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HARVARD UNIVERSITY.

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MUSEUM OF COMPARATIVE ZOOLOGY.

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TRANSACTIONS

OF THE

AMERICAN PHILOSOPHICAL SOCIETY

Bie iD AV EAC bese TAs,

FOR PROMOTING USEFOL KNOWLEDGE.

VOL. XIX—NEW SERIES.

PUBLISHED BY THE SOCIETY.

Philadelphia:
MACCALLA & COMPANY INC., PRINTERS.
1898.

CO INTIT TEL INGIES Ova WO GOS

PART I.
ARTICLE I.

A New Method of Determining the General Perturbations of the Minor Planets. By William McKnight
Ritter, M.A. .

ARTICLE II.

An Essay on the Development of the Mouth Parts of Certain Insects. By John B. Smith, Sc.D. (With 3 plates)

PART II.

ARTICLE III.

Some Experiments with the Saliva of the Gila Monster (Heloderma suspectum). By John Van Denburgh, Ph.D. .

ARTICLE Ivy.

Results of Recent Researches on the Evolution of the Stellar Systems. By T. J. J. See, A.M., Ph.D. (Berlin).
(With 2 plates)

ARTICLE V.

On the Glossophaginz. By Harrison Allen, M.D. (With 10 plates)

ARTICLE VI.
The Skull and Teeth of Ectophylla alba. By Harrison Allen, M.D. (With 1 plate)

PART III..

ARTICLE VII.

The Osteology of Elotherium. By W. B. Scott. (With 2 plates)

ARTICLE VIII.

Notes on the Canidve of the White River Oligocene. By W. B. Scott. (With 2 plates)

ARTICLE IX.
Contributions to a Revision of the North American Beavers, Otters and Fishers. By Samuel N. Rhoads. (With 5

plates)

199

237

267

417

Aes

BME ate

DEC .6 1996 Pho ee eA GE TONS
DEbdLrS eta

AMERICAN PHILOSOPHICAL SOCIETY,

HELD AT PHILADELPHIA,

FOR PROMOTING USEFUL KNOWLEDGE.

VOLUME XIX —NEW SERIES.

PAR bak

Fi
ArvicheE I.—A New Method of Determining the General Perturbations of the Minor Planets. By
Wiliam McKnight Ritter, M.A.

ARTICLE II.—An Hssay on the Development of the Mouth Paris of Certain Insects. By John B,
Smith, Sc.D.

Hhiladelphin:
PUBLISHED BY THE SOCIETY,
AND FOR SALE BY
Tue American PuHinosopHicaLt Society, PHmapELPHIA

N. TRUBNER & CO., 57 and 59 LUDGATE HILL, LONDON.

1896.

DEC 16 1896

TRANSACTIONS

OF THE

AMERICAN PHILOSOPHICAL SOCIETY.

ARTICLE I.

A NEW METHOD OF DETERMINING THE GENERAL PERTURBATIONS OF
THE MINOR PLANETS.

BY WILLIAM McKNIGHT RITTER, M.A.

Read before the American Philosophical Society, February 28, 1896.

PREFACE.
In determining the general perturbations of the minor planets the principal diffi-
culty arises from the large eccentricities and inclinations of these bodies. Methods
that are applicable to the major planets fail when applied to the minor planets on
account of want of convergence of the series. For a long time astronomers had to be
content with finding what are called the special perturbations of these bodies. And
it was not until the brilliant researches of HANSEN on this subject that serious hopes
were entertained of being able to find also the general perturbations of the minor
planets. HANSEN’s mode of treatment differs entirely from those that had been pre-
viously employed. Instead of determining the perturbations of the rectangular or
polar coérdinates, or determining the variations of the elements of the orbit, he regards
these elements as constant and finds what may be termed the perturbation of the
time. The publication of his work, in which this new mode of treatment is given,
entitled Auseinandersetzung einer zweckmiissigen Methode zur Berechnung der absoluten
A. P. S—VOL. XIX. A

6 A NEW METHOD OF DETERMINING

Stérungen der kleinen Planeten, undoubtedly marks a great advance in the determina-
tion of the general perturbations of the heavenly bodies.

The value of the work is greatly enhanced by an application of the method to a
numerical example in which are given the perturbations of Egeria produced by the
action of Jupiter, Mars, and Saturn. And yet, notwithstanding the many exceptional
features of the work commending it to attention, astronomers seem to have been de-
terred by the refined analysis and laborious computations from anything like a general
use of the method; and they still adhere to the method of special perturbations devel-
oped by Lacraner. HaANsEN himself seems to have felt the force of the objections
to his method, since in a posthumous memoir published in 1875, entitled Ueber die
Stérungen der grossen Planeten, insbesondere des Jupiters, his former positive views
relative to the convergence of series, and the proper angles to be used in the argu-
ments, are greatly modified.

Hint, in his work, A New Theory of Jupiter and Saturn, forming Vol. IV of
the Astronomical Papers of the American Ephemeris, has employed HAwnsEn’s
method in a modified form. In this work the author has given formule and devel-
opments of great utility when applied to calculations relating to the minor planets, and
free use has been made of them in the present treatise. With respect to modifica-
tions in HAwsen’s original method made by that author himself, by Hit and others,
it is to be noted that they have been made mainly, if not entirely, with reference to
their employment in finding the general perturbations of the major planets.

The first use made of the method here given was for the purpose of comparing the
values of the reciprocal of the distance and its odd powers as determined by the pro-
cess of this paper, with the same quantities as derived according to HANSEN’s
method. Upon comparison of the results it was found that the agreement was prac-
tically complete. ‘To illustrate the application of his formule, Hansen used Egeria
whose eccentricity is comparatively small, being about ;4;. The planet first chosen
to test the method of this paper has an eccentricity of nearly +. And although
the eccentricity in the latter planet was considerably larger, the convergence of the
series in both methods was practically the same. It was then decided to test the
adaptability of the method to the remaining steps of the problem, and the result of the
work has been the preparation of the present paper.

HAnsEN first expresses the odd powers of the reciprocal of the distance between
the planets in series in which the angles employed are both eccentric anomalies. He
then transforms the series into others in which one of the angles is the mean anomaly
of the disturbing body. He makes still another transformation of his series so as to
be able to integrate them.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 7

In the method of this paper we at first employ the mean anomaly of the dis-
turbed and the eccentric anomaly of the disturbing body, and as soon as we have the
expressions for the odd powers of the reciprocal of the distance between the bodies,
we make one transformation so as to have the mean anomalies of both planets in the
arguments. These angles are retained unchanged throughout the subsequent work,
enabling us to perform integration at any stage of the work.

In the expressions for the odd powers of the reciprocal of the distance we have,
in the present method, the La Place coefficients entering as factors in the coefficients
of the various arguments. These coefficients have been tabulated by RuNKLE in a
work published by the Smrrnson1An InstituTIon entitled New Tables for Determin-
ing the Values of the Coefficients in the Perturbative Function of Planetary Motion ;
and hence the work relating to the determination of the expressions for the odd powers
of the reciprocal of the distance is rendered comparatively short and simple.

In the expression for A’, the square of the distance, the true anomaly is inyolved
In the analysis we use the equivalent functions of the eccentric anomaly for those of
the true anomaly, and when making the numerical computations we cause the eccentric
anomaly of the disturbed body to disappear. This is accomplished by dividing the
circumference into a certain number of equal parts relative to the mean anomaly and
employing for the eccentric anomaly its numerical values corresponding to the various
values of the mean anomaly.

Having the expressions for the odd powers of the reciprocal of the distance in
series in which the angles are the mean anomaly of the disturbed body and the
eccentric anomaly of the disturbing body, we derive, in Chapter II, expressions for
the J or Besselian functions needed in transforming the series found into others in
which both the angles will be mean anomalies.

In Chapter IIT expressions for the determination of the perturbing function and
the perturbing forces are given. Instead of using the force involving the true anom-
aly we employ the one involving the mean anomaly. The disturbing forces employed
are those in the direction of the disturbed radius-vector, in the direction perpendicular
to this radius-vector, and in the direction perpendicular to the plane of the orbit.

Having the forces we then find the function W by integrating the expression

5. Papo

aw do
al dr?

aE -° iG

in which -A, and B are factors easily determined.

8 A NEW METHOD OF DETERMINING

From the value of IW we derive that of W by simple mechanical processes, and
then the perturbations of the mean anomaly and of the radius-vector are found from

pe Oe = nf W .dt

y being a particular form for g.
The perturbation of the latitude is given by integrating the equation

C being a factor found in the same manner that A and B were.

It will be noticed that in finding the value of . dz two integrations are needed ;
in finding the perturbation of the latitude only one is required.

The arbitrary constants introduced by these integrations are so determined that
the perturbations become zero for the epoch of the elements.

In all the applications of the method of this paper to different planets the cireum-
ference has been divided into sixteen parts, and the convergence of the different series
is all that can be desired. In computing the perturbations of those of the minor
planets whose eccentricities and inclinations are quite large, it may be necessary to
divide the circumference into a larger number of parts. In exceptional cases, such as
for Pallas, it may be necessary to divide the circumference into thirty-two part s.

In the different chapters of this paper the writer has given all that he conceives
necessary for a full understanding of all the processes as they are in turn applied
And he thinks there is nothing in the method here presented to deter any one with
fair mathematical equipment from obtaining a clear idea of the means by which astron-
omers have been enabled to attain to their present knowledge of the motions of the
heavenly bodies. The object always kept in mind has been to have at hand, in conve-
nient form for reference and for application, the whole subject as it has been treated by
HANSEN and others. Thus in connection with HANseEwn’s derivation of the function

IV, to obtain clearer conceptions of some matters presented, the method of BRuNNow
for obtaining the same function has also been given. In some stages of the work
where the experience of the writer has shown the need of particular care the work is

TITKE GENERAL PERTURBATIONS OF TILE MINOR PLANETS. 9

given with some detail. And while the writer is fully aware that here he may have
exposed himself to criticism, it will suffice to state that he has not had in mind those
competent of doing better, but rather the large class of persons that seems to have
been deterred thus far, by imposing and formidable-looking formulz, from becoming
acquainted with the means and methods of theoretical astronomy. In the present
state of the science there is greatly needed a large body of computers and investiga-
tors, so as to secure a fair degree of mastery over the constantly growing material.

The numerical example presented with the theory for the purpose of illustrating
the new method will be found to cover a large part of the treatise. The example is
designed to make evident the main steps and stages of the work, especially where
these are left in any obscurity by the formule themselves. As a rule, the formule are
given immediately in connection with their application and not merely by reference.
It has been the wish to make this part of the treatise helpful to all who desire to
exercise themselves in this field, and especially to those who desire to equip themselves
for performing similar work.

The time required to determine the perturbations of a planet according to the
method here given is believed to be very much less than that required by the unmodi-
fied method of Hansen. Nearly all the time consumed in making the transforma-
tions by his mode of proceeding is here saved. The coefficients b are much more
quickly and readily found by making use of the tables prepared by RuNKLE, giving
the values of these quantities. Doubtless experience will suggest still shorter pro-
cesses than some of those here given and thus bring the subject within narrower limits
in respect to the time required. If we compare the time demanded for the computa-
tion of the perturbations of the first order, with respect to the mass, produced by
Jupiter, with the time needed to correct the elements after a dozen or more oppositions
of the planet, computing three theoretical positions for each opposition, it is believed
there will not be much difference, if any, in favor of the latter.

Again, when we wish to find only the perturbations of the first order, experience
will show where many abridgments may safely be made. And whenever the positions
of these bodies are made to depend upon those of comparison stars whose places are
often not well determined, it will be found that the quality of the observed data
does not justify refinements of calculation.

One of the things most needed in the theory of the motions of the minor planets
is a general analytical expression for the perturbing function which may be applicable
to all these small bodies. Thus if we had given the value of aQ in terms of a periodic
series, with literal coefficients and with the mean anomalies of the planets as the argu-

2X5 JES Sk A\VOlke IIDG 18.

10 A NEW METHOD OF DETERMINING

dQ : es :
ments, we would at once have a a by differentiation. And since

G2 dQ
i ahi?

only two multiplications would be needed in finding the value of aan

, whose expres-

sion has been given above.

In the present paper we have dealt only with the perturbations of the first order
with respect to the mass. The method has been employed in determining those of the
second order also for two of the minor planets ; but as those of Althsea, the planet em-
ployed in our example, have not yet been found, it was thought best not to give any-
thing on the subject of the perturbations of the second order, until the perturbations of
this order, in case of this body, are known.

The writer desires here to record his obligations to Prof. Edgar Frisby, of the
U.S. Naval Observatory, Washington, D. C., and to Prof. George C. Comstock,
Director of the Washburne Observatory, Madison, Wis., for kindly furnishing him
with observations of planets that had not recently been observed; to Mr. Cleveland
Keith, Assistant in the office of the American Ephemeris, for most valuable assistance
in securing copies of observed places. And to Prof. Monroe B. Snyder, Director of
the Central High School Observatory, Philadelphia, he is under special obligations for
the interest manifested in the publication of this work, and for continued aid and most
valuable suggestions in getting the work through the press.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 11

CHAPTER I.

Development of the Reciprocal of the Distance Between the Planets and its Odd
Powers in Periodic Series.

The action of one body on another under the influence of the law of gravitation
is measured by the mass divided by the square of the distance. If then A be the dis-
tance between any two bodies, this distance varying from one instant to another, it

3 : : Wes :
will be necessary to find a convenient expression for (5) in terms of the time. If

r and 7’ be the radii-vectores of the two bodies, the accented letter always referring
to the disturbing body, we have |

N= 4+ r? — 2rr A.

If we introduce the semi-major axes a, a’, which are constants, and their relation

a’ S
a=, we obtain

= (3) + () @—2 (2) (2) (1)

HT being the cosine of the angle formed by the radii-vectores.

Let the origin of angles be taken at. the ascending node of the plane of the dis-
turbed, on the plane of the disturbing, body. Let I, Il’, be the longitudes of the peri-
helia measured from this point; also let f, 7’, be the true anomalies. The angle

formed by the radii-vectores is (f’ + I’) —(f + Il); and the angles f + 0, f+ WW,
being in different planes, we have

H = cos (f + II) cos (f’ + Tl’) + cos Zsin (f ae iN Fsimy (G7 > UI), (2)

I being the mutual inclination of the two planes.
To find the values of I, Il’, J, let ® be the angular distance from the ascending
node of the plane of the disturbed body on the fundamental plane to its ascending

12 A NEW METHOD OF DETERMINING

node on the plane of the disturbing body. Let y be the angular distance from ascend-
ing node of the plane of the disturbing body on the fundamental plane to the same
point.
If x, 7, are the longitudes of the perihelia,
, 2’, the longitudes of the ascending nodes on the fundamental plane adopted,
which is generally that of the ecliptic, we have .

W=a—Q-—49, Woaw—Q—r. (3)

The angles ®, y, 2 — 9’, are the sides of a spherical triangle, lying opposite the
angles 7’, 180 — 2, J, -

2, v, being the inclination of disturbed and disturbing body on the fundamental
plane.

The angles J, ®, 7, are found from the equations

sin $ sin $ () + ©) = sin $(Q — 2) sin $(¢ + 7)
sin 3 J cos} () + ®) = cos § (Q — Q’) sin 4 (¢ — 7) (4)
cos $ Jsin $ (Y — ®) = sin $ (Q — 2) cos $ (¢ 4 72)
cos § Lcos$ (J) — ®) = cos $ (Q — &) cos § (¢ — 7)

In using these equations when Q is less than 9’ we must take 4 (860° + Q — 9’)
instead of $ (2 — Q’).

We have a check on the values of £ ®, J, by using the equations given in Han-
SEN’s posthumous memoir, p. 276.

Thus we have

cos p. sin q = sin 2. cos (Q — &’)

COS p. COS Y = Gos U

cos p. sin 7 = cos 2. sin (3 — %)

cos p. cos rT = cos (83 — &’)

sin p = sin?’ sin (8 — 8) \ 6)
sin J sin ® = sin p [

sin J cos ® = cos p. sin (? —

(2 — q)
sin sin (} — r) = sin p .cos (2 — q)
sin J cos (Wy — r) = sin (7 )

( )

cos I = cos p. cos (@

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 13

To develop the expression for (): we put

cos J/.sin 1’ = sin A, sin I’ = & sin Kj, )
cos Il’ = k cos K, cos Icos Il’ = k, cos K,,J

and hence

i= cos f.cos f’.k cos (Il — K) + cos f. sin f’.k, sin (Il — K;)
— sin f.cos f’.k sin (11 — A) + sin f. sin f’.k, cos (11 — 44).

Introducing the eccentric anomaly «, we have

a . A a .
cos f = — (cos e—é), sin f = —. cos. sing,
é being the eccentricity, and ¢ the angle of eccentricity ; and find

._. H= cos «.cos &.k cos (11 — K) — cos ¢. ek cos (11 — K)
— cos e.¢k cos (11 — K) + eek cos (11 — KX)
+ cos «.sin ¢.cos ¢’.k, sin (II — A,) —sin ¢’.é. cos ¢’. k, sin (Il — A)
— sin e.cos ¢.cos @.k sin (1— A) + sin e.é.cos p.k sin (1— XK)
+ sine. sin e’.cos ?. cos ? .k, cos (Il — K)).

re d ‘ A A\2
Substituting the value of ~, me fT in the expression for (=) we have

(=). = 1+ a’*— 2e.cos « + & cos *« — 2aeek cos (Il — I)
+ 2a¢k cos (11 — K) cos ¢ — 2ae’ cos @. k sin (II — FV) sin ¢
— [2a7e — 2aek cos ((I—K) + 2ak cos (Il — K) cos «
— 2a cos o.k sin (I] — XX) sin €] .cos
— [ — 2ae cos 9’. k, sin (11 — K,) + 2a cos > cos 9’. k, cos (11 — Aj) sin ¢
+ 2a cos ¢’.k, sin (11 — 4.) cos e] . sin &

+ a? é€?. cos 7’.

Putting 71, Go, 72, for the coefficients of cos ¢’, sin +’, cos */, respectively, and 7, for
the term not affected by cos «' or sin ¢’, we have the abbreviated form

er = 7) — 71. Cos &’ — By. sin & + yz. cos *e’. (7)

14 A NEW METHOD OF DETERMINING

: : 4
In this expression for ce

of the disturbed body; y. is a constant and of the order of the square of the eccen-
tricity of the disturbing body.
In the method here followed the circumference in case of the disturbed body will

2 . °
) »% Y and 9, are functions of the eccentric anomaly

be divided into a certain number of equal parts with respect to the mean anomaly, g.

. : 2 2 360° 360°
The various values of g will then be 0°, = Bs a6 5 8b ene —1. - :

For each numerical value of g, the corresponding value of ¢ is found from
g =e&—esine.

Before substituting the numerical values of cos <¢, sins, for the n divisions of the cir-
cumference, the expressions for 7, 71, 9, will be put in a form most convenient for

computation.
Let
asin la) 20 g — 2ak cos (II — XK ) (3)
p.cos P = 2a cos ¢' k, sin (II — K,),
and
HSjeom le |)
See )
yi =f.cos F; J
we find

By =fsin f= 2a. cos p. cos 9’. k, cos (11 — K,). sine + pcos P. cos e — ep. cos P
yi =f COS i= (24? —psin P). cose — 2x. cos p.ksin (Il — K). sine + ep.sin P.
And from these equations we find, since

f.sin (#— P) = f.sin F'cos P —f cos FP’. sin P
J .cos(#— P) = feos #’.cos P + fsin F’. sin P,

f.sin(#— P) = [2a. cos ¢?. cos 9’. k, cos (11 — Kj). cos P
+ 2a.cos ¢.k sin(I—). sin P]. sin e + [ _ 202% sin P| . COS E—EP
f. cos (#’— P) = [2a. cos. cos 9’. k, cos (Il — K,).sin P

/
. — ° é
— 2a.cos ~.k sin ((I—FK). cos P]. sine + 2a". .cos P. Gos é.

THE GENERAL PERTURBATIONS OF TITLE MINOR PLANETS. 15

[f we now put

vsin V= 2a.cos¢.ksin (II— KX)
vcos V= 2a.cos.cos 9’. k, cos (Il — K,)
wsin W = p— 2a’. nee
weos W= v.cos(V— P)
w,sin W,= v.sin( V— P)
w, cos W,= 2a’. a cos P,

|
(10)
J

we get

J.sin(#— P)=w.sin(e + W)—ep

f.cos(f#— P) = w,. cos (e+ IV,). (QU)
Further, if we put
R=1+0°?—2a’.e, (12)
we have
Yo = R— 2e.cose + €’. cose + ey,

or, ¥) = R—2e.cose+e.cos*s +e .feos F. (13)

We find the value of y, from

The constants, *, A, hk, Ay, p, P, w, W, w,, W,, 2, are found, once for all, from
the equations given above. For every value of « we have the corresponding value of
Jj and F from equations (11); hence, also the values of fsin #, fcos /, which are the
values of @) and 7, Equation (13) furnishes the value of y) by substituting in it the
various numerical values of <, as was done for 3, and y,. ‘The value of the coefficient

: ‘ A\2 :
y. being constant, we thus have given the values of (“) for as many points along
a

the circumference as there are divisions.

16 A NEW METHOD OF DETERMINING

We can put

Ay? :
() = 70 — 71 Cos e’ — By. sine’ + 72. Cos .e’

in the form

(2) = [C—q. cos (¢ — Q)] [1—q- cos ( — @,)], (14)

in which the factor 1 — q, . cos (e’ — @,) differs little from unity. For this purpose, if
we perform the operations indicated in the second expression, and then compare the
coefficients of like terms, we find

y= C+ ¢q-qsin Q.sin Y,
v1 =¢.cosQ+q,.Ceos Q,
¥2=7-M-cos(Q + Q)
Po=q-snQ+tqu-.CsinQ,
0=sin(Q+ Q,).

The last of these equations is satisfied by putting

Q ==.

