to be pruoccupied
by Jlenodon v. Meyer, and Tiliinothcriiini Leidy niiisl supersede i(.

THE MAMMALIA OF THE UIXTA FOIJMATIOX. 515

7th and lOtli (Fig. 2), The centrum is opisthocd'lous and considerably excavated at
the sides ; the antero-posterior and transverse diameters are al)out the same ; the
lower half of tlie spine is preserved, it is a stout triangle in section indicating a great
length and strongly oblique inclination; the zygapophyses are almost horizontal. One
of the 1 u m tj a r centra preserved is considerably longer than the abovc-descrilied
dorsal, and has a stout keel ; the spine is broad (Fig. 4) and grooved posteriorly ;
the zygapophyses are rounded and vertically placed.

These vertel^ral characters closely repeat those observed in Palceosyops, except
that the cervical centra, while nearly double in height and breadth, are only a tritle
longer. In Titanotherinm the cervicals are not further shortened, but retain {\\v pro-
portions seen in JJiplacodon ; they are, if anything, somewhat longer. The dorsal
spines are still more elongate, forming a great hump which was incipient in Pithvo-
syop.^ and probably well developed in Diplacodon.

Two ribs are pi'cserved which belong respectively in the anterior and middle
region of the chest. They are much lighter and more rounded than in Titanothcriinn.
The complete rib from the midregion has a subquadrate section in the up])er thiixl of
the shaft, and an oval section in the lower third ; it does not show the extreme flat-
tened oval seen in Titanotherium ; the length, not allowing for curvature, is 71 era.,
showing that the deplh ol' the chest was about thii'ty inches.

Platk IX.

Scapula (Fig. 12). Both scapuliB are preserved, but the superior and lateral
borders are incomplete. The coracoid process is a stout tuberosity. The glenoid
fos.sa is an elongate and rather shallow oval. The spine ascends very gradually from
the neck and passes without an acromial process into a deep recurved ridge ; in a
midsection the spine is much expanded along the border and ovcihaugs the post-
8cai)ular fossa. The })ai-ts preserved indicate that the scajmhi was lofty, with a
rounded supraseajnihir lioidci- unlike the sonu wlial angular lioidi r of l^itaiiot/icrium
and without any di.stinct indentations such as are seen in lihinoctros.

Ifumerus (Figs. 11 and 11a). The humerus lacks the head, lesser tuberosity
and bicipitaj groove. There is a stout deltoid ridge terminating in a prominent re-
curved hook. The shaft is iwisted upon itself as in Ilhinoceros, that is the major
diameter passes obliipiely from the outer posterior lo the inner anterior siile. The
supinator ridge is less distinctly marked than in lihiuoceros and less prominent than
in Titanot/ierium. The pronator ridge is also rugo.se but not ])rojecting. The supra-
trochlear fo.ssa is deeply excavated and the tiochlear surfaces are somewhat oblique
to the main axis of the .shaft.

516 THE MAMMALIA OF THE UIN^TA FORMATION.

Ulna and radius (Fig^. 18). These bones are complete, and taken together indi-
cate a rather long and slender forearm. Tlie proximal portion of the radins covers
the entire nlna, and the shaft crosses to the ental side. The relative diameters of the
distal fiicets are about as five to two. The ulna has a very stout rugose olecranon ;
the posterior border pi'esents a single concave curvature to the distal extremity ; the
midsection of the shaft is triangular with a deep groove upon the anterior face. The
radius in midsection of the shaft is suboval anteriorly and flattened posteriorly;
the facet for the ental condyle of the humerus has the same transverse, but much
deeper antero-posterior diameter than that for the ectal.

The Manus. The carpus is entirely wanting. The metacarpus and a few of the
phalanges are preserved. They show a high stilted tetradactyl metapodium of the
"digitigrade" type, that is, with the jihalanges resting entirely upon the ground.
The distinctive feature of the foot is seen iu the subequal size of the second and fifth
digits, which brings the working median axis between the third and fourth digits
instead of through the middle of the third. The carpus was thus undoubtedly of the
tyi^e intermediate between Pala^osyojjs and Titaaothermm as restored in outline. The
second metacarpal has proximally (Fig. 13?;) a pear-shaped facet for the trapezoid,
a magnum facet extending to its full depth ; it also overlaps mc. iii. The third meta-
carpal has a subquadrate magnum facet and triangular unciform facet, overlapping
mc. IV by separate anterior and posterior facets. The Iburth metacarpal has a corre-
sponding mc. Ill facet ; the uncifoi-m facet is fractured j)osteriorly. The fractured
proximal portion of mc. v shows a narrow concave unciform facet and a lateral facet
for mc. IV. The keels are entirely confined to the posterior surface, as shown in a
distal view of mc. ii (Fig. Via). The measurements are as follows : Length, mc.
in — 18 cm., mc. v — 13 cm. Depth o f p r o x i m a 1 f ac e t s , mc. iii — 3.5 cm., mc.
Ill — 3.3 em. Breadth of distal facets, mc. in — 4 cm , mc. v — 3.4 era., mc.
II — 3.4 cm.

T\\c2)elv!s (No. 10,393). The marked characteristics of the pelvis are the great
length of the ossa-innominata as compared with their breadth. The accompanying
restoration of the pelvis, as viewed from above, shows the parts which are preserved
in Di])laco<lon in comjiarison with the pelves of Titanotherium and Pakeo^i/ops. The
ilia do not expand immediately above the acetabula as in the Miocene genus, but
there is a long and rather slender neck as in Falaiosyops, beyond which they expand.
The acetabular border presents a short sharp curvature, and is relatively much shorter
than in Pahmsyops ; the ischial border {!h.) is much longer, with a gradual curva-
ture. It follows that the suprailiac border apiiarently presented outwards more than
directly upwards and forwards as in /'aht'osyops and Titanotherium— a result of the

TUK MAMMALIA OF THE UINTA FOUMATION.

517

unequal growth of the acetabular and ischial borders. The acetabular rim is much
fractured ; from its upper border a ridge (the pubic border) exteuds beneath tlio
ilium, parallel with the ischial border to the sacral surface. Below the acetabulum
the ischium has a triangular section ; it then expands in a plane directly perpen-
dicular to that of the ilium. There are no remains of the pubes.

Di.u.. 3.

DiAO. 4.

DiAO. 5.

Posterior view of llic pelvis of (3) Palaoynjia X I'j, of (4) Dtpla(od<m x t'j. of (5) Titanothcrium X i',.
11, ilium ; Is, isdiiiira ; P, i)ul)is ; ti, ib, ab, borders of the ilium ; a, acetaliulum ; of, oblnmtor foramen.

The fem7tr. This bone is represented by the somewhat crushed shaft (No.
10,39.')), and a well-pi-eserved distal half apparently belonging to individuals of tin-
same size, both belonging to the loft side ; they are combined in Fig. 5. The head
and great trochanter of the former specimen are wanting. The lesser trochanter is
represented by the base of a stout ridge, which indicates that it was much more
prominent than in Rhinoceros. The third trochanter, judging by the base, was still
more prominent and strongly curved forwards. It stood higher upon the shaft than
in lihinoceros. The lower portion of the shaft with the facets are ver}' similar to
those of the Rhinoceros, cxcei)t that the patellar trochlea is longer and more de('i)ly
excavated; the condylar facets have a greater fore and aft extension, and the inter-
condylar notch is deeper.

Another specimen in which the head and great trochanter are preserved shows
that the head was well out of the lint- of the shaft, with a dee]) pit for the round liga-
ment. The great trochanter stands upon the level of ilu' head.

The tihla (No. 10,390) isabout five-sevenths the lengtli -.f tin- femur (Fig. 6). The
cnemial crest is moderately prominent. There is the usual triangular section of the
shaft just below the crest passing into an oval section In llir lower thiid. Tlu' prox-

A. r. s. — vni,. xvi. 3v.

518 THE MAMMALIA OF THE UINTA FORMATION.

imal and distal faces are too much worn to admit of description. Another specimen
(Fig. 6a) shows a prominent internal malleolus and strong spine.

The tarsus (Figs. 8-8Z)). The well-preserved astragalus and calcaneum are of
great interest. Their relations to those of Palmosyops and Titanotheriam are fully
discussed in the next section. The pi*incipal features of the calcaneum are the
extremely narrow, deep and elongate tuber calcis, which has an unusually flattened sec-
tion ; there is also a distinct fibular facet, and the calcaneum also forms part of the
tibial trochlea. The three astragalar facets, the ectal, sustentacular and inferior, are
entirely distinct. The astragalus rests upon over one-third of the upper surface
of the cuboid. These elements of the ankle joint indicate that Dlplacodon was of
much less bulky proportions than Titanotherium.

Comparative Measurements of the Skeleton, in, Cm.

Palaosyops. Diplaeodon. Titanotheriam.

(?) Fifth cervical vi'riclini, antiTO-posterioi- 3.2 4 6.5

" " " transverse 4.3 6.7 9

(?) Ninth dorsal vertebra, antero-posterior 3.7 6.5 6.5

transverse 4.7 5.7 10

Lumbar vertebra, antero jiosterior 4.7a 6 7

transverse 89a 6.3 10

Total length of scapula, estimated 27 60e 72a

Length of humerus 30.5 45e 67.2a

" " radius 25 35 65a

" third metacarpal 11 18 24a

Width of carpus 6.7 10

Depth" " 4 8e 11.2a

Total length of innominate bone 41 61 78a

Length of femur 33 5 46e

" tibia 27.5 33

Width of tarsus 0.5 8.5 14.5a

a = appro.ximate measurement. e = estimated.

The PalceosyojhH measurements are taken from the middle size species (P. pahc-
dosus), which predominates in the "Western Bridger basin, and while taken from dif-
ferent individuals are probably very nearly correct in proportion. Tlie Diplaeodon
measurements, with the exception of the femur, tibia and tarsus, are also from por-
tions collected together and belonging to one individual. The Titaaotherlum verte-
bral measurements are from a single individual, but the other parts are taken from a
number of individuals. The P. major of the Bridger is fully one-fourth larger than
the above si)ecies.

ISECTOLOPUUS S. & 0.

Isedolojjhns is a small tapiroid, slightly lai'ger but with shorter limbs and feet
and more spreading digits than the Uinta 2\/)>lopi(i^.

THE MAMMALIA OF THE UINTA FOKMATIOX. 519

In tlie line of genefa to which it belongs the cliaractors of the premolars in
respect to the gradual assumption of the niolai' pattern are vei y important. These
are wanting in the Uinta specimens, but the Bridger species, which was formerly
referred by us* to Helaletes (IT. latidens) is closely related to Isectolophus, if not gen-
erically the same, and shows double internal lobes upon both pm. ' and pm. \ The
second upper premolar probably had two external lobes and one internal lobe. The
distinctive features of the dentition, so far as known, are seen in the conic postero-
external lobe of the upper molars, in the third lobe of the last lower molar and the
absence of a diastema in the lower jaw. As pointed out in our preliminary descrip-
tion, these features place Isectolophus in or near the true tapir line. This conclusion
is sustained by the discovery of the complete carpus and of the cuboid. The carpus
is of a more primitive type than that of Triplopus, since the magnum and cuneiform
surfaces of the lunar are subequal, as in Ili/rachi/us, while in Triplopus the magnum
sui-face is much reduced, leading to the Hyracodon type. A still more important
feature is the size of mc. v, as shown by its articular facet upon the cuneiform, in-
dicating that there were four functional toes. The cuboid, when compai-ed Avith that
of Triplopus, is broad and reduced in vertical diameter.

Generic characters. Dentition, c. 1, p. I, m. s. No diastema behind the
canine. Fourth and probably thiid premolai's in both jaws submolariform or with
double internal lobes. Molars in both jaws resembling those of Tapirus. External
lobes of superior molars (paracone and metacone) strongly convex and equal in size.
Last lower molar with prominent third lobe. Manns and pes. Digits, 4-? 3.
Lunar with subequal magnum and unciform facets. Cuboid broad with an extensive
astragalar facet.

Isectolophus amiectenSjf nobis. The postero-external lobes of the superior molars
are produced beyond the junction of the j^osterior transver.sc crests. A strong cing-
ulum surrounds the crown.

/. (Ileldletes) lalidens,* nobis. The postero-external lobes are opposite the junc-
tion of the ])ostenor crests. There is no cingulum.

The Biidger species may subsequently be loiiinl to represent another genus, but
in the absence of the complete dentition of Isectolophus, it can only be separated l)y
the characters of the molar crests, which are hardly generic. The structure of the
premolars removes this genus from Systemodon Cope.

• K. M. Museum Bullelin, No. 1, tS7.S, p. 54.
+ Proc. Am. Phil. Soc. Sept. 2. 1987. p. 260.
} E. M. Jliiseuni Hull-lin. No. I, 1878, p. 54.

520 THE MAMMALIA OF THE UINTA FORMATION.

ISECTOLOPHUS ANInECTESTS.
Plate X.

This genus is represented in the Princeton collection by portions of four individ-
uals, which apparently belong to the same species, and by the type of a second species
from the Bridger. The type specimen (]N^o. 10,400) consists of the second premolar and
first and second molars of the maxillary series and the last lower molar and portions
of the last premolar and first molar of the mandibular series. The second specimen
(No. 10,401) preserves the first upper molar and fragments of the premolars and the
fourth lower premolar associated with a portion of the ramus ; also a complete car-
pus and portions of the metacarpus ; also the cuboid and metatarsal elements, in
addition to the vertebral and limb fragments. The third specimen (iSTo. 10,393) is a
mandibular fragment containing the fourth premolar and first and second molars.

Dentition.

Molars. In the type specimen (Wo. 10,400) m. ^ and m. '' are preserved of the
superior series. The external face is continuous but strongly ti'ilobate (Fig. la) with
a pronounced external cingulum and prominent anterior accessory cusp. The third
lobe of m. - is strongly produced beyond the junction of the posterior crest (Fig. 1).
The transverse crests are rather low and obtuse, curving oblicpiely backwards. At
the head of the main valley, directly opposite the median external lobe, is a low but-
tress. The inteiuial cingulum is prominent, extending upon the anterior and poste-
rior faces of the crown. The first molar is a smaller tooth of very similar pattern.

The third lower molar (Fig. 2) has a prominent creseentic heel. The anterior
crests are stout and slightly oblique ; the outer face shows a creseentic prolongation
inwards, which is rather faint in the middle crest but is strongly produced in the an-
terior crest, the crescent thus formed being strengthened by a buttress. This
buttress is a marked feature of the first and second lower molars, as seen in another
individual (No. 10,391)). In the latter the crowns are unworn (Fig. 11), the crests
are subcrescentic ; the buttress is very distinct in the concavity of the anterior crest
and this crest extends inwards, forming an anterior valley.

Premolars. With the tyjje specimen there is a superior premolar (Fig. 1) in
which the internal lobe is single, with a slightly paired crest extending to the outer
lobes. This tooth is almost unquestionably the second })remolar. Associated with
the third specimen also there are portions of the external lobes probably belonging
to pm. '".

The fourth inferior premolar is preserved in each specimen (Figs. 2 and 11).

THE MAMMALIA OF THE UTNTA FORMATION. 521

The anterior crest is lofty, bilobute, with a distinct anterior valley and buttress; the
posterior crest is low, especially at the postero-internal anj^le. As shown in Fig. 11,
the second and third premolars were bifanged, while the first premolar had a single
fang and abutted closely against the canine as in I. laUdens.

The Skeleton.

Axial. The vertebral and other portions of the skeleton, associated with the
second specimen (No. 10,-101), indicate that Ihey belong to a young indiv idual. All
the verteliral centra, lacking the epiphyses, and the contours of the posterior faces of
the carpals being incompletely developed. The vertebrm belong to the posterior dor-
sal and lumbar region ; the centra are much flattened vertically and slightly keeled.
Fragments of the cranium and infi\ioi'l)ital region are present, but arc not character-
istic.

Ap2)e)icUcular. Fore limb. The .sea[)ula has a suboval glenoid lace, with a
raised anterior border and the base of an ai)parently prominent coracoid process.
The distance between the outer border and the rise of the spine is very narrow, in-
dicating a rather short neck. The proximal portion of the radius presents a limited
lateral extension, with rather deep antero-posterior diameter. These proportions,
taken in connection with the rather broad carpus, indicate that Isectolophus had a
short fore limb.

The Carpus. The most important portion of the skeleton is the nearly complete
right carpus with portions of the second and third metacarpals attached. This
is the first carpus discovered belonging to any genus in this line of descent and
with the cuboid fully establishes the systematic position of Iseclolophus in the tapir
line, as already inferred from the peculiarities of its molar teeth. The vertical diam-
ctei- of the carpus proper is considerably less than the transverse, the series retaining
in great measure the rather spreading character of the iiriinitive type. The first and
second rows fully alternate and interlock. The scaphoid has a subquadrate anterior
face; the radial facet has a broad posterior extension. The trapezium facet is worn
away; the trapezoid facet is excavated and is only a trille broader than that abut-
ting against the magnum. The hinar has a symmeti'ical radial facet with a promi-
nent posterior hook; the inferior face presents subequal magnum and unciform facets,
with closely similar angles of inclination ; this is a very distinctive feature;. The
cuneiform has rather a limited transverse extension of the idnar facet, with a small
pisiform facet; the lunar and unciform surfaces are subequal. The trapezium is not
preserved, but is indicated by a wide facet upon the trapezoitl and upon mc. it. The

522 THE MAMMALIA OF THE UINTA FORMATION.

trcqjezoid is small, subquadrate, with convex scaphoid and metacarpal facets and sub-
equal trapezium and magnum facets. The magnum has a pentagonal face with sub-
equal scaphoid and trapezoid facets; the remaining three sides articulate— o, with the
lunar ; this facet, although narrow in front, extends back to the full depth of the
lunar; also presenting a narrow unciform facet posteriorly ; 5, with the extension of
mc. Ill ; c, with the horizontal face of mc. iii and the extension of mc. ii. The
unciform has a broad cuneiform facet, and oblique lunar facet ; the inferior face pre-
sents subequal facets for the extension of mc. iii and the upper flxce of mc. iv.
The facet for the fifth metacarpal is narrow upon the anterior face and is presented
obliquely outwards and backwards. The full extent of this impoitant facet cannot
be ascertained, owing to a partial fracture and the immaturity of tlie individual.