The remaining equations then take the form

Ne = mee ce > ee |

|
a Nn |
B= (¢—a-C).sing |

The expressions

GS OQ) = by sb 2 1)
q. cos OS ee (16)
Gh. Csi C) = & |
Gia C..cos O77) a

pen the relations expressed by the second and fourth of equations (15‘, where
— /Yo als Gi
We haye now to find expressions for the small quantities £, 7, ¢ found in these
equations,

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 17
Equations (16) give
q-q- Csin’?Q= (8+ &).-£.
The equation
y= C—_.qnsin’Q
then becomes
(Yo + S)S = (Go + &)& (a)
From (16) we haye, also,
PO C= (nae Qe a> Waa
from which, since y,=q.q, and C= y, + ¢, we obtain
Oar 6) 9% = (Gnae BE Se (aaa (b)
Equations (16) give again
(yi —n)& = (Go + &) n. : (¢)
When ¢ is known, & is found from (a) ; and the difference between (a) and (0)
Got) 62—5) =O: 5%). (d)
gives 7 when ¢ is known.
The equations (a) and (c) give
Bo +4 (yo + 6) 5 = (Bo + 28)"
Bo+2E= 71-33

and hence

Bi 4 t+ )S=ye.5
A. P. S.— VOL. XIX. C.

18 A NEW METHOD OF DETERMINING

Deduce the values of 8) + &, y, — 7 from (a) and (d), substitute them in (¢), we find

G — (oS
eo Sra Ma eas

The last equation then takes the form
O= 77-5 — Bo (¥2—$) —4 (70 +S) (Y2—S).S- (¢)

This equation furnishes the value of £; and with ¢ known, we find &, 7, from equations
already given. The three equations giving the values of the quantities sought are

Sar Soa) S se lA +P ban = B70 oo) ho Or y2=0)
BP Boo = (yar S)G =) (Cr)
n—y1.n+ (Yo+%) (¥2—4) =)

Finding the values of ¢, £, 7, from these equations, and arranging with respect to y.,
preserving only the first power, we have

Belt

bea =- Bo: ye :

a (9)
jae te ty i

= oom

hae? ies Ae?

Substituting these values in equations (16), they become

q7.5n0Q = Ge ee Yo

q.cosQ = y,— Fie ga an
q Csin Q= ao (ait rr 2

gq. Coos Q= Esty,
noting that C= y, + ¢.

If more accurate values of ¢, &, 7, are needed than those given by equations (gq),
we proceed as follows :

Substitute the value of ¢ given by (g) in the second term of the first of equa-
tions (/), we find, up to terms including y,’,

Pome ReMi Coats » : _ fo: (bts By 18
GS= 7 2° V2 ap abe Ge aE APP” Yo — 4. Yo: ( )

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

The last two of (7) give also

ea Gals Os
= Bo les
C eee Difeo = (Ove
r= (x ae ates)
Nn Nn

Introducing the values of f, F, given by (11), putting

L=72+4.y2. 7p. cos °F

Ua=y—4.y2 A . sin °”

we have
G7 eine
so that
C=y+ %.sin*f.
Moreover, since
v2— 6 = x . cos,
we find from the expressions for &, 7, given above,

Boek 6 jane nein
m—n=f.n'.cos Ff,

if
g=14+2—(4)
a1 (4)

Substituting these in the expressions for gsin Q, gcosQ, they become

gsin Q=/f.&. sm
gq cos Q = f. 7’. cos F.

(19)

(21)

20 A NEW METHOD OF DETERMINING

The value of g, is found from

= 23
n= (23)

The quantities g, g,, Q can be expressed in another manner. The equations (22)
give
ig Q= a .tg #
7
Ga fia SI Bet fi ely aCOS els

from which we derive

Q =F ao a sin 27 =— $ G = .sin 47’ -+ ete.
ql SEG '
log. = log. f + 3 log. (&". sin *#’ + 7” cos °F).

Since y and y” agree up to terms of the third order, the equations for & and 7’

give
ef RON ac ie)
Sp i Ve vie
or,
rar = ue io Me 3)
ra = th a ae (2%. as ) cos 2”
Further
£? sin °F 4+ 7” cos*H=14 2 - (yz. sin °*L— 7’ cos °F’) — (4)
and
4 log. (&° sin >2’+ x” cos “PY = Fs C (x sin’ “F— yx cos “F’)

= “(x sin °L’—y’ cos “F’)°—4$ (A)

Substituting the values of y, y’, C, given before, we find

at i Yo 12 es 9
(Xx sin “H— 7’ cos °#’) = ey ae + S cos 2
fo 7 es
Ce) cos Af

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 21

The equation y, = ¢.q; gives
log. y. = log. g + log. q
Putting

looag = loon f= y,

we have for q,

log. g, = log. 7 — y.

Writing s for the number of seconds in the radius, and 2) for the modulus of
the common system of logarithms, we find

Q=F+2
log. g =log. f + y (24)
log. q: = log. i —y

in which

ga eee + Z,,) sin 27+ s (2 — 1.) sin 4F a
p= Np i iy a dn) cos 277—A, Cue = +) cos 47
And for C' we haye from the first of (15)
C=y7 +y72-sin >Q. (26)

By means of the last three equations we are enabled to find the values of
Q, 9; Gi, C, with the greatest accuracy. The equations (17), where not sufficiently
approximate, will, nevertheless, furnish a good check on the values of these quantities.

All the quantities in the expression for (e) are thus known; and substituting their

values corresponding to the various values of g, we have the values of ie =) for the

different pomts of the circumference.

22 A NEW METHOD OF DETERMINING

Using the values of C, q, @, Q, just found, Hint, in his New Theory of Jupiter
and Saturn, has given another expression for ) which we shall employ.

To transform
(2): = (C—q. cos (¢ — Q)) (L—m. cos (¢ + Q))

into the required form we put

C= 1B

= sin x, = sin 7%

Gi
a=t93%, b=tWan (27)
_ seC 2%. 8eC2 %
NORE

Then

a) = O[1—sin x . cos (¢ — @Q) | [1—sin y.. cos (e+ @Q) |

_ C[ sec? by (1—sin x . cos (e’ — @)) | [sec? $41 (1 — sin x: . eos (¢’ + Q)) |
Ce, sec” by see’ 3%
_ Of 1+ ty? by — 2ty 4 cos (e’ — Q) | [1 + ty? by, —2lg yn cos (e’ + Q) |
ta “sec” by sec’ 2Y,

Substituting the values of a, 6, NV, we get

(2)" = _W* [1 + @— 2a cos (¢— Q) | * [1+ — B cos’ + Q]? (28)

We compute the values of a, b, NV, corresponding to the different values of g, and
check by finding the sums of the odd and the even orders, which should be nearly the
same. If we put
[1 + a — 2a cos (¢ — Or = [4 6° + 6. cos 0+ 6 . cos 29 + 6°. cos 39 + ete. |
[1 + 0 —2b cos (e + ®) | = [3 BO + B .cos(¢+ Q)+ B”. cos 2 (e+ Q) + ete. |

where s = ”, = ¢ — Q, we are enabled to make use of coefficients already known.
9?

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

For 2. cos 0, write x + = and then we have

[1 + a — 2a cos 6] ° — [1 4 @—a (e+ ye
= [1—ae|~ [a—“]-

Expanding we have

1G So ee ee a a’a? + ete
online: Ss a ss) Ia? s stl s+2 @ s stl s+2
[1 al Sea aoa gee ee ae ae 3
s 19 (aes ! 5
+* a a ae aie

Peep eae
a (2 ae ee Sa eee] (w +‘)
el ee Ge) ae!

s stl st2\2s+3 s+4 (¢ Ve =)
sees Be aa . Be + ete. wr

s s 1 s+2 03 ei Size 2 Sap 2 S=- 8 6
ee ge a) Se a

Ag + ete. | (2° +.)

|
i
5

But « + ~ = 2cos6, a? + = 2.c0s 20, a + ©, = 2.cos36, etc.,

24. A NEW METHOD OF DETERMINING

and hence

s s+2 ,.,8 stl s4+2s5+3 4
Se SUES BY a os aia Se ne ae G
i? 3 Lae 2 3 ee

EE OP eg ae]

3 $ ep-bi g+9 s @ts ».,8 stil sts sta ., Vee
youa? st1 242 gy | ee api Sbt Shs std (29
ath @4® gt s-L4 5-46
4 eo ee eS Se, Sr +? a + ete, |
oO

and generally

O—® 8 Sarl @=Fe=1 ss 218 Shel Saree 9se ob 4
b = 2.5 eyo ; ee ae; Pe et te |

Since s = —, we find from these expressions the values of the 6 coefticients for
>?

different values of 7.

Runxk sz has tabulated the values of 6 in a paper published by the Sm1rrHsonran
Institution. Thus the value of

[1 + a? — 2a cos (’e—Q)]~2
is obtained with great facility.

The value of [1 + 6° — 2b cos (¢ + Q)}2 is found in the same way.
We now let

- aa 2 5 IN a cos *Q ) (30)
sO=21 WN. B®. sin2.1Q)
And hence have

CU TENGE.

C= NP Be cos.

Ce) = eo A oO), sin AAG)

iain i 15°), COS 40)

ei IN Jina)

etc.= ete.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 25

Multiplying the series [4 6 + 6. cos 6 + b®. cos 20 + 6”. cos 36 + ete. ]
by [5 B® + B® cos (e+: Q) + B®. cos 2(e + Q) + ete.],

noting that 6 = Q@— ¢, and arranging the terms with respect to cos7#, sin 7,
we find

@'\ — i KO) AO 1 D) {O
(“) = 130, 6 +B, 459.6

A
4 fb. © + (B© +b) cD + (6 +4 B®) c] cos 6
a |f + (6 — 6%) s® + (6 —}®) s@] sind

+ [62.6 + (6M + B®) c& + (6 + 6) c®] cos 26 (31)
tall + (6 — B®) s+ (6 — 6%) 8] sin 26
+ [6%. 6 + (6 + B%) c + (6 + B®) e®] cos 36
FE $24) 8-4 (HY —B) 9°7] sin 30
a= ete. ete.
Now let
i; cos K; — £9, 6 aL (G52 = Hem) ree) + (Gee) + Be) ce) ) (32)
k,sin K; + OBEY) 5 4 GH) 5 J
and we find
(“) = k; [cos K;. cos 20 + sin K;. sin. 76]
= k,cos (#0 — K,) = k;. cos («@ — ve’ — Ki). (33)

Subtracting and adding the angle 2g, this becomes

If we put

(©) 9 a:
A, . = 7 i, C08 [(O.=9) = all |
5 : (35)

Ae esi (0) eel ]

A. P. S.=— VOL. XIx. D.

26 A NEW METHOD OF DETERMINING

n being the number of divisions, we find
R (c) 8 GC) ree
() = A,,.cost(g,—eé,) —A,,.8int(g,—€&’,) (36)
If now, for the purpose of multiplying the series together, we put
(c) (c)

A,, => C,,.cosvg + >. Co (37)

(s) (c) (s)

A, =2>S,,.cosvg +> S,,.sinrvg

we have

G) = Ps C,, cos vg +> C,, sin vg] cos 7(g—e’)—[= S.. , cos vg+> e ‘sin vg | sint (g—e’)

(38)
Performing the operations indicated we get -
e e (¢) (¢) e e (c) . .
=> cos (tg —te’).C;,, cosryg = =4C,,cos[(t+vr) g—te’ ]+35 4C,, cos [(t@—v) g—ae']
° . (s) . (s) . . . (s) . e .
=> cos (4g —ze’).C;,, sinvg= =4EC,, sin[(¢+v)g—ce']—s>4 3 sin [(¢—v) g—“#’ |

e fe . (s) e (8) . . . e
— sin (7g—ze’) S,, sinvg= ZS, cos[(¢+v) g—te’]— S518.,¢ cos [(¢—v) g—%’]
Summing the terms we find

(‘)'= SE1(0,, + S,.) ) cos | (¢=F v)g g—te |F4SS(C,48,) ) sin | (Fv) g—te' | (39)

Thus

(s) (c)

(¢) (ce)
A,=4 8. + S,, cos g + S,. cos 29g + ete. |

io) s = SS, » GOS VG + SS. , sin 7
+ S,, sing + S. sin 2g + ete. j J I:

Hence where vy = 0, each series is reduced to its first term.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 27

In the application of the very general formulz care must be taken to note the
signification of the various terms employed.

In case of

3] no

Ki. . COS [2 (Q.— 9x) — K,,. |

2 : : si

A, =" h,..sin [1 (Q—9.) — Kid,
oe ° O=N0 n s

n shows the number of divisions of the circumference; and we divide by ; in form-

ing k,, to save division when forming the coefficients ¢,, S,.
The index and multiple 7 shows the term in the series

1p +4 B cos (e’ — Q) +b. cos 2(¢ — Q) + b. cos 3(¢ — @) + ete.

The double index 7, x shows the term of the series of La Place’s coefficients and

the particular point in the circumference.
The index » shows the general term of the series expressing the values of

(¢) (s)
ix) When we give to » values from » = 0, to the highest value of » needed in

in)

the approximation.

2 & z .
In ~.&,., 0(Q. — 9.) — Ki for each value of 7, there are » values of each
n yk) «9 ?

quantity.

© GO CO 2 ©
The next step is to express the n values of PA eAn As As) A>, ete, respec-
tively in terms of a periodic series. And since these quantities are functions of the
mean anomaly g, if we designate them generally by Y, of which the special values are

i Cate a MVS Ma are ee arct NC, 25 > y
we have

Y = he, + c cos g + © cos 2g + ete. ) (40)
+s, sing +s, sin 2g + ete. ) re,

The values of ¢,, s,, in this series are found from the ~ special values or 3%

28 A NEW METHOD OF DETERMINING
From

(s)

(e)
A, ,or A, =$q+ 6, cosg + © cos 2g + ete.
+ s, sing + s, sin 29g + ete.,

abe (c) (s)
and similarly, for every other value of x in A;,, A;,, we have a check on the values of
C,, S, In each series. Thus if in case of sixteen divisions of the cireumference we
take g = 22.°5 and find the value of the series, the sum of the terms must equal the

© ©
value of A;,, -A;,, corresponding to g = 22.°5. And this check should be employed
on each series, using that value of g that gives the most values of c, and s,. If 7

; (Onn) ‘
extends to z= 9, we have ten separate checks for the values of A; ,, A;,., respectively.

In the equation

Y=3e + ¢,.cos g + ¢.cos 2g + c;,. cos 3g 4 ete.
+ s,.sing + s.. sin 2g + s;. sin 3g + etc,

if the circumference is divided into twelve parts, each division is 30°. Then for the

special values of Y we have
¥, = te + & + 6 = ¢ + ete.

Y, = $q + 4. cos 30° + ¢,. cos 60° + ¢ cos 90° + ete.
+ s, sin 80°+ s, sin 60° +s, sin 90° + ete.

Y, = q+ «.cos 60° + c,. cos 120° + c, cos 180° + ete.
+s, sin 60° + s,. sin 120° + s, sin 180° + ete.

Y,, = q+ ¢,.330° + ,.cos 800° + ©; cos 270° + ete.
+ s5,.330° -+s,.sin 300°+ 8, sin 270° + ete.

In the same way we proceed for any other number of divisions of the cireum-

ference.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 29

Now let
Qe wy | @) =m ¥
(let SS ae Ge (GA) Ge
Coe) ee eC) Se
(6.W=¥%+% G)=%—Pa
Then

3(s,+ 85) =|) + Gy) ] sin 30° + (8)
3(s:— 33) =| (2) + (tp) | cos 80°

6.8, = (%)—(G) + Gy):

The values of these coefficients can be easily verified by finding the values of
each one from the sum for all the different values of Y’ as given in the series for

DE Atle) epi (eM ne eer
When we divide the circumference into sixteen parts, each division is 22.°5. We
find the values of ¥, Yi, Y.,.... Yis, as in the case of twelve divisions. To find

the values of ¢, and s,, in the case of sixteen divisions, we put

CO gare 8) Se
(Loe) ie ase 285 () = 14 16
(Beal) = Y2+ Yio (Zr = ¥,— Vu

(c= SE V5 Gy = Na oe

30 A NEW METHOD OF DETERMINING
.4)=(0.8) +(4.12) @©.2)=(0.4)+4 2.6)
=(,9) 26.8) €.3jS0.54 6,9)
(2.6) = (2.10) + (6.14)

G.0) = Goll) -4 (15):

eS
iS ©

Then
A(c + 2. ¢) = (0.2)
Ale) —2.¢) = (1.3)
4(e,-+¢) = (0.8)—(4.12)
4(ce,—¢) = {[(1.9)—(.18)]—[(8.11)— (7.15) | cos 45°
A(s, +s) = §[(1.9)—(.13)|+[(8.11)—(7.15)]} cos 45°
4(s,—s,) =(2.10)— (6.14)
8.c,= (0.4) — (2.6)
8.s,=(1.5)— (8.7)

Aleit) | = (4) G4) — Gs) cos 452

4(¢q,—c,) = L@ _ (5) | cos 22 .°5 + [GD — (4°5 )| cos 67 .°5
Ale +65) = (t)—| (Ps) — Gy) | eos 45°

4(e,—e,) =| (4)—(z5) | sin 22.°5—[ (,8,) — (5) | sin 67.°5
4(, +s) = L@ + (5) | sin 22.°5 + [a rae a sin 67 .°5
A(s,— 5) =|’) + Gip)| cos 45°=- G aA

A(s, +s) =| (4) + Gs) ] cos 22.°5—| (8) + Gy) | cos 67° 5
A(s,—ss) =| (5) + Gi) | cos 45°— (G4).

When the circumference is divided into twenty-four parts, each part is 15°.

Let

(0.12)=¥%+ Y¥, (0.6)=(0.12)4+ (6.18) (2)=(0.12)—(6.18)
C= 4 ale) 1) E\=O1)=—@.)
(2.14)= ¥,+ Yi, Ca) 2) OR Cae)

(il By= 42, GansG. neoan 23) a y= Geil) — O23)

Then

Further, let

Then

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS, Bl

6(q + 2.¢.) = (0.6) 4+ (2.8) + (4.10)
6(q—2.¢) = 1.7) 4+ (8.9) + (6.11)
6( + Go) =(@)+ [@) ws Gay sin 30°
6(a—eu) =[C) — Gi) ] 208 30°

6a+4¢) =(0.6)— [(2 .8)4 (4. 10) | sin 30°
6(4—e) =| (1.7) + (6.11) | sin 30°— (3.9)

6)s+sy) = @ + (4%) | sin 30° + (4)
Gos) = @iee (347) | cos 30°
G(s, + ss ) A eG cos 80°
6(s,—s,) = |(2)— Gir) | cos 30°
12.6 = (¢) —@) + Go)
12.5,= (CAG) a= (Ge)

(; ts) = . os Vis
Go=¥ — Yu
aye Yn — ¥,,

6(6, + eu) = (as) +[G) — G0] c0s 80° + [G) — Gis) ] €08 60"
6(¢,—¢,) = LG) —G 1) | cos 15° 4 [()—- Gy] cos 45° + [ Ge — (75) | cos T5°

6(¢,+¢)=
6(¢;— ca ) =
6(¢, +6) =

6(¢;—¢, ) =
6(s, + s.) =
6(s,—s,) =
6(s; + 8) =
6(s;— 8s) ) =
6(s;-+s5,) =

Gp=Ge)s (y)

+Gs ae AOS: —| (3) ax (2) |— |G) — (#5) |? cos 45°
(qr) ae )— G8 2) | cos 80° + Gs) — Gr] cos 60°

(33,) — (Gis) | sin 75°

[Gs + G4) ] sin 15° + (G8) + Ge) | sin 45° +
[2p + (9) ] sin 80° + (Ge) + Gy) ] sin 60° +

| Gs 1G) sim 14° — (3) — Gr) sin 45° 4 [
[ Gi) + G's) | sin 75°
ap GS)

§(G4) + a eee cos 45°

ge) = GE) + G8

[ Gs) + G4) ] cos 15° —[ ( Gh) epi cos 45° + | (3) + (ix) | cos 75°
6(s; s— 8 )=|(2 ae (Gs $) | sin 30° —| (445) + (x) | sin 60° + (4%).

32 A NEW METHOD OF DETERMINING

When the circumference is divided into thirty-two parts, each part is 11°. 25
Let

(OH hte, @O8j)eGOihe(s2) @.64=0.8)£6.
Ch s 42%, Cosa) 69.28) C8) =C.9 46.
(2.18)=¥,+¥, (2.10) =(2.18)4+ (10.26) (2.6)=(2.10)4 (6.

. : @.H)=C.id)+@.

G5 3l)—= v8 Ya (7.15) 2 (0-23) G5.3l) 2) = 04 hee

(1-3) = 4.5 )+G:
OsCij (2) Gsi0.8j)—¢.
OSG Ci25) O=a@.oj)—6.
(q) = (2.10)—(6.
Je) = (7.23) — (15.81) (2) = (8.11) —(7.

Then

8 (e+ 2.6) = (0.2) + (1.3)
8 (¢o— 2.¢5) = (0.2) — (1.3)
(
(

8(e+¢y) =(%)4 [ (2) —( f;) | cos 45°

8 (¢,— 1) = [ (4) — (qs) | cos 22.°5 + [G4 — (,5;) | cos 67 5a
8(a+ Ce) = (4)

8(G—es) = [(a- (3) | cos 45°

8(%+¢) =(%)—| Gr) — Gp | cos 45°

8(¢—eC») = 4) —(5) | sin 22.° 5 — [Ga — (5s) | sin 67.°5
16.¢; = (0.4) — (2.6)

8(s,+sy4) = [@® + (75) | sin 22.°5 + [G+ + (;°5) | sin 67 .°5
8@—su) =(G)— (8) | cos 45° + (;4;)

8(s, + 82) =[(4) + () eos 45°

8(%— sp) = (%)

8 (s+ So) =[ (4) + Gs) | cos 22.°5—| (,3-) + (435) | cos 67 .° 5
8 (s.—S») =[ (as) — (a) | cos 45° — (74).

THE GENERAL PERTURBATIONS OF THE

Further, let

And besides, let

A
B
A
B

(tr) — G4)

= | (2s) — GA]

[G@)— G5]

=| Gs) — Gi) |
= [Gn — Gi).