The second and third metacarpcds have subequal shafts with a rather flattened
and slender section, indicating that they were not long and that the digits were
rather short and spreading. The second metacarpal has a lateral trapezium facet, a
concave upper face for the trapezoid and an extension for the magnum and mc. in ar-
ticulations. The third metaeai'pal besides the above and the deeply concave magnum
facet has a lofty extension and abutment against the unciform. The fourth and fifth
metacarpals are missing ; the latter was borne upon the oblique unciform facet ; the
rearward extension of the unciform which foi-ms the principal support of the toe in
the tapir appears to be undeveloped. The proximal phalanges probably belonging
to these metacarpals are preserved ; they are elongated and rather slender.

The only portion of the tarsus represented is the cuboid, which, however,
pi-esents important diagnostic charactei-s. It is very short and cylindroidal. The
superior face is subcirculai-, with a large hooked calcaneal facet and a compara-
tively extended facet for the astragalus. The outer surface is grooved for the flexor
tendons. The lower facet apparently supported mt. ii and a portion of mt. iii. The
vertical compression of the cuboid indicates that the navicular and cuneiforms were
much flattened. The distal portion of mt. in is broad with a prominent posterior
keel, while mt. ii is considerably narrower with an oblique face.

The distal portion of the tibia shows a prominent malleolus and posterior spine.
The astragalar grooves are slightly oblique, and decidedly shallower than in Epi-
htp]?us.

Special Measurements, Skeleton.

licclohphus, No. 10,401. M.

Cuboid, vcrlical diiiiiu'ter Oil

" Uansvt'ise " 0095

Distal face oft il)ia, transvtrsc 014

" " " antcro-poskrior 020

Carpus, viTliciil iliaiiiolcr, iiiuxiTiium -019

" transvuisc ■• •' 02b

THE MAJIMAIJA OF THE UINTA FORMATIOX. .')2'{

Other measuremenls may be obtained from tlic plates, which aiv caivfiilly ihMwii
to scale.

Dentition, Comparatice i/easurement».

Upper Upper m' m' Lower Lower m^ tn,

series, mohirs niitpos- trans- series, molars, antpos- antpos-
pm'-ni'. m'-m*. ttrior. verse. pm^-m, Wi-Wj. tcrior. terior

hectolophus .047* .016 .018 .088* .05» .014* .023

annectens.
Isectolophus .036 .012 .013 .081 .042 .012 .018

latidens.
Systemodon 063 .032 .OlO.i .012 .073 .035 .010 .016

tapirinum.

Tlie Systematic Position of Isectolophus.

Before discussing the relation.s of Isectolophus., we may review the grounds for
referring this genus to a position in or near the direct tapir line : (1) As far back
as the Wasatch and the overlying Wind Kiver periods, we observe, as recently staled
in full by Copcf that a division in the lophiodont forms is indicated by the shape of
the postero-e.Mternal lobes of the superior molars ; in one line they are conic, in the
second or side line they are flattened. The former leads to the true type of tapir
molar ; the latter extends into another line of genera such as II>/rachyus, in which
tapir characters were partly retained in the feet, but in which the teeth were i-apidly
departing from the tapir pattern ; (2) another feature, to which Marsh first called atten-
tion as characteristic of this line, is the presence of a third lobe upon the lower molars ;
Qi) a third feature is the absence of any considerable diastema behind the canine.
So much for the dentition ; the distinctive foot characters are more fully discussed
later, they are : (4) In the carpus, the retention of subequal maguuin and unciform
facets upon the lunar, and the large mc. v facet upon the unciform ; (5) the exten-
sion of the cuboid upon the astragalus, anteriorly and posteriorly.

The single tapiroid mark in Ilelaletrs is the small third lobe of the last lower
molar ; in the llattened postero-external cusjis, as in J/yraclii/us, and in the high com-
pressed tarsus resembling that of Tiiplopu.% with its narrow astragalo-cuboidal con-
tact, this genus widely departs from the tapir line. Isectolophus, on the other hand,
has all these marks, and the points in which /. annectens varies from T. latiden^s are
all ill the direction of closer approximation to the recent tapir.

In Systemodon of the Wasatch (Cope Collection, see Tert. \'ertebrata. I'l. J^\ I.
Fig. 1-2), the third and fourth upper premolars have two external lobes and a

* Approximate r.-limiitr.

fThc I'irimodac'tyla. Am. N'ol., 18S", p. MK).

524 THE MAMMALIA OF THE UINTA FORMATION.

single internal lobe. The third has a single transverse crest, and the fourth has the
second crest just developing. In the true superior molars, the convex postero-
external lobe is slightly posterior to the junction of the posterior transverse crest
(metaloph), and there is a prominent cingulum embracing this lobe. In IsectolopJius
latidens of the Bridger, which is distinguished from Systemodon by the double in-
tei-nal lobes of pm.' and pm.^, the postero-external lobe is also slightly posterior to
the crest, but in the Pi'inceton specimen (/. latidens) there is no cingulum. Both
these forms have slight diastemata in the dental series. In I. annectens the same lobe,
which is homologous throughout w^ith the primitive metacone of the tritubercular
molar, is greatly produced beyond the junction of the posterior crest. The external
cingulum reappears, so that in this respect, the genus reverts to Systemodon, but pre-
sents such a marked advance even upon the I. latidens molars in other respects that
it must be considered a much more recent type. The third and fourth premolars
probably show a more distinctly developed pair of internal lobes than in I. latidens.
The inferior molars present the tapirine buttresses in the valleys and inflection of the
anterior crests which ai-e wanting in the Bridger species, but are occasionally seen in
the Bridger varieties of Ilyrachyus. Another developmental feature is the steady
growth of the anterior accessory tubercle, which is so marked in the tapir. The
comparative measurements of the teeth in these thi-ee genei-a show that, with the
development of the molar pattern and metamorphosis in the premolars, there was a
steady increase in size, extending proportionally in all parts of the dentition.

It thus appears that the primitive tapirs were well distinguished from the other
lophiodonts very early in the Eocene period, and the tooth and foot structures charac-
teristic of the recent forms were acquired even more rapidly than in the parallel line
of primitive horses.

The Miocene successor of IsectoJoplms is undoubtedly represented by the single
three-lobed lower molar from the White Kiver beds, which Dr. Leidy has referred to
Lopldodon occidentalis. By analogy with the premolar evolution in all other perisso-
dactyls we may anticipate that this tapir will be found to have three premolars like
the molars {Mesotajyirus) .

TUIPLOFUS Cope.

Syn. : Prothyracodon, nobis, Proc. Am. Phil. Soc, Sept. 2, 1887, p. 2G0.

This genus was discovered by Prof. Cope in the Washakie or Upper Bridger
strata, and we find that it is very abundant in the overlying Uinta. The mandibular
dentition so closely resembles that of the smaller species of Ilyrachyus described

TllK MAMMALIA OF THE UIN'TA FOiniATIOy. 525

from the Bridfjer, that it is i)Ossihlc' that some of the hvtter belong to Trlplopia. As
Cope has pointed out, this genus is near the J/t/rachi/us-IJi/racodon series, as shown
by the teeth and the reduced condition of the fifth digit. It was an extremely light
and graceful animal, with long skiulcr limbs and feet of the proportions seen in
Jifesohipjnis, and about the height of tlie smaller domestic sheej). The manus and
pes are extremely elongate, the podials being hiterally compressed, the metapodials
spreading very little, much less apparently than in either IlyracJn/us or Jlijracoihut.
The peculiarity of the manus is that the lunar rests almost entirely upon the unci-
form anteriorly. In the pes the cuboid is elongate and articulates with the astragalus
by a very narrow joint. The genus is distinguished from Hijrachyus by the Hatten-
ing of the external molar crests and their backward extension, also by the advanced
reduction of mc. v. From Ilelaletes, by the single internal lobes of the i)remolars
3 and 4. There is a wide diastema behind the canine and the last lower molai- is
without a third lobe.

This genus is represented in our collection by jxirtions of five or more individu-
als; in every case the specific determination is uncertain and the generic reference of
one specimen is somewhat doubtful. The t\'pes include the Prollniracodon hdenne-
diitm (Xo. 10,40.'3) and IluracJn/us ohiiquidens (No. 10,402) of our preliminary bul-
letin. The former type, together with several individuals which are represented by
skeletal fragments, is nosv referred to the same species as the latter; the latter, a
superior molar, m.", evidently belongs to Trqdojms, and has associated with it a lower
jaw which either lacks or has a rudimcntal first premolai', a good specific distinction
from T. cuhitalis Cope. The type of P. intermedium cannot be distinguished from
T. cuhitalis, except by indirect association with the T. ohliguiden.s type.

Generic characters. Denlit ion, i. [!, c. ], pm. ;| I, ra. if A diastema behind the
canine. Fourth and third premolars submolai'iform ; upper third and fourth pro-
molars with single internal lobes. External crests of superior molars llatlened.
Third lower molar bilobed. S k u 1 1, meatn^ nn/litoriiis e.vternus closed by bone infe-
riorly. Tymj)anic expanding into a bulla. Posttympanic and paroccipital processes
distinct. No postorbital arch. Manus and pes. Digits 3-3. Fifth metacarpal
rudimental. Lunar resting principally upon unciform. Astragalus with slight
cuboidal contact.

.TniPLorus obliquii>en.s S. A: O.

I'l.ATK XI.

7'. olifit/uiden^s. Syn. : fli/rdc/n/ns ithlit/xidens, nobi.s, Proc. Am. Phil. Soc, .^ept.,
18S7, p. 2.^0; probal)ly also, Protlniriicitdmi. int-rnifdiuin, nobis, /oc. ''/V., p. 2(30.

A. !■. S. VOL. XVI. 3o.

526 THE MAMMALIA OF THE UINTA FORMATION.

Specific characters. The transverse crests of the superior molars continuous;
the first inferior premolar rudimentary or wanting.

Dentition.

The superior premolar-molar series (Fig. 6 ; type of P. intermedium, Ko. 10,403),
consists of pm, -m.". The external crests of m.^ and m.' are wanting, as well as the
transverse crests of pm.'. The external crests of the two premolars are subquadrate
with two symmetrical vertical ridges (Fig. 6«). The anterior ti-ansverse crest of
pm.^ is strongly recurved, forming the single internal lobe, against which the slender
posterior crest abuts. The entire inner face is surrounded by a prominent cingulum.
The peculiarities of the molars are, the elevation of the transverse crests, the con-
cavity of the postei'o-external crest and faint marking of the vertical ridge. In m.-
and m.^ the external crest is produced widely beyond the posterior transverse crest,
but in m. ' the extei-nal crest is very oblique and barely overlaps the posterior trans-
verse crest, leaving no jDosterior fossa. These teeth closely resemble those of T.
cubitalis as described by Cope,* but are slightly larger.

Meastireme?its. M.

Svperior molars (10,403).

Third premolar to third molar inolusive , 064

Second molar, antero-postcrior diameter 015

" " transverse •' Ol.j

Fo\irlh premolar, antero-posterior diameter 01

" transveree " 0135

Inferior molars, etc. (10,403).

Median incisors to third molars inclusive 102

Length oC the premolar-molar series 076

" " molar series 042

" " postcanine diastema 013

" " sym]ihysis 035

Least width ol' " 016

Antero-posterior diameter third superior molar 017

Transverse " •' " " 014

The second specimen (10,402) is the type of T. oUiquidens. It is intermediate
in size between JJijrac/H/us agrarius and II. nanus. There are two mandibular rami
(Fig. 7) with the symphysis complete and portions of the premolar-molar crowns ; also
a fragment of the maxilla Avith m.' and m.- and the complete crown of m.' (Fig. 10).
The latter tooth resembles that of the specimen- above described so closely, that it
enables us to identify this s])ecimen, first ascribed to Hyrachyus, as belonging to

♦Tertiary Vcrtcbralji, p. («!.

THE MAMMALIA OF THE UIXTA FORMATIOX. 527

Tii]}lo]ni.-^. The fimgs of three subeqnal incisors and a modiuin sized canine are pre-
served, behind which is an extended diastema, followed in the left jaw by the minute
alveolus of pm/ which may have been retained in sila at a less mature age. In the
right jaw no trace of this alveolus can be seen. If this premolar is found to be in-
variably missing, this may be given as the generic distinotion of Prollnimcodon.
The second premolar is rather small, the third and fourth are submolarifoi'm. The
three molars are nearly equal in size with the ordinai-y lophiodont pattern, the ante-
rior crest is well incurved in the worn condition of the crown. The mandible is stout
and well rounded, contracting greatly opposite the diastema. The measurements
indicate a heavier jaw and laiger animal than the type of T. cubitalis. The last
molar has no ti-ace of the third lobe, but there is a considerable space between it and
the ascent of the coronoid process.

The third specimen (10,397) is of unusual interest, since it contains, associated
with a few teeth, many portions of the skeleton and the complete tarsus and carpus.

The Fore and Hind Limhs.

The fore limb of T. cubitalis has been fully described by Cope, op. ciL, p. 081.
It is remarkable for the unusual elongation of the fore arm and of the manus. The
head of the humerus is laterally compressed, with marked fore and aft extent;
the distal trochlea is unusually deep and extensive, indicating great play at the elbow
joint. The distal portion of the scapula has a very long neck, and the spine
rises very gradually ; the scapula was probably high and narrow. The pi'oximal
facet of the radius is very deep antero-posterioily. The ulna is remarkable for the
great extension of its olccraTion ; distally it is apparently more reduced than ui T.
cubitalis.

The extremities of the bones of the hind Ivnb are mostly preserved, and indicate
that the proportions were long and slender. The femur has a long patellar facet and
a deep intercondylar pit. The tibia has a narrow but prominent cnemial crest and
double spine ; the distal face has a deeply excavated astragalar trochlea which is
decidedly oblique. The iibida has a fairly stout distal extremity, and is throughout
entirely distinct from the tibia.

The carpus (Fig. G). The scaphoid is very stout with a deep radial facet ;
inferiorly it presents a minute facet for the trapi'ziiim, an oblique trapezoid facet, sub-
equal with the slightly larger and horizontal magnum facet. The lunar has a
nearly horizontal unciform facet and vertical ental facet which rests partly against
the scaphoid and has a slight magnum contact. The c u n e i for m is very thin, and
has a narrow ulna facet extending into that Ibr the pisiform. The trapezium was

528 THE jrAMMALIA OT THE UINTA FORMATION.

extremely reduced. The trapezoid is wanting. The anterior face of the magnum is
subquadrate, since the ectal lunar and unciform facets are in the same slightly oblique
line. The unciform is remarkable for its depth; superiorly it has subequal
cuneiform and lunar facets ; inferiorly it has subequal mc. iii and iv facets and a
small facet for the rudiment of mc. v. The metacarpals have sharp distal keels
confined to the deep posterior faces. The phalanges are deep and elongate.

The tarsus (Fig. 9). The main features of both the carpus and tarsus are de-
scribed in the section on the feet of the Perissodactyla. In addition the following fea-
tures may be mentioned : The sustentaculum of the c a 1 c a n e u m is very narrow ;
the cuboidal facet slants obliquely downwards as in the Artiodactyla. The metatar-
sals have the proportions seen in MesoMiyp^is,

Measurements may be obtained from the natural size figures upon Plate XI,

Tim Systematic Position of Triplojrus.

Prof. Cope (Tert. Vert., p. 678; Am. Naturalist, 1887, p. 909) has placed Trij}-
lojms in a distinct family. He says : " The entirely rudimentary character of the
fifth metacarpal, which with its digit is so well developed in Hi/nicJryus, places
TrtplojJits in another family and in a distinct line of descent. I think that it must be
regarded as one of the forms connecting the lophiodonts with the rhinoceroses." We
cannot accept this conclusion. In the discussion of the manus and pes in the Hyra-
chyus-Uyracodon series, on a later page, it is shown that the feet of Triplop'm are in
every detail intermediate in structure between those of the above Bridger and White
River genera. The teeth are also transitional. In the upper molar scries of Triplo-
jynn, from the thiid premolar to the last molar inclusive, the external faces have a
striking resemblance to those of Ilyracodon, while the wearing surfaces of the same
teeth are close to those of Hyrachyits. The departure from the Hyrachyus pattern is
seen, first, in the decided elevation of the crests and disappearance of the postero-
external lobe or ridge ; second, in the rhinocerotic shortening of the posterior trans-
verse crest of m. '■ ; both these changes are in the direction of Ilyracodon.

These characters suggest our considering the genus Triplopus as a transition
form. !None of the known species, however, exactly fill the requisite transition
features. T. nih'tah's and 7T oUiqnidens are apparently to be considei'ed aside from
the main line, on the following grounds : The fore ai-m is proportionally much longer,
the metapodials are more compressed, the lateral elements of the pes especially are
more reduced, than in any possible diiect ancestor of H. nehrascencis. A smaller,
lighter species of Ilyracodon is, however, represented in the Harvard collection by a
carpus which while slightly larger than that of T. ohliquidens presents a very similar

TUK MAMMALIA OF THE UINTA FORMATIOX. 529

stage of reduction of the lateral metapodials. This is further evidence that some
species of the Lower Miocene genus will be found to approach Triplojjus much more
closely than // nebi-asceiicis does. The ujjper molai's of the known species of Trip-
loj)us entirely lack the rhinocerotic "anticrochet" seen in /////rrnyf/o//. and are also
devoid of the strong external cinguluin.