=| Gs)— Ge]
= G5) + |G) — Gb]
=G;) — (Gy) — G)]
| sin 11°.25 4

‘| cos 22°.5

| cos 33°.75

= |G) + G2) |

(35) = Yy— V4
Gip) = 35;
(5) = ¥,

cos 11°.25 + [ (5)

MINOR PLANETS.

cos 78°.75

rin) INO [ (35) —

cos 22°.5

Gis) |

(5) | sin 78°.75

(a) — (2) ] e08 67°.5

sin 22°.5 — [ Gs) = (oh |

sin 67°.5

cos 33°.75 + [ +) — (4) cos 56°.25

sin 33°.75
cos 45°

cos 45°

[ G4) — G4) | sin 56°.25

[( tee) ae (5) | sin 78°.75
] 08 11°.25 —[ (a's) + (e's) | cos 78°.75

|sin 22°.5 + [(%) + 44) | sin 67°.5

[ + 10) | cos 67°.5 —
[ Gs) + G8) | sim 88°.75 + [Gi + (4) | sin 56°.25
( or) ar (4 4 )| cos 56°.25

cos 45° + (8;)

= [ (ts) + (42 | cos 45° — (ge

A. P. 8.— VOL. XIX. E.

34 A NEW METHOD OF DETERMINING

Then

8(¢,+¢5) =A” + A’

8(¢4—«¢;) =A-+ A”

8 (¢; + 3) = BY’ + B’

8 (c¢;— ¢3) = [A— A” + B+ B’] cos 45°
8(¢,+¢.)=B”—B

8(G— en) = [4— A” — (B+ B’) | cos 45°
8(¢,+¢)= A” —A’

8(¢,—G® ) = B—B’

8 (s, + 55) = C+ C”

8 (s:— 55) = CO" +

8 (83 + $3) = | D + Di —(C— Cc”) | cos 45°
8 (s3 — 83) = D’ + D”

8(s;+ $1) = [D+ D” + C—C"] cos 45°
8(6¢,— sn) = Di — Dp”

8(s; +s) = D—D’

8(s,—s )=—CO'"+ C..

The expressions for the determination of the values of c, and s,, just given, are
found in HansEn’s Auseinandersetzung, Band I, Seite 159-164.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 35

CHAPTER II.

Derivation of the Expressions for Bussew’s Functions for the Transformation of
Trigonometric Series.

Ge 20 ° . °
-The value of (5) given thus far is found expressed in a series of terms the argu-

ments of which have the eccentric anomaly of the disturbing body as one constituent.
But as the mean anomaly of both bodies is to be employed, it will be necessary to make
one transformation ; and the next step will be to develop the necessary formule for this
purpose. HAwnseEv, in his work entitled Entwickelung des Products einer Potenz des
Radius Vectors ct cet., has treated the subject of transforming from one anomaly into
another very fully ; what is here given is based mainly on this work.

Calling ¢ the Naperian base, and putting

= GUE. | = Gua".
we have
yy’ = (cose + /—1 sin e) (cos e+ /—1 sine’);
also

yy” = (eoste+ f—I1 sine’) (cosv’ e+ Y—1 sin?’ é’)

= cos Ge— 7 e') | f= sin Ge —7 &’').

Denoting the cosine and sine coefficients of the angles (¢e—7 «’) by (G75)
and (7, 2’, s) respectively, the series

F=>(41,c) cos (¢e—7 & )—=ES V—1 (4,7, 8) sin (¢e —7e’) (1)
ean be put in the form

FH=1353 §(47,c) -V—1 2,8) } yy”. (2)

36 A NEW METHOD OF DETERMINING

In a similar manner we get

= £55 G2) = I= (Gi®))o 2 (3)
where
aon
We have now to find the relation between y and z.

Let

g = the mean anomaly,
and ¢ = the eccentric anomaly.

Then from
= e—esineg,
introducing »4/ —1, we get

(i SS Qala aie eal

Since
B= ine ==,
we find
gV—l=eV—1—S(y—y").

Now from

yao"
we obtain

GNM = ae. 2,

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

and
= (y—y7) = log. (2 O—¥).
Thus
g V/—1 = log. z= log. Gc2 Gq us)

and hence

YC (Ce)
From

z=y.c 29-9"),
we have

PG te).
and

yf =e ve ZU) |
Let 5 be denoted by 4; then

C- 7) io Aa. y . ch " ye,

and
ex9—-Y7) = GY cH Bey”,
But
chy cr a(l—m.y + yp — Pe y + oy! Fete.)

Ase) 2 ie ee
(1+ ha.y7 ae ol te peel! + rear y+ ete.)

(4)

(6)

(8)

(9)

38 A NEW METHOD OF DETERMINING

and

% - e ye. )2 5 5373 5 Fs 4
Oo wr! = (L+ta.y +57 +53. Sy! + etc.)
a 4.0.

+
‘e e- 22 5 3 8 5 y4 )A Es ?
(l—ia.y “tS Yor ae Ot eed ‘+ ete.)

Performing the operations indicated, we have

a ee Hea es
NO te roe + pose Fete.)

he2s [Poe Joe ge oa
(4a — ra + pes — jogs ete. ) (y =4)

(+ ty —aas + peas Fete) (ety

hii

)
(+ ee re re a= ete.) (y— 4)
Cee)

hm jm ee? nis -
als : (1 ant a 1.2.m-+1.m-+2 FP ete. )y

tes a2 vA

-é -1
OD (yy y= 29 2
2 =1— 77+ eee EEE: -++ TapaeE fame ete.

43)8 a5 75 ve

(+ Di ra ) 293 — 12234 = ete.) (y =)

Se ee

(jee a)
(toe OS) ey)

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

. . ° e é =
As we may write h in place of 7, we have, thus, also given the value of c’2(¥—Y")

Now put

chy

sy—-y)

(yy) = ae J_in oO"

(<2) (—m)
nz

+00 (m)

—— ae Dane Wc

Then, from the preceding developments, we see that

(—m) m (m)
Sinn = ( ees 1) : Tin )
(m) m (m)
Jin = ( eae 1) Sin p)
(—m) (m)
—hr = ha

Again

(—my

(0) (=

ae Sin - = Jin =F J_,

1) (=2) (=3) a
np OOF spin oO) “ae dino? “se ies

(1) (2) (3) +
qe wine y She J 1+ ye + Sn. y + ete.

Sea (Ne (0) eb) O. ©).
aes) ls 0 7] = Jinan =F Sin 9 Yy =F Jin 0 OF + Jin 0 y° ain ete.
Cas eo). oS) .
ap Cin 00) = shiny a0h Sachin e0h a Ke
O +a _(—m) ne 5
Comparing the values of =_, J_, .y~™ and c-"2—y")
we have
(-1) (1) h378 Aes hi ts
In =In =lA— aay 1 eg erga cE ete, fory”,
(1) (1) hz hope hin ;
cat Tin = Tin = hrA— 72.2 ate 923. «12.92.37.4 == etc., for Y>
ae ©) We nis neas ‘
Fin =In = ig E23 + pose + ete, fory”,
(2) (2) A222 hye AEE A
Jon =In = iz vost poe t ete, fory,

ete. = ete. =

etc.

(10)

(11)

(12)

(13)

40 A NEW METHOD OF DETERMINING

(m)
Tn -y™and chxY—y™)s

we get the same expressions for y” and y~”.

@ (2)
We see from the values of J,,, J, , ete. found

+o
oo)

Comparing the values of >

above, that the

term is
(m) hinym him+2_jm-+-2 Aim+4 Jm+4
hr = Sn (2 2-2 SS ete.
1.2...m 1?.2..m.m--1 V.2?...m.m—+-1.m-+-2
eee We hin
ri a ~ Im+1 + 1.2.m+1.m+2 a ete.)
Further, we have
I; e 4 (m) ;
eg —c va(Y—y diy" Shs Oma

and, by putting m= h—7,
this becomes

sa iy 0 yf!
Let
+ 0 (h)
a ae (Oh OP

|
ee
Y= > aatlees neo}

Multiplying the second of these equations by z~". dg,
we obtain

+o (2)

yz" .dg=> FP, .dg.

Integrating between the limits + 7 and — a,
we have
(7) 1 +7

PS y- 2%". dg

Qa —T7

general

(14)

(15)

(16)

(17)

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

From
z= 0"! = cosg + /—1 sing,
we have
dz = (—sing + /—1.cosg) dg;
also
z/—1= /—1 cosg —sin g.
Therefore

dz=2 /—1 . dq,

and (17) becomes

In like manner we find

e . (h) e (2) e
Comparing this value of @, with that of P,, we obtain

©) (i) (i)
ee io ll Leas at De Le

A. P. S.— VOL. XIX. F.

(18)

(19)

42 A NEW METHOD OF DETERMINING

Thus we have, between the mean and the eccentric anomaly, the relations

(h—1) ;
gl — wh. i y |

ea ae (20)
y= 7 Um 2 |
Tn the application of these relations, since
: (7) ;
Dy ape SIPs» e aie
the expression for /’ is changed from
F=4333 §Gt/c) -v—1 Gi, 8s) yy
into
= ISS Ei,e) Vl @H,a)) oS Pee
The other value of /” is
F=433 $3 ((4,4,c)) —v—1((Gh,s)) y'. 2.
A comparison of these two values gives
Oo Ry ee) aaa a UO 9)
((4, 0) == 2) IPSs (47,6) = 2.5, Su (4, 7, €) (21)

In transforming from the series indicated by (7, 7’, c) into that of ((z, /’, c)), it is

evident that h’ is constant in each individual case, and 7 is the variable.

Thus we find, beginning with 7’ = h’,

y (WW) iil (—(Wt—=1)

) :
(Gl! .e)) = Me shine (G50, @) Se = 3 din (2, h’—1, c) + ete.
h h

eel Gi—(-+-1))
— ial ite Siu @ (fe IL, Cc) + ete.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 43

To transform from ((4, h, c)) into (2, 7, c)
we have
(—i') _(W=7)

(2,0,@0) =O (2, lo c)) = Dhow KG h’,¢)).

Here, 7 is the constant, and /’ the variable; and for the different values of /’, begin-
ning with h’ = 7,
we find

(0) ((’—1) —7))

(40,6) =JSyv (4,0 6)) + Serax ((47—1,¢)) + ete.

((/+1)—v))

> Fussy ((%, v +-1, ¢)) + ete.

The expression

(mm jm nei aii re
= weet (1 soe _— : + ete.)
1. .2..m 1.m-+1 1.2.m-+_1l.m--2 1.2.3.m+1.m—+2.m-+3

(m)
enables us to find the value of -/,, for all values of m.
A simpler method can be obtained in the following manner :

Patting c's -Y) in the form

(1) (-1) (2)

aie) ges © (—2)
JEP? = Y—S,.-¥! +5.c-Y tJ .-y + ete.

— é e€
hy hy

we have, for the differential coefficient relative to y,

e ae nety—y) (1) (2) (1) - be (2) A
he(lt+y) c's HJ, 2.+2.5,..yrete +S ..y?— 2S, ey ete.

If we multiply the second member of the first equation by h{ (1+ y~), we have
an expression equal to the second member of the second expression, and by comparing

the two we find

(22)

44 A NEW METHOD OF DETERMINING

Let
(m)
Je
[prada ean Ts
yn (23)
hey
then
(m) (m—1)
= fOr
hey pee
From this general expression we find
pe ca
= aD
hs ie
(2) (1) (0)
I pS k pa aah ni (24)
hy hy hy
= Go = Ee
(m)
: : g hs + sates
From the values here given, since —Gey 1s put equal to p,, we have, by increas-
hs s
ing m by unity,
(m-+1)
hs ae
may — Pm: Pri

Putting a = Tm, equation (22)
Com

takes the form

Pm » Pm Ste 1 = Tn = Pm:

From this we find

1
), = —
Pm Tn— Pm+1
z 1
Tn — — 1

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

We also have

1 —
(25)

Pm = Tn — Pin+19

a form more convenient in the applications.

(
The general expression for J, . 1s
Coy

(m) (0)
Te =e Pi- Pa Ps--- Pos (26)
2 i
where
(0) oat, E I!
Te =1—p+ pa pm + ete, (27)

if we put l= Ad.

From the expression
oe (ty Puta A Ww)
(GUs@) = Sizan (EO) => 7 dine (GH)
it is evident that when h’ = 0, or when both 7? and #’ are zero, this expression cannot

be employed.
To find the values for these exceptional cases let us resume the equation

When hf = 0 we have

46 A NEW METHOD OF DETERMINING
The equation
1
a a =O)

gives

ee = ons (1+ y~) dy. (28)

Hence

@ l grt

== i—1 € ae e i-2
ah we eee oe
When p is a whole number
+ny—1
ie or, ay = 0;
J oony=1

except when » =1, when this integral is 27./—1.
Hence it follows that

When 7 = 0, we have

Using the expression

pe (2) « s > (=e) e 8, (EF) . :
(G hie c)) = SJPag(G050) = an (6058) + ap (GU 12)
(—t-+-1)

se Jeg (2, v— it. C),

we have
((0, 0, c)) =(OF0K0) eae 2n/ (0, 1, ¢)

for the constant term, the double value of this term being employed.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

For h’ = 0, we have

(GOe) = LO =w de) = 27 GE abe)
(LO, 9) == (G08) = 20 GG 1h 5) yy Gal)
(ZO,e) = @Oo— 27 Co) — 2) CS 10)
((2, 0, s)) = (2,0, s) — (2,1, s) — a’ (2,—1, s)

ete. ete.

In what precedes we have put

and obtain

g = the mean anomaly,
é = the eccentric anomaly,
c = the Naperian base,
Bu
= Clas

a yf". ols ¥—-y »

ee 2G ae
yi = 2. oY y’),

he(y—y) + : ‘ ewe
where ¢2U—Y") ig expressed in a series, the general term of which is

we hig! ney
hr oe (1 rae ee ) ram
l.m--1 1.2.m--1.m-+2 2.3.m--1.m--2.m-43 SE Ces) Gy)
Thus
| I hin hers
h h mam

B= 0 APOE (A — ee

y lm+1 1.2.m-+-1.m-+2 1.2.3.m—--1.m+2.m+3 35 ON 10)

We have also put

and since

Offre py +00 _(—m)
eh Y ) — Dan ea . Oe

(m)

ca YY *) =s" my 00S

(—m) (m)

Jin — Sin ?

48 A NEW METHOD OF DETERMINING

have found
(m)
gh — Thx 2 ima : OP
(i=)
=In -Y;
if
m=h—i.
Again supposing
; +o (h) ‘
— 2
= O) . Of
+o (é)

y' = 2s JP, ° Zz

we have found

Thus we have

(h—t)

ee ;
S=dhx oh

(h—2)

= Sn [ cos de + sin ve »/ =,

fase RA
y = h Jin °&

(h-~)

— sdhn [ cos hg + sin hg v= :

Hquating real and imaginary terms, we have

z 7 h=o (h—1)

cos te = =. 2,» Um - cos hg,

(29)
h=n (h—)

sin ve = 2 Ji, « Bin hg.

h=-o

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 49

We notice that

x) (-1)
dE ) S = — 36,
(0)
0 == alk
For all other values of 7
(i)
We a=)!

If a large number of the -/ functions are needed they are computed by means of
equations (24) to (27), as shown in the example given in Chapter V.
If we wish to determine any of them independently we have from

(m) _— pmjm we na i he, 28
Ne aL Tem ) L.2m-lm+2 — 1.2.38.m2lm2m+3 = ete. |,
fe Ge
Fe= li-F Gta a a S|
ie = ae pie oe ect | ) (30)
mote D Peta
oe be

In these expressions we have written for @ its value 3¢.
(m)
Since h has all values from hk = +o to— o we find any value of J, by at-

tributing proper values to h.

From equations (29) we find the values of the functions cos Ze, sin ze, in terms of
cos hg, sin hg, and the J functions just given; always noting that when 4 = 0, we
have only for 7 = +1, — $e as the value of the function.

We can employ equation (22) when only a few functions are needed, or as a

check.
A. P. §.— VOL. XIX. G.

50 A NEW METHOD OF DETERMINING

It may be of value to have y’ in terms of z" and the J functions.

ond of equations (20) we have

(0) (a) (2)
yPs=—AaAt+t+ J, .2 +43,.2 +43,.2 + ete.

(2) oO . (Or
== dh oF linn ain Oe:

(0) @ (2)
yt=—Aat J .et+4dy.2°% 4+ 4d, 2% + ete.

(2) (3) (4)
2

— J,.zg —td,.2? —t4d,.2? —ete.
- (1) (0) | iO .
Op —2SJ,.2 +2d,.2 + 2J,.2 + ete.
@ Oates eae
—2J, .2*— 2S, . 2° — 2d, .27* — ete.
ze Pace bee ee) Maa. ch He
ye = — id, 27 + 3dn. 27 + 2S3,.27 + ete.
(3) on oO
— fd, .2 — gdn.27 —2J3,.2° — ete.

Then from
y' + y~ = 2 cos ve
=o S2 Jl. si ve

we find the values of cos «, sin ¢, cos 2¢, sin 2e, ete.
In case of the sine, as for example when 7 = 1, we have

y—y i =2/ —I1 sine; butinze—2z1=2/ —1 sing,

From the sec-

we have the same factor, 2 »/—1, in the second member of the equation.

From

r= a(1—e cos €)
we find
7

(=) = 1 — 2e cos ¢ + & cos

e: = 1+ 2e cos « + 3¢ cos *e + 4¢ cos *e + ete.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 51
2
For (") we have

Gy = 1+ de?— 2¢ cos e + de cos 2e

But

d [(r* : de 9 .
"\ = % 5 = 22 sin
a (3) 2e sin e (1—e cos « ) ay sin ¢,

and
“ (0) Oa - (1) 8), (2) o>.
am eel wh oh sing +3[ Jy + J, | sin 29+ 4 | Jn + J;, | sin 3g + ete.
Multiplying by 2e.dg we have for the integral of i (“)

” 26 (0) 2) Qe (a) (3) j Qe (2) (4)
= 0 J, th cos g——| Joy. + J, | COS A J; +3, |cos3g— ete.

where c= 1 + 3e’.

By means of (22) this becomes

r\2 (1) (2) (3)
(“) =1-+ $e— 4J, cos g —4J2 cos 2g — 4; cos 3g — ete.

In case of (om we have
3é . cos «= $e (1 + cos 2e), 4¢ cos *e = ¢' (3 cos ¢ + cos Be),
be’. cos *e = 3e' (3 + 4c0s 2c + cos 4e), 6e?. cose = 38° (10 cos e + 5 cos 3e + cos 5e),
Té° cos °e = x5e° (10 + 15 cos 2e + 6 cos 4e + etc.)

and hence

(“)°H1 +438 + A + We + ote.
+ [2e + 3e’ + $8 + ete.] cos «
+ [3e + 20¢! + 195¢° + ete.] cos 2e

+ [é + 2% + ete.] cos 3e

+ [Be + 42e'+ ete.] cos 4e

52 A NEW METHOD OF DETERMINING

Attributing to 7 proper values in equation (29) we find the expressions for cos ¢,
cos 2:, cos 3z, ete. We then multiply these expressions by their appropriate factors and
thus have the value of ¢ Ne. Wer

+o _ (—2)

@\= = | ae 19, | (=) = Bue FR; cos 1q-

ic)

(2) (—2)
The following are the values of R; and FR, to terms of the seventh order of ¢.

au 3 5 7
Lig SS Bee EO ag?” sb ae
(2) 5 ‘ ,
fy = —te + 1e— Ae
©
a 1333 9 81 7
Li Bae be = ae
x 4 6
Ry, = —te'+
se 25 op BOR A
—— Oe) f 7
Ti; = — Pa 7e ap aes’
(2) ae
R, — — soe
> 2401 77
fi, = — geivaye
re 1 4 5 6
Rh = j= = 1+ 64 344 Le + ete.
V1—2 4 8
Gee 88 1 6595 | 2675¢7
— 38 >) 9 5 pf
Ei, = 2e+ Fe + FR + Ze ane

ly SO Be -p Be
peice 103p4__ 88 76

1 == ga GF yarn?
Oe 1097¢5__ 1662147

5 192
ie 12236

6 — 160 ©
jhe 472787

1 = “agg

See HansEn’s Lundamenta nova, pp. 172, 173.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

)

(2) (—2
We add also the differential coefficients of ;, ; , relative to e.

dk, 3
= v0é
de
IR,
a —— 8e2 __ _5 ¢t 7 6
= = 2+ 36 — 3 + ane’ F ete.
ae.
2 — D8 il)
A = = Oa Be eS SE
e
dk,
— 3 p2 5 pt i; 6
es Ae ee ee
de
dR,
4 = —2é + 4¢ Fete.
de
dR,
ores 4 5 6
i — _13te 4 t8hte' Fete,
IR.
GLb, = D7
= —2le ete.
de 40 S=
dk,
fp eats 168076
We = — x30406 = ete.
etc. = ete.
ip,
= e+ dé + 436 + 1956’
é
aR
i — 2 4 25,0
= — 9, 4 e€ + 325 ¢ + ie é
é
Te,
a = 43 |. 6399
Zo be + 46 +- $3¢
Rae
— OY 2 5 4 F 6
= — 39 125¢! + 215 1¢
é
ane
in NS BS
= +276 ne
de 4()
(—2)
dR, — 5485¢t__ 11634768
Ae 4608
(—2)
dR, — 36696
d —: 8
é
(2)
aR, — 830911,6

54 A NEW METHOD OF DETERMINING

The value of = found by integrating a(“) = 2e.sin e.dg, is

2 (1) 2) (3)
r
5 1 + 3e — 4J, cos g — 4J,, cos 2g —4-J;, cos 3g — ete.

(2)
In terms of the #,; functions,

(2)

2 (2) (2)
“=1+3€—R, cosg — R, cos 2g — R, cos 3g — ete.

Again, since

Oj __ @
apa oe
we have
2 aes Sir V1l—e* dg
Let
+0 ee
F=J + =, Crsin2g ;
then
- =1+ ae 7C; C08 219,
and hence
(—2) 1. C;
ki; a V1 Gi

The coefficients represented by C; designate the coefficients of the equation of

the centre.

* THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 5d

Using the values of the C;, coefficients given by Lz Verrizr in the Annales de

V Observatoire Impérial de Paris, Tome Premier, p. 203, we have

f—g =[4G) —2G)?+ $6) + WG) + PE (H)"] sing
“F >Gy— (Gp) ae ag) se a ap i | sin 2g
ts Gy — 4G) + SG) — 24 G)+ ete. | sin3y

Gece ce eG) eect, 9 || sini4y
+ [1982 (ey — pet Cy + tpn (4)! | sinbg
qe: it G) === G) +, ete: | sin 6g
=e [432328 (Gy SS eP | sin 7g
+ [2H? G) ] sin 89
r [4 Lee Ge) | sin 9g

Converting the coefficients into seconds of arc, and writing the logarithms of the
numbers, we have for the equation of the centre,

4
.
4
+
Mi
i
cs
ae
4

| 5.9164851 (5) —5.6154551 (5)° + 5.5362739 (5)° + 5.787506(5)' + 6.25067 (§)° |sin g
[ 6.0133951 (4)? — 6.179726 (5)! + 6.067753 (4)° + 5.59571 (5)°| sin 2g

| 6.252272 (5) — 6.6468636 (5)° + 6.690089 (+)’ — 6.22336 (¢)’| sin 3g

| 6 5491111 ($)'\— 7.093540 (5)'+ 7.27643 (4)*| sin 4g

[ 6.875105 (¢)'— 7.533150 (4)' + 7.82927 ($)°| sin 5g

[7.225760 (4)'—7.96973 (4) | sin6g

| 7.587638 ($)’ — 8.40484 (5)"] sin Tg

[7.95944 (4)°] sin 8g

| 8.38880 (5) | sin 9g

56 A NEW METHOD OF DETERMINING

CHAPTER III.
Development of the Perturbing Function and the Disturbing Forces.

By means of the formule given in the preceding chapter, the functions pals),

acre (4)"; etc., can be put in the desired form. The next step is to determine the com-

plete expression for the perturbing function, and also the expressions for the disturb-
ing forces.

If # is taken as the measure of the mass of the Sun, and m the relation between
the mass of the Sun and that of a planet, the mass of the planet is represented
by mk’.

If x, y, 2, be the rectangular codrdinate of a body, those of the disturbing body
being expressed by the same letters with accents, the perturbing function is given in

the form
a=.) 2a]
Now
IS = (Wa) =e (=) a (C= 2)
=p tr? Dr’. A;
hence

oO= [bas 2]

ItmtL4 rl?

If a © is regarded as expressed in seconds of are, and if we put

SST nes oa CG). (eo

1m
we have

GQ = Me Gi— @):

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

Finding the expression for (ZZ) first by the method of Hansen, we let

A= = .k.cos (I—K), hi =". COS p. Cos 9’. k,.cos (II — K)
1=*,.cosp.k.sin (I—K), U=*,. cos 9’. k,. sin (11 — K)),

and have, if we make use of the eccentric anomaly,

(17) = h.cos oe cos f’ —ch(“)*.cosf’ 1. sin €. ( i)? COSN i

+ 1.cos aa) pins ats op) )\.. Sle +h’. sine GC) _ sin

cos ¢! 7 COS ¢ 7 cos ¢!
Putting
\ 2
(2) cos f’ = y’,.cos gy’ + y’,. cos 29’ + y’;.cos 3g’ + ete.
a’\2 sin f’ ; : Fi , : , pees ,
(“) oa a= Une RING) ae 0’5. sin 2g’ + 4,’. sin 3g’ + ete.
- ¢
we find

(1) = $ (hy’, —'8,) cos (— g'—e) + 3(ly’, —U0',) sin (—g' —e)
—ehy’, cos(— gs) + el’, sin(— g's)

+ ally’ +884) cos (gy — 2) + 3(/'1 +15) sin( g’—e)
; (1)

+ 2(hy', —h'8’,) cos (— 2g’—«) + 2(ly’.— U8’,) sin (—29' — «)
—4.ehy’,cos(— 2g )+ 4.eld’, sin (—29’_ )

+ A(hy’s + h's'2) cos ( 2g'—e) + Aly’. + U8’,)sin( 29’ —e)

+ ete. + ete.,
where
(0) (2) (0) (2)
— y

04 = Jy, nN 9 DP i IN? === CAN
(1) (3) (1) (3)

= 3] Sa » |» 2 = 3 Jn eae: » |

ete. ete

A. P. S.— VOL. XIX. H.

58 A NEW METHOD OF DETERMINING

‘When the numerical value of (#7) has been found from this equation we trans-
form it into another in which both the angles involved are mean anomalies. For this
purpose we compute the values of the -/ functions depending on the eccentricity, ¢, of
the disturbed body just as has been done for the disturbing body. The values of the

(0) (1)
J functions can be checked by means of the values of J,,, J,,., given in ENGEL-

MAN’S edition of the Abhandlungen von Friedrich Wilhelm Bessel, Erster Band, seite
103-109, or by equations (30),.
Thus by means of the equation

(m-+1) a (m—1) en (un)
TN hr =< hd A

(m) (0) (1)
we are enabled to find -J,, if J), J), are known.
It must be noted that the argument of BrssEL’s table is 2.5, or 2.hd, or he.

Go)
Thus if it is sought to find the value of -/,,, we enter the table with 2.2 or 2e as the

argument.
When we need the functions for h from h =—1toh=4, we must find the
4 a 4 HO 2 (—2)
yalues of a se» ad es ates the een and — td.

(1)
The values of 3. J.

(0) (3)
and J, we take from the table. ‘To. find J. - we have
oy z

e
oy

oe (1) 2 Fe
a ne 4.5) 4g
(4) 2 of 1 7°
Pe ae eae le 46 1 45 |
(2)
For J... we have
as
(2) (0) 1 (1)
SI og SS td eae) et
385 385 3 85
(2)
And for J, we have
z
(2) O 7
I, — J, ale
z z mo

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 59

The expression for (JZ) can be put in a form in which both the angles are mean
anomalies. ‘Thus, resuming the expression for (/7),

(Ca Fi cosie Gr cos f’— ch Gr cos f’—1.sin « (yr .cos f'

7 a\2 sin f’ a\2 sin f’ 6 a\2 sin f’
+U.coseé C) F su — a (“) a= £ +th’'.sine. @) a

7 cos ¢ r cos * cos g’’
in which
a
hk = —.k.cos (i—K)
h' = E- . ' k TI TK ee | vcos V
Vv = —. cos p. cos 9’. k.cos (I— Kj) = gu.
p C 5 1 v sin V
(=i COS) Pr. k.sin (11 —K) = fu. aie
a -
U eee “yaa , k 9 (11 Pabek K,) ee | p cos Ie
= ae CUS Gp : ,- Sin 1) = 3u.—=,

: a’\2 a\? sin f”
we find the expressions for (<) cos f’, (5) as as follows. We put as before
¢

@) ws" = Dar COs gy aie Y's cos 29’ + Y's GOs 3g + ete.

'\2 sin f’ 5 ‘ : F BPS cu eae,
(*) am = 8 sin g +0: sin 29’ + 0’, sin 8g’ + ete.

c 0 7 , O
If we differentiate (, cos f relative to q’ we have

A
d(G.cos 7’) = cos fl dr’ eis Tid sin f’ Cia sin /”
dg’ a ala al” fala cos v?
; dr’ ae’ sin f’ df’ @®
since —— cae —_— 7 + COS
dg cos g dg ip
and hence
a? G cos f’) a!

ees SS cose

Similarly, in the case of

is (i) ,
But a cos f’ = cose’ — €,

Hence

Now

From the values of cos é’

- COS ni

Pe snin i/”
; COS @

We now

A NEW METHOD OF DETERMINING

r’ sin f’
pon. we have
cos g
a (G sin i Poesia if
dg” \a' cos ¢ 7? cos ¢’
r sin f’ .
and * f = sine’.
a’ cos ¢
a G cos f” ) 7 @.cose
Sk, a 12 cos T ee 12 9
dq” 2 : dg”
2 (7 sin f’
1? (S =) _. @? Sin 7” __ &. sine
dg” > 7? cos ¢ dg

COS &’

sin ¢’

Bec
0) (2)

[woot oh

assume

jh + Ee) cos g’ + tlio = Te cos 2g’ + ete.

i (1) OA.
[ee = ee sing’ + | Av + “he | sin 29’ + etc.
and sine’ we have

"| cos 9’ + 2 eee abt cos 2g’ + 3 [ Ja — Say_| cos Bg’ + ete.

| sin g’ + 2 lez + i | sin 29’ + 3 [wee ++ Toe | sin 3g’+ ete.

(i—1) pte 1 i—1) a,
x ae a Jin ik 0; =i ees Si ]
1 (v’—1) (7’+1) 1 (v/—1) fh +1)
(/
i [. va —Siny ii ver | Jim +m i

Comparing these expressions for y’;, 6’), with those found in the expression for

sin fe
“cos

, given above,

we see that the relation between them is 7”

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 61

The expressions for cos ¢, sine, are the same as those of cos ¢’, sine’, if we omit
the accents.
Hence if we perform the operations indicated in the expression for (47), we have

ay (2)

Tr a
= 30” [hy yy + Wd; Wy] cos (41g — Vg!) — 30? [By Ely 0'v] sin(Lig—’g’) (2)

é and v’ having all positive values.

Attributing to 7 and 7 particular values, we find, noting that 6, = 0, and 0’, = 0’,

(1) = 3 [heyy + W004 Jeos( g— 9')— 3 [byi t+ lydi] sin( g— 9’)
+ $3 [h.nyi—h'd,0, ]cos(—g— 9’) — [By — 7184] sin (—g— g’)
+ gh. cos ( — J)— flys, sin( — g’)
+ 2 [h. yy’, +h’. 8,82] cos( g—2q’) — 2 [h.by’.4+ U'y,8'2] sin( = g — 2g’)
+ 2 [h.yiy’2—h’. 30’. cos (— g — 29’) — 2 [1.d,7’.— V'y,0’.] sin (— g — 29’)
+ 2h.yuy's Os ( — 2q') — 20. yoo sin ( — 29’)
+ 3 [h.yy’3+ h’.60'3] cos( g—3q’) — $[0.by7/3+U.70'3] sin( g—38g’)
+ ete. — ete.

+ $ [h.yoy'1 + W’.6,0.] cos( 2g— 9’) — 4[bb.y. 40.7281] sin ( 29 — 7’)
+ 3 [he yoy + h'.630',] cos (— 29 — 9’ 3 [Ld —U.y20',] sin (— 2g — 9’)
+ ete. — ete.

The numerical value of (#7) given by (1) must first be transformed into a series
in which both the angles involved are mean anomalies before it can be compared with
the value given by the equation just found.

If we find the value of (H) from the preceding equation, it can be checked by
means of the tables in BrssEv’s Werke.

The expression for « (“) is known; and with the expression for (/7) just given,

we obtain the value of
Ce On" i (“) —().

The next step is to obtain expressions for the disturbing forces.

62 A NEW METHOD OF DETERMINING

Let v the angle between the positive axis of X and the radius-vector measured in
the plane of the disturbed body, here called the plane of X Y. The differential coeffi-
cient of the perturbing function © relative to the ordinate Z perpendicular to this
plane is found by differentiating © relative to z and afterwards putting z = 0.

Thus from

m J rr’ i
(Oa Le |;

ae cle

1tm

AO = (Gaal Pe (Qa P se (ee)
r - r? aoe ory’ #,

we find
dQ m [- Wy 4p &: |
dv 1+ m 2) ky > ihe
dQ mm [— 1 Gate A
dr ~~ Vem E 4 ) Pl?
mm 1 9 CB
dQ = aoa [— = Ua =|
dA dH dA dA Z
== = Hf = A— = r—?H, == = — =,
dv dv’ dr 2 dz 4
Hence
dQ m’ 1 1 ery
du 1+m Ls a wr id
dQ m’ il m’ 7
TLidh € Raye (Asem = aller A yn
GQ __ mm 1 1 a iapieg ae ois Ay j
@ = tole ja |sin 2.7 sin (f’ + I’)
where

HY = sin (f +11) cos(f’ + TI’) — cos J cos (f+ UH) sin (f’ + I’),
2 = —r’.sin Jsin (f’ +541’).

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 63

As before the origin of angles here is at the ascending node of the plane of the dis-

turbed body on the plane of the disturbing body, and the plane of reference is that
of the disturbed body.

E ; : dQ dQ
If we differentiate the expressions for ip? - yp we find
1 OZ

o @2 AQ ml

Sa pO aT TNS
dp le ~ Bl vr Hf)

eo -=o) ipa nae

1=Em \ 2 Tage ° oP
r a ee : = (7? — rr’ HZ) sin Tr’ sin (f’ + 1’)
a = a ( 7 =) sin Z.r sin (f+ 1)
x os =— ties : = (7° —rrH) sin 7.r sin (f+) + S
ae —— =e : = .sin “J. rr’ sin (f+ I) sin (f’ + I’) + aon (4-3) cos L

To eliminate H from some of these expressions we find from

2 nat Al 2. )
Y= + r?— err’ . A,

that

: dQ
The expression for Uae then becomes
ar

dQ a rey? 1 r i |
Oise 1m 248 2A te

From the value of A’ we have, further,

PoaTp Tl | eee 1
i = uae DE?

64 A NEW METHOD OF DETERMINING

and hence
C2 3 om (pi 1 : : 3
=o Bre SG ree seit ‘ f _/ ‘ 7 IV
drdZ 2 1+m [ ae al eu a su (f 1 )
HQ nv i 1 : : dQ
7 ete ea sib [a — — | sin J.rsin TI —
drdZ’ 4 iba AP | (f+ ) da’

dQ
becomes

the latter of which, by means of the expression for 77,

RD Vo SS 1 | 2 . Sab ss Po
= 2 aap 7 3x| Sm Ir sin (f + I) im I va Sin (PE TD)

The expression for A’ also gives

(P—rr Hy (ry 7S fe i
OM Sr ah 44 2H 44?

by means of which we find

5 AQ _aQ m | 3(r?—r*)? r? 1 m r
ee = eal eee oe le

ip Ts 1m 44 1+m 7”
If we put, for brevity,
(T), = = sin [ ey sin (iF? si Il’)
(ry = sin (2) (2) sin (FED

(= z cos 2 (2)

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 65

the expressions which have been given for the forces, together with the perturbing
function, are

=—po(s) 2 Zain Ge ery

9 3 sin r’ . 2
+ $ua2(4) S22 7 sin (f’ +11’)

¢ oe | = Buai(“)’ 2 =. = sin *( f’ + Il’) — u(“)

aa (77) =0e(G) eG sin (F + 1)

dQ a\SF 2? 1. 75) sin ip .
Call ZV Se A) is egy ee r
ae Dee 4 a”? a2 a sin Gi oe Il)

— Sua? (“)" pesintd = sin (f+) —(Z)’
aa (<,) =— Buai(*)”, eo: - “sin(f + Il’ )= sin( f+ 11) + ua (4 cos FT _ (dD

The form given to these expressions is the one best adapted to numerical compu-

tations; and the equations are readily derived from the preceding in which the magni-
tudes occur in linear form.

Thus from

2 m’ py? 1 -
pe = [=e — an Fz]
dr 1+m 24° 24 r?

A. P. S.— VOL. XIX. I.

66 A NEW METHOD OF DETERMINING

we have

where, as before,

In a similar manner all the other expressions for the forces have been derived.
When we compute only perturbations of the first order with respect to the mass

we need the perturbing function
CO SS (4) — fH
and the forces
dQ 4 Ee "|-3 ON .
are, = eS) Lam oa] — eG) —

qe — — pa2(“)" ae 2 sin (f+ Il’) + (1).

The other forces are only needed when we take into the account terms of the sec-

ond order also with respect to the mass.
An inspection of the expressions for the forces shows that besides the functions

(> wal(G) » wai)

we need expressions for the magnitudes

r!\? i 7 git I 7 6 , , sti JT Rs a
G)o gm a gin F 41), = 7 ain (f+ ID;

(BD), (i, GBe, Gay”.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 67

‘When these are known we multiply the function ua’ (ey by

r\2 ean sinI 7. , , sinir. .
Le —i. eal =. osin(f +1), eT ain (f + I),
(a cosl .

a’ ? a b)
: 5
the function ua (=) by
@ |r lr]? 3 sinlr’ . ; ar \ene Ig
alae pale oa “sin (f+ WW) [7 — 2 a;
aie J PR ay ae 3 sinl 7 7 il

pe E stale a) em (Gg? Sone

We will now find the expressions for (7), (7)’, (7)”, and for the various factors
just given, that are the most convenient for numerical computation.

We have
(1) = “sin (2). sin (f +1’).
Putting, for brevity,
b = — *, cos ¢’sin J cos Il’

6.) = /sing/esinplil
3

and noting that

a/\? sin f’ (0) QF. @- LOA.
() ae ms = Fe + J, ] sin g’ + ole = The sin 2g’ + ete.

(2) cos ie eee Tet | cos J! ++ a Fl cos 29’ + ete.

- 68 A NEW METHOD OF DETERMINING

we have

(0) (2) (0) (2)
=o (+a lene a o Loe les

(a) Or). qa) (3)
+26 [ Jar oly | sin (— 29’) + 28’ | J, — J _| cos (— 29)

(3)
(2)

(2) (4) (4)
+ 8B [ Tye + Sou | sin (— 8g’) + 8’ [ey — Fay | 08 (— 89’)
+ ete. + ete.

The value of (Z)’ is found from

(Ty = “sin (2). 7 sin (f + 0).

From
_=l— 2 as,
a
we find
a’ = , \—3
6) = (= excos ea
a
Expanding,

ANS oil pene ;
(5) = Se + (de + 22e + ete.) cos g
+ (%e”-+ Ze* + etc.) cos 29’
+ 43e" cos3q’ + 231e cos 49’ + ete.;

which, for brevity, we write,

Gy = ~ + 2p; cos g’ + 2 p, cos 2g’ + 2 p; cos 3g’ + ete.

But

, ‘sinof (0) (2) Es () (3)
~ r= [EA +4, |sng+4 Ba + JS, ] sin 2g + etc.
(2) (1) (3)

7a (0 3
“cos f = —e+ [ee pete cosg +4 Jn. — Jn | cos 2g + ete.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 69

Putting
— = . cos @ sin J cos II, = a sin J sin,
© @ 0) (2)
a= oh a oy) a, Se Oh
(1) (8) (0) (2
ye eos | Beer Bs cer
ete. GUC,
we have
(Ly = — 3 be. po
+ l. 9.6, sing +- L,. Po. 71 COSY

+1.p,.d.smn( g— g') 4 h.pieyicos( g— yg’)
—l.,.6,.sin(—g— 9’) +1.—:% cos(—g— g’)

— 2lep, cos ( — g') (4)
+l.p..6, sn( g—29') +h.—.y, cos( g—2g9’)
—l.p,.6, sin(—g—29') + 1.2.7, cos (—g— 29’)

— 2he.p. cos ( — 29’)

+ ete. + ete.

For (7 )” we have the expression

Ci) Econ te( Se

Putting
pope = cos J, and using the p; coefficients as for (Z)’,
we have
(Dy = © 4 1. pcos (—g’) + hy. pr c0s(—29') + ete. (5)
To obtain an expression for the factor [G)— = a | it is only necessary to

have that for (Ne

70 A NEW METHOD OF DETERMINING

In terms of the eccentric anomaly we have, at once,

Tr 2 9 ©
(“) = 1—2ecose + € cose
= 1-4 $6 — 2ecose + 3’ cos 2e.
Substituting the values of cos «, and cos 2¢, we have

r\2 5 a) Q) (3)
(2) = 1+ 3¢—4J, cos g— 4-J., cos 2g — 4-J;, cos 3g — ete.

To find an expression for the factor 8 Lis z sin (f’ + Il’), for brevity, we let

sin I sin J

= . cos 9’ cos IT’, — .sin Il’,
a a
: 7’ sin f’ Y 5
and from the known expressions for a sae A cos jf’, we get
gia JT Fo ; (0) (2) s (1) (3) :
= (S00 (jae IN) = [ee + Sy, | ¢, sing’ + 4 [ey =5 Ja ¢, sin 29’ + ete.

a a

(2)

(0) (a) (3)
— $2G + ee —vJ, |e,cosg’ + le ae | c, cos 2g’ + ete.

In the same way, if

: ae
CG = Sin T cos @ cos il, e, = 2+ sin UH,
we find
Shall ~P _s : : _(0) (2) : a) 6) :
— poly =p) = J, +J, |essing + $| J +2 | sin 2g + ete.
a ~

6
athe (0) (2) @ _6) (6)
— $ee,-+ [a —J, |c¢,cosg+3 | ahs + Jo. | c,cos 2g + ete.

By means of the expressions for the factors

r\2 f in I yr’. % j in I - S 5
(Z)i, — ; 5 sin (f’ + Il’), Pelee : . sin(f + I),

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

just given, we can form those for

Sie 1 =
4 La? 0) a a
@ Gin de HF 5 ; rl? 1 2
7 oe gl (f +11) | 5 |
1 af2 A
g mt | sin’ (f +1’)
3) simi 7 Tr? iL 72
A aD la ae |
sin 7p ee iF oo ; .
3 oe Sl Pe) SG Slay)

72 A NEW METHOD OF DETERMINING

CHAPTER TY.
Derwation of the Equations for Determining the Perturbations of the Mean Anomaly,

the Radius Vector, and the Latitude, together with Equations for Finding
the Values of the Arbitrary Constants of Integration.

HLANSEN’s expressions for the general perturbations are

dW, 9
_ dz + ¥* |dt

Megs = at + Jo + mo f | Wo +

= ae Ae:

dk, — if = dQ
her’ sin (@ AF 7) C08

aM ah} 2 (0) G0) Se [cos (f —o) —1] ‘4

+ 2h, ©. sin (f— 0) r(=).

In this chapter we will show how these expressions are derived from the equations

of motion, and from quantities already known.
The equations for the undisturbed motion of m around the Sun are

iF 9 Xx
= PUM) = =0

“1 +e 1+m)4=0

<_+#d+m)5=0

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 73

The effect of the disturbing action of a body m’ on the motion of m around the
Sun is given by the expressions

2 (uv —x ce qe2(y—y y ey ae 2!
tee Eg 2x) Ee — a):

Introducing these into the equations given above we have in the case of dis-
turbed motion

ix 2 NBEO tat Shae aoe =)
aa t+ (1 4 m) —; = TV Hi a

dy 2 ( yo > yefy—y _ y (1)
ae + k#(1+m) = =m k ee ¥)

a bs

1 9 / > Zz i) 9 z'—2z tf
oi + kh (1+ m) 2m ae —— =)

The second members of equations (1) show the difference between the action of
the body m’ on m and on the Sun. The action of any member of bodies m’, m’’, m’”
ete., can be included in the second members of these equations, since the action of all
will be similar to that of m’.