Another feature- of Diplajyu'^ should, however, be mentioned, which may serve to
justify Prof. Cope's views of its separate systematic position, that is, the apparent ex-
pansion of the tympanic into a small auditoi-y bulla. The meatifs anihtarins exteiiivs
is completely enclosed bi'low, whereas both in llijrachyus and ILjracodim it is widely
open.

We probably have in T'rrploims a highly specialized cursorial type in which there
is a slightly more marked tendency to monodactylism than in the predominant spe-
cies of cither llijrach ijm or ILjracodon. A recently completed restoration of II. ne-
brascmcis from materials in the Princeton and Harvard Museums shows that the
Miocene genus retained substantially the proportions of jryrachijus tximhis as seen in
the complete skeleton discovered by Cope.

EPIIIIPPUS Martili.

This genus is represented ijy the lower jaw and numerous portions of the skele-
ton of a single young individual (No. 10,405). The species cannot be determined.
The specimen is provisionally referred to E. uintensis Marsh.

Generic characters. Dentition, i. ^, c. j, pra. |, m. j|. The third and I'ouith
upper premolars are like the molars, and the corresponding lower premolars have the
double antero-internal cusps (a a Riitimeyer). The ulna is complete and distinct
fiom the radius. The lunar rests equally upon the magnum and unciform. Digits
4-3.

EiTiiiri'iis UINTENSIS Marsh.

Plate XI,

Dentitiou. The lower jaw is shallow, tapering to the narrow, slender symphysis.
The incisor fangs (Figs. 3, 3r0 are equidistant and subequal. The canine is conside-
rably larger. The first premolar has a simple pointed crown and single fang, 'i'he
premolars 2-4 are missing. The second molar has the characteristic reduplicatiun of
the antero-internal cusps. From the antero-external cusp ( protoconid ) a sharp
ridge extends downwards and inwards ; there is a well-defined external cinguluin.
The third molar shows the same characters with a strong posterior cingulum. The
superior teeth are not preserved.

530 THE MAMMALIA OF THE UINTA FORMATION.

Fore limh (Figs. 4, 4a). The distal portion of the humerus shows an extensive
trochlea and deep anconeal pit, but no perforation. The proximal portion of the
radius entirely conceals the ulna ; the shaft is arched forward, strongly convex in
front ; the ulna descends along the flattened posterior surface and appears at the
side in the lower third of the shaft. The ulna has a small pisiform and narrow unci-
form facet. The chai'aeteristics of the carpus are fully described elsewhere and
well shown in the figures. The third metacarpal does not show the increased size
we should have anticiiJated in this genus, but would probably be relatively much
larger in an adult individual. The fifth metacarpal is about three-fourths the length
of the second and fourth. The phalanges are rather short and stout. The ungual
phalanges spread distally. The raetapodial keels are very prominent on the posterior
face.

Hind limh. The tibial shaft is preserved, but is not characteristic. The tarsal
characters (Fig. 5) agree closely with those of Hyracotlierium, and are fully described
elsewhere. The astragalus has a very deep groove, narrow navicular facet, narrow
cuboidal contact, and three distinct calcaneal facets. The ealcaneum has a distinct
fibula facet and rather short stout tuber; the sustentaculum is very narrow. The
third metatarsal is very dee]3 and much stouter than the second and fourth, which are
strongly rotated backwards. The second metatarsal apparently articulates laterally
with the ectocuneiform.

Taken altogether E^jihippus preserves more of the primitive characteristics of
Ilyracotherimn than we should have anticii^ated. These are especially seen in the
caipus. The tarsus shows more marked reduction of the lateral members. The
fibula is probably still complete.

TIIK MA^OIALIA OF THE UINTA FORMATrON. 531

Part IY.

I?Y HENRY FAIRFIELD OSBORN.

THE EVOLUTION OP THE UNGULATE FOOT.

Ill the Uinta and undeilying Washakie beds we meet with, at least, five distinct
series of Perissodactyhi, representing the recent Tapiridai and Equidje, also the
Titanotheriidtii and Hyracodontidi^, which became extinct in the Miocene, and Amyiio-
dontidte which may have given off the true rhinoceros line. The foot bones of each
of the series are fortunately represented wholly or in part in the Princeton collection
and present interesting transitions to the Bridger types of feet on the one side and the
White River on the other. Many notes upon the latter have been derived from the
study of the collection in the Museum of Comjiarative Zoology through the courtesy
of Prof. Agassiz. The writer is also greatly indebted to Prof. Cope for free access
to his collection which is so rich in Lower Eocene forms.

A careful comparison of these earlier and later forms showed it to be i)ossible to
distinguish, in nearly every instance, the separate elements of the feet in each seiies
by a number of inconspicuous but thoroughly diagnostic marks. This led the writer
to a study of the minor characteristics of the foot bones in the recent and extinct
Perissodactyla and the earlier Ungulata in geueial. Kovvalevsky has given us a
model for such research in iiis Anchitherium memoir, in which structural modidcation
is constantly viewed from the functional standpoint, while we are greatly indebted to
Cope for his numerous essays upon the broad lines and philosophy of the transitions
from the taxeopod to the recent ungulate foot. The purpose of the present contribu-
tion is to indicate the chief characteristics of the foot bones in each of the phyla
diverging from the primitive Taxeopoda, chielly in their bearing upon phylogenelic
questions and the laws of modification of foot structure. The subjects may be con-
veniently treated in the following order:

I. The foot stnicture in the ancestral Taxeopoda or Protungulata and tin- niodili-
cations which take place in the ungulates in general in the evolution from the
primitive type.

532 THE MAMMALIA OP THE UINTA FORMATION.

II. The main features of the mamis and pes in the Hyracoidea, Amblypoda,
Proboseidia, Toxodontia, and in Macrauehenia.

III. The chief and minor characteristics of the manus and pes in the various series

of Perissodactyla.

IV. The taxonomic value of the primitive and secondary articulations in the manus

and pes.
V. The laws of modification of foot structure.

I. THE PRIMITIVE FOOT.

The more recent types are connected with the remarkably simplified feet of the
Puerco ungulates by Phenacodm ; without this link it would be presumption to
describe the feet of such genera as Periptychus, Uctoconus, and Memscotherium, as
ungulate, since, excepting in the terminal phalanges, they resemble the plantigrade
carnivore foot far more closely than those of any living ungulate — possibly excepting
Hyrax.

The pes of the Puerco Penptychus and Pantolamhda are fully known. The
former corresponds so closely with that of Urf<us, we may anticipate a similar homo-
logy in the manus, especially as the entire structure of the upper portion of the fore
limb is bear-like. Our actual knowledge of the primitive taxeopod manus is at
present inferential from Plienacodus in comjjarison with the more recent types in
which taxeopod characteristics persist — as in most cases we can readily detect and
eliminate secondary structures.

General cliaraciers. The primitive manus and pes of the lowest Eocene ungu-
lates had five spreading digits bearing spatulate terminal phalanges marked by a me-
dian groove. The first and fifth digits were shoit. The separate elements of the
podium and metapodium were articulated serially, the only interlocking joints being,
as in all mammalia, beneath the scaphoid and unciform, cuboid and navicular. In
standing, the entire sole rested upon the ground, the foot forming a right angle with
the lower leg; the limited astragalo-tibial facets in all the Puerco forms leave no
doubt upon this point, and we have a persistent example of this type in Hyrax.

The pp:s. In the pes both the tibia and fibula wei'e supported upon the astragalus;
a f i b u 1 o - c a 1 c a n e a I facet is frequently assigned as one of the characteristics of
the primitive ungulate foot,* from the fact of its appearance in so many difterent
phyla. Among the primitive Ungulata this facet has thus far been observed only in
Meniscotherium, it is wanting in all the other Condylarthra. (1) In this genus it has

* See Marsh, Diiiorcnila, C'li:n:icli'rs of llic J'rotiingiihild, \). 171.

THE MAMMALIA OF THE UINTA KOHMATION.

533

a trochlear character, the liljiila resting upon the calcanenm throughout extension, as
in Macrauchenia and the ArtiodactyUu (2) In the primitive Equidto and in the Titano-
therida-, the fibula rests upon the calcanenm in extreme tlexion. In ancestors of
the latter family we observe this facet developing evidently as an adaptation to groat
body weight. (3) The third type of libulo-calcaneal facet, as seen in the Proboscidia,
(?) Dinocerata, and Toxodon,* in which the ankle motion is limited and the (ibula
rests broadly upon the calcanenm, is also probably a secondary adaptation to weight.
As in several instances this facet has been acquired secondarily, as it is wanting in the
majority of primitive taxeopods, and is not essential to the perfect working of the
plantigrade ankle joint — the evidence as to its primitive character is certainly not
conclusive.t

A B.

DiAOS. fi-7. — Tliiid linil) of Periptychus rliahdodon Copo, from original in the Cope collection. A, Front view of
pes, X \. D, Siili- view orriL'lit himl linil). X I- Tlir rc.-torfMl and conjectural portions are in dolicil lini-j. <if, Aslmg-
alar foramen.

The trochlear groove of the astragalus faced upwards, when the foot was
prone, and with its slight concavity and obtuse edges allowed considerable lateral
movement. The antcro-posterior movement was evidently very limited, for this facet
in all the Pucrco genera is small ; it was indented by a foramen probably transmitting
the flexor <l>f/Horiim cmnmunis. The importance of this astragal a r foramen as

• " In Taxodon tlie fibula articulation is of unusual size." Cope, Proc. Am. Phil. Soc, .\|>ril, I8.SI, p. 'lOS.
f Wc are spcakins; of primitive unf;ulales, not of primitive Mammalia in general.

A. P. S. — VOL. XVI. 3p.

534 THE MAMMALIA OF THE UINTA FORMA TIOIST.

a characteristic of the Ungulata and primitive Mammalia generally has been hitherto
ovei-looked. Cope has observed it in Bathiiodon^ and in Peript[icJius ;t Marsh has
described it as present in about one-half the Uintailierium asti'agali.J Lemoine§ has
found it in all the Cernajsien mammals, which aie of about the same age as the
Puerco. It is also present in Ectoconus, MevJ soother lum, Pantolarnhda, and in many
if not all of the Puei'co creodonts. The astragalus had a short neck articulating
distally mainly with the navicular by an oval and strongly convex facet, and, while
in several Puerco genera (^Periptychns, Pantolamhda), it has a considerable ecto-
lateral facet for the cuboid, this facet was probably absent in the most primitive
forms. Another important feature was the articulation with the calcaneum by two
facets only, the ectal (posterior) and sus tentacular (see i-*/(ertaco(:Z«s, Diagram
10).

The tibiale. || A careful comparison of the tarsus of PeriptycJius and Den-
droliyrax has brought out the interesting fact that there is a broad ental facet upon
the astragalus in some of the Taxeopoda for the tibiale similar to that in the Hjn-a-
coidea. This facet, which Cope has recognized as the tibiale or " internal navicu-
lar " in PeriptycJius, is also observed in Meniscoilierium, Pantolambda, and the Puerco
mammals generally. It has been pointed out by Baur in Uiatatherium, by Cope in
Bathmodon, and should undoubtedly be added to the list of primitive characters of
the jjes.

The calcaneum resting upon the ground with its sustentaculum bore the whole
weight of the leg transmitted through the astragalus, articulated distally by a
concave facet with the cuboid. The cuboid supported mts. iv and v, the three sub-
equal cuneiforms articulating respectively with mts. i, ii and iii. A very constant
feature was the relative shortening of the mesocuneiform (Phenacodus), so that
the proximal portion of the mt. ii was inserted between the endo- and ectocunei-
forras. A similai- facet may be present between mt. iii and the cuboid ; these ectal
facets being analogous to those upon mcs. ii and rii in the manus in not being of a
supporting character; in other words, there was no true displacement between the
podium and metapodium.

The Manus. There was in all probability a cent rale occupying the same posi-
tion as in Ilyrax, for this clement has now been found both in the Hyracoidea and

* Ti:rliaiy Vc-rtclirata, p. 551.

t Op. cit., p. 399: "This Ibraiucn is foriufil by the closing of llio usual tendinous uoteh, " etc.
I Dinoeerata, p. 148.

§ Various pa]iers upon tlu' Cernaysien fauna.

1 Baui-, On llu' Morpliology of llie Tarsus in tlie Mammals, Amerieau Naturalist, .Tan., 1885, p. 349. See also,
Marsh, Dinoeerata, p. I-IG.

TlIK MAMMALIA OP THE UIN'TA FORMATION. 535

Prol)oscidia.* The distal row of carpals was probably much broader than the prox-
imal, and the metapodials, as in the pes, were without any lateral facets.

General Modification of the Fee^.

The {irimitive structure, as Cope has shown, is ai)proximatcly preserved in the
plantigrade Jli/rax, showin<^ that all the secondary articulations in various un<^ulates
were adaptations to digitigradisni. There are four stages in the foot elevation :
First, the plantigrade, already described; then the s e m i p 1 a n t i g r ade,
seen in the feet of the Proboscidia and Dinocerata, with short slanting metapodials
well raised proximally. Kutiraej'er has proposed ''digitiplantigrade " for the reduced
types with elongate vertical metapodials in which the phalanges rest fully upon the
ground {Ccmtelus). As this condition is practically analogous to that in one division
of the Carnivora, it may be more simply described as d i g i t i g r a d e. There arc of
course many steps between this condition and the u n g u 1 i gr a d e foot, in which
the terminal phalanges alone rest upon the ground (^Eipim^).

The universal result of elevation of the wrist and ankle joints was displace-
m c n t in some form, from the serial ordei', adapting the foot bones to the new inci-
dence of impacts and strains. Then came reduction of one of the lower leg bones
and of the lateral digits. With one exception, seen in the proboscidian manus, the
departure from the serial tyi)e was upon a single principle : First,t the ii and in
metacarpals and metatarsals a c q u i r e d late r a 1 s u j) [) o i- 1 i n g
facets upon the second row of p o d i a 1 elements, usually u})on
the outer side. Second, the bones of the upi)er row of carpals and
tarsals formed articulations with the more octal eleniuiits of
the second row.

Til the general modilications of the Maxl'S it is important to note that nu-s. ir
and III invariably accjuired supporting facets upon the octal side with tiie magnum
and unciform respectively. Excej^ting in the scmiplantigrade orders the reduction
of the first digit was very lapid, it being already absent or functionlcss in the
Eocene Diplarthra. The extension of the lunar was the next step in displacement ;
spreading upon the trapezoid in the Proboscidia, or upon (lie unciform, by its own
giowth or that of the unciform beneath it, in all other ungulates. This is inci])ient in
P/^eHacorfM.-* and extensive in the Arablypoda. In other phyla tlu- displacement ol

• Riilinieyer, Uuber einige Buzieliiingcn zwischen ilcn SiiiigcUiicrslnmmcn Alter iind Neuer Welt, Zurich, KVSM.

tSccMnrali, nimxiTaHi, |>. is:i, ^ 3 . Tlint Uic mctn|>ni1liil dhpliirfinrnt wiis iimrr |iriinilivc timn llw |><>ilml In
shown both in P/ien'foilu» iind /lyr<ir, in which mk-9. ii und III aliut laliriilly iigiiin^^l il.' m .-iniiii iiikI iinrilorni, while
the carpals arc serial.

536 THE MAMMALIA. OF THE UINTA FORMATION.

the scaphoid aud hinar was probably concomitant." In the earliest known Diplar-
thra the scaphoid has a facet upon the magnum of the same width as that of the un-
ciform beneath the hmar — these facets increase, pari passu, until we reach the most
specialized forms. In all the Perissodactyla the law of ectal displacement of the
metacarpals is maintained ; after mc. i disappears the trapezium persists ; as mc. ii is
reduced it invariably retains its entire hold upon the trapezoid ; mc. iii sj^reading
only to the ectal side (Kowalevsky). In the ada])tively reduced Artiodactyla, how-
ever, mc. Ill acquires a broad trapezoidal facet, forming an exception to the otherwise
universal rule.

The general modifications of the Pes are still more interesting. In the astragalus
the flexor foramen disappeared, being for a long time represented by a groove ;
the primitive foot undoubtedly had a limited grasping power, the flexion being dis-
tributed between the metapodials and phalanges ; this was replaced by the greatly in-
creased play in the ankle joint and the development of the muscles inseited in the tendo
acliillis as propellers. The ti'ochlear groove deepened, limiting the motion to the fore
and aft direction (Cope) ; the navicular facet became saddle-shaped and then flat, ex-
cepting in the artiodaetyl phylum ; the sustentacular facet divided into two, which may
be called the "sustentacular" and "inferior ," producing three distinct cal-
caneal facets.f The displacement of the astragalus and cuboid by reciprocal growth
was almost universal, having but two exceptions, in the Proboscidia, where the navic-
ular extended upon the cuboid, and in the lines of Perissodactyla, which tended to
monodactylism. The calcaneum in several series acquired a fibular facet, the tuber
was elongated and the sustentaculum extended inwards. While rarely in contact
with the navicular anterioi'ly (as in the primitive horses), the calcaneum in some

"* Upon this i)oinl hangs the question whether the diplarthrous toot ever passed through the amblypodous stage,
as held liy Cope. In the absence of any direct palfeontological evidence, Schlosser (Beitriige z. Stammesgeschiehte der
Ilufthiere, p. 6) has strongly insisted upon the direct derivation from the Condylarthra upon the ground of the dis-
similarity both in foot and tooth structure. His objections, though well taken, indicate a misunderstanding of Prof.
Cope's ta.\onomic principle of selecting genera at a certain stage of evolution in foot or tooth structure, to form a family
or order, irrespective of actual descent or relationship. When, upon this system, the derivation in question is main-
tained, no genetic relationship of the Perissodactyla aud known Amblypoda is thereby implied, but that at one stage
in the evolution of the diplarthrous foot the lunar rested widely upon the unciform, as in Coryphodon, the scaphoid
not reaching the magnum. Tliis jiroposition .seems to us highly improbable, for we must suppose tliat the lunar first
extended upon the unciform while retaining its entire magnum facet, then it became reduced while the scajihoid ex-
tended upon the maguuiu— this hypothesis is rendered improbable by the fact that in all the early Diplarthra the
scapho-magiiuiM and lunar uncitbrm facets are subeciual. The proboscidian manus is an e.xact counterpart to the
amblypod manus, and shows that tlu' latter is to be considered not as in nii intermediate Imt as in a linal stage of de-
velopment.