The second members can be put in more convenient form if we make use of the
function

(Ove nv’ (' kei eee)

tee WA Pe

Differentiating relative to «

CO an ( 1 dd =)
dx AB ale Fas”

But since

we have

Gia x —x
dz” Ame’

A. P. S.—VOL. XIX. J.

74 A NEW METHOD OF DETERMINING

and hence

In the same way we derive the partial differential coefficients with respect to

y and 2.
The equations (1) then become

d2
dx

“2 4 eA+m)2 =P +m)

“4A +m)4 =e (1+ m) | (2)

= + (1 +m), =k (1+m) =

Let X, Y, Z, be the disturbing forces represented by the second members of

equations (2),
R, the disturbing force in the direction of the disturbed radius-vector,
S, the disturbing force, in the plane of the orbit, perpendicular to the disturbed

radius-vector, and positive in the direction of the motion.
If f be the angle between the line of apsides and the radius-vector, the angle be-

tween this line and the direction of S will be 90° -+ f. We then have
Ka—Ssnj, v= S cos j-
In case of 2, we have

R= xX ee,

and for S,

From these we find

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 75

If we wish to use polar coérdinates we have

d° = Roos f — 8 sin f
daz :
2 5
= = Rsin f + S cos f,
From
=P COS ih, TST sm,
we find

dx = dr cos f — rdf sin f
dy = dr sin f + rdf cos f
a= dr cos f—rd7f sin f — 2dr df sin f — rdf? cos f
@y = drsin f + rd°fcos f+ 2dr df cos f— rdf? sin f

From the expressions for dx and dy we find

dy cos f.—dxsin f= r df
dx cos f.+ dysin f = dr,

and hence
da 1 Ge de
— ™ —-— .—~ sm — GO
ae peg hs te Oey
do 1 ag ae
—= 2,60 == ii
dy pa COR ae a BID

from which we see that

R=VA+m) 2 saPa+m)i@

If we multiply the expression for d’w by cos f, that of dy by sin f, and add,
we obtain

ax cos if a @y sin f — Gp df.

76 A NEW METHOD OF DETERMINING

In a similar manner we find

dy cos f —d@asin f =r df + 2dr df.

Operating on equations (2) in the same way, we have

S 3 COS f + 5 sin f + Eee = X.cosf+ Y.snf=F

a cos f — sin f = Y.cosof—X sn f=S8S

Comparing the two sets of equations, we have

ref, odr af 3 1
ap 0 Sap ar =h (1+ m),

(3)

ir ay” ed+tm) — 7.
a aa

The second members of equations (1) and (2) are small, and in a first approxi-
mation to the motion of m relative to the Sun, we can neglect them. The integration
of equations (2) introduces six arbitrary constants; and the integration of equations
(3) introduces four. These constants are the elements which determine the undis-
turbed motion of m around the Sun. Having these elements, let

a) the semi-major axis,

nm the mean motion,

go) the mean anomaly for the instant ¢ = 0,

é the eccentricity,

$) the angle of eccentricity, —

nm, the angle between the axis of « and the perihelion,

v, the angle between the axis of x and the radius-vector,
jo the true anomaly,

é the eccentric anomaly.

These elements are constants, and give the position of the body for the epoch, or
fort=0. Let us now take a system of variable elements, functions of the time, and
let them be designated as before, omitting the subscript zero, and writing x in place

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 17
of m. The former system may be regarded as the particular values’ which these
elements have at the instant ¢ = 0.

In Elliptic motion we have

nt + go= e—esine
r cos f = acose—ae
rsin f = acos¢sine -
Dan
an? = k’ (1 + m)

Now let mz be the mean anomaly which by means of the constant elements gives
the same value for the true longitude that is given by the system of variable elements.
Further, let the quantities depending on mz be designated by a superposed dash, and

let the true disturbed value of r be given by the relation r= r (1+ 7).
We have then ;

Ne = E€—eQ sine
r COS f = A COS E — AE
r sin f = a cos oy sine
Ve vi + 1
ayny = kh? (1+ m).

We will now first give BkRuNNow’s method of finding expressions for the pertur-
bation of the time, and of the radius vector.

Neglecting the mass m, multiplying the first of equations (1) by y, the second
by x, we have

ye = {( Ya— Xy) d+ ©

C being the constant of integration.

Introducing

_ x cS a
cos f = *, and sin f = =

78 A NEW METHOD OF DETERMINING

into equations (2), neglecting the mass m, we find

2, Fe} f
Ore k?. cos f 2Sey

a th 7

@y 4. Wsinf _ y (@)
dé! pe
We have also
= = age
a) = la joe + reosf. os
and hence
ee = es
or
eS = ( (Ye — Xy) dt + C;
and

ae = f Sr.dt+ ©.

In the undisturbed motion we have
Gin ——
rT). oe =k Py

po. being the semi-parameter.

Hence
Po r= { Sr. dt + kv/ po

ON Oe

THH GENERAL PERTURBATIONS OF THE MINOR PLANETS.

From these relations we derive

ie i
oe oe in {Sr . dt,

and also

es oss af Sr. dt
VP ky/ po) 1/p

If we eliminate + from equations (4), noting that

» af pee ak a
fe = ih Se elt
di NOE pe rae oe

we have

— ee =2(|[¢= a EGP | at

fy _ See =/((r= ue _ Sr | dt,
neglecting the constants of integration.

Since r =r (1+ v), we have also
a=ax(l+»), y=y(1+>).

The equations (7) then become

= a da ksin f _ sin f
eee ia hinge =f (X—=E. 8r)at

— d ») YY. _ Beosf _ cos f :
Dore let er ae =J (V+. Sr) at

From the equations

L = My COS E— AL, Y= Ay COS Hp SIN ky

(5)

(6)

(7)

(8)

80 A NEW METHOD OF DETERMINING
we have
dx = —d sin «de
dy = d COS >. COS ede.
é
Then since

dg = ” de, ee, “dy, “oe he as
dz hr? Vv.

using the values of sin «, cos ¢, in terms of sin f, cos f, we find

dx ae ke sin f dy __ cos fe,
Ge = 1/ Det dz ~~ V Po
And these give
bsinf _ _ da V Bo
VP sae VP
k cos f __ dy VP. key
vp ae yp VP
— dy YR her _ (% Sr dt
dz VP VPs Pp

The equations (8) then become

= ao da dz “V/ Do sin f 2
~+ =[d+% ver] = == S42 Gaya

~—dy st Ge V Po — : cos fe, :
Yue dz [ato ) dt eS eye) ae

(9)

MEN qo : 5
the constant — Ye being included in the integral.
J

pe will now voor equations (9), and for this purpose we multiply the first

by 2 7, the second by oe “ , and noting that

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 81

we have

== sen Lt {(x- sae Sr\dt+ ae ee jens aah) Sr dt (10)

Now multiply the first of (9) by y, the second by «, putting for | Ps its value
V2
given by (6), noting that

we have

sae nar Sr)dt
(
2 (11)
+ poe f (H+ Eee. sryac

c Czas
We can write Spe the form
;

a ue v) = —( +7. 24¥.%

lt dt
We have

na df dz df a
aby) = eS Ss Se LS oe
(EY) ae al CE GR GG ?s

df 2 AS) oD

df = pe eo COS Po, UN = Ay Nh.

dt r

Making use of these relations we find

de
di (C55) Pyne

ale
and for 5, given above we have

Zz lz y
@ = Al +0). 2 — Ve + Ve,

A. P. S.—VOL. XIX. K.

82 A NEW METHOD OF DETERMINING

The equation (11) is thus changed into

a= 1— Jo f(1 +24 H) srar— 2! J (x! sryat

dt ky Po key/ Do
(12)
Te COE os Sir ee WA
1a ele ) a apy V Po
The equations (10) and (12) can be put in briefer form.
Let
Rex Bs VS s:
Sass A b) (Bi 5 -
se 1D
Then
a COSTAE (5 Lv Gye
ae J X,dt += J Y, dt,
(18)
dz 1 ) Qy >
cae ee UEN. Bio gy a (Pe eee 7
dee loa mA (+ ) Seat Ed tS || Y.d
The values of «, y, found in these equations we get from
cA + 5% 1 ce os
x= a + —"(2—t) + 4.5 (e—t) + ete.
dy . Wy, \2 CS)
Y=Yot Fi ee t) +4. ae (g—t) + ete.
! df
From the expressions for = ; a , we have also
Az Zz
cos f +e, __ ue — Ki 10% :
Gay LW € ee de ic t)) UC:
(15)
sin f 1 fds, PI fgg ;
Sa a oe CO ce

The quantities given by equations (14) and (15) are found in equations (13)
without the integral sign. They can be put under the sign of integration and regarded

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 83

as constant if we designate all magnitudes in these factors dependent on ¢ by a Greek
Jetter.

We thus obtain

( = 1 aa 9 at te A
eS t) — reais fa + a =) Sr dt — A J (26, DG 2) ali
we ee tea) thee (16)

- =| (2S i ee
dt ke y/ por dt dz ke 1/ Po © dz?

These equations include terms of the second order with respect to the mass. If
we put

vs ae (ere ve pond ce Wee sre jr

we get

ngz = net + gy tm | W + +r dev] at
f (17)
v= N—43( [9% 4+ 0 be] at

In equations (17) g is the mean anomaly for £=0; JV is the constant of inte-
gration in the value of ».

From the value of JV given above, we have

IW 1 Pi 2 es ae
i ae (Lae ‘) Sr — —=(X,.v— Y,.8).
dt hy/ po VP hy/ po °
Now since
X = cos foe — — gin Joa ~
¢

) 2
= oes d
; ar < if

df
12
a
dr
Si 1 dQ

84 A NEW METHOD OF DETERMINING

neglecting the common factor k (1 + m),

we have

dw 1 ( VPo\ de 2 do = Ry Se esa
all oe) eer meee ak
dt key Do dy ky/ Do dr eee y r sin f df ;

2 dQ. q Uae F 2 sin f dQ (cos f--e,) dQ |
—— ae 2 ae | SBE ee (cos eo aa
r ky/ po (7 sin nf a cos f) bor ki/ Po L p wie tks p df 5

And as
v =psinoa, c= 0 COS,
this becomes
a = [ (—1-2 a a2 2 sino. cos f. “2 +2 .psinosin f. 2
dt ky/ po Vp? df df Say.
+ 20 cosa. sin f. ey 2p . Fy 0080 008, f + 2p Sine sin 7
aL yp a f ae. sl . 9 COS @ a)
= yy (32) ie +208 (0) oP £04 9? oa (f— oie
QZ cosa (jf gf = 2e,.” cosa =|
p P df
But
26,9 cos a. — 2p.” i= = oe “(6 p COS 0 — py) = ps8:
also
= 5 k= se

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 85

Hence since k? (1 + m) is included in X, Y, R, S, we have

is 20. hh 5 ds
— = fh, [ze cos (f —o)—1+ tee (cos (f—o) — 1) | af

(18)

+ 2hop.sin (f—o da

If we write h,. a cos *@ in place of & in equation (18), we have the same ex-

pression for = as that given by HANSEN.

Equations (17) and (18) are fundamental in HansEn’s method of computing the
perturbations. We will now give Hansen’s method of deriving them.
Using the same notation as before, we have, since

a _ 1+ecosf
;

cos *¢
also
r cos? g,
Geeele=e\costac
hence
ra _ 1+ecosf COs *¢,
dae cos ch lemencosias

Using f + ™— x in place of f, and developing, we get

r.a __ r+reos f.e cos (y—m)+r sin f.e sin (y—7)
Pity A, COS “Gy

Let us put
esin (y— 7%) = 1 C08 "hp,
(19)
€ cos (y—™) = £ cos*q + 43

since €é = sing, we have

C08 "p = Cos) (1 — 2ey E — cos *y &* — cos “hy 7”).

86 A NEW METHOD OF DETERMINING

With this value of cos*?@,and r= a, cos “py — &7 cos J,
we find

Pr. __ dCos*e,—e,.71 cos f --r cos f (E cos *¢,--e)--r sin -4 Cos *¢,
Pi My) COS *Qy

__ a cos*g,--r cos f. cos*g, +r sin f.7 Cos *¢g, ,
Ms COS *¢g, (1—2e,E—cos *¢, & — cos *gy7 2

and hence

ai saritdh, aiken
1+&—.cos f-+-7.— . sin
mh a :

7. 1— 2e,—cos’¢,€’— cos’,

From
GO Ui, df dz
a tb @a ° de”
and
df __ ky/p(.--m)
ae r ?
we have
df a
di. — %- » - COs >.
In like manner we find
lf 2 ;
— n=". cos op.
dz }
We have therefore
dz

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 87

n . ( “a 9
If we put |» =1-+ 34, substitute the values of ——, and cos *, we get
) VT.

r D U0. TONG)
él ance : ooo 2

(1—2¢6,&—cos *¢,5’— cos *¢97")?

(20)

dz
qa =U + 6)

Further, in the case of », we have

1lt+v= =

=~
Then since

OG 2 9 n
Oi = Osos 5 = (1 + 5b),
0

and
cos * 2 £2 Da
£ = (1— 26, £ — cos oy &? — cos 7"),
COS "gy
we have
: 1—2e,E—cos *¢,.’—COS “gy.
G7) = Panne: 26 rare
(1+ —cos f+ —sin f.7)* (1--6)8
CP Ay
If we let
a - iP °
Al = 2 eat +—.sinf.7,
a My
B= 1— 2¢,& — cos *@, &? — cos *y 7”,
i GeauOy
hy a BS ?
we find

diziares: Ai ') = Fea
ae = (ae) fap (Lar?) = Fae

88 A NEW METHOD OF DETERMINING

From the latter we have

(a) =1-20 + FS + +8. a
Hence
Gy oS) Ca,
Fie tae
If we put
Wee i <2 a mn 2. “6 S cos f + a. ne sin f,
we have

& 14 W+h(e).

lnc
We have yet to express |, in terms of the elements.

From
ie SO 5 ‘ 9 cos ?
Sil 3Q2 — 24 cosy. sy SS
COS 79,
and from
iy Oe
ie GB”
Vi
1) S—= 5
No
we have

h = n y COS ¥o
a “cos ¢ ’

(21)

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 89

or
kh an cos %
ly COO Cie,
If we put
he
Y cos @,’
we have
h re an
~ ¢os g

These values of h and h, being substituted in the expressions for WW, - is found
expressed in terms of the elements and of », in a very simple form. To find the rela-

: dz 0
tion between a and v, we use the equation

2. B °
(1 + v) —— A*(1-+0)! Z

e e h
and as this is also equal to eee

“dt

we find

ie tk 1
dt sh (A+)?

(22)

For the purpose of keeping the formulz simple and compact, HANSEN makes use
of the device of designating the time, and the functions of the time other than the
elements, by different letters.

Thus for é, r, «, f, 2, v, & Y, we write,
T, 0, Ny @, 6, B, & v, respectively.

Whenever we integrate, these new symbols are to be treated as constants, noting
that the original symbols are used after integration.
Ao TE: SATO, SAD, Th.

90 A NEW METHOD OF DETERMINING

If in equation (21) we introduce 7 instead of ¢ we shall have

dg cae j h, B 2
Sait +5), (23)
where
ro h hy h P = h rn ee
Ul Shs — 7 sb D8 = ots m ob 2 yo ES SN.
; h, h hy ay h, Qy
We have also

dé at: h, (24)

de ~ nh(l+p)?”

The codrdinates of a body vary not only with the time but also with the variable
elements. In computations where the elements are assumed constant, that part of the
velocity of change in the coérdinates arising from variable elements must, evidently,
be put equal to zero. Coérdinates which have the property of retaining for them-
selves and for their first differential coefficients the same form in disturbed as in undis-
turbed motion, HANSEN calls ideal codrdinates.

If Z bea function of ideal codrdinates, it can be expressed as a function of the
time and of the constant elements. Thus let the time, as it enters into quantities
other than the elements, be itself variable and, as before, designated by tr.

The function dependent on ¢, 7, and the elements we designate by A. Then

ay da
Gig — Ghe 2

dL = (--\at

where the superposed dash shows that after differentiation 7 is to be changed into ¢.

Let us write the equation (24) in the form

+ BY = 7

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 91
Differentiating relative to 7, we have

at
GH dv’ ;

The differentiation of (23) also relative to t gives

CGO WENAG
d@ d& dt

hy 2B dp

aly ib” (1-Ep)8 dz :

Eliminating = by means of (24), we have

de dW 28 dg
Lr ee de 1+8° dr*
dt ;

Substituting in the expression for ae we have
LT

dp dw

dt ” Ge

Since » is an ideal codrdinate, we get from this

es 8 if (© at, (25)

IV being the constant of integration, and the dash having the same signification as
before.
This expression for » is a transformation of that given in the equation

1 — 2e,€ — cos *g,.€’— cos *¢y.7?

ITtyv= G@LEpyi@e mera coseen
a

“im 7)
ay
Since z is also an ideal coordinate, we have from (23)
es is ea
raj = rob + G+ mf 1 W +5) \ dt (26)

g being the constant of integration and being the mean anomaly for ¢= 0.

92 A NEW METHOD OF DETERMINING

When we consider only terms of the first order with respect to the disturbing
force, ¢ changes into 7, and we have

rie Una es tf W, dt
ae (27)
J

yo N—}h (=) dt
where
Wy = 27 — #1427 £2 cos +27 .n./ sino, (28)

and p and are functions of +, being found from

Mt +G = xy7—EASiINY
p COS @ = A COS Y — A &

psIN@® = a COs sin 7.

Also in the last two terms of W,, . is put equal to unity.

‘When terms of the order of the square and higher powers of the disturbing force
are considered, ¢ cannot be changed into t. In this case let

Nyt = MT + J + ndz.

Likewise let
M6 = MT + Jo + 206

where

no¢ is a function of 7 and t.

According to Taylor’s theorem we have

W = Wi+ 2 064 4S 80 + ote.

the value of W, being given by (28).

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 93

We then have

GUY GN WwW, 1 dW, .
dg a 3 dz: 206 SP Sa ie 06 + ete.

Retaining only terms of the second order, the equations (25) and (26), replacing 6¢
by dz, give

MZ = MtE+ 9+ to f [Wot . de + | dt
yaaa f(s 2M sda

The equation (26) has been put in simpler form by Hix. For this purpose from (21)
and (22) we have

(29)

Mt) ere=$—a4m

Hence

Developing the second member and adding JW, we have

M2 = Mt + Go + NH f Se dt. (30)

The next step is to express sale and = in terms of the disturbing force. From (19)

we find

TY
|

~~ e082, C08 (% = 7%) — COS",

4 = ——.sin (y— m).

94 A NEW METHOD OF DETERMINING

Using these values of £ and z, and pcos @ = 4,cos*$)— p, in equation (28), we
find

20 20 h
W, = —" — -heeos (y —m—o) + —*+_ -h— 2 — 1.
” Taga, COS 7, G=a=es hqa, COS*¢, h

Since

k= an __ kj/ltm

cos ¢ VP

we have from the expression of / already given,

Fi+m)
(amar a

re
di

By means of

h _ Gar
~ cose ”
we may transform the expressions
du a
= —.. G08
di ®
dr an .
—— = é sin
dt cos ¢ I;

into

r. —h=cos (f —o) .he cos (y — 2 —) + sin(f—o) .hesin (y —™— @)

+ =sin (f —o). he cos (y —2%— 0) — cos (f —@) .he sin (y —™—«@)

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 95

Multiplying the first of these equations by cos (f— a), the second by sin (f —o),
and adding the results, we have

he cos (y%—7™%—o) = (re —

—h) cos (f—0) +2 sin (f—a).

Substituting this value of h.¢.cos (y—7,—«) in the expression for W,, noting
that

1 pe
hyd, Cos*g, — #(1-+m)?

we have

7 — _2hop _ pl. 2h,.p fey OT,
y= Fim) Reon a) TT 7 + Baum .sin (f- 0)

(eas (Ft) ST

cay? J, COS? go

Differentiating relative to the time ¢ alone, 7 remaining constant, and haying care
that all the terms of the expressions be homogeneous, we have

dW, 2hop dv 2hp a&r
= —o)r ‘ ae
aE ae (f—2) = + Batm in (f—o).—, a
re ++ dh
= cos’ ¢ 5, le aS Gh a) 14 SFR
and
dh 04m) @e PF de
aan a) dae OED
Y ha —
dt
Substituting

96 A NEW METHOD OF DETERMINING.

we have

Wy — f 2h’ rr dQ
Te = hy {2 cos (f—0) —1 + , [eos (Fa) —1]} (#2)

(30)
+ 2h sin (f— 0) » (47)

: P dW. See :
This expression for 7, is the one used by Hansen in his Awseinandersetzung.

It is given in a much simpler form in his posthumous memoir, and as the latter is the
form in which we will employ it, we will now give the process employed by HansEn
to effect the transformation.