I It is intciTstiiig to note the close paralh'lisiii to tlic-^e adaptations to digit igiadism in the Carnivora ; we oliserve
an entirely analogous division in these facets and evolution of tlie anUlc Joint.

THE MAMMALIA OF THE UIXTA FORMATION. 537

series acquired a posterior navicular facet. The reduction of the entocuneiform fol-
lowed that of mt. i. The lil'th digit was much more rapidly reduced than in the
manus. Independently of these reductions, mt. ii and mt. in formed oblique sup-
porting facets with the ectocuneiform and cuboid respectively analogous to the
"alternating " metacarpal articulations, l)ut the metatarsal articulations were much
more variable than the metacai'pal. As a general rule metatarsal displacement anJ
growth was to the ectal side, the exception being in the Equidio, where mt. in
extended rapidly over the mesocuneifoi-m.

The inetajjodials. In all the perissodactyls, primiti\e artiodactyls, and inadap-
tively reduced artiodactyls we iind, as above stated, but oue alternating type
of metacarpo-carjjal articulation ; the adaptively reduced artiodactyls develoj) a
second tyi'e of metacarpal articulation. In the jjes, beginning with the "serial"
type of Phenacodus, we find that genera tending to nionodactylism develop what
may be called the " p 1 a n e - s e r i a 1" type, in which there is for a period no lateral
spreading (J/i/raco(/ieriu7n). The "reverse" type is exemplified in AjjJielojys and
T. iiulicus, in which the second and fourth metatarsals both acquire ectocuneiform
facets. The alternating type is developed in the Titanotheriam tarsus and some
other Diplarthra precisely as iu the caipus. Each of these types has, of course, its
functional significance.

"We may now consider some of the special characteristics of the manus and pes
in the five side lines. While observing many features which point back to the com-
mon taxeopod stem form, there are few or none which indicate any nearer mutual
affinity than this.

TI. THE HYEACOIDEA, AMBLYPODA, PROBOSCIDIA, Etc.

1. IIyuacoidea.

An impoitant fact is brought out by the compari.son of the feet of /////".' and
Detulvohijrax. Tlie manus and pes are absolutely plantigrade. In both, the caipals
are arranged in strictly serial order, as described by Cuvier and Cope, although the
metacarpals .show decided lateral displacement. But in the tarsus of Ifi/r<ix the
calcaneum rests exclusively upon the cuboid, and, as in riiradcodax, the caleaneo-
cuboidal articulation is actually in .some ca.ses below the lc\ cl of the astragalo-navicu-
lai-.* In i'e/i(//o////?Y/a; (//. arWe?/s, Princeton Mu.seum), also from the Cape, the

* Tlicso rpliilinn.4 witc foiiml liy rcmovinjt llio skin nnd fascia from llir fi>ol of a ilricd <i|>oi-imcn in the Co|«" Col-
li'flinn. I)c Bliilnvillc'!) flftnn-- nf //. r>//irn*u nml // tyriarun !«li<iw tln,'.»«' lii<x-ls <(ii llu' •^uni.- Irvol.

538 THE MAMMALIA OF THE UINTA FORMATION.

astrajT^alus has quite a broad cuboidal facet. The striking difference
in the pes of these two genera bears directly upon the question of the taxonomic
vahie to be attached to these articulations.

Other features of the feet are the intei-locking tibio-astragalar joint, the probable
presence of a tibiale, the centrale in the carpus, the reduction of mts. i and v in the pes.

2. Amblypoda.

Cope has described the feet in this order as " plantigrade" (" Tertiary Yertebrata,"
p. 507). This term, which has also been generally applied to the proboscidian feet,
should now be restricted to the types in which the entire foot rests on the gi'oiind.
The Puerco PantolamMa* was probably plantigrade. The position of the meta-
podials in the Pantodonta and Dinocerata as indicated in the figures of Marsh
(" Diuocerata," p. 184) was analogous, the wrist and ankle joints in each being well
raised from the ground. Oiu- own study of the tarsals of Coryphodon has, however,
convinced us that the pes was almost plantigrade, while the manus was semi-planti-
grade.

The metacarpals ii and iii are displaced, alternating upon the magnum and unci-
form. The intercai-pal displacement is unique ; the lunar has a broad foothold upon
the unciform, while the scaphoid has either no magnum facet {Cory pi odon, fide Co-^o)
or rests by a considerable posterior facet upon the magnumf as in some species of
Uintathenum. The cuneiform rests partly upon mc. v.

In the tarsus Baur called attention to the fiicet probably for the tibiale upon
the astragalus of CorypIiodonX and Uintatlierium. The displacement is extreme, the
astragalus extending the entire breadth of the foot in front, covering both the navicu-
lar and three-fourths of the cuboid. The astragalar foramen is frequently present in
Corypiltodoii (see Cope, loc. cit.) and in Uintatlierium (see Marsh, "Dinocerata," p. 148).
The calcaneum has the primitive ectal and sustentacular facets, and the fibula occa-
sionally came in contact with it {Uintatlierium, Marsh, oj). cit. p. 152). The meta-
tarsal articulation is of the " I'everse" tyi)e, mts. ii and iv articulating laterally with
the ectocuneiform. The Amblypoda with unreduced digits, very short spreading

* riiiccd by Copt; in the Auililypocla.

f III his Uefinition of the Amblypoda, Prof. Copi; says (" Tcrliary Vortcbrata," p. 378) : "Scaphoid supporlcd liy
trapezoid and not by magnum, wliicli willi uncilorm sujiiiorls tlic lunar." Sec also, Amer. Naturalist, Ni>v., 1884,
p. 1110. While this definition may ajjjdy strietly to Coryphodon, it cannot to Uintatlierium. Of three carpal series in
the I'rincclon Museum, two show a broad scaphoid facet uiion the jiosterior half of the magnum, the third has none
In Marsh's " Dinocerata," the lithoirrajihic figure on Plate LIV is incorrect ; the wood-cut, Fig. 153, is approximately
correct ; the scajiho-magnum facet is not indii-alcd in Fig. 111?, lint is well shown in Plate XXXIV. Fig. 5, see page 113.

^ Cope, however, seiianiles Coryphodon by the ab.sence of a tibiale, from Bathmodon, in which it is present.

THE >r\MMALlA OF TIIK UINTA FOIJMATIOX.

539

metapodials and peculiar carpal displacement, a foot type adapted to animals of heavy
bulk, really present a striking analogy to the Proboscidia and an intermingling of
primitive with acquired characters.

rhrmirodiis

JIaslcflon

Den c/ro/tymx

Dl.vr.. ft. — The (lisplacoracnt of the bones of the manus in the Ungulata, I. Phenacoiui firimtctu*, from oritcinal
in the Cope rolleclion. Coryphodon hamatus, after Marsh. Dendrohyrai arboreus, from original in the Prinrcton col-
lection, .ffaerauc/ienia pnliichonica, ader GcrvMs. Mastodon Americanus from originalia Princeton collection. The
oblique arrows indicate the direction of displacement.

3. Pkobosctdia.

"Weithofcr* has recently directed attention to the unirpie depart iire from tlic 1a\-
eopod condition of the carpus in this oixlcr — the lunar s[)reading uj)un the trapc/.oid,
i. e., to the ental side, instead of upon the magnum. In M(txln(lon arvevnens/'s, Ele-
phcis meridionalis and E. antiquum from one-fourth to one-liftli of tlie inferior face of
the lunar rests upon the trapezoid. We find the same condition in Mastodon Amen'canus.

This author attributes this to the enlargement of the ulna and its extension upon
the lunar r.'*. the radial enlargement in all other ungulates. This displacement is
varial>le, since it actually recedes and sometimes disappears (<>]>. n'f.. p. "l") in I he
adult recent forms {E. Indims, E. Afrlcamis). Burmeister moreover has figiuedf
the carpus of a specimen of M. ILimhohllii, in which the lunar pj)rcads both upon
the trapezoid and magnum. Another primitive feature of the carpus is the recently

• Einigc Btmcrkiinjren fiber den Cnr|)ii8 der Probosrjdicr. Morph. .Jahrb., 1888, p. liOr.
f Ann. del. Miis. Publ. d. Bucn. Ayrcs, Plate XIV, p. 287.

540 THE MAMMALIA OF THE UINTA FORMATION.

discovered ceiitrale.* The distinctive features of the tarsus have long been recog-
nized, viz., the extension of the navicular with the astragalus above the cuboid, and
the fibulo-calcaneal facet.

4. TOXODONTIA.

Piof Copef has found a striking resemblance to the proboscidian pes in Toxo-
(lon. The astragalar and fibular facets of the short c a 1 c a n e u m are subequal and
face upwards, the cuboid facet is on the inferior face of the extremity of the calcan-
eum and faces downward. The astragalus has probably no cuboid facet; its
navicular facet is plane and truncates the bone somewhat inferiorly as Avell as distally.
The metapodial elements are robust and short, the distal metatarsal keels are poste-
rior and rudimental ; the proximal faces have lateral facets indicating a slight dis-
placement. The number of digits is uncertain, the lateral ones are reduced in
length, but robust. " The posterior feet were more truly jjlantigrade for the extrem-
ity of the calcaneum reached the ground." The instep was raised, forming an arched
angulate type of foot. If this description is correct, this foot is more primitive than
that of the Proboscidia.

.I. Mackauchenia.

The feet of Macraudlienia have some striking peculiarities. The lateral displace-
ment in the carpus and tarsus so characteristic of all the Perissodactyla is absolutely
wanting in the tarsus and aberrant in the carpus. As Burmeister pointed out in his
most recent paper J (p. 263) a gap exists in the usual point of junction of the calca-
neum, cuboid, astragalus and navicular. The astragalus is entirely separated from
the cuboid and even from the calcaneum, the navicular participating in this
separation. Ilis description and figures strikingly suggest the arrangement of the
tarsals of the dog in this region. The calcaneum has but two astragalar facets,
the sustentacular and inferior having coalesced, as in some species of Pahaotliermm,
liliinoceros, and the Hyracodontidaj ; its distinctive feature is the extensive fibular
trochlea, very similar to that in the Artiodactyla, and functionally unlike those
observed in the primitive Equus and Titanotherium series. The c a r p u s is also

* Kiitimeyer, op. cit., pa.nc 11, loot-note.

fNotc on the atnictnre of the I'osterior Fool in Toxodon. Proo. Am. Philos. Soo., April, 1881, p. 403.

:t Neuc Beobiicliliin.ijun un MiicranclK'niii Pattichonica. Nova Acta der Ksl. Loop. -Carol. Dculsclu'n Akadoniie
dor Naturforschcr, Bd, XLVII, Nr. 5, (ip. 239-207 (Nachtriigliclie Beschreibung dor Fusswurzel von Macrauchenia).
Also, Dcscriiicion do la Macrauchenia Patachonica. Ann. del. Mus. Piilil., Buenos Ayres, Tom. i, p. 32-GG.

Tin; MAMMALIA OF THE UINTA KOKMATION. 5il

unique, in the fact that wliile the scaphoid has acquired an extensive magnum facet,
the hinar is extremely small ; the magnum is large and not only excludes the huiar
fi-oin the uncifoi-m, but actually has a cuneiform facet. Burmeister describes facets
for latei'al digits upon the pes.* The metapodials are strongly keeled.

Schlossert lays great sti'ess upon the jJrimitive character of this animal as shown
in the serial disposition of the foot bones and the epicondylar foramen of the humerus.
In his earlier paper (Zool. Anz., p. 083), he placed this genus with the Perissodactyla;
later he separated it: "Fasst man alle diese Merkmale zusamraen, so kann man kauin
entschliessen Macrauchenia den Perissodactylen bcizuziihlen " (Morph. Jahrb., Bd.
XII, 188(3. p. 22). Upon this it may be observed that the arrangement of the carpals
is not serial, as he intimates, but has a peculiar adaptive displacement. The fibulo-
calcaneal facet may be considered a secondary character (sec p. 533). We can in
fact only conjecture the phylogenetic position of this genus, with its extraordinary
mixture of characters. Upon the whole it is more nearly related to the perissodactyl
than to any other phylum, and is possibly an offshoot of the primitive mesaxonic
stem, separating from the main line at a period corresponding to the Wasatch.

111. THE PERISSODACTYLA.

When we examine the Diplarthra as a whole, after reviewing the unrelated and
widely divergent groups which have already been considered, in which the displace-
ment of the carpals and tarsals presents little or no analogy, we find in contrast a
very striking unifbi'mity in the displacement of the upper elements to the outer side,
*'. e., in the carpus, the scaphoid a i' t i c u 1 a t e s with the magnum in the
same degree that the lunar a r t i c ula tes wi t h the unciform; in the
tarsus, the astragalus invariably extends upon the cuboid. This
is the universal law of displacement which is subject to variation in degree only. It
characterizes the stem forms of every family, ami is secondarily modilied by growth
arising from special causes.

We may begin with a tyi)e in which the displacement is least extreme, and will
here consider only the Perissodactyla.

* Op. pi/., p. 205, "01>i;leii-li iliirrnurhenia nur dn-i rnllstandigt- Zi-hi-n .-im jnleni Kussc lic^il/.l, so siiul iloili Urstcn
von zwci ftniliTen, wenigHK^ns am Ilintcrfu^so vorlmmlen gfwiscn. wic ilic Cii-lcnk rucollcii iiin A^'lragiilns innl Nnvini-
larc, glcich wic am Cul>oi(lfiim iKiweiscn."

t Ziir Stivmm<>u'>-<l)ifhli- iI>t Iliillliicri'. Zool. Anzcijr.r, X.>. 210, 1885, |i. 68:1. Al»<), Bi.ilnigc /.in K<nnini«
ilrr Stiimmcsgcschiclilr il<r Hiirtliiire. .Mi>r|ili. .lalirli.. Rami XFI,

A. r. s. — VOL. xvr. 3(^

542

THE MAMMALIA OF THE UINTA FORMATION.

1. The Tapir Series.

In Isectolophus (Ko. 10,401, Piinc. Coll.), from the Uinta beds, we have a genus
which is in or near the line of the true tapirs. It is represented by the almost com-
plete carpus and a single element of the tai-sus, the cuboid. (See full previous de-
scription, also PI. X, Fig. 3.)

The carjjus. The elements of the carpus are broad relatively to their height.
The scaphoid has a horizontal trapezoid facet and slightl}' oblique magnum facet ;
thus its main weight is borne by the former bone. The lunar is lai-ge and has a
slightly larger unciform than magnum contact, but it may be said to rest equally
upon these bones. The cuneiform divides the upper surface of the unciform with the
lunar. The magnum articulates with the scaphoid, lunar, trapezoid, unciform, and
distally with the metacarpals II and III. The carpo-metacarpal articulation is of the
usual alternating order. The unciform has inferior facets for metacarpals iv and v.

The tarsus. The cuboid is subquadrate in shape when viewed from in front and
from above and proximally has a small astragalar facet.

No intermediate forms have been described. The manus and pes of the recent
Tapirus indiciis do not show a very wide divei'gence from the type above described.
The scaphoid is enlaigod posterioily and besides its trapezium facet has subequal
and horizontal tiapezoid and magnum facets. The lunar has a relatively larger
unciform facet, but still rests directly upon the magnum by a horizontal facet which
is variable in width. The cuboid is subquadrate and partly supports the astragalus.
The tarso-metatarsal articulation varies in ditierent species. In T. indicus both mts,
II and IV abut against the ectocuneiform.

/inhelonSy

DiAC. 9. — Tlif (lisplafcmonl of llie bones of the niiinus in llic Un.ffulata, II. Su» scrofa, illnslraling tlif typical
adaptive disiilaci'iiu'iit in the Aiiiodiictylii. TiUtnothvrium {MnimUis). fiont original in ll;uvanl collection; AiieJiit/ie-
rium aurelianeme, after Kowalevsky ; AphHops, fioin oriijinal in Harvaid collection. The oblique arrows indicate the
direction of dis|ilaccnient ; the double \i'rlical arrows indicate the niesaxial line of the foot.

the mammalia of the uixta fokmation". g-dj

2. The Equus Series.

The equine series is the most complete known, and is well distingulslied from
the others by the fact that the eifects of lateral displacement in the carpus, which are
well advanced in the Eocene membcn-s, although never extreme, are subsequently
counterbalanced, and practically lost sight of, in the growth of the bones in the
median line resulting from monodactylism. In the tarsus the displacement is very
limited and is not similarly retrogressive. It is surprising to note the constancy in
size and shajje of the small astragalo-cuboid articulation from the Wasatch llijraco-
theriurn to the recent horse; secondly, the early appearance of minor eijuine niMrks in
the manus and pes. The disjilacement or diplarthrism is thus limited in both feet.
The impoitant secondary characteristic is the backward rotation of the lateral ele-
ments of the carpus, tarsus and metapodials.

Caupus. The distinctive feature is the fact that the scaphoid eaily gains a
somewhat broader foothold upon the magnum than the lunar has upon the unciform.
This may be attributed to the early enlargement of mc. iir, and ectal growth of the
magnum. The lunar in the Eocene members retains subequal magnum and uiieiluiiu
facets (as in Iseclulophuf^), but the carpus is laterally compressed. Ii: ilu- more
recent forms the magnum, by its rapid growth, extends beneath the lunar, and the
latter bone is thus brought back to its primitive position and rests in some species
exclusively upon the magnum.

I/ijracothenum and Epiliiiypus. There is no marked dillerence betAveen the car-
pus of //. venticoUun and of E/)i/iippus, excepting the advanced reduction of mc. v.
In ^pifn'pjuis (Xo. 10,401, PI. XI, Fig. 4), the bones of the upper row are elongate.
The seal) ho id '•'' directed backwards and rests distally u[)on the trapezoid and
magnum, ])y sube([ual horizontal facets. The trapezium contact is very small. The
lunar has the high and slender pi-oportions seen in Triplopiix^ but ihi- ilisial facets
for the magnum and cui\eiform are subequal and incline obli(piely towards each other.
The cuneiform is narrow and slender. The trapezium was evidently small, articu-
lating with the scaphoid, trapezoid and mc. ii. The trapezoid is much Hattened. The
m agn u m supports the lunar anteriorly and posteriorly. The u n c i Co r ni has an
infero-lateral facet for the fifth metacarpal. The foot is practically tridactyl, jnc. v
being well retracted. The metacarpal articulation is of the a 1 1 c in a t i n g type.

The carpus of Mcso/iijijms has never been described, mir thai nl' any American
species of Anrhilhnrium. Referring to Kowalevsky's Memoir (I 'I. 11. I"'ig. la), wc
observe that the mag u u m extends beneath three-fourths of the lunar and the same
proportion of the scaphoid, yet the metacarpal articulation i-< unaltered, mcs. ii and

544

THE MAM:mALIA of the UINTA FORMATION.

Ill forming " alternating " lateial contact with the magnum and unciform respectively.
Kovvalevskj, however, desciibes mc. iii as spreading jjosteriorly beneath both the
trapezoid and unciform. Thus the spreading of the magnum has already oftset the
progress of the more primitive displacement. A farther step is seen in HqfjJCirion.

If De Blainville's figures are correct ((r. etjiius) the recent species, £J. caballus,
E. Burcliellii and E, asinus, present three stages in the growth of the magnum be-
neath the lunar. In the former there is a broad contact hetwocn the lunar and unci-
form, much as in Eipparion; in the latter there is none at all, the lunar resting wholly
upon the magnum, as in the pi-Imitive serial type.

/'h

eruu

Pholo.

yp

ama,r.-

A/esoh-

Anch"'

Hi

n-

i^.-

DiAG. 10.— The evolution of Uic astnigalar and calcaneal facets in the Hyi-acothcriiim-IIipparion series. . Phenaco-
dmprimaviis. Cope collection ; Ilyracolheriam ventieolum. Cope collection; PUolopJius (Oioliippus) major, Princeton
collection; P. (Triplopus) amf/ron^ra, Cope collection ; MesoJdppus, Princeton collection; A7ichithtrium aurelianense,
Princeton collection ; Uipparion meditcrraneum, Princeton collection. The upper row represents the distal view of the
astraijalus and calcaneum ; the middle row the posterior view of the astrairahis ; the lower row the anterior view of
the calcaneum. a', a", a'", octal, sustentacular and inferior astragalo-calcaneal facets, c', c", c'", ditto calcanco-
astrajralar facets. ch,fb, n. cuboidal, fibular and navicidar facets upon astragalus and calcaneum. f, fossa in the
astrasalo-navicular facet.

The reduction of the metacarpals is too familial- ground to require description

here.

Tarsus. In the tarsus we fortunately have a complete series from Hyracotherium
upwards. "We hegin with a tridactyl pes, although in the Wasatch species 77.

THK MAMMALIA OK THE UINTA FOHMATION. 545

vaUdns (Marsh), there is a rciiinaiit ol' int. i.* In the Eocene genera the separate
elements are ver}' nairow and vortifaily elougatc ; hilor the elements become broad
and spreading. A primitive and persistent feature is the triangular cuboidal facet
upon the astragalus (see Diag. 10). Another marked characteristic, early devel-
oped, is the reduction in size of the ''inferior" astragalo-calcaneal facet and its wide
separation from the " sustentacular." An importaut equine mark is the depression or
fossa in the astragalo-navicular facets ; this is a slight valley (r) in the Bridger spe-
cies of Hyracotherium, which is variable in size but constantly increasing in the
ascending series, terminating in an extensive fossa, in Equus.

There are, moreover, a number of minor tarsal features not always persistent but
of great value in diagnosis; the three astragalo-calcaneal facets (c, c", c'") are always
separate, the trochlear groove is deep and sharp. In many Eocene species the navicu-
lar is inserted anteriorly between the cuboid and astragalus and thus develoiJs a dis-
tinct calcaneal facet; this feature does not persist. In all Eocene specimens the cal-
caneum has a distinct pit for the fibula, in extreme flexion ; this is usually facetted;
it reappears in Mesolitppu^ and sometimes persists in AnrhilherinmA-

Tiic metatarsal aiticulations pass through three distinct types. In the Eocene
genera we find the "alternating" type, in the Miocene the " plane serial," replaced
later by the equine type.

Ilijracotherium venUcolum, Pliolophus and Epihippus, the Eocene genera, may be
considered together. The first species from the Wind River is distinguished by the
absence of the fossa in the astragalo-navicular facet (Cope, ''Tcrt. Verl.."' p. 045); this
fossa is more or less distinct in five small and two large astragali {P. pHmilus and
P. major), in the Princeton collection, from the overlying Bridger, and very distinct
in the P. (Tri/ilopus) amaroram o( the still more recent Washakie (Co|)eColl.).
The fibulo-calcaneal pit is present throughout, but is not invariably facetted. The
naviculo-calcaneal facet is not present in JI. venticoluiu ; it appears in P. inajor and
P. amarormii, but not apparently in P. pwniliis. The highly coinpi-cssed tridactyl
character of the foot is retained throughout. The mts. iii and iv articulate exclu-
sively with the ectocnnciform and cuboid respectively. Mt. ii articulates proxinially
with the mesocuneiform, also laterally with the culo- ami (•cldcuntifniin-i. These
relations are seen in //. renticohiin and in P. rnnjor. In /'. (unaronnn, mt. iv articu-
lates laterally with the ectocnnciform. This sjKcies is also distinguished by the
flatteninsr of the navicular and cuneiforms and relative shortening of tin- cuboid.

•Mnrah, Am. .lour. He. and Arts, Nov , 1H70, |>. 401.

f This Tacft Ls well mnrkcd in a calpaiiciim Inlely received in cxcli.ingc fron) the Pi»ri.H .Mii.iciim, nmrked .1. mire-
lianentt, from Sansnn. Soc Olag. 10.

54G THE MAMMALIA OF THE UINTA FORMATION.

Through the Eocene series, the latei-al toes are not more rolled backwards than in
H. venticolum.

In MesoMpjms, the most marked development is seen : first, in the flattening of the
cuneifoi-ms ; the entocuneiform is united with the mesocuneiform, but does not sup-
port mt. u ; second, the tarso-metatai-sal articulation is absolutely of the plane serial
type; third, the lateral digits and corresponding tarsal elements are rolled backwards
so that the outer elements are almost or quite in contact posteriorly, the cuboid
touching the entocuneiform. The fibulo-calcaneal facet is always distinct. In the
astragalus the ectal and sustentacular facets become nearly confluent; the astra-
galo-cuboidal facet is reduced to a small narrow triangle placed in front. The seven
astragali examined (Princeton (JolL, ISTo. 10,255), apparently represent two species,
in one of which, the smaller, the fossa upon the astragalo-naviculai' is faint ; the
larger species shows this fossa very distinctly (Prince. Coll., No. 10,250). This
may, however, be not a specific but an age chai-actei'.

The serial metatarsal articulation is an extremely interesting feature, when we
compare this tarsus with that of Ily racotherium and PliolopJms ; it precedes the growth
of mt. in ui)on the cuboid in AncMtherimn, an altogether difl:erent process from the
lateral displacement seen in the above Eocene forms.

It is not necessary here to go over in detail the ground made familiar b}' the
memoirs of Kowalevsky and Forsyth Major,* of the subsequent evolution of the
pes; in fact, the material from American strata for such a purpose is not at present
accessible.

3. The IIyrachyus, Triplopus, Hyracodont Series.

There is an intermingling of i-hinocerotic and equine characteristics in the teeth
and feet respectively of this series, which give it an especial interest and separate it
widely fi-om both the above stocks. It has nothing in common with the tapir series
either in the structure cf the teeth or feet, and unlike both the tapirine and equine
lines, little is known of the sub-Biidger forms, and there is no parallel European
phylum.

The general characteristics of the carpus and tarsus are the vertical elongation
and lateral compi-ession of the elements of each and the close union of the metapo-
dials in some of the later forms, which, while tridactyl, renders the foot in some
forms functionally monodactyl.

* AL-conlini; to Kowulcvsky (Anchilh., PI. II, Fig. 20A . . ), Uio octo- and mcsomiipilbrms unite in AnchUherimn,
liut ;ire distinct in Hipparion (Antlinu-iillici-., ]). 104). According to Forsyth Majm- (IJoit. z. Gcsch. d. toss. Pferde. II
Til., p. 62), the union of c-n. 1 imd 3 is tin' lulu and of cii. 2 and 'A tliu fafc exception.

THE MAMMALIA OF I'llE UINTA FOUMATIOX, 547

Carpus. In tlie tetradactyl Huraclujus we find nil the distinctive features of
the carpus in tliis series are practically attained, these resulting from the great lateral
displacement. The scaphoid has a very narrow trapezium facet and broad and
decidedly oblique trapezoid facet; it rests broadly upon the maijnura. In the
lunar the radial facet is shallow; it is compressed below, with a narrow oblique
anterior facet ujaon the magnum and broad subhorizontal unciform facet. The
scaphoid and lunar thus rest principally upon the magnum and unciform. The cune-
iform presents a small ulnar facet, this bone being rather slender. The trapezoid has
a broad trapezium facet. The magnum has a narrow and oblique lunar facet. The
unciform is a very characteristic bone since its vertical is greater ihau its transverse
diameter. It nevertheless has a narrow facet for mc. v, which, according to Cope, is
considerably the most slender.

Taken altogether the vertical and transverse diameter of the carpus is aV)out the
same.

In I'rijtlojnts the vertical diameter of the cari)us is slightly greater than the
transverse, the metapodials and carpals are somewhat rolled backwards so that the
anterior face of the manus is sti'ongly convex. Every feature in which the manus
departs from that of ILjrachyus is in the direction of greater elongation and lateral
displacement. The scaphoid is narrow and has a very oblique trapezoid facet ;
its main weight descends upon the magnum. (The small trapezium and the trapezoid
are missing in the Princeton collection.) The 1 u n a r rests entirely upon the unciform,
its contact with the magnum anterioi'ly is by a vertical facet, variable, never in the
nature of support, sometimes entirely wanting. The c u n e i f o r m, like the scajihoid,
is rotated backwards. Judging by the facets upon the mc. ir and the scaphoid, the
trapezoid was laterally compressed. The vertical diameter of the unciform exceeds
the transverse ; it still bears a considerable mc. v facet.

Ill I lijracodon, the carpus repeats the structure of TriphjjH/s upon a larger scale.
The proportions of the metapodials are approximately the same ; the proximal por-
tion of mc. V persists but is very much reduced in size. The magn u m lias ivl.i-
tively increased in size, but still has a very limited contact with the lunar anteriorly.
The trapezium is small and has no contact with mc. n. The vertical diameter of
the unciform much exceeds the transverse; the cuneilbrm is also high and vei-y nar-
row. The degree of backward rotation of the lateral elements evidently varies in
the different species.

Taiwus. In the astragalo-cuboidal articulation, ill the niimluT and disposition
of the digits and in the shape of the terminal phalanges, this series closely' approaches
the equine series, and. in some eases, can be distinguished onl}' by very careful

548

THE MAMMALIA OF THE UINTA FORMATION.

examination.* The diagnostic features of the members of the Htjracliyus series are
the relatively flat or obtuse keels of the trochlea and the frequent confluence of the
snstentacular and inferior calcaneal facets of the astragalus. The same facets are
invariably separate in the equine series, beginning with Ilyracotherium. The Hijra-
chyus calcaneum occasionally shows a pit for the fibula, but this does not bear a facet
as in nearly all the equine series.

ra.lonZ'

H/ra^chr

Trinlon-

Hyra^cod-

DiAO. 11. — The astragalar and calcaneal facets in the nyracUyus-IIyracodon series. Paloplotherium minus,
Princeton collection. IJyrachyus (sp. inilet.), Princeton collection. Triplopus ohliquidena, Princeton collection. Hyra-
codoii major, Princeton collection.

Hyrachyus (No. 10,199, Princeton Coll.). The cuboidal contact with the
astragalus expands j)osteriorly, but is very narrow anteriorly in some species
{H. j)rinceps\ bringing the navicular into contact with the calcaneum precisely as in
some species of Ilyracotherium {Triplojms amarm'um) ; as above stated there are

* I liavc referred the Triplopus amarorum of Cope, from the Washakie beds (uppermost Bridger), to the eijuine
series.

THE MAMMALIA OF THE UINTA FORMATION. .'349

but two calcaneal fticets, the Inferior and susteiftacular being confluent ; the variants
in these facets, seen In- comparing a large number of astragah, arc found to be :
(1) The greater or less union of tliesc fixcets, looking towards a more primitive Ibrin
in which they were probably separate ; (2) the presence of a small ridge, r, at the point
of junction, looking towards the well-developed ridge in the later forms (//^racWo/i).
The c a 1 c a n e u m presents a very long slender tuber, three astragalar facets and an
extremely narrow and deep distal cuboidal facet. The cuboid is corresjwndingly
narrow and elongate, in the small species compressed in the middle, and with no trace
of a facet for mt, v and little or no contact inferiorly with mt. iir. The ectocu-
n e i f o r m is elevated and has a lateral facet for mt. ir. The m e s o c u n e i f o r m
is small : according to Cope, "the en t o c u ne i for m is large, flat and posterior
in position." *

TrijAojms (Xo. 10,397, Prince. Coll.). Although the pes as a whole is very nar-
row and slcndei', the trochlea is much shallower than in the equine series (Diag. 11,
Plate XI, Fig. i>); the cuboidal contact narrows anteiiorly to a point, but the navicu-
lar is excluded from contact with the calcancum; the inferior and susteutaeular
facets are cither confluent or closely api)io.\imatcd and vary in the development
of the ridge (r). Tlie calcaneum has a narrow and oblique cuboidal facet, a
long latei'ally compressed tuber and narrow sustentaculum ; there is no trace of a
fibular facet. The cuboid is compressed in the middle as in Jli/racJu/us, elongate
and with a relatively small distal facet for mt. iv, indicating that this digit was
niiicli reduced. The navicular is very deej) and laterally com])re.sse(l. The meso-
cuneiform and cntocuneiforin are wanting in the Princeton collection. The ccto-
cunciforra articulates with mt. hi only, indicating that the metapodial aiticulation
was absolutely plane; the lateral toes were rotated backwards precisely as in Anchi-
iherium.

IIijraco(h>n\ (No. 10,31)7, Princeton Coll.). lu this genus we liiid an e.\aet rejie-
tition of the tarsus of Ilyrachijiis in considerably heavier form than in IVi/iIopits, and
with few modifications. The a s t r ag a 1 u s has the same limited cuboidal contact ;
the three calcaneal facets arc nearly confluent, the "sustentaiMilar" and '"iuferior"
are fidly so and a sharp ridge at their point of junction interlocks will: a corresiujud-

• Tertiary Vcrtebmta, p. '>,59 ; U. niminii.

f On the Skull <) tlif Eofx-nt; I{liliiori;ro», Orth loynodon, siiul tlic Kcliition of llii>» Qfnim to ollirr Urcmhcrs nf tin'
Group. £. M. Museum Bull. No. 3, 1833, p. 17, "llymrodon was a sli'ndvr, lonK-1>>nl><'<l and Hlj^'litly Imili aniinat
Willi a long ni;<k and delicate hi'iul. Its prnportionii were IIiohc of a llorsr rallicr than of a Kliiiiori-rog." p. 20, " Fir
short, it wa.-< a fursoriul llhinn<;i!ro.«, and all il.i mndiflmlions went towards n<l:iptin[; it lo nwifl iMconiiilinn • • •
One can hardly help believing that had this line |H!rsLsti-d, it would have resulted in a iinidigitui type, junt ax the triduc
lyl Anfhilfurium of the Mioeene has tcnniiintcd in the Horse." — W. B. Scott.
A. P. S. — VOL. XVI. '.hi.

550 THE MAMMALIA OF THE UINTA FORMATIOIn.

ing depression in the calcaneum. Tlie navicular is shallower than in Triplopus,
resembling closely that of Hyracltyus, and showing facets for three cuneifoi-ras.
The cuboid is elongate and compressed in the centre as in the above forms, and
articulates distally with mt. iv only. The e c t o cunei f or m has a lateral facet
infei-iorly for mt. ii (unlike Triploims), thus reverting to Hurachyus. The m e s o-
c u n e i f o r m is correspondingly shortened, while the e n t o c u n e i f o r m is flattened
and i-otated backwards as in Hyrachyus (IsTo. 10,005, Princeton Coll.).