Substituting first the value of h, omitting the dash placed over certain quantities,
noting that in the posthumous memoir ¢ takes the place of o, and remembering that
we are here concerned only with terms of the first order with respect to the mass, we
have

Ss | 22 cos Go) 1+ a [cos (f —o)—1] ew

VA 1—é?

an Po 7k e dQ
te ass oa (f o) (4)

From the relation
p = al —e’)—ep cosa

we have

Pp von ep COS w

a(1—e*) a(1—e*) ©

. : dw 5 : —
An inspection of the value of “q_ Shows that its expression consists of three

parts, one independent of 7, the other two multiplied by p cos o, and p sin a, re-
spectively.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

Put

dW _ d& dY
= dt

5 ole
( cos @ + je) oe Binkon
a I dt a

de a = [ cos f e cos f 1 | ( =) aesinf ( (
n.dt V1—e r a 12? =P ie df aR = of
ME 5 4 | (ee a. (cos f--e) le a r(
oe Vie r ae df neues
DE BX DEG | eae a sin f ie = a cos f r (
jock; — /l—é 7p =e df Z :
But
4 2 = s 5 a 1
a ay eee cos f ¢ eae ! =a
dg i ry 1 (l—e')3 (ee
dr _ ae sin f
i a
dia (@ 1 )
aE : ae ioe ) sin Fs
Ghp
ao ee cos f;
hence
ne dQ
ndt 3a ee )s
Ce 2 f dQ 1 dQ
oe Ge) ye
ndt é dg V ie" df
oo
Wale LE de]
Again from

Gp =a) Gal

A. P. 8.— VOL. XIX. M.

(a) (i)
dr dg

98 A NEW METHOD OF DETERMINING

we have

(2) ay (7°) a =n (2) resin f
df) Nil) Me dr/ a(1—e?) ©

Eliminating = from the expression for — , we have
na

dV 2 | a (1—e?)— 7? 7d r sin f dQ
= ala) ae)

jel; ~~ ae dg a/1l—eé dr

In the same way we find

CP 2 P 2 sin a cos f e sin ?f
mile | i sys Wee ()— r vI—e+ V1=

resin*f =
a (1—e’)? we )

But if we employ the relation

as r recos f
— ad) * aie)

f /1—e’, of the preceding expression, the whole term becomes

r cos f e re dQ
alle (1—e? yo ‘1—e? a a ail ae )

. @ COS
in the term,

Using the equation
0=—recosf—r+a(1—e),

multiplying by

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS

adding to the preceding, it becomes

[ r cose a 2e | (“)
=— Oe ©
a y/1—e Wg Ne dr
Further, we have
cos f e sin?f resin ?f
m= Vl1—é a (1—e?)3

Gd TP .. r’ sin
== || = Sun =-cosf /1—@
dg Lis eas a (1— re lak Oy Ay ae
Reducing this expression in the same manner as employed before, it becomes
2rcos f+ sae

7? sin
dg _sin Ha aia l= ale

Multiply this by dg, the last expression for Fae becomes
recos f+2 ae

dQ
= (7) ar (
a 1/ 1—e* ar] ”

d¥ 2 2rcos f+ 3ae
= ——— g a
1—e?’ Jj ay/1—e* J dg

the integral to be so taken that it vanishes at the same time with g

dY dv
> ndt? ndt’

Substituting these values of <.—
dW dz dY¥ /p j :
ndt — ndt a ndt e GUS @ a e) +° ndt a sin 2

this expression can be made to take the simple form

7) + Bar (|)

dw
= adele

in which
; fe 2psinw 297 *
f ( cos f

a’ (1—e’) —1

a y/ l—e?

: .4 (2% c08 o + 3¢) ae

r sin f 2p sino (F cos +2¢) bo

1 2p
J G cos @ + 30) ———— ——
a a / 1—e* a7 \—e*

(31)

+ 3e) dg }

100 A NEW METHOD OF DETERMINING

Since
@). 7 9 rsin f
(POdg — @@/l= Ee”
a 7
= — 2- cos
a e.de J
we have

(oe pal
Be To anal ee - San ley ae ; 8e | oP

These expressions for A and B can be much simplified.

Thus from
"=1+}¢—(c—} 2) csg = G2 @) eran — 3 é cos 3y—“ cos 4g— eles,
2

tnt . p
and a similar expression for —, we get
- a”

ad.

ae.dy =(2—{) a ie

d.
a .de

— 3¢e = —(2— 2) cosy,

a = @—) sin g + (e— )sindy +26 sin 3g + 3€ din dg 4 ein,

f [ 58 | dg = —(2—?e’) sing— (5 — °) sin 2g —— sin 3g — ; en 4g —ete.,

be— (2—7) cos g — (6) cos 27 —* cos 8y —“ cos 4y,

—4e= —e —(2— 2 €) cos g— (e—3¢) cos 2g — 26 cos 3g — 2’ cos 4g.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

From which we obtain

A =—3 +(4-+ 26’) cos (y — g)

+(e+ -) cos (y — 29)
—(5e +

2
WS

e
+5
3)

25e°

; ) cos y

cos (y — 3q)

cos (y — 49)

+o +20)

B=—(24+ &)sin(y—g) |

—(e + “) sin (y—2q)

7 3

—(e + =) sin y

—* sin (y—39) (

= ze sin (y —4q)

eae
aE 5, Sin (y + 29)

101

(32)

These are the expressions of A and B whose values are used in the numerical compu-

tations.

When we have the coefficients of the arguments in which y is + 1, and —1, we

obtain the coefficients of the arguments in which y is + 2, with very little labor.

é dW :
Let us resume the expression for a , that is,

dWiee
ndt ~~

‘3 a *
Since - can be put in the form
a”

dQ
dg

Aa(

A and B having the values given before.

Yr
—~ = >R™ cosk
a w q,

— — >" Rin kq,
é

Tr Ose
2— cos f _

) + Bar Se)

CP d Ri)
de de

cos kq,

102 A NEW METHOD OF DETERMINING

and

SY (e ) Bef da = Sie sin tg +8 9 — Beg:

But since

m= 1 + ge — (2e— fe’ + sige?) cos g — (he — ge! + aige*) cos 2g
a?

— (4é — e’) cos 3g — ete.

we have

Hence the integral just given is simply ae sin kg.

A and B can then be written

1 a(1—e’) —r? Q2psinw dR) ,
A=—3 al (26 oso 3e ) 5 —
eas - +P Te aoe ie oe kg |
2 psi dR)
a p OF, hoe Fh Attlee 9 |
B= 5 [ (22 cosa + 8¢) Sk R sin kg ofle\ cos kg e

Putting

2
{= > R™ cos x y,

we have likewise

Oe = Te a oe 2° sino = v 1a Ss 2 RO Sinz
—COS @ Les — aR aR cos XV 5 7 oo = edy —e — 7 Y:

Introducing these values of 2 ” cos o, and 2° sin @ into the expressions for A and

BL, after integration relative to y we can write W in the form

VSS oo ae) (xy 4+ Bt ) ;

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

where

Bt = ty + ey,

Uand V being two functions depending alone on ¢.

Putting x = + 1, and —1, we have

(1) (1)
Gs ee ye ey
de
d RY R® 1,
(1) = ae ei:
Qa =
de e 4
and hence
aY 1 aD a® — gD
C= dk® ? v= R®
de Ce
Thus we find
[ dR) dR)
de RW de RW =
Oi AC eae Gari | GE — 2 oe jes
de de
or putting
dR«)
A de Rie)
a 9d RO + x aR®
de
dR)
de Rix)
Ql) = 9 ER® —2.4 Ro 9
de

we have

a® = aM + 9M ag,

103

(33)

104. A NEW METHOD OF DETERMINING

The values of 7) and 6“ are readily found from

f=14+3 @—(2e—té4 74 6)cosy—(se —1 eH + gs &) cos2y
—(4¢é— &) cos 3 y — ete.
> ee Neosiziy.
We have

BOSilpee
R® =—(2e—Fe 4+ ge &)

)) — iL 4 6
RO=—Gé—fe+ we)
R=—Ge—he )
Cy = ete.

d RO

Ge de
d RY 2 Ba 6
5 ae (CO a a)
GLO So, Ih 8

de S—(0@— 3 e+ ee)
ad k® 2 4

Ge 7 (Zé ay cz Ze)
GO 2 BAB
de aa G € ¢ )
ete = Cte:

For 7” we have

(2) Crs? Car 3)

G
" —G=1e4%9 C

SS Og ca, Oar Oe @ a ee)

(9 Se he ade, (34)

For 6 we get at once

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 10

In a similar way we have

(0) = BP Sa ey
il = ts @ T258 > 7

Col
a

(35)

In case of the third coordinate we also compute the coefficients of the arguments

having no angle y from those having + y. For this purpose, putting x = 0 in the
expression for a’ we have

d PO) ad RO a -+- a)

OF SS eS Oe) (—1)
Cy = We apart CLE 1g = 4 (4 + o );

where

9 de
n —
B d R®
de

For 7) we then have

(0)

72 =—(Be+ Ye + etc.). (36)

Perturbation of the Third Codrdinate.

Let 6 the angle between the radius-vector and the fundamental plane,
@ the inclination of the plane of the orbit to the fundamental plane,
v —o the angular distance from the ascending node to the radius-vector.

We have then
sin 6 = sin 7 sin (v—o).

If we use for 7 and o their values for the epoch and call them 2 and Qo, & being
the longitude of the ascending node, we have

sin b = sin % sin(v— Q)) + 8;
s is the perturbation.
Thus we find

s = sin 7sin (v—o) —sin 4%, sin (v— Qj).
A. P. S.— VOL. N. XIX.

106 A NEW METHOD OF DETERMINING

Putting
p=sin?i sin (oc — Q)) , g =sin 7 cos (o — Q)) — sin %,
we find
s= qsin(v—Q)) — p cos (v—Q)).

Instead of s, let us use

and we have

n= 7gsn(@—Q,)— £ pars (=):
Q FE: a ;

Introducing 7 and calling #& the new function taking the place of u, we have,
putting © + 7 for v, 2 being the longitude of the perihelion,

a@E dgp . dp p
=a 7, 22 (© =F %)—= Qy) = =F @, COS (@ + 7 — Qo).
d dp : : i s
To find and = we will employ the method given by Watson in the eighth

chapter of his Theoretical Astronomy.

Thus « and (@ being direction cosines we have

A SO =— B O3

also
2 = rsin? sin (v—o).
But
T = 7 COS ¥, and ¥ = Tr sin v.
Hence

2 = —xsin? sins + y sin? cos o,

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 107

and

a=—sinzsino, (6 =sin7zcoso.

The values of p and qg then are given by the equations

p= —a cos 2, — sin Qo,
g = — asin 2 + 6 cos Q — sin
from which we have
dp da. dp
a = — GCOS Qo Fh SIN {2 Fe
dq 3 da dp
Ros SIN {2 as + GOS a:

From the equation z, = a « + @ y we have, first regarding a and ( as constant,
then regarding # and y as constant,

dx 5 dy
Giseetes

dz, da. Ap oe
al = 0 FF SP I) ap SO
Differentiating the first of these, regarding all the quantities variable, we have

az da dx dp dy au p GY
= oa
dt’ dt dt dt dt dt? dt?

Z, being the component of the disturbing force parallel to the axis 2, and X and
Y the other two components, we have

Z=aX+ 6 V+ Zcosz.
Writing for X and Y their values

vx
di?

+h (1+m)”, ated +m)s,

108 A NEW METHOD OF DETERMINING

and reducing by means of

Z=an+ By,

we have
ped ZL He
A= “+ (+m) * + Zecosi,
or
V2, on ae i
de = Oe + be = Z GOS 2.

V2,

Fae given above,

Comparing this with the other expression for ~

we haye

da da dp dy __ Z 2
Fie Tein Tach Gia a
0 ° dz, 0
From this equation, and the value of ak since
¢

d 2S Sep
2 yi =kv pl =e

we find

da.

Fi =—hrcosisinv Z,

da hrcostcosv Z.
a

d
Substituting these values in the expressions for = and “! ae

we have

dp dp

7 =hrcostsin (v— 2) Z,

= = hr cost cos (v— Qo) Z.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 109

a i . dR
Introducing these values into the expression for =
a

we have

a = Pt cost cos (v — Q) 7 8n (@ + m—2) Z

—hrcostsin (v — 2) ~ Cos (@ + m)— Q)) Z

=—hreost

[ sin @ COS (v — Qo — (m— 20) | Z

—hrcosi% [ cos @ sin (v — &— (7% — &0)) | Z

dQ
= fhpeans = sim (@— 7) ==.
Hsin(o—f) 4,
é ke y/1 em ke y/1-m
Introducing » = mean mips = E :
Vv P
we have
ak ] Tr O dQ 4
a p P :
: = —£ im (©— Sk :
ndt \/ 1—e’ a aac (o f) g AIF a (37)
Let
il Pp
CSS = =n (@— 77) e
V1—e a ae (o—f);
then

dk 9 dQ
— — o> || == Iho
cos 7.ndt C ee

To find an expression for C similar to those for A and B we have, first,

1 po. i p Jape
oe sO sie. conf" cos. - sin f |.
\/1—e’ La, a My a

110 A NEW METHOD OF DETERMINING

= e ° iP Tm e a ° . °
Substituting the values of —cosf, —sin f, given before, and similar ones for
a a s

® cos o, © sin w, we find
My My

Ca ( d.p* ) (<7) mr. (oe) ()
ma: aedg ¥ \agedg/ \ade/*

My le

Substituting the values of these factors we obtain for C’ the expression

C=(1—te) sn(y— g)

— (8¢ — 3) sin y

+ (Le— 32 &) sin (y — 29)
+ § @ sin (y—8y) ee)
+ $e sin (y—4y)
—7,¢ sin (y + 29)

. 5 d du
Haying found the expressions for and :
nat ndt . cost

we have, finally, for determining the perturbations, the following expressions :
noe =n f W dt,
dw
n f a dt,

— = (Oa (>).

tos 7 dZ.

|
n=

° : dw
Two integrations are needed to find ndz. We first find W from Fae then, form-

pa Re an) td . oS ge
ing W and — $ ip from JV we have néz and » by integrating these quantities. In

: : dw . . F
the integration of aa We Sive to the constants of integration the form

ky + k, cos y + k sin y + 9 k, cos 2y + 7°) & sin 2 y + ete.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 111

aw
— we have
dy

Then in case of —

tol

+ 3k, sin y — 3k, cosy + 7 k, sin 2 y — » kh, cos 2y + ete.
In the second integration we call the two new constants Cand J, and the con-
stants of the results are in the forms

C+ kh nt + k sing —k, cosg + $7 k, sin 29 — $ 7° kh, cos 2g + ete.

ol—

Nf —4k,cosg— tk, sing — $7” k, cos 2g — $7 hk, sin 2 g — ete.
In case of the latitude the constants are given in the form
AQ +isng+l,cosg + 7 1, sin 2g + 7° L cos 2g + ete.

The constants are so determined that the perturbations become zero for the epoch
of the elements. Hence also the first differential coefficients of the perturbations
relative to the time are zero. We substitute the values of g and g’ at the epoch in

4 u d : a ; 9

the expressions for ndz, », — , oa (ndz), ete., including in g’ the long period term.
cost” na

Putting the constants equal to zero, and designating the values of néz, v, ete, at

the epoch by a subscript zero, we have the following equations for determining the

values of the constants of integration:

C+ ksin g —k, cosg + $7 k, sin 2g — $ 4k, cos 29 JL ef, at (ndz)y = gy

= 7 Gh, ;
k + k,cosg + &sing + 7° k, cos 2g + 7° k, sin 29g + ete. + — (nbz) = 0

ndt
N — 1k, cosg—ik, sin g— $7 k, cos 2g — 4 7 & sin 2g — ete. + GO); = 0
2 9 1
+ dk, sin g— tk,cosg+ 7° hk, sin 2g — 7 k, cos 2g + ete. + a (7) = 0
i+ %sn g+becosg + 7” 1, sin 2g + 7° L cos 2g + ete. 4 i= Jo = (I

d ni
1, cos g — l, sin Gs 7” 1, cos 2g — 7 1, sin 29 + ete. + ( : y. 0)

ndt \eost

112 A NEW METHOD OF DETERMINING

To find k, and &,, we derive from the preceding
k, [ cos g —e-+ 7) cos2 g + 7 cos 3 g + ote. | aL fh, [ sin g + 7° sin 2g + ete. |
d
—3 44+ 6(v),+4 fe (ndz)) = O

k, [ sin g +27 sin 29+8 7° sin 38 g + ete. | — k, [ cos g + 27° cos2g + ete. |

The value of JV is found further on.

Having &, we find i, from

ch, 84, +3" (nbz), + 6 (rp = 0.

ko
We have »
hb=—el, N= —2h—Zh—3Z,

where Z% is the constant of W.

Let us find the expressions for the constants WV and K, A being the constant of
: : c ° : h
integration in the expression for 3 ; 6

The equation (22) we can put in the form

dz hy : a hy hy
fo ee ee
Rae ay + (3p 4y* + ete.) = 27 (5

The differentiation of nz relative to the time gives

- =1+h+4+ Z,+ periodic terms,
where Fh, = — 32.7162, in the case of Althea, and 7, the part to be added when

terms of the second order of the disturbing force are taken into account.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 113

The expression for y is

vy = IV + periodic terms.

; h ; ; ; Ii
The approximate value of ie being 1, the complete expression for the integral of d 7
U 1

is given by
hy

1 = 1 + k; + periodic terms,

k; being the constant of integration.

2 = B h h zs ees 3
Putting (3»* — 4r° + ete. = — 2» C — 1) = JV,-+ periodic terms, and substi-
U U
s A 6 F lt > 6 : dz
tuting this expression, together with those of » and , in the expression for uw?
U

have, preserving only the constant terms,
I = y, (k; —k —4— %, se Vad

It is necessary now to find the value of &; in terms of the constants. If in the
7 Gai F ; : 9 9
expression for ae given by equation (18) we write for p , its equivalent a cos “o

— & p cos w , we will have

dW, = hy} 2° co (f—0) 1-2, ee Oe) Ty es ‘e) dt

hg! di, COS *g ° hy? a, cos*y, J \df
+ 2h) p sin (f —o) ( \dt.
We also have
a =h =) dt.

Selecting from the expression for dJV, the terms not containing p cos o and
p sin @, we have

dW,=—h, (1 +27.) oe

A. P. S.—VOL. XIX. O.

) dt.

114 A NEW METHOD OF DETERMINING

If the eccentric anomaly is taken as the independent variable we have for the

complete integral

W.=kh+ hk cos 4 + k, sin 7 — hy ine de 27) (F) dt.

Introducing the true anomaly instead of the eccentric, we have,

cos w + e : sin w cos
>» sny = —___,

since cos 7 = oS
a is

1+ ecosw
= ky ke : h’ d2
Wo=khtek + {9 608 © -+- pepsi — hh J (142 =) (FF) dt.

Neglecting the terms having cos w and p sin w we have in JV the constants h
and €) hy.

ho
The integral of dj; is

h, om dQ
ait hth fi) a
hg
From the expression for d ; we find
h Wh? 7 dQ
dji=—i,l(g)&

a : 5 h ,
Integrating this, making use of the value of ;', and adding the constants, we have

h h

Qo = ==
My It

0 h? dQ
=14h)+eh—h f (1+ 2.) (ae dt.
And since the quantities under the sign of integration do not have any constant terms

Wwe can write

yh te

foe == 1 +k + ek, + periodic terms

h 5 0
a = 1 eis + periodic terms

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

. hy s : : : “
Since G — 1) is a quantity of the order of the disturbing force we have

es ee =i) . —— (a) 4 etc.,

h h h

from which we get

Now putting
hy 2 hy 3 eee
Ge — 1) = (e — 1) + ete. = H, + periodie terms,

substituting this expression and those for

oh fx ho

Fos ape aaa i Ae

the preceding expression for

gives, preserving only constant terms,
kj = —4(kh + ek,) +2 Af.
Introducing this value of /; into the expression for WV it becomes
N=—}4(4h, 4: ek, +384) +4(8V,4+ 20,—3Z,).
Preserving only the terms of the first order we have

N=—}(4k + ek, + 3%).

; = 6 Sats h
To find the value of A, the constant of integration in case of 6 7» We have
L

: =—1-+ K + periodic terms,
"0

116 A NEW METHOD OF DETERMINING
also
f = 1+, + periodic terms.

From these we get
~— 1p ele w= Fh,
Hence

I= — hk, + H, = 3 (hy + ey) + 3 HL;
or, neglecting the term of the second order,

K=3(k +e).

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. Wi hs

CHAPTER YV.

Numerical Kxample Giving the Principal Formule Needed in the Computation
Together with Directions for their Application.

ALTH HA 119. JUPITER.
g = 332° 48’ 53.2 G4 = 63.9 on 48.6
a le 542) wm =12 36 59.4 |
= 203" bol: 1894.0 Y =o 22 1894.0
*= 5 44 46 | (roll)
@¢= 4 36 249 go = 2)7 45) 50-2
nm = 855’.76428 nm’ = 299” 12834
log n = 2.9323542 log n’ = 2.4758576
log a = 0.4117683 log a’ = 0.7162374

The epoch is 1894 Aug. 23.0.

The elements of Jupiter are those given by Hitt in his New Theory of Jupiter
and Saturn, in which the epoch is 1850.0. Applying the annual motion of 57.9032
in x’, of 36’.36617 in &’, to Hriw’s value of z’, and of 9’, we have the values given

above. The mass of Jupiter is ;>57/ 575. The elements of Althzea are those given
in the Berliner Astronomisches Jahrbuch for 1896. The ecliptic and mean equinox
are for 1890. ‘To reduce from 1890 to 1894 we employ the formule of WATSON in
his Theoretical Astronomy, pp. 100-102.

v=it+ 7 cos (Q— 9)

Q=Q+ UU —t) “ — 7 sin (Q —8@) cot .2

dl ; 9
nm =n+ (¢—t)> +nsin (2— 6) tau $2

118 A NEW METHOD OF DETERMINING

where
§ = 351° 36’ 10” + 39”.79 (¢ — 1750) — 5.21 (¢ —t)
n= 0.468 (v — 1)

These expressions for 2’, ’ and 7’, can be used for the disturbed body as well as
for the disturbing body by considering the unaccented quantities to be those given,
and the accented quantities those whose values are to be found for the time, /.
HARKNESS, in his work, The Solar Parallax and Its Related Constants, using the

° ¢ P dl
most recent data, gives the following expressions for 0, 7, and ae when referred to
at

1850.0:
6 = 353° 34 55” 4- 32’.655 (t — 1850) — 8”.79 (t — f),

n = 0.46654 (¢ — 1850),

“= [50.2362 + 07.000220 (¢ — 1850) | (” — 4).

Letu ==

n! ?

we have then
we = 0.34955
2u = 0.69910
du = 1.04865
4u = 1.39820
Su = 1.74775
6u = 2.09730
Gio, Guec,
Hence
1 —3u = — .04865 ,
2—6u = — .09750.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 119

This shows that the arguments (g—3q’), and (2g — 69’), have coefficients in the
final expressions for the perturbations greatly affected by the factors of integration.
In case of the argument (gy — 3g’), we should compute the coefficients with more deci-
mals; also those of (0 — 39’) and (2g — 3q’), since in the developments the coefficients
of these affect those of (g —34q’).

From
sin 5 Z.sin § (¥ + ©) =sin$ (Q— Q’)sin} ¢@—7)
sin I. cos 2 (¥ + ®) = cos $(Q—Q’) sin $ (¢— 7’)
cos 5 J. sin 5 (¥ —®) = sin} (Q— 9’) cos 4 (7+ 7)
cos 5 I.cos 4 (© — ®) = cos 4(Q — 2’) cosh (7 4 7)
where, if §2’ > &, we take § (860° + 9 — 2’), instead of 4 (Q— 9’), we find

B=116° 15’ 36.7
De Il 80 sey
Shes 1 0 S583

An independent determination of these quantities is found from the equations

cos p sing = sin?’ cos (Q — 8’)
COS p COS g = Cos 7
cos psinr = cos?’ sin (Q — Q’)
cos pcosr = cos (2 — 2’)
sin p = sin?’ sin (Q — 9’)

sin Jsin ® = sin p

sin cos © = cos psin (¢— q)

sin Zsin (¥ — r) = sini p cos (¢ — q)

sin Icos (J —r) = sin (t—q)

cos I = COs p COS (t— 4).

120 A NEW METHOD OF DETERMINING
From
Il =a —2 —®

inl’ = 7! — OY —

we have

esl God 55/2 Ue —— OG OCeAe. ee:

_ Then from

k sn K = cos Jsin Il’
k cos kK = cos II’
ie, Si [kG = sin IT’

k, cos K, = cos J cos Il’

p sin P = 2a? — 2uk cos (II— EK)

p cos P = 2a cos 9’ k, sin (11 — A)
vsin V = 2a cos > ksin (11 — KX’)

v cos = 2a cos 9 cos 9’ k, cos (1I— 45)
w sin W = p— Qa? sin P

weos W= v eos (V — P)

w,sin W,= vsin( V — P)

y
w, cos W,= 2a?* eos P,
é

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 12]

we find

IK NBO Is OAS log k = 9.999614
Kee — lbOMeol T4 log k, = 9.997849

3 Ari) log p = 9.932748
Sa) © 2.4 log v = 0.601463
W=266 4 39.5 log w = 0.605196
W,=266 15 380. logw,= 0.601352

Then from

We Orn Oe Che ay apn Ole Clas

we have

log R = 0.702855, logy, = 7.976024.

The values of the quantities from Il to y, should be found by a duplicate compu-
tation without reference to the former computation, since any error in these quantities
will affect all that follows.

We now divide the circumference into sixteen parts relative to the mean anomaly,
and find the corresponding values of the eccentric anomaly # from

g= H—esn LH,

where ¢ is regarded as expressed in seconds of are. Substituting the sixteen values

of ¢ in the equations
fsin (7 — P) =w sin (HH — W )—ep

f cos (Ff — P) = w, cos(H+ W,),

we obtain the corresponding values of f and /.

A. P. 8S.—VOl. XIX. DP.

122 A NEW METHOD OF DETERMINING

Then in a similar manner from

C=y + y.sin*Q

logqg= log f+ y

Yo V2 a : 3 Yoo, Ee
L=s (4 see ) sin 2 74 s( av — js) sin4

2 f?

where s = 206264’.8, log”, = 9.63778,
we find the values of Q, C, log ¢, «, and y.

Thus we have found all the quantities entering into the expression

(-) = (C— qcos (E’ — Q)) (1—F cos ( 2 + Q)).

Instead of this, we use the transformed expression

({) =" (1 + —2acoa(H—@Q)) * (1+ —2e08(B' + Q)*,

and have, for finding the values of V, a, b°, the equations

a = sin y

? — sin val

a=tgsyz

b= tg 5 Xa
Va

Vara G

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 125

To find the value of C, . we put

(1 + a— 2a cos (Z’ — Q))~ ve +, “cos( B—Q)+ cos 2( #—Q)-+ ete. |

)

(1+ 6 — 26 cos (H’ + Q)? =[t2. nes cos (4 + + Q+ By cos 2 (#’ + Q)

ls oO
Hm

“bE ete. |

For finding the values of the coéfficients in these expressions we use RUNKLE’S
Tables for Determining the Values of the Coefficients in the Perturbative Function of
Planetary Motion, published by the Smithsonian Institution. With the sixteen values
of a as arguments we enter these tables and find at once the corresponding values of

q@) (2) (3)

(0) by br by Ge - (O) GP .Gb) Gea «@)) ae
b 1 , then those of A OOD etc., ete. 5 Ae bs > bs ae bs , ete., ete., where 6" is found
a*
from the sixteen values of 0? = eaae

Since 6 in (1 — 26 cos (’ + @)) is very small it will suflice to put

|
Ou
a

Nilo

Then from

iz)
I|

n (i)
zN B,,cos271Q
z

ro)

n (i)
NM Be sina
2
5 a
we have, in case of u (),

(1)
= il ee SS Va )-
La, iN, nO = — qe NV bcos2 Q, 3 o4% = cd bsin2Q;

124 A NEW METHOD OF DETERMINING

and, for ua? =)

3 (1) 3
iz NV 3bcos2Q, $8, = 7,wN 3bsin2Q.

We divide by 8 to save division after quadrature.

() (2) @)
With these values of c,, s,, and the values of the coefficients 6,, we find the
7 z

7
7 Uy
values of k,, ;, from

3 (7) (0) (i-1) (+1), (1)
k, cos K; = 0, ¢, + (6 + 5b, \e
Dy

TN
z B

(=i )) (+41), (1)
bs )

For 7 = 0, we find /) from

, (0) (0) (it) CY)
ko — 3 Dr Cn b,, Cn
2 2 my

. a
Then in ease of tu (“) from

A,, = }m'sk,cos [1(Q—gq)—4\]
A,, = tm'sk,sin [0 (Q—g)—&A,],

where m/’ is the mass of the disturbing body and s = 206264.’8 ;
and from

A,, = 3m sak, cos [i (QY— g) — Ki]

A,, = tm soek,sin [¢(Q—g)— A],

(a)

: 3 (c) (s) z Zs
in case of uwa> ( “) , we find the values of A,, and A,, for the 16 different points of the

circumference, and the various terms of the series.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 125

A,, = >C,,cosvg + >C,,sinvg
(c) Om

A, = 2S,,cosyg + >S,,sin vg,

i,ve

(O38 )= 4% Xs (2) = Yo— ¥,
dona 7.8% (i) Sie
GIO) =e Ke Aj = B=,
(AUS) Ss Ee G)=V%— Vs

(0.4) = (0.8 ) + (4.12)
(1.5) = (19 ) + (5.13)

(2.6) = (2.10) + (6.14) (0.2) = (04) + (26)
(3.7) = (3.11) + (7.15) (1.3) = (1.5) + (3.7)

4 (Gy + 2e,) = (0.2)

A(G—2o) = Gs)

AGE @) = ©@8)=Ge)

4(e— o) = §[ (1.9) — (5.13) | — (0) —@aB)]/ cos 45°
4(s + 8) = $[ (1.9) —(5.13) | + [ (8.11) — (7.15) |{ cos 45°
(Ge) — (210) (G14)

8c, = (0.4) — (2.6)
8s, = (1.5) — (3.7)

126 A NEW METHOD OF DETERMINING

The values of ¢,, s, must satisfy the equation

+ s,sing + s,sin 2g + ete.
. Oo 0 (7) .
2 answering to? in b,, and x being any one of the numbers, from 0 to 15 inclusive,
23

(¢)
into which the cireumference is divided. We use ¢, s, as abbreviated forms of C,,,,

(s) (c) (ec) (s)
C;,,, ete. Having found the values of ¢,, s, from the 16 different values of A), A,, i,

(Cc) (s) (ec) (8) a > (4
Aly, Aly 5 6 9 lay 4b, lootiin ioe m (4) and war ( “)s we have the values of these func-

tions given by the equation

a (c) (s

n (s) c =
(3) = $33 (C= S,,) eos [GF ») g 1B" F £35 (C,,4 S,,) sin[ GF») gE |

The values of the most important quantities from the eccentric anomaly £ to ¢,,
s,, needed in the expansion of w (“) and «a> Cr are given in the following tables,
first for uw (5) , and then for wa? fe 3 when not common to both.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 127

Values of Quantities in the Development of « (3) and woe(“).

H H+ W H+ W, Hr — P Br
SF 2 Mh Ona ® fF. MM So 7 Li ; Ol aie
( 0 0 0.0 266 4 39.5 266 15 38.0 266 21 17.2 399 24 44.9
( 2A JA ALD, 290 28 48.7 290939) 4272) 290 8 1.8 23 11 34.8
( 48 26 37.2 314 31 16.7 314 42 15.2 313 40 58.4 46 44 95.4
( Tl 52) 2429 331 bT 4.4 dae) ON 2.9 Bax) Ges Be) Oe) HY (0,33
( 94 35 14.0 0 39 53.5 0 50 52.0 309) 41 13 92 44 98.3
( 116 36 51.7 22 41 31.2 DY 2) DS) MN BG ish 1d) 253428
( 6 138 4 29.4 44 9 8.9 44°90 7.4 | 43 47 3.8 136 50 30.8
( 159 8 19.6 65 12 59:1 65) 23) 57:6 65 8 48.4 158 12 15.4
( 180 0 0.0 86 4 39.5 86 15 38.0 86 13 41.4 WG) aly Bet
( 200 51 40.4 106 56 19.9 NOT) Ska 107 15 14.8 200 18 41.8
) 221 55 30.6 128 O 10.1 128 11 8.6 128 28 47.5 221 32 14.5
243 23 8.3 149 27 47.8 149 38 46.3 150 8 27.6 243 11 54.6
265 24 46.0 171 29 25.5 171 40 24.0 172) 23 51.4 265 27 18.4
N33 Ball 194 12 14.6 194 93 13. 195 27 19:4 288 20 46.4
dll 33 22:8 Pike ats) 233 217 49 0.8 218 43 0.9 311 46 27.9
335 35 55.8 241 40 35.3 24 di 33.8 242 28 57.5 | 330 32 24.5
| 1613 47 17.9
| 1438 47 18.6
Log. f. y x 0) Log. q. Log. C.
i Oo Pm
(( 0.612427 —.001251 | — 12.2 359 24 32.0 0.611176 0.706582
( 0.612078 —.000860 | +431.5 23 18 46.3 0.611218 0.706349
(2 0.609315 —.000081 | +598.0 46 54 23.4 0.609234 0.705534
( 0.605242 —-.000981 +390.0 70 3 36.3 0.606233 0.704403
( 0.601312 + 001292 — 58.6 Ce dish WELT 0.602604 0.703241
( 0.598569 --.000846 —476.9 let By! So!) 0.599415 0.702241
( 0.597310 + .000091 —626.7 136° 40 4.1 0.597401 0.701493
( 0.597194 —.000956 —435.1 158 5 0.3 0.596238 0.701011
C& 0.597621 —.001322 == 15,7 179 16 52.7 0.596299 0.700788
( 0.598109 ——.000997 + 408.7 200 25 30.5 | 0.597112 0.700494
( 0.598532 —.000152 618.1 22) 42) 32.6 0.598380 0.700021
0.599177 +.000TTT + 496.6 243 20 11.2 0.599954 0.699872
0.600584 —-- 001278 + 96.7 265 28 55.1 0.601862 0.700504
0.603163 +.001032 —363.1 288 14 43.3 0.604195 0.702020
0.606734 --.000148 —600.1 3811 36 27.8 0.606882 0.704038
0.610302 —.000825 —452.4 335 24 52.1 0.609477 0.705810
4.823835 at 3 — (5 1613 47 174 4.823838 5.622201
4.823834 = 9 == (hy a yee Sy ets) 4.823842 5.622200

128

Values of Quantities in the Development of «(4) and war or

A NEW METHOD OF DETERMINING

g x VG Log. b. Log. a. a. Log. WN.
(e) 7 // i; ie

(0) 3 93 45.3 1 51.83 7.063818 9.701484 0.502902 9.695669
@ 1) 3 96 41.3 7 54.78 7.063792 9.701945 0.503437 9.695880
( 2) 53 14 15.6 7 59.97 7.065778 9.699988 0.501173 9.695892
( 8) 52 54 33.7 8 3.30 7.068781 9.696876 0.497594 9.695837
( 4) 52 28 55.6 8 1.35 7.072405 9.692804 0.492951 9.695616
( 5) 52 6 81.2 8 10.95 7.075601 9.689226 0.488907 9.695421
( 6) 51 53 41.2 8 13.23 7.077613 9.687169 0.486597 9.695400
(Gn) 51 46 50.0 8 14.55 T.OTSTT4 9.686068 0.485364 9.695430
(8) 51 49 41.2 8 14.49 7.078721 9.686526 0.485877 9.695629
(9) 52 0 52.3 8 13.57 1.077913 9.688321 0.487889 9.696120
(10) 52 18 36.9 8 12.12 7.076635 9.691160 0.491089 9.696905
(11) 52 36 21.2 8 10.34 7.075061 9.693986 0.494294 9.697532
(12) 52 49 37.5 8 8.19 7.073153 9.696093 0.496699 9.697631
(13) 52 58 10.6 8 5.58 7.070825 9.697448 0.498251 9.697141
(14) 53 5 12.5 8 2.58 7.068133 9.698559 0.499597 9.696354
(15) 53 13 54.4 7 59.70 7.065534 9.699932 0.501109 9.695743
ay TT.553183 3.956815 77.569096
x 17.553803 3.956845 77.569088
© @ @ ©) @ Q

g Log. te, ILOS, 5 Oy, Log. ts, Log. b, Log. b Log. b,

0) 2 2 2 21 a

( 0) 8.792579 6.16064 4.475270 0.332110 9.748094 9.329969
() 8.792790 5.98934 6.02920 0.332186 9.748669 9.331018
@) 8.792802 4.985510 6.16173 0.331867 9.746235 9.326571
@3) 8.792731 6.05070n 5.97267 0.331369 9.742375 9.319511
( 4) 8.792526 6.16734n 5.14693n 0.330730 9.737346 9.310298
(5) 8.792331 5.98219n 6.05562n 0.330182 9.732946 9.302224
( 6) 8.792310 4.93934 6.173782 0.329872 9.730425 9.997590
(% 8.792340 6.03383 6.016142 0.329707 9.729076 9.295111
C8) 8.792539 6.17549 4.575070 0.329776 9.729636 9.296143
( 9) 8.793030 6.05359 5.99045 0.320045 9.731836 9.300183
(10) 8.793815 5.23282 6.17067 0.330477 9.735329 9.306586
(11) 8.794449 5.948120 6.07618 0.330914 9.738805 9.312970
(12) 8.794541 6.16466n 5.36611 0.331246 9.741407 9.317738
(13) 8.794051 6.07296n 5.942029n 0.331460 9.743078 9.320808
(14) 8.793264 5.237420 6.16200n 0.331637 9.744461 9.323327
(15) 8.792653 5.97789 6.04134 0.331858 9.746165 |° 9.326448
} 2.647715 77.912926 74.508222
x 2.647721 77.912945 74.508268

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

Values of Quantities in the Development of u(“) and wa?(“) .

129

(3) (4) (5) (6) (7) (8) (9)
g | Log. b, Log. b, Log. b, Ios yOnen|) oss 0) |) Losvdy "Loe: b,
2 2 z z z 2 z
( 0) 8.954999 8.60017 8.2570 1.9215 7.5915 7.2654 6.9426
(I) 8.956515 8.60214 8.2594 7.9244 T.5947 7.2691 6.9468
( 2) 8.950082 8.59373 8.2490 7.9120 7.5804 7.2528 6.9286
( 3) 8.939865 8.58036 8.2326 7.8926 7.5578 7.2271 6.8997
( 4) 8.926521 8.56292 8.2110 7.8668 7.5280 7.1932 6.8617
( 5) 8.914818 8.54760 8.1921 7.8444 7.5020 7.1636 6.8285
( 6) 8.908100 8.53882 8.1812. 7.8314 7.4870 7.1466 6.8094
(Co) 8.904506 8.538411 8.1754 7.8244 TAT89 7.1373 6.7991
( 8) 8.906000 8.53606 8.1778 7.8273 7.4822 7.1411 6.8033
(9) 8.911861 8.54373 8.1872 7.8386 7.4953 7.1561 6.8201
(10) 8.921142 8.55588 8.2024 7.8565 7.5160 7.1796 6.8464
(11) 8.930392 8.56797 8.2172 7.8742 7.5367 7.2031 6.8728
(12) 8.937298 8.57701 8.2285 © YSIS 7.5520 7.2205 6.8923
(13) 8.941742 8.58283 8.2855 7.8960 7.5618 7.2317 6.9048
(14) 8.945388 8.58760 8.2415 7.90380 7.5700 7.2410 6.9152
(15) 8.949898 8.59349 8.2488 teSilaly 7.5800 7.2524 6.9280
Dy 71.449530 68.55219 65.7484 63.0060 60.3071 57.6402 54.9995
a! 71.449597 68.55223 65.7482 63.0063 60.3072 57.6404 54.9998
3 ‘- @) (On rari) @) (2) (3)
g |Log.t N | Log. te, | Log. ts, | Log. 46, | Log. 6, | Log. 6, | Log. bg
2 2 2 2 2 2
( 0) 8.183917 5.42374 3.1383 7n 0.280319 0.417421 0.200612 9.961097
(Ib) 8.184550 5.25307 5.29293 0.281000 0.418474 0.202090 9.963016
( 2) 8.184586 4.24928n 5.42550 0.278120 0.414013 0.195824 9.954877
( 3) 8.184421 5.314307 5.23627 0.273612 0.406981 0.185917 9.941987
( 4) 8.183758 5.430287 4.409877 0.267827 0.397890 0.173060 9.925223
( 5) 8.183173 5.244547 5.31797n 0.262860 0.390004 0.161858 9.910585
( 6) 8.183110 4.20163 5.43607n 0.260054 0.385513 0.155458 9.902210
Ga) 8.183200 5.29621 5.278520 0.258559 0.383116 0.152039 9.897732
( 8) 8.183797 5.43847 3.838057 0.259184 0.384116 0.153464 9.899598
@ 9) 8.185270 5.31804 5D: 25490 0.261621 0.388024 0.159038 9.906900
(10) 8.187625 4.49962 5.43747 0.265530 0.394254 0.167901 9.918485
(11) 8.189506 5.21681n a 34487 0.269488 0.400515 0.176758 9.930076
(12) 8.189803 5.433640 4.63509 0.272484 0.405223 0.183435 9.938754
(13) 8.188333 5.340477 5.20953n 0.274429 0.408267 0.187732 9.944350
(14) 8.185972 4.50257n 5.42714n 0.276036 0.410773 0.191265 9.948948
(15) 8.184139 5.24121 5.304667 0.278037 0.413885 0.195644 9.954643
2) 65.482568 2.159554 | 3.209203 | 1.421019 | 79.449192
a! 65.482592 2.159606 3.209266 1.421076 79.449289

A. P. S.—VOL. XIX. Q.

130

A NEW METHOD OF DETERMINING

3
Values of Quantities in the Development of « (4) and uor(“) 5
i (4) is (5) KG) (7) (8) (9)
g Log. bs Ihog.b, | og. 63 Log. bs Log. b, Log. b,
P 2 2 2 2
( 0) 9.70884 9.4484 9.1822 8.9118 8.6383 8.3621
( 1) 9.71121 9.4512 9.1854 8.9155 8.6425 8.3665
( 2) 9.70116 9.4393 9.1716 8.8998 8.6247 8.3471
( 8) 9.68524 9.4203 9.1496 8.8747 8.5965 8.3158
( 4) 9.66450 9.3955 9.1207 8.8418 8.5595 8.2747
( 5) 9.64638 9.3739 9.0956 8.8131 8.5273 8.2389
( 6) 9.63600 9.3614 9.0818 8.7968 | 8.5089 8.2184
( 7) 9.63043 9.3549 9.0735 8.7880 8.4991 8.2077
( 8) 9.63276 9.3576 9.0766 8.7914 8.5030 8.2119
(2) 9.64181 9.3684 9.08983 8.8058 8.5191 8.2298
(10) 9.65617 9.3856 9.1098 8.8287 8.5449 8.2585
(11) 9.67052 9.4028 9.1292 8.8515 8.5705 8.2868
(12) 9.68125 9.4156 9.1440 8.8684 8.5893 8.3078
(13) 9.68816 9.4937 9.1537 8.8791 8.6015 8.3213
(14) 9.69382 9.4305 9.1614 8.8882 8.6118 8.3329
(15) 9.70087 9.4389 9.1711 8.8992 8.6240 8.3464
x 77.37450 75.2339 73.0471 70.8269 68.5804 66.3134
a 717.37462 75.2341 73.0474 70.8269 68.5803 66.3132
g | Log. k, | Log. k, | Log. k,; Log. k, | Log. &,| Log. &, | Log. k, | Log. k,
( 0) 8.824187 8.54492 8.12562 7.750420 7.89550 7.0523 6.7168 6.4105
( 1) 8.824302 8.54433 8.12588 7.151220 7.39678 7.0540 6.7190 6.4054
( 2) 8.823605 8.53875 8.11916 7.742693 7.38634 7.0416 6.7046 6.3714
( 3) 8.822665 8.53172 8.10982 7.730361 7.37091 7.0232 6.6832 6.3298
( 4) 8.821701 8.52543 8.09963 7.716100 7.35261 7.0007 6.6565 6.2932
( 5) 8.821143 8.52236 8.09246 7.705215 7.33807 6.9826 6.6349 6.2764
( 6) 8.821183 8.52360 8.09009 7.700585 7.33130 6.9737 6.6239 6.2809
( 7) 8.821397 8.52470 8.08981 7.699023 7.32855 6.9698 6.6187 6.2913
( 8) 8.821810 8.52671 8.09164 7.701551 7.33151 6.9732 6.6226 6.3027
( 9) 8.822444 8.52829 8.09567 7.107159 7.33895 6.9824 6.6337 6.3093
(10) 8.823323 8.52965 8.10077 7.715298- 7.35002 6.9965 6.6506 6.3129
(11) 8.824009 8.53059 8.10550 7.723069 7.36070 7.0100 6.6669 6.3147
(12) 8.824233 8.53159 8.10915 7.728940 7.36874 7.0202 6.6793 6.3196
(13) 8.824055 8.53359 8.11238 7.733450 7.37462 7.0274 6.6879 6.3342
(14) 8.823809 8.53721 8.11622 7.738311 7.38053 7.0345 6.6960 6.3608
(15) 8.823826 8.54164 8.121158 7.144423 7.38795 7.0433 6.7062 6.3901
2 70.583851 68.25726 64.85258 | 61.793910 | 58.89655 56.0927 53.8503 50.6520
ral 70.583841 68.25722 64.85260 | 61.793920 | 58.89653 56.0926 53.8505 50.6512