In the pes of Pdlophtlierium we have a striking resemblance to that of Triplo-
2ms, and of the Eocene horses except for the greater reduction of the lateral toes.
This is simply an example of parallelism, for the teeth* show no affinity to eithei- of
these lines. The lateral digits are as far reduced as those of HijJjiarion.f

4. The Rhinoceros Series.

A. — Acer cither mm. B. — Aplieloj^s.

The comparison of the carpus and tarsus in the hornless rhinoceroses of the
American Miocene shows a wide divergence between the Aceratlierium (lowermost
Miocene) and Aphelops (uppermost Miocene) series. The Acerntherium feet are ex-
tremely elongate, with the toes closely compressed, reminding one strongly of the
feet of the pseudo-i'hinoceros, Hyracodon, from the same beds. The more recent
Aplielops is generally considered a successor, but shows a widely different type of

*Schloss(;i- (Beitriige z. Stamraesg. d. Huft.) points out the anomalous mixture of primitive and secondarj- char-
acters in tliis genus. Tlie premolars are reduced iu number, f, but simpler than the molars, while the molar crowns
are more advanced than those of Palicotherium, showing cement.

f Lydekker (Cat. Foss. Mamm., Part IV, p. K!) follows Flower in uniting PuloploiJieritim minus and PaliBotherium
(see Cat. Roy. Coll. Surg., Part II, Mammalia, p. 390). The ground assigned is the presence of a " complete transi-
tion between the two types." On precisely similar principles we should unite Eyracotherium venikolum with Mesohip-
pus ; nay more, because the American genera undoubtedly belong to one line of descent, whereas the two French
genera, judging by the extremities and teeth, belong to different lines. Further, is nomenclature actually simplified
when a genus is made so comprehensive as to extend over long geological and wide geographical areas, embracing
such a variety of forms that it becomes necessary (vide Rhinoceros and Palaoiherium, opera cilata) to arrange the spe-
cies into groups? We think not. Genera can no longer be defined in the Linnajan sense. Lamarck, a century ago,
anticipated the difticulties which would arise when the gaps between Cuvier'sand Button's genera were filled up. The
range of variation which should be included in a single genus is largely a matter of arbitrary judgment. Never has
the problem been more perplexing than among the unbroken series found in the western Tertiaries. The writers, for
tlie present, have adopted the rule long since employed by Marsh and Cope, viz , diflorences of degree, size, propor-
tion, may be talven as specific ; dill'erencos of kind (in the number of teeth or digits) or of actual form («. g., tlie as-
sumption of the molar pattern by the premolars), may I)e taken as generic. This rule, when applied, for example, to
the horse series, is found to work admirably — each of the subdivisions of the Eocene is characterized by a genus — and
by a number of species varying in the direction of the lower and higher forms. Judging by the thickness of the
strata, these subdivisions represent long periods of linu^

THE MAMMALIA OF IIIK UINI'A KORMATIOX.

551

foot, which is hiijhl y " modernized.'" The relations of these genera to the European
forms is somewhat nncertain, so that the following observations apjjly especially to
the above American jronera.

yimynod-

Aenilh"-

J/i/wi

/-

i '/>

DiAO. 13.— The astrag.ilar and calcaneal facets in Amyaodon, Aeernlhtrium and Aplielops. All species indcler-
minate.

A. — AcEiJATUEUiUM (Coll. Mus. Comp. Zoil,). The general characteristics of
the feet arc lateral compression and vertical elongation of the carpal ;iml tarsal ele-
ments and the elongate character of the metapodials. The caiyus shows the same
degree of lateral displacement seen in the tapir. The scai)hoid is subqnadrate,
resting by a broad and slightly oblique facet upon the trapezoid. The lunar has a
broad unciform facet, but retains a small horizontal ficet u|)on the uncilbrm. The
cuneiform extends widely backwards. In the second row, the trapezoid supports mc.
II ; the magnum is a stout bone, subqiuidrate anteriorlv, projecting beneath the lunar
posteriorly with the incipient stout pivotal peg, which is so largely developed in
Aj)hdops. The third metacarpal is less enlarged lli.in in Ap/iclupa ; the second and

52 TH1-: MAMMALIA OF THE UIXTA FORMATION.

fourth are slightly smalioi- ; the unciform facet indicates that me. v (missing in this
collection) had ahout the same relative size as in the tapir.

The tars'us. The astragalus rests by a rather nai-row facet upon the cuboid ;
this facet is equally broad posterioi'ly and anteriorly, as contrasted with the Hyra-
codon and equine series on the one hand and the Palicosyops series on the other. The
arrangement of the calcaneal facets is shown in Diag. 12. The calcaneum has a long
slender tuber expanding into a knob. The cuboid has its vertical diameter esceed-
ing the ti'ansverse, unlike all the more recent rhinoceroses. The articulation of the
third and fouith metatarsals with the ectocuneiform and cuboid is of the " plane
serial" type, but mt. ii abuts against the ectocuneiform.

B. — In Ajyhelops the carpus and tarsus are very bi-oad and spreading and show
the extreme of diplarthrism. In both, the metapodial articulation is of the primitive
" altei-nating '' type. The toes sjii-ead widely, being reduced to three in each foot.

Carpus (see Diagram 8). The scaphoid is very broad, covering the whole ante-
rior surface of the magnum, while the lunar rests anterioi'ly upon the unciform alone,
posteriorly upon the pivotal jDrocess of the magnum. A remarkable feature of this
bone is its unique articulation with the magnum hook posteriorly. The great devel-
opment of the middle toe and relatively increased size of the magnum, renders the
scapho-magnum surface broader than the lunar-unciform. Facets for the trapezium
are seen ui)on mc. ii and the trapezoid. The trapezoid and magnum facets of the
scaphoid are subequal and hoi'izontal, directly transmitting the weight. Mc. ii abuts
proximally against the magnum; mc. iii is much larger than the lateral metacarpals
and has a very broad unciform facet. There is no trace of a facet for mc. v.

This type of carpus is precisely similar to that of R. unicornis and Ii. hioornis,
as figured by De Blaiuville.

Tarsus. The astragalus is broad and low with a shallow trochlea, and broad
cuboidal facet, which is as large as that upon the calcaneum ; a characteristic feature
is the ental extension beyond the three distinct calcaneal facets. The calcaneum is
chai-acterized by a short tuber caleis and very slender sustentaculum; the three astra-
galar facets are sepai'ate and rather small. The transverse diameter of the cuboid is
greater than the vertical. The mesocuneiform is much reduced. The mts. ii and iv
have lateral fiicets upon the ectocuneiform and cuboid respectively of the "alternat-
ing" type.

This type of tarsus is found in H. unicornis and M. hicornis, which are, however,
less broad and spreading. These recent genera differ from Aphelups and resemble
Acerat/wrium in the mode of metapodial articulation, the mt. iii not possessing a
cuboidal facet. In fact the cuboidal facet upon mt. iii in Aplielops is apparently unique.

the mammalia of the uinta formation. 563

5. The Paljeosyops, Diplacodon, Titanotherium Series.

The general characteristics of the feet in this series arc the following: The car[)iis
is broad and llat, and the lateral displacement is limited; there arc four spreading and
fully functional toes. In the pes on the other hand the cuboid extends rapidly
beneath the astragalus, the lateral displacement being very rapid. The calcaneura
lias a fibular facet. The terminal phalanges arc flat and spreading distally.

Carpus. In each of the carpal elements in the genera of this series we invari-
ably find (excepting in tlie hinai- of some of the Bridger species) that the transverse
diameter exceeds the vertical, in a ratio increasing as Ave ascend. The lunar can
always be recognized by its nearly subequal magnum and unciform facets, and the
unciform by its broad facet for mc. \.

Pakfosyops (No. 10,013, Princeton Coll.). There are two distinct types of
feet which may be here referred to Palceosi/ops, but cannot with certainty be deter-
mined specifically. One belongs to a large and heavy form with short, stout digits
(possibly 7^ major Leidy, or P. validens Cope), the other to a lighter, more graceful
animal witli long, rather slender digits (possibly P. pahidosus Leidy). There is a
third type, intermediate in size, between the above, but showing the heavy propor-
tions of the large type. The scaphoid has subequal magnum and trapezoid
facets, and, in its somewhat oblique position, transmits its weight equally to both
elements. The lunar has a dee]) radial facet; it is wedge-shaped inforiorly ; the
direction of its magnum and unciform facet varies in the two types from a sharjier
to a more obtuse angle ; thus, in the lighter forms the magnum facet is small and
strongly inclined, Avhile in the heavy forms the same facet is larger and more horizon-
tal; corresponding variations are found in the superior facets of the magnum;
taken together they determine the degree of transmission of the weight through the
magnum and uncifoim I'cspcctively. The t r a p e z i u m sustains no direct pressure,
but ailiculates with the scaphoid, trapezoid and mc. ii. The trapezoid receives
half the weight of the scaphoid. The u nci form is \ei\ broad, resting inferiorly
upon the mc. ni-v. The variation^ in the metapodials between the heavy and slender
types are stiongly marked.

Diplacot/iiit. The scaphoid, lunar and cuneiform of a carpus have been collected
from the Uinta which agree so closely with the corresponding elements in the slender
type of Pahiiosijops above described, that they require no further description. They
belong either to a small sjiecics of Z'/yj/rtcoc/w^, much smaller lliaii the iKdalnsoi
Marsh, or to a persistent species of Pnhwsijops. The only departure from the above

554

TUB MAMMALIA OF THE UINTA FORMATION.

type is in the slightly increased massiveness, which might well represent individual
variation.

The metapodials of a right manus of Diplacodon are well preserved ; they are
long, relatively slender, and spreading, interlocking closely proximally and keeled
upon the posterior distal faces.

Titanotlierium (Coll. Mus. Comp. Zool. of Cambridge). The growth of the lunar
and magnum are the distinctive features of this cai'pus. The lunar presents two
distal subequal facets for the magnum and unciform respectively. The c u n e i f o r m
is relatively somewhat reduced, while the scaphoid is enlarged and rests by two
nearly horizontal facets upon the trapezoid and magnum.

Blb£OS.-

Biji/di

Tltanoth'.

DiAO. 13. — Tlie astrngalar and calcantnl facets in the Pala-osyops TilanoUifciuim series. Falceosijoiis major, Prince-
ton collection ; Diplncodoii ? elatus, Princeton collection ; Titanotli,erium (sp. indct.), Princeton collection.

Tarsus. The sjieclal characteristics of the tarsus, in addition to those above
mentioned, are the wider separation of the three astragalo-calcaneal facets and reduc-
tion of the inferior facet ; the astragalo-cuboid facet is an elongated triangle, broad
in front, narrowing posteriorly ; the astragalar trochlea extends backwards upon the

THE MAMMALIA OF THE UINTA FORMATION. 555

calcaneuin so that the tibia articulates with the latter in Diplacodoa and Tllanolhe-
ritun. The fibiilo-calcaneal facet in Pala'osyops increases to a considerable size in
Titanothevlum. The cuboid and other elements of the second row of tarsals are
rather l)road. The metatarsal articulation is secondarily of the alternating type.

Pahvosyops. The astragalo-calcaneal facets together present a broad surface for
the cuboid and navicular, between the bones is a gap which is bridged b}' the cuboid.
The astragalo-cuboid facet is slightly nari-ower behind than in front. The c a 1 c a -
n e u m has a small fibular facet, and distally presents a broad oval face for the cuboid.
The cuboid is subquadrate in proportion, much as in the tapir sei'ies, and presents a
sharply defined astragalar facet ; distally it rests upon mt. iv, and presents a variable
facet anteriorly for mt. in. The ectocuneiform is also subquadrate anteriorly and not
only suppoi'ts mt. iii, but has a broad facet laterally for mt. ii. The mesocuneifoim is
thus somewhat llattencd. The entocuneiform is not preserved in the Princeton
collection.

Dijdacodon. The tarsus (Plate YIII ) is represented merely by an astragalus
and calcancum which are somewhat crushed, but nevertheless present impoitant
transition characters. Thus the filnilo-calcaneal facet is much larger ; the tibia ex-
tends from the astragalar troehUa upon the calcancum ; the tuber calcis is very long
and laterally compressed. In the astragalus the trochlea is dee}); the three cal-
cancum facets are nnu h reduced ; the cuboidal facet is broader anteriorly and nar-
rower posterioily. There is thus an intermingling of Pahvosijops and TUanothen'um
characteristics. Viewed together, however, the.se bones are elevated and laterally
compressed, indicating a high and rather narrow pes. With the metacarpals above
described, we may infer that Diplacodon had gained greatly in height upon Pakfio-
st/ops without having acquired any of the massive proportions of TlUtiiotlierium.

Titfinotheyiinn. The tarsus is veiy broad and fiat, tlie metapodials spreading
widely. lathe astragalus the trochlea is shallow, the inferior calcaneal facet is
greatly reduced and is confluent witli the cuboidal facet; the latter is broad in front
and tapers to a point posteriorly. The gap between the astragalus and calcancum,
observed in Ptdipoyiyops, is here greatly enlarged, the cuboidal contact with both as-
tragalus and calcancum is intcirupted by slight surface depressions, analogous to tlie
"navicular fossa" in the horse. The tibia and Hl)ula both have calcaneal facets. By
the broadening of the cuboid the navicular is greatly rcdiu-ed. 'i'lic cuneiforms are
ttattencd. Mt. in has a bioad contact with the cuboid, ami ml. ii with the ecto-
cuncilbrm.

556 THE MAMMALIA OF THE UINTA FOUMATIOlsr.

In a general review of the above series, we observe practically the same type of
intercarpal articulations in all tetradact}'! and tridaetyl forms irrespective of the geo-
logical 23eriod to which they belong. Thus the tapir manus is a persistent example
of a type developed independently, at different periods, in Hyracotheritim, Hyracliyus,
Acerathermm and Palmosyops. The tridactjd rhinoceros, with Tr/plopus, Hyraco-
don and PalcwtTierium present another type. The significance of this parallelism is
discussed in another section.

In the tarsus we find a striking analogy between the equine and Hyracodon
series. In another series may be grouped all the remaining Perissodactyla.

Thei-e is actual disproof of Kowalevsky's generalization as to the bearing
of the spreading of the metapodials upon the survival of the fittest, iu the non-
spreading of the metapodials at an important stage in the development of the horse
series (see p. 546). In fact we can trace no connection between certain foot struc-
tures and survival in perissodactyl phyla. We must either with Schlosser (^op. cit., p.
121) regard the inadaptation of the teeth to the changes in the character of the food
as the cause of extinction, or rather look at the organization as a whole in its rela-
tions to the changes both in food and to the rapid increase of Carnivora and com-
peting Herbivora.

IV. THE CLASSIFICATION OF THE UNGULATES.

This detailed comparison certainly supports in a measure Riitimeyer's* designa-
tion of the foot bones as a " verschiebare mosaik," and raises the question as to how
far these articulations can be employed in classification. With all the direct con-
tradictions in Cope's system urged by Riitimeyer and others, which are here pointed
out, tliore still remains the universal principle that in every step of transition from
the primitive plantigrade foot towards digitigi-adism, some form of displacement
from the serial order, or interlocking of i)arts, took place. This proceeded not at
random, but along certain definite lines, conforming in general to the succession in
geological time and large existing subdivisions of the Ungulata, and of great value
as expressing their relations to each other and to the primitive stock. "VVe cannot,
therefore, as the distinguished Swiss palaiontologist advises us (p. 18), rest content
with the lines drawn by Cuvier and Owen, because the derivation from the plaui igradc
taxeopod ungulate was not recognized in their system, and the underlying principle
of Cope's system, since adopted by Marsh, and many of the younger school of palaa-
ontologists, rests upon a secure basis of fact. The real problem is, therefore, to ex-

* Ut'bcr ciiiiiii' Bczicluiiiiii'ii '/.. d. SiiiiiritliitTs'.aiiimrn ullrr ii ncucr Wult. Zuricli. 18S8.

TllK MAMMALIA OK THE UINTA FORMATION. Tj-l^

cluck' tlic variable elements, and asceitaiii those which are constant in the ordinal
divisions ; of course, no sharp lines of division, such as wci-e apparent to the oldei-
naturalists, exist as we descend in the mammalian series.

Fii*8t: We observe that while the metapodials play an impoi-tant part in the dis-
jMacement, the separate metapodial articulations are not available
for taxonomic purposes in the larger divisions above the Protungulata. In
the carpus there arc practically but two types of articulation, the "seiial" in the
absolutel}' primitive and in some later foi-ms, and the "altornatinp;'' in all but the
highly speciaii/ed "adaptively reduced" Aitiodacty la. There are no stages which
are of more than family value.* In the tarsus the primitive serial type is i-etained
almost nnmodilied in the Proboscidia and Hyracoidea, and in the early equine and
Hyrachyus series it passes into the "plane serial," and into the alternating in the
Paheosyops series.f The I'emaining Diplarthra show great variation in the mode of
metatarsal articulation, so that there is no single type characteiistic of all the genera
of the Tapirkhv, or Hhinocerotidai ; Aphelop>i, Acei'athenum, Atelodus and Wiinoceros,
each has a different type of metatarsal ai-ticulation.

Second: In the tarsus Prof. Cope's definitions and principles
will not strictly apply, first, because the astragalus extends over the cuboid
in some of the earliest and not in some of the more recent Mammalia, and this dis-
placement is thus by no means distinctive as between the lower and higher types.
The astragalus extends upon the cuboid in one or more members of the " taxeopod "
Hyracoidea (Dendrohi/rax) and Condyhwtha (Perijdi/chns), nud in all the Amhly-
poda, to a much greater extent than in many of the diplart-hrous Perissodactyla (sec
the Equus, Hyrachyus, Paloplotherium series, p. 548).

As Kiitimeyer points out, thei'c is no parallelism between the diplarthrism oltlic
fore and hind feet — diplarthry may be rapidly developed in one while taxcopody per-
sists in the other. This arises from the fact that the mechanical problems of adapta-
tion to the unguligrade position are dissimilar in the fore and hind feet owing to the
ditl'ercnce both in the arrangement of the l)ones and in the motions. Among the Hy-
racoidea, Ilijrax is a taxeopod in both feet, while Dendrolnjra.v has a taxeopod manus
and diplarthrous pes. In tlie Proboscidia the manus is nearly serial, somelimcH
completely so, while the pes is invariably di|)larthrous, although well distinguished

• Kowiiluvftky'it "(utiptivi; rt'fliiction " nrlsoa intlfpcndcully In llie IJufioilnntii miil ScloiiiKlDnlii.

\ There are some other mnrkcd lllii<<lmtlrin» of ihi' vuriiihln nntiirc of the poiliiil nrilmlrttlons. Thi; conMucrKT of
one or Uie other of the three a8tra);.t1o-eak-anea1 facets is sul)jcct lo speciflc viirintioiis ; rompnri' R. incititui ami /?. Ifp-
torhinui (De Blninvillc, Ostco!»r.). Company the aslmgalo-eulioiilal fai'els mi Tiipirut amiruanui m\i\ T. indicuM. The
lunar mngntim fiicel ilifTers widely in Etpiui hiirehtlii, tab'iUii* iinil atiniit.

A. r. s — VOL. XVI. 3s.

558 THE MAMMALIA OF THE UINTA FORMATION.

by the movement of the navicular above the cuboid. Cope has himself recognized
the difficulty arising in the Amblypoda from the union of limited diplarthrism in
the carpus with extreme diplarthrism in the tarsus. In Macrauclienia we find a pecu-
liar displacement in the carpus and no asti-agalo-cuboidal contact in the tarsus.

The fibulo-calcancal focet may be regarded as a secondary character, since it is
developed independently in so many phyla — as a trochlear joint in Memscotheriwn,
Macrauclienia. the Artiodactyla ; as a supporting facet in the Proboscidia, Titano-
theridaa and Hyracotheridje. It cannot be employed in ordinal classification [vs.
Cope, Tert. Vert., p. 378).

Third: The definitions based upon the carpal series cannot
be availed of strictly as employed b}' Cope.* For (1 ) in some
of the Amblypoda (Uintatherium) the scaphoid has a broad postei'ior facet upon the
magnum, although of less extent than the lunai' unciform facet. (2) In the Probos-
cidia the lunar frequently rests upon the trapezoid, and sometimes upon both the
trapezoid and unciform. (3) In Maerauchenia the lunar is entirely excluded from
the unciform, the cuneiform resting upon the magnum.

A comjDarison of the entire evolution of the carpus and of the tarsus shows that the
former, while moi'e complex, is subject to more definite laws, and is therefore more
available for purposes of classification. The carpal characters are constant and dis-
tinctive, but not exactly as interpreted by Prof. Copef — we must look at the manus
as a whole, including the metacarpals. The tarsal displacements can be employed for
subsidiary definition only. It has thus been shown that Cope's whole system breaks
down if Ave attempt to establish a parallel between the evolution of the cai-pus and
the tarsus in relation to displacement.

"With these limitations the })rinciple of classification of the ungulates by foot
structure loses the universal application claimed for it by Cope, but is by no
means invalidated to the extent held by Riitimeyer.J Giving the first rank to the
characters of tlie manus, and second to those of the pes, the folloAving modifi-

*'lVitiary \'cilclii-.il;i, p. 37G.

f Tfrliary Vcrtelirnta, p. 377. "In Uie first place I find the diversity in the structure of the carpus to be greater
in the relations of the magnum and scaphoidus than in the relations between the unciform and the lunar. In other
words the trapezoides and magnum are more variable in their proportions than is the cuneiform. » * » I therefore
view the relations of these bones as more characteristic." In the succeeding section, V, this view is .'-liown lobe
untenable.

t Op ciL, p. 18.

TIIK MAMMALIA OF THE UINTA FOKMATION.

550

cation of Cope's table is
for brevity :

A. — a. Carpals serial.
I. Taxkopkda.

1. Priinilive. Pro t u ii gul a ta.

c. Plantigrade.
e. Tarsals serial.

b. Metapodials serial.

2. I n t c r m e il i a t e. Condylar

thra.

c. Sulidigitigrade.
«. Tarsals serial.

b. Metai)odiBls displaced.

3. Recent. Ilyracoidea.

c. Plantigrade.

e. Tarsals serial or disi>laccd.
b. Metapodials displaced.

l)i'oposed ; several subordiiial characters being omitted

B. — a. Displacement of first upon sec-
ond rote of carpals not uniform.

b. Metapodials displaced.
n. Amblypoda.

c. Semi-planlisrade, plantigrade.

d. Lunar upon cuneil'orm.
«. Astragalus upon cuboid.

lir. PitOBOSCIDrA.

e. Seuii-planligrade.

d. Carpals serial, or lunar upon

Inipezoid.

e. Astragalus and na%'icular upon

cuboid.

IV. (MACnAUCHESIA.)

c. Digit igriide.

d. Scaphoid mid unciform upon

magnum.

e. Tarsals serial.

C. — a. Displacement of first upon second
row of carpals uniform.*
b. Metapodials displaced,
e. Digili'jrade to unguligrade.
e. Tarsals more or less displaced.

V. Dll'LAItrUKA.

1. Pe rissod act y la.
/. Mcsaxonic.
e. Carpal disidaccnient extreme in

reduced forms.
e. .Vstragalo-cuhoidal lacet Hat.
3. A r t i o d a <U y 1 a.
/. Paraxon i(;.
a. Carpal displacement limited in

reduced Ibrms.
e. Astragalo cuboidal facet gingly-
moid.

The position of the To.vodontia and of Macrauchenia is doubtful.

The three subdivisions above employed (A, B, C) bring out what appeals to l)e
a fundamental distinction between Orders IF, III, IV and V. The significance of
the diverse carpal displacement in the Artiodaetyla and Perissodactyla is shown in
the next section.

V. T[IK TRINCIPLES OF DTSPLACEMKNT.

The evolution of llic luamis and pes ol' the iiiiL-iilatL's inrliidLMl llic I'uliouiiiir
proces.ses : (I) Elevation fioin tiu' [)lantigrade to the digitigrade position ; (2)
growth of certain tlcments and icd ii e t i o ii of otiiers, including the loss of lat-
eral parts ; (3) d i - p 1 a c c iiic ii I of I lie elements of the podium and inefapodium from
the primitive .serial arrangement; (4) coalescence of parts primitively distinct.
All these processes interacted upon each other, and so far as they can be distinguished,
took place, for the most part, in the order named. That is, elevation usually preceded
the metatroj)hic or unequal growth of parts and the elongation or vertical growth of
the ineta|todi;il elements involved the mutual displacement or iiiterloeking of the
podials and metapodials. The etlects ofthe.se proces.ses were in the nature of adapta-
tions correlated with changes in the entire body which were induced by the habitat,
manner of feeding, character of the ground, necessity of speed liotli in ranging for

*Scc this Memoir, p. .">ll. Tlic scaplio-magnum = lunar-cuneiform fiicet.

560 THE MAMMALIA OF THE UINTA FORMATION.

food and escaping enemies. Kowalevsky* and Riitinieyerf have treated these factors
very fully. Then there were the changes in the movement of the limbs with refer-
ence to the axis of the body ; for example, in the evolution of the horses Kowalevsky
has directed attention to the atrophy of muscles which rotated the fore limb, resulting
finally in a diiect fore and aft motion. Another factor was the transmission of the
weight unequally through the bones of the lower leg, as one or other of these ele-
ments o-rew or became reduced. Finally there were the influences which the reduc-
tion of the lateral digits in the artiodactyl and perissodactyl series exerted upon the
central digits and upon the podial elements by modifying the principal lines of "im-
pact and strain." '

The Manus.

It has been shown that the modifications of the manus are more complex and yet
proceed along more definite lines than those in the pes, and in spite of the great
variety of influences exerted we can discover certain fixed laws of modification.

1. Relations of growth, reduction and displacemeyit. As displacement changes
the vertical relations of the three rows of podial elements to each other, it follows
that there must be concomitant growth in some elements and reduction in others,
otherwise some of the lateral parts would be left without support. That the dis-
placement of an element, however, involves a principle distinct from mere growth
and is an actual shifting or "rotation" process, although not to the degree main-,
tained by Cope,J is shown by a number of facts. Pii-st, as to the chief factors in
each process. Growth is more directly brought about by vertical pressure, as seen in
the niaguum of the Equidte ; and displacement, by lateral strain, as seen in the shift-
ing of the metapodials to the ectal side of the carpus. Displacement is apparently
arrested in such a series as Ilyracotherium, where the fore limb acquired an orthal
movement and, the entire vertical pressure passing through the median toe, the
growth of the magnum counteracted the incipient displacement of the lunar upon the
unciform. We cannot explain the lateral facets of metacarpals ii and lu in the pen-
tadactyl Amblyjjoda and Proboscidia as due merely to growth of the elements dis-
placed, for the digits are strikingly equal in size ; they both extend to the ectal side,
and what mc. iii gains upon the unciform it apparently surrenders to mc. ii upon the
magnum. Finally, there is every reason to agree with Kowalevsky that the reduc-

*Monogr. der G. Aiitlirucotbcruim, Paltuontograpliica, Bil. XXII, 1S7:\ jip. KU-T).

t Op. cit., pp. 17-34.

}Thc PcrissoiUiclyla. AnuTicaii Naliiralist, 1887, p. ',)8G.

TIIK MAMMALIA OF THE LINT A FORMATION. 5(51

tion of the lateral tlij^its in the ungulate foot is largely the direct result of elevation
to the uuguligrade position which rendered useless the shorter lateral digits of the
plantigrade foot. Yet every step in reduction of the toes inlUienced the growth and
displacement of the more proximal elements.

It can be demonsti-ated that these three processes were concomitant, and while
resulting directly or indirectly from the same causes, wei-e not all'ected in the same
degree by them. The problem may be expressed in this way : What were the prin-
cipal factors in each of the above processes, and how did these processes interact.

2. Theories of modijicaiion. As Kowalevsky was unaware of the primitive
serial arrangement of the podial elements, his studies were chiefly directed to the
phenomena of growth in the metapodials and the manner in which in the adaptive
and inadaptive artiodactyls and in the perissodactyls the second and third metacar-
pals spread from their respective carpals to the adjacent elements.*

Kyder}- latei- pointed out the ettect of impacts and strains upon (he phenomena
of reduction of the digits.

CopcJ the first to recogni/.e the common development of the ungulate foot from
a primitive serial type, was also the first to advance a "displacement" theory: " As
the foot is descending towards the ground it is with the distal part of the leg rotated
from within outwards. The rotation of the foot is promptly arrested at the moment
of its contact with the ground, and the efiect of this arrest is to produce a torsion of
the leg, and a pressure from within outwards of the proximal or moving element of
each articulation against the distal or fixed element."

An adequate theory for the displacement phenomena must, however, explain the
shifting of the metacarpals and metatarsals, as well as that which takes place between
the two rows of carpals and tarsals. The latter always takes place (excepting in (he
Proboscidia) in the same direction. I'lof. Cope's theory has thei'cfore to meet the ap-
parently fatal objection that the arrest of rotation to the ectal side would be first felt
at the junction of the metapodials and podials, and if it were o[' tiie character he de-
scribes, would coiUinually tend to separate these joints and displace them (o the eiital
side, whereas v/e find them provided with facets to resist pressure towards the ectal
side.

It is a significant fact, recently pointed out by Weithofer,!^ that in all ungulates
in which the radius is larger than (he uln.i, the proximal row of cari)als is displaced

• Monog. dcr Gait. AntbraroUieriiiin, IHTH, p. 142.

fOnUie Laws ot" Diirital liccliirlinn. Aniericnn Nnliirali^l, 1H77, p. *W^.

tThc PerisaoOurlylii. American Niiluralist, IH-ST. p. 980.

§ Einigc BcmcrkiingcD iibcr (IcD Carpus dur Probosc'iOicr. K Aiil. WcillinliT, Murpli. .lulirl"., 1888, p. 508.

562 THE MAMMALIA OF THE UINTA FOKMATION.

to the ectal or ulnar side of the distal i-ow ; in the one order (Proboscidia) in which
the ulna is the larger bone, the proximal row is moved, if at all, to the ental or radial

side.

3. Relation of the size of the radius and ulna to displacement. It thus appears
probable that the transmission of the main body m"- e i g h t through
the ectal or ental side has caused displacement between
the first and second row of podial e 1 e m e n t s r e s po ct i vel y t o
the ental or ectal side. But this is clearly not the only ftictor ; if it were,
this displacement should be proportional in the Diplarthra to the enlargement of the
tibia and radius respectively ; this is certainly not the case either in the carpus or tarsus.

In the early horse series (II>jracotJienum), in which the radius is enlarged and
the ulna reduced more rapidly than in any other, the ectal displacement of the lunar
is not so uniform nor so rapid as in the Rhinoceros series in which these bones retain
approximately their primitive proportions. Compare the Triplopus carpus, with the
scaphoid resting chiefly upon the magnum and the lunar resting wholly upon the
unciform, anteriorly, as a type of extreme displacement, with that of Hyracotheriam
or Epihippus, in which the displacement is slight. The latter presents much less
divergence from the taxeopod type, although the ulna is proportionately more reduced.
"We meet still gi'eater difficulties in the pes, where, as seen by the contrasts in the
members of the Equus, Hyracodon and Rhhioceros series, the extension of the astrag-
alus upon the cuboid is seen to be entii'ely independent of tibial enlargement. Nor
is this extension a resultant of weight, as proven by the diversity of the cubo-
astragalar articulation in Dendrohyrax and liyrax. We reach the conclusion that
while the di i-ec t i on of the displacement may be determined by
the trans m i s s i o n o f the m a i n weight through the ental or
ectal side, the degree of displacement is not proportionate
to this enlargement, but is modified by other causes.

4. Growth and reduction take place by metatrophism, the separate elements of
the foot being hypertrophied or atrophied by increased oi- diminished use, resulting
in a transfer of nutrition, one element growing at the expense of another. We have
to deal with the adaptations to both vertical pressui'e and lateral strain, since the
motion of the limbs, of the foi'c limb especiallv, is not in a perfectly direct line. It
may be assumed that the parts grow most rapidly which are subjected to the maxi-
mum resultant of these vertical and oblique forces. As a strong confirmation of the
theoi'y of displacement here advanced, it may be mentioned that from a study of the
resistant facets the writer reached independently the same conclusions as to the mo-
tion of the fore feet as those drawn from the study of instantaneous photographs.

THE MAMMAF.IA OP THE UIN'TA FORMATI()>f. 561}

5. We are indebted to Prof. Harrison Allen's* studies of the eelebi'ated Muv-

«

bridge i)hotograiAs for the following laws of quadrupedal motion : As the fore foot
descends it passes under the body (p. 51), i.e., inwards; striking the ground, it
arrests the torsion of the trunk as the shoulder is turning forwards towards the oppo-
site side (p. 57) ; it strikes upon the outer border (p. 8S) and leaves by the inner
border (p. 50), so that the pressure of the body is boine from without inwards across
the foot; as the foot is raised the sole is everted (p. 50). It follows [nobis) that to
strike upon the outer border the sole must be slightly inverted as the foot is lowered;
secondh', in so far as the trunk torsion is transmitted to the feet, the feet must be
rotating upon their own axis from the outer to the inner sidfe. Let us first see how
this theory of foot motion agrees with the metapodial displacement.

G. Tlie metapodial displacement is pi'actically uniform throughout the Ungulata
in the carpus, but very diverse in the tarsus. This difference is undoubtedly duL- to
the fact that the fore foot is for support, the hind foot for propulsion. The latter
moves forwards and backwards in a nearly straight line, thus the meta])odials are
vai-iously displaced in accordance with slight lateral variations in this orthal move-
ment. Jn tlie manus the second and third metacarpals invariably have ectal facets
upon the magnum and unciform lespectively. The only variation is in the adaptively
reduced Artiodactyla in which the third metacarpal spreads also upon the trapezoid
(Diag. 9). This disposition of the facets is perfectly adapted to resist the strain
upon the metapodials as the foot swings inward in. descent, this motion being univer-
sal among the primitive types. Thus the ectal displacement of the metapodials be-
gins with the elevation of the wrist joint and precedes the inteicari)al dis})lacement
(see Phenacodus, Diag. 8). It does not advance but appears to he retaided by the
elongation of the metapodials, as in the pala^othere and equine line, as tin; lateral motion
diminishes. The extension of these carpo-metacarj)al facets is largely a matter of
reciprocal growth, i.e., the unciform, for example, extends inwards as the thiid ineta-
cari)al extends outwards (compare Diags. 8, i<).

7. Growth of the scaphoid and unciform. As a result of metacarpal displace-
ment and of digital reduction, the unciform, already supporting the tourlh and lillh
digits, received a portion of thi' third digit. As the lil'tli digit persisted longer than
the first, both the motion of the foot striking upon the outi'i- i)order and the trans-
mission of the direct weij^ht throu<rh three diyfits contributed the maxiiuum vertical
and lateral strain to tlie ectal side of the second row of carpals and especially to the
unciform, while in the bones of the first row the cntal side, the scaphoiil and lunar,

•The Muyljridgc Work al tlic Uiiivi-rsily of Pennsylvunlii. MiiluriitlM lor a .Mciimir u|ion Aniin^il [.■!■ .uni.iiriii. Ily
Hnrrison Alkii. I'hila., 188H.

564.

THE MAMMAIJA OF THE UINTA FORMATION.

received the maximum pressure nud lateral straiu. At the same time, the eetal or
ulnar side of the upper row and ental or i-adial side of the second row received a
diminished pressure and strain by the reduction of the ulna and loss of the first digit
respectively.* At all events the scaphoid grew outwards and the unciform grew
inwards in all the Diplarthra ; they thrust the lunar and magnum apart over the
unciform and beneath the scaphoid respectively, in many cases without any actual or
relative increase in the size of these elements, as demonstrated by the extremely,
small size of both the magnum and lunar in the greatly displaced carpus of Triplo-
pus or Ilyracodoii. Thus what appears to be a rutation of the bones of the uppei-

rhermcodiis

//asiodon

Den c/roh vf^hx

DiAO. 8.— The displarpiiient of \\w. bones of Ihe manus in the Unsuleita, I. Phemicodus primii'i;m, from original
in the Cope collection. Coryphodon hamalus, after ]\Iarsh. Dendrohyrax arborcns, from orii;inal in the Princeton col-
lection. Macrauchenia palmhonica, adar Gcxxsns. Mastodon Americaniis from original in Princeton collection. The
oblique arrows indicate the direction of displacement.

row upon the second row, is in reality the growth of the scaphoid and unciform to-
wards the median axis of the foot. This is cai-riod to its extreme in tlic tridactyl
Perissodactyla, in which the mesial borders of the uncifoi-m and scaphoid are almost
ill contact in the mesaxial line, i. e., in the line carried througli the middle toe.

This explanation is still inadequate, because we find, first, that under cer-
tain conditions the lunar shows a marked tendency to expand, and secondly,
the mesial growth of the scaphoid and unciform ceases.

* A convenient illiHU-a',ii)n of the elVccl of digital rodncliDU upon the c,iri).ds was rem irkcd liy Dr. Scolt in Hie
reduction of the scaphoid in llic inadaptively reduced Artiodactyla— the scaphoid resting upon the trapezoid and nio
trapezium supporting tlie fnnctionlcss second digit only.

THK MAMMALIA OF TIIK UINTA FOKMATION. 5G5

8. Reduction and disj)lacement. There is unqucstiuii:il)ly an intimatL' c':)nncoti(ni
between the reduetioii ol" tlio latei'al toes and displacement. Kydci* has poiiitod out
that where the strains are ctjnally distributed among- all the digits, tiiere is rarely any
specialization of toes, and that in general the reduction of the digits is in direct ratio
to the increase of mechanical strains and the frequency and severity of impacts.

Copet has recently assigned displacement as a cause or as preceding digital
reduction. This supposition is rendeied improbable by the numerous instances of
reduction without displacement, for instance, the first digit of the manus and fii-st
and fifth digits of the pes in ITyrax ; secondly, by the numerous cases of extreme
reduction, as in the Pecora, with limited disi)lacement.

Xow if we consider the manus wilh reference to the correlation of certain types
of reduction and displacement, without reference to phylogeny, we find six types:

I. The first type is theoretical, and will be found among tiie Puerco Condylar-
thra. In it there is no reduction or displacement. J Plieaacodus is the nearest known
representative, in which, however, there is initial reduction, metapodial displacement,
slight intercarpal displacement, and a marked tendency to the niesaxonic type. J I.
The second type embraces the Amblypoda, witli the lunar strongly and the scaphoid
slightly or not at all displaced. There is no distinct median axis. III. The third
or proboscidian type ditlers from the last only in the direction of llie lunar dis-
placement, which is usually upon the trapezoid. The median axis is not wei.1 defined.
IV. The fourth type is shown in a typical condition in Titanot/ier/'iai/, also more or
less marked in all tetradactyl Perissodactyla, as well as in all tetradactyl and
didactyl Artiodactyla ; it represents a stage in which the pcrissodactyl loot
approaches the artiodactyl, the thiid and fourth toes spreading upon either side
of the mesaxial line; the displacement of the caipals is apparently arrested,
the lunar retaining some foothold upon the magnum. Ail tridactyl and monodactyl
ungulates pass through this stage. Y. The fifth type is invariably associated with
tridactylism, incipient {Jfijraclnjns) or advanced (A2)lielops), in which displacement
is carried to an extreme and the mesaxial line jjasses directly through the third digit.
VI. The sixth is associated with monodactylisni and entire loss of lateral motion, in
which the primitive displacement is counteracted l»y the growth dI' l iif magnum.
\\\. The seventh is represented in Macraucheni'i, the lunar showing extreme reduc-
tion, both the scaphoid and unciform articulating with the magnum.

* On llic Laws of Digilnl Hr'tliK'iion. AmcrirAn Naliinilisl, 1X77, p. 00;t.

f "This rolaiimi lins resulted, sooner or lulcr, in the loss of the first <liglt." Aini-ricin Niitumlist. 18M0, p. Oho.
J The pes of I'uriptyfhui is so closely similar to that or the bear. Oii! nmniis may be assumed to I"' i>l' il"' -amc
type.

A. P. S, vol.. XVI. .'Jt.

566

THE MAMMALIA OF THE UINTA FORMATION.

■tfenodus. AnrliMieriuin /Ijlhelops^^

First. In Types II and III the unreduced, nearly isodaetyl and semiplanti-
grade form?, the Amhlypoda and Proboscidia, the unspecialized spreading toes trans-
mit the strain from eveiy side. We observe a marked increase in the
size of the lunar. Among the Proboscidia the lunar eithei' rests upon the
magnum only or upon the magnum and trapezoid ; this is the common condition ; or
upon the magnum, tiapezoid and unciform (see page 539).

Second. In all the genera of the reduced types, excepting the highly spe-
cialized Types VI and VII, it is a remarkable fact that the scapho-magnum and
lunai- unciform facets ai'e subequal. It appears fi-om a comparison of the vast num-
ber of forms included in Types IV-VI, that reduction, altering as it does the entire
system of digital strains, has had some definite influence in limiting or widening the
extent of these facets, and that Prof. Cope has put the effect before the cause in
stating that displacement precedes reduction. Schlosser has rightly reversed this
relation.*

The pei-manent tetradactyl condition in the perissodactyls of Type IV, like the
ai'tiodactyl didactylism is paraxonic, i. e., the greatest vertical strain is not trans-
mitted directly through the median toe but between the third and fourth toes, this,
with the lateial strain coming from the functional fifth toe, in some manner limits the
growth of the scaphoid and unciform, and in many instances results in the spreading
of the lunar in both directions ; it will be observed that in nearly every case the
lunar is directly in the mesaxonic line. AVhere tiie four toes sjjread, as in Titano-
tlierium, the metapodial strains must be somewhat analogous to those in the Probos-
cidia and Amblypoda — at all events the grow th of llie lunar in each instance suggests
such an analog3%

As this condition of the lunar occurs in widely separated phyla and in a targe

*0p. cit., p. lis : "Gleiclizeilis Jiiil tVwsi;r [{ciluUlicm iki- SfilciizcluMi liiuU-1 iuuli ciin- AcjuUTiiiig- in ilcr Anonl
liimg iler Cariiiiticii uiul Tiusalicn stall, '■

THE MAMMALIA OF THE UINTA FOUMATIOX. 567

number of forms, it cannot be due to mere coincidence, but must be the result of cer-
tain modifications of the pressure and strain transmitted to the carpus in i)araxonic
types. The loss of the Literal toes in the highly reduced Artiodactyla. in which the
pressure is most jjerfectly divided between mts. iii and iv, from Eocene to recent times,
has not been accompanied by any alterations of the relations of the lunar to the mag-
num and unciform (see Anoplotherium and Cervus), apparently because there is no
alteration in the axis of the metai)odium. But in the Perissodactyla the reduction of
mc. V, as can be readily observ'ed in the rhinoceroses (see De Blainville, Osteographie,
R. tetradactylus of Auvei-gne, of Sansans, and li. sumafrensis, utiicorms, bicornh), is
accompanied step by step by the displacement of the lunar from the magnum. Lateral
compression of the carpus has nothing to do with this displacement, because the
tapir, which is even now in the transition stage between Types TV and Y, has a
much narrower carpus than Aphelops. The tapir manus is now in the stage of evo-
lution which was passed by the rhinoceros in the Miocene period.

It follows from these cases that I'eduetion per se does not influence displacement,
but only in so far as (in the Perissodactyla) by releasing the carpus from the pressure
of the V digit it brings the major axis directly througli the centre of the third digit, as
seen in Type Y. This type thus embraces a large number of Ibiins, in which reduc-
tion of the fifth digit has been very rapid. The writer can find no exceptions to the
law that mesaxonic tridactyl types of all the Tei'tiai-y pei-iods show the extreme of
diplarthrism. *

Dr. Scott has called the writer's attention to a third correlation of the lunar and
reduction, as found by a comparison of the lunar, sca|)hoid and unciform in tiie
adaptive and inadaptive Artiodactyla. In the inadaptive types (Euteledon, Ano-
plotherium, etc., see Kowalevsky) the lunar is enlarged at the expense of the sai[)h-
oid ; the latter resting upon the trapezoid, which in turn suppoi-ts only the rudi-
mentary second digit, the third digit foi-miug no trapezoidal union. In the adajHive
types, on the other hand, the third digit acquiring a trapezoidal facet, the scaphoid is
somewhat larger and the lunar less enlarged.

TAiisrjs.

The above observations api)ly chielly to the carpu>;, bill |)i('ciscly the same |)riu-
ciples operate in the tarsal displacements.

•Tlie manus o( PaUiolheriitm medium, us figured liy KowulevnUy (Piiln-onloRrniiliica, Hd. XXI[, Tnf. VII). prt-!tcnl»
an exception. But reference to Do Rlainvllle's flgure anil to a cast of the original foot recently presented to Princeton
by llic I'ari'i Museum, shows that Kowalevsky's flgure is incorrect. There i.s no ejtiensivp lunar niagniiin facet, !«ucli
as he represents.

568 THE MAMMALIA OF THE UINTA FORMATION.

The rapid reduction of the fibula accompanying the orthal movement of the limb
hastens the displacement of the astragalus, in advance of any displacement in the
carpus.

The growth of the asti'agalus upon the cuboid and the cuboid beneath the as-
tragalus are reciprocal. The former may be compared to the scaphoid and lunar dis-
placement, the latter to the growtli of the unciform beneath the lunar, i. e., towards
the mesaxial line.

The line of association of reduction with certain forms of displacement observed
in the manus has a parallel in the astragalo-cuboidal disj^lacement. It will be ob-
served that in the artiodactyl or paraxonic types there was a primitive disposition to
form an astragalo-cuboidal facet, adapting the middle axis of the astragalus and
calcaneum to the middle axis of the metapodium, which here passes between the sec-
ond and third digits. On the other hand the Hyracotherium, Paloplotlierium and
IlyracodiM mesaxonic types, as they all tend to monodactylism, show an arrest of
displacement, owing to the growth of the third digit producing conditions similar to
those in the monodactyl carpus ; while genera of the iso-tridactyl mesaxonic type,
the tapir, i-hinoceros, etc., all show extreme astragalo-cuboidal displacement exactly
parallel to that in the carpus.

Conclusions.

The laws of adaptation of the serial plantigrade foot to digitigradism may be
summarized as follows :

1. Displacement is effected by the gi'owth, arrested growth, or reduction of
different elements, and takes place in the direction of the greatest lateral strain, being
most rapid in the elements which are subjected to the maximum vertical impact and
lateral strain.

2. The direction and degree of intercarpal displacements are adapted
t o t h e g r a d u a 1 alteration of the major axes in the bones of
the fore arm and of the metapodium respectively, as brought
about by reduction, and tend to maintain these proximal and
distal axes in the same vertical line.

3. The initial displacement, however, preceding and independent of reduction, is
the e c t a 1 movement o f t h e m e t a p o d i a 1 s , adapting these elements to resist
the strain of the "stroke" upon the outer border, as the foot extends downwards and
inwards.

4. Til ihe unreduced isodactyl types, the strain ol" the spreading metapodials

THE MAMMALIA OF THE UIXTA FORMATION. 569

converges to the centre of the carpns, without a defined median digital axis, and the
lunar spreads to the ectal or cntal side according as the respective growth of the
radius or ulna alters the major axis of the fore arm.

5. In the Diplarthra the major axis of the fore arm passes through the radius,
and through the thiid digit (Mcsaxonia) or between the second and lliiiil digits
(P a r a X o n i a). The outwaid displacement of the entire ui)per upon the lower row
of carpals is apparent, not real ; the magnum and lunar are arrested in growth, and
the lunar remains almost directly in the mesaxial line ; what actually takes place is
the ental growth of the unciform and ectal growth of the scaphoid towards the
mesaxial line, thrusting the magnum and lunar apart.

(i. This growth is affected by the i-eduction of the lateral digits in so far as this
alters their relation to the major axis of the metapodium. Where reduction leaves
the major axis between the third and fourth digits, this growth is arrested ; where it
leaves it directly through the third digit it is extreme ; where the third digit alone
transmits the main impact, the displacement is neutralized by the growth of the ele-
ments which directly support this digit (/. e., magnum and cuiK'ifoi'm 3).

THE MAMMALIA OF THE UINTA FORMATION. 571

EXPLANATION OF PLATES.
Plate VII.

Protoreodox PARVrS.

Fig. 1. Skull, side view ; la, base of skull and superior molars ; 16, brain from above ; »», sylvian g) rus : \,
lateral : hi, median gj'rus.

2. Inferior molars, internal view ; 2<i, crown view.

3. Porlion of c a r p u s with first digit.

4. Humerus, anterior view of distal end.

5. Tibia and fibula, anterior view ; 5a, distal view.

6. Pes, anterior view, of a larger individual.

7. Phalanges, of fourth digit.

8. -Metatarsal 1 1 , medial side of proximal end ; c^, facet for mesocuneiform ; <•,, for entocuneifonn.

Leptotraoulvs proa^tjs.

9. Lower jaw, side view, combined from specimens shown in Figs. 10 and 1 1.

10. Lower jaw, inner view, showing pm. 4 and m. 1 ; a, crown view.

11. Second and third lower molars of another specimen, inner view ; a, crown view.
13. Ulna and radius, pro.ximal end.

13. Scapula, distal end ; 13a, glenoid cavity.

14. Metacarpal iv, anterior view ; 14a, proximal end ; u, facet for unciform.

15. Portion of tarsus, anterior view.

10. Cuboid.

(Figures natural size.)

Plate VIII.
Dii'LACODON, see page 514.

FxQ.\-\a. Fifth cervical vertebra, anterior view, lateral view.
2. Dorsal vertebra, between d. 7 and rf. 10.
3-4. Lumbar vertebra, laleml view, posterior view.

5. Femur, anterior view ; 5a, external view ; 56, distal view.

6. Tibia, anterior view ; Ga, distal view.

7. Left OS innominalum, lateral view, x i-

8. Calcaneum and astragalus, front; 8a, external ; 86, posterior view of astragalus; c. distal view.

9. Carpal 9, proximal row ; probably belonging to a young individual ; u, proximal surliices. Reference to

Diplacodon uncertain.

Plate IX.

Drri.AronoN. see pagi-s 515-10.

Pig. 10. 11 na and radius, external view ; 10a, distal view ; 106, anterior view : lOc, internal view.

11. Humerus, posterior view ; 11a, external view.

12. Left scapula, outer surface, X J ; a, glenoid fos^a.

13. Right man us, x 1; the cari'als are restored Inuii Paltrntynpt and Titannl/ieriiim ; the nu-liirarpiil!<

arc slightly foreshortened ; b, mcs. ii-iv, proximal view ; a, mc. iv, distal view, x ',

14. Superior molar, probably to'.

15. Axis, belonging to a smaller indiviilual ; .tide view.

572 THE MAMMALIA Or THE UINTA rOK3IATION.

Plate X.

IsECTOLOPHrS ANNFXTESS, SOC page 520.

Fiij. 1. S u [1 e r i o r molars of left side ; la, external view of same.

2. I n f c r i o r molars, crown view ; 2a, external view of m^.
11. Inferior series of a younger individual (No. 10,399).

3. L e ft c:i rpu s, front view, with Ihe metacarpals partly restored.

4. Ilia; lit cuboid, external view ; 4n, anterior view; 4i, superior view, showing at (astragalar) and cc

(calcaneal) facets.

5. P h a 1 a n ;;■ e s probably belonging to the second and tliirtl digits.

6. 51 ctatarsals, distal portion, probably of in and iv ; a. dislal faces of same.

7. Tibia, distal face.

8. Proximal vii-w of radius and ulna.

Amynodon ixTEiiMEDirs, see page .'308.

10. Palate and sup c r i o r d c n t i t io n ; 10a, side view of superior dentition.

Meson Yx vintensis.

9. Inferior ji r e m o I a r - m o I a r series, internal view.

Plate XL

PlESIARCTOMYS SClUIiOIDES.

Fig. 1. Skull, from above ; la, base of skull ; IS, side of skull ; If, crowns of superior molars ; UX, crowns of
inferior molars and incisor.
3. Tibia, side view ; 2a, h u m e r u s , side view.

EpiHirrrs tintensis, see page 529.

3. Lower jaw, from above ; 3a, side view.

4. Forearm and man us, front view ; 4a, side view.

5. Tarsus, front view.

TitiPLOrus OKLKitTiDENs, scc page 525.

6. Superior molars, crown view ; fir/., external view.

7. Inferior mo 1 a rs and alveoli, typo of T. obliquidetis (No. 10,402).

8. Man us, front view.

9. Pes, front view.

10. Tlnrd superior molar (No. 10,402).

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