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

Values of Quantities in the Development of «(4) and ua*( ate

131

g | Log.ks Log. ky || Ky JEG ISG K, K; Ky
/ i? / / J

( 0) 6.0606 5.7378 — 0.6 —04 —038 — 0.3 3 0.3 0.3
(1) 6.0636 5.7413 +20.3 +12.9 +-11.4 E111 10.6 aa + 9.5 + 8.3
(2) | 6.0454 5.7212 JE) ET SRR —t5T16,0) “igen exe SEO | Ss
( 3) 6.0178 5.6904 +-18.4 11.7 10.2 10.0 EO + + 92 + 8.8
( 4) 9.9830 5.6515 —28 —18 —16 —1.5 3) 1.5 1.5
( 5) 5.9541 5.6191 —22.7 —14.5 —12.7 —12.0 8 11.4 11.0
( 6) 5.9391 5.6019 —29.8 —19.0 —16.7 —15.7 3 : 14.5 BBE
(C®)) 5.9316 5.5934 —20.7 —13.2 —11.6 —10.9 5 —10.1 — 9.7 — 8.9
( 8) 5.9364 5.5985 — 0.7 — 05 — 0.4 — 0.4 A —0.3 —03 -- 0.8
(9) 5.9512 6.6151 +19.5 +12.8 10.9 L 10.2 8 + 9.4 + 9.0 + 8.2
(10) 5.9737 5.6405 +29.1 +18.6 +16.4 15.3 a) : 14.1 +13.3
5.9959 5.6656 +-23.4 +14.9 +13.1 +12.3 lL +11.9 +117 +11.3
6.0124 5.6842 + 4.5 + 2.8 + 2.5 L 2.4 | 2.4 + 2. + 2.3 + 2.2
6.0251 5.6968 —17.0 —10.8 — 9.5 — 89 = 88 — 8 — 86 — 84
6.0341 5.7083 —28.1 —17.8 —15.7 —14.7 —14.3 ‘ 13.6 13.0
6.0468 5.7224 —21.0 —I13.4 —I11.8 —11.0 —10.6 9.8 9.0

45.3439 Sah) ee ee ee aS

45.3441 b = a @ ds. 8 = ofl
g | Log.k& Log.k lLog.k lLog.k; Log. k, Log. k Log. k& Log. k,
( 0) 8.465272 8.60289 8.38621 8.14674 7.89481 7.6341 7.3679 7.0975
(Gp) 8.466247 8.60407 8.38777 8.14874 7.89694 7.6369 7.3712 7.1013
( 2) 8.462637 8.59849 8.38030 8.13935 7.88563 7.6238 7.3561 7.0843
( 3) 8.457236 8.59018 8.36903 8.12505 7.86829 7.6033 7.3326 7.0577
( 4) 8.450550 8.58006 8.35509 8.10719 7.84645 T.5TT4 7.3026 7.0237
(Gay) 8.445362 8.57214 8.34391 8.09259 7.82837 eDooO9 1.2776 6.9950
( 6) 8.443224 8.56872 8.33868 8.08545 1.81922 7.5446 7.2645 6.9800
(C1) 8.442508 8.56750 8.33651 8.08224 7.81495 7.5395 7.2581 6.9726
( 8) 8.444020 8.56954 8.33902 8.08521 7.81840 7.5433 7.2623 6.9771
(G9) 8.444679 8.57452 8.34564 8.09354 7.82847 7.5551 7.2760 6.9925
0 8.453274 8.58206 8.35573 8.10632 7.84401 7.5734 7.2971 7.0165
8.458368 8.58906 8.36522 8.11851 7.85895 7.5912 7.3176 7.0400
8.461465 8.59345 8.37153 8.12680 7.86927 7.6036 7.3320 7.0564
8.461922 8.59532 8.37468 8.13126 7.87506 7.6105 7.3405 7.0660
8.461886 8.59651 8.37704 8.13471 7.87957 7.6163 7.8472 7.0739
8.462852 8.59905 8.58088 8.13992 7.88616 7.6242 7.38564 7.0845
68.69172 66.90360 64.93175 62.85706 60.7165 58.5297 56.3095
68.69184 66.90364 64.93185 62.85719 60.7166 98.5300 56.3096

132 _ A NEW METHOD OF DETERMINING

Values of Quantities in the Development of u (4) and uor(“).

g Log. ks; Log. ky | A; K; Kk, (Q-g)-K, 2(Q-g)-k, 3(Q-9)—K;

/ / / | oO. i fo) y yy

( 0) 6.8240 G50 |) Ol —().1 —0.1 359 25.1 3858 49.5 358 13 55.0
(1) | 6.8280 6.5522 +4.4 +-4.4 +4.4 || 0 28.5 1 24.6 Dut B70)
(2) | 6.8092 6.5317 +6.0 ++6.0 --6.0 Wo 2535) 3 31.0 By Pi ay!
( 3) 6.7795 6.4988 +3.9 +3.9 +3. 2 Way?) AL Byay5) to BO 33.9)
( 4) 6.7414 6.4566 -—0.6 —0.6 —().6 2 46.3 5 28.8 Sa eae
( 5) 6.7093 6.4209 —4.7 —4A —4.7 2 NTS Oy) ake) 7 26 37.6
( 6) 6.6921 6.4016 —6.2 —6.2 —6.2 2 9.8) 3 se) 5 16 57.0
( 7) 6.6837 6.3923 —4.3 —4.3 —4.3 0 55.7 1 23.2 1 5 Boo)
( 8) 6.6887 6.3976 —(0.2 —0.2 —0.2 358) TLS 358 34.3 357 Ol 3.3
@9) 6.7058 6.4165 +4.0 +4.0 +4.0 Bo 36.2 B55) BS Se) By) BI)s5)
(10) | 6.7327 6.4463 +6.1 +6.1 +6.1 356 13:4 3538 6.5 349 51 14.9
(11) 6.7589 6.4752 | —-5.0 -+-5.0 -L5.0 355 26.8 301 20.5 SA 2 Oe!
(12) 6.7773 GASH |) SLT) +1.0 1.0 305 «24.4 850 55.0 346. 24 12.8
(13) 6.7883 6.5081 || —3.5 —3.5 —3.5 Boe Ist B01 40.2 347 23 40.2
(14) 6.7976 6.5187 —6.0 —6.0 —6.0 351 «4.6 353 30.7 BO) By BUS
(15) 6.8093 6.5317 —4.5 —4.5 —4.5 |) 358 15.9 3563.1 353 56 22.5
By 54.0630 51.7961 0 0 0 | 1793 47.8 ISI V22)> 326
a 54.0628 51.7957 + .3 + .8 + 3 || 1483 47.3 1497 2) 59:

g A(Q—9)—K,5( Q—9)—K; 6( Q—9)— K, 1 09), 8( Q—9)— K3 9( @ - 9) — Ky
(o) / (eo) / e) / (e) / e} y

( 0) 857 38.5 Sf 0) 356 27.5 30D) 2a 855 16.7 354 41.2
Gp) 3 Bos) 3 53.2 4 42.5 5 31.8 6 21.1 7 10.5
( 2) 2204 S) Ina 11 12.4 UB YB Il 2) 1G AHA
(3) 10 4.4 12 38.3 15 12.2 17 46.0 20 19.8 22 53.6
( 4) 10 55.5 13 39.0 16 22.5 IQ BA) 21 49.5 24 33.0
(@5)) 9 50.6 12) 15.0) Wal Bw) ly &§) 19 28.4 21 52.8
( 6) @ 5.8) 8 35.6 10 15.3 11 54.9 3) 3425 15 14.2
(Gig) 2 30:9 BS By) 3 40.1 AS ART 4 49.3 5) 25)
( 8) 357 ~— 8.0 396 24.9 Boe) LIL. T 354 58.6 354 15.5 353 32.4
(( $)) 301 31.8 BAG) Dit 347 23.6 845 19.5 343 15.4 341 11.3
(10) 346 34.9 343 17.8 340: 0:7 336 43.7 333 26.7 330) 956
(11) SAD mOED) 338 58.9 334 49.3 330 39.7 326 30.1 322 20.5
(12) 341 53.2 Solmeooel So Grllil 328 20.0 323 48.9 BIG) ie)
(13) Bytes © otf 388 52.3 yey! | BXOLY) 330 21.5 326 66.1 321 50.7
(14) 346 40.5 342 16.6 3389 52.7 336 28.8 Baa 4h.) 329 41.1
(15) sol 50.4 349 44.9 347 39.4 345 33.8 343 28.2 B40 2207
py 174465

aft 1384 6.0

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 133
Ti i =
n the expansion of u ( aE
(c) (c) (s) (c) (s) (c) (s) (c) (8)
0 245 1 2 2 3 3 A, 4
7 7 7 "7 1 "7 ame. eee 7

( 13.13109 6.9027 —.0701 =+-2'6281 —.0539 | +1.10745 —.03418 | --.4889 —.0201
( 13.13458 6.8933 -+-.0571 2.6294 --.0647 1.10917 -++.04356 4901 —-.0262
( 13.11352 6.8033 -+-.1712 2.5849 --.1588 1.08348 +.10356 AT51 --.0615
Cz 13.08513 6.6912 --.2633 9.5254 + .2176 1.04890 -—-.13827 4553 --.0809
( 13.05615 6.5922 3192 9.4646 +.2364 1.01333 --.14604 43538 --.0840
( 13.03939 6.5457 --.3187 2.4959 1.2150 0.99004 —-.12935 4994 | 0733
( 13.04058 6.5584 —-.2479 2.4172 +1543 0.98367 --.09095 4190 —-.0509
( 13.04700 6.5880 --.1067 2.4198 1.0585 0.98375 +.03339 4190 +-.0184
(8 13.05942 6.6190 —.0816 2.4317 —.0606 0.98937 —.03712 4218 —.0211
( 13.07850 6.6377 —.2779 2.4464 —.1863 0.99667 —.11189 4249 —.0633
d 13.10500 6.6498 —.4389 2.4645 —.2979 1.00593 —.18002 | 4287 —.1023
el 13.12573 6.6578 — .5301 9.4816 —.3742 1.01487 —.22886 | 4322 —.1310
al 13.13248 6.6727 —.5359 2.4991 —.3995 1.02497 —.24789 4373 —.1431
(1: 13.12612 6.7090 —.4658 2.5224 —.3693 1.03984 —.23954 | 4463 —.1354
dd 13.11967 6.7727 —.3458 2.5559 —.2907 1.06142 —.18555._ | 4600 —.1090
qd 13.12018 6.8478 —.2074 2.5954 —.1791 1.08668 —.11537 | 4760 —.0683
104.75791 | 53.5708 —.7340 | +20.0460 —.5531 8.26962 —.34421 | 3.5661 —.1992
104.75663 | 53.5705 —.7354 |+20.0463 —.5531 8.26992 —.34409 | 13.5662 —.1992

g 5 5 6 6 7 7 8 8 9 A,

Sly lad tad vr dA JI} I IH/ II vad

(GO) eee een O eae aS: =" 00630. |e 20505) 003GNul) 0226) 0019) a= 0L0j 0010
(C10) 2223 +.0151 1027 +-.0085 0498 +.0048 0226 +.0025 0108 --.0014
( 2) 2138 -+.0350 0978 -+.0194 .0451 -+-.0105 0211 +.0057 0099 —-.0030
( 3) 2028 —-.0454 0916 +.0249 0401 —-.0128 0192 —-.0071 .0089 —-.0038
( 4) 1916 +.0465 0856 —-.0252 0365 —-.0126 0176 —-.0070 .0080 +-.0037
(5) 1848 -+.0401 0821 —-.0215 0356 +.0109 0167 +.0059 .0076 —+.0030
( 6) 1832 +.0277 0815 —-.0147 0368 —-.0078 0166 -+.0040 0076 +.0021
( 7) 1833 +.0099 0816 —-.0052 0384 -.0028 0168 +-.0014 00TT +.0607
( 8) 1847 —.0116 0823 —.0062 0394 —.0035 0169 —.0017 .0078 —.0009
(9) 1860 —.0346 0826 —.0185 0388 —.0102 0168 —.0051 0077 —.0026
(10) 1870 —.0561 0827 —.0301 0372 —.0160 0166 —.0083 0075 —.0043
(11) 1880 —.0722 0827 —.0389 0354 —.0199 | .0163 —.0108 0072 —.0056
(12) 1904 —.0793 0837 —.0429 0350 —.0216 | .0163 —.0120 0072 —.0062
(13) 1956 —.0756 0867 —.0411 0369 —.0210 0173 —.0116 0077 —.0060
(14) 2041 —.0613 .0918 —.0336 0414 —.0180 .0190 —.0096 0087 —.0051
(15) 2140 —.0387 0978 —.0214 .0468 —.0120 0210 —.0062 0098 —.0033
z 1.5765 —11050| --.7077 —.0598 | +3219 —0318 |7421467 —0168 | 42.0674 —.0087
x 6 | 11.5768 —.1106 | +.7078 —.0598 | +.3218 —.0318 | -++.1467 —.0168 |-+.0674 —.0086

134. A NEW METHOD OF DETERMINING

3
In the expansion of wa? Ga) 3

(9) | (c) (s) (c) (s) (c) (8) (c) (s)
g 0 A, A, A, A, A, A; 4 sly
AA I I wah /I aA // AA aA
( 0) 23.3520 | 132.0569 —0.3301 | +-19.4613 —0.4009°| +-11.2092 —0.3464 +6.269 —0.258
( 1) 23.4045 32.1423 0.4199 19.5273 0.5272 11.2569 +-0.4603 6.300 +.0.347
( 2) 93.2107 31.7192 +1.0033 19.1618 +1.2486 10.9731 -+-1.0737 6.096 --0.802
( 3) 29,9239 81.1043 -—-1.8503 18.6375 --1.6470 10.5748 -+-1.4097 5.813 -+1.041
( 4) 22.5737 30.3821 +-1.4503 18.0367 --1.7240 10.1342 -+-1.4580 5.516 +-1.063
( 5) 22,3086 29.8387 +1.2952 17.5937 1.51122 9.8190 +1.2644 5.310 +0.912
( 6) 22.1960 29.6180 —-0.9110 17.4156 +1.0505 9.6988 +-0.8734 5.239 +0.626
( 7) 29,1595 29.5473 0.38342 | 17.3564 --0.3782 9.6618 -+-0.3118 5.219 0.292
( 8) 22.2368 29.6867 —0.3713 17.4552 —0.4367 9.7264 —0.3654 5.259 —0.204
( 9) 29, 4949 30.0100 —1.1187 17.6808 —1.3068 9.8617 —1.0915 5.331 —0.786
(10) Q2.T157 30.5036 —1.8033 18.0224 —2.1155 10.0630 —1.7762 5.436 —1.285
(11) 22.9837 30.9679 —2.3042 18.3471 —2.7150 10.2558 —2.2962 5.5386 —1.667
(12) 93.1482 31.2707 —2.4810 18.5835 —2.9616 10.4121 —2.5144 5.627 —1.839
(13) 23.1725 31.4193 —2.3026 18.7500 —2.7837 10.5580 —2.3763 5.139 —1.148
(14) 23.1706 31.5386 —1.8212 18.9291 —2.2155 10.7412 —1.9027 5.895 —1.409
(15) 23.2299 31.7564 —1.1097 19.1791 —1.3716 10.9764 —1.1843 6.091 — .882
yy 182.6038 246.7758 —3.44293 147.0656 —4.1071 82.9580 —3.5000 +45.337 —2.564
YY | 182.5968 246.7862 —3.4356 147.0719 —4,1125 82.9644 —3.4985 + 45.339 —2.563
(¢) (s) (c) (s) (c) (s) (ce) (s) (c) (s)
g 5 5 A, 6 A, A, A, A, 9 A,
Wt aA 1 iA // Wit aA I aA t/
Coy) 43440 Soi | 41863 115: |) 41.000 —l079)> | 2539 —0a4 +.282 —.027
(( Jl) 3.458 +0.240 Wasa SAL a7 1.005 +.098 -535 --.060 .283 +.036
( 2) 3.318 --0.550 WS <b Bae 944 + 297 A9T +.134 .260 -+-.076
( 3) 3.130 +0.706 1.660 -+.453 868 -+.279 450 --.167 231 ==.098
( 4) 9.937 0.713 1.540 —+.453 197 —--.276 409 --.164 208 +.095
( 5) 2.812 +0.606 1.467 --.381 Hib b) ==.232) Jt SETS 196 +.078
( 6) 9.772 0.413 1.448 --.960 W148 2 157 .280 +.092 195 --.053
( 'h) 2.766 --0.146 1.446 -+.091 -150 +.055 086 +.032 197 +.019
( 8) 2.789 —0.175 LAS) IIo) OY OS 2089 —.039 1199 —=028
( 3) 2.824 —0.522 1LAT4 — 399 .160 —.199 og) INT 197 —.067
(10) 2.870 —0.855 1.491 —.540 159 —.326 30) — 92 108} =—JUU!
(11) DEO —— lal 1.505 —.705 a 5b) 9) Pil 187 —.144
(12) 9.963 —1.235 LH 133 SB kB .382 —.280 188 —.162
(13) 8.042 —1.179 1.582 —.153 .803 —.457 404 —.272 201 —.158
(14) 3.164 —0.957 1-670 —615 ES OOS 446 —.227 QI BS
(15) 3.312 —0.604 1eui5) —30il! 1940) 943 A95 — 147 259 —.087
y 24.953 —1.723 | 12.780 —1.094 5561639 6480 | =pe428— 1392 | SSIeTs2 939
DH 94.959 —1.792 12.783 —1.095 6.641 —.660 +3.415 —.395 Sil 25)

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 135

ibs (c) (s) (c) (s) . ;
The Quantities $C; , 3 5S;, , 2S; , arranged for Quadrature in the Expansion of

Ae) 23
(4)
t 4 ©
7=0 01 i G=33 A 4=5 41=6
[ (c) 1 i Us 1 1 1
| Cyo |-+-4[209.51454]] +453.571 +20.046 | 1896978 | +3.566 | +1.576 +707
v=0;
(c)
| Sio = 1B5 —.553 —.34414 —.199 —.110 —.060
l
( ©) =
| Cy +.25653 +.548 +.382 | +.99949| 4.199] -+-.07L| +.038
(s)
| Sir +1.706 JOS | aLeogy. || suas Te b ass) 7 Sere
v1} (s) |
Oh —.25027 —129 —046 | 01129 | +002 | +.005| +-.006
| (c)
| Sia 1.022 +.017 | +.00807 aE 003 | 2.001 000
©)
| Cre -++.00463 4.957 +.096 | 1.05847 -+.038 1.024 +.013
(s)
| Sis — 170 003) (ee eeorsss | oly |) 00m) 004
fe (s)
| Oe 4.12279 4.128 4.080 | -+.04667 18) seas ||| Sear
(©)
| Si ———1 065 Hieig4e) |e tos0es) In wee O18 mo10N | | 2.006
1%
{ (c)
| Cis -+-.08070 4.020 4.007 | --.00662| +.005| +.002| +.001
pees
| Sis —.003 4.002 | +.00216 +002 4.001 001
Vv=9i (8)
| C.. 4.05945 4.041 4.993 | -+.01319| +.006| +.003| -+.002
(ce)
| Si.3 000 —.001 —.00217 —.002 —.001 —,.001
U =
a) ;
| Cis +..00037 +.001 +..00030
| (s)
| Sis 000 4.00052
DA)
| Cia +-.00055 000 +.00076
|
| Sia —.001 —.00103
ae

136 A NEW METHOD OF DETERMINING

Abs (c) (s) (c) (8) : 3
The Quantities $C; , $C; , 48, , $8, , arranged for Quadrature, in the Expansion of

eG)

7=0 a=1 62% |9=3 eH |o=HS 220 0ST 0S 7=9
f (c) Ut i WW 11 1 MW a W 1
‘A 40 |4-$[364.6002]) +.246.7810] +-147.068] +82.9613|+45.338|-+-24.256|-+12.781/-+ 6.640|-+-3.419/11.751
Y=}
| (c)
| Sio —3.4388] —4.110) —3.4992| —9.569| —1.729| —1.095| —.654) —.3929] —.99
{ (c) }
| Cir -+-4.3500| +4.6277| 43.973] ++2.8862| 11.956) 41.953) +.771/ +.461) +.970] 4.154
(s) |
Sis 7.8438) 9.378] +-7.9505| +5.816| +3.910| -+2.488|-+1.514) +.898] +.521
v=], | ! |
Cin —1.8014) —1.1511] —.801] —.8643|- —106) +.017| +.062) +.078] 4.058) +..049
(c) | |
Sia +.1015} +.104| +.0731/ +.043| 1.024) +.011| —.008] +.003) +-.001
nO
Ci —.2566) +.0899/ -+.994/ +.3888/ 41.384) +1.397| 1.959] +.193] +.134| +.086
ee
LG -+.1010 296 3297| +.309) +.939/ +.173/ +.116] +.078| +.047
y=) 6
| Cie +-1.1803| 41.1209} -+.883/ +.6281/ 4.418] +.966| -+.169| --.093/ +.058) +.031
(©) |
|S +..3367 400 3459/1 .955| +.170| 1.106] +.065| +-.034| —.018
( _©@)
| Gs JLB) He SILA 099 0809} +.066| +.049| 1.035) -++.024] +.013] +.01
(s)
\ Sis —.0170 000] +.0059} +.019) -+-.015 015| +.015] -+.013] +.008
v=3) ©
| is +5132) +6602) +.817| 4.2097] 4.130) +.076| +.043| +.020) +.019] +.00
| @
| ie ==0138|==030|) 210344 agsa oor) =to90) 2005s orn ien005
f i
| Gia +0177] —-+.0085 003 .0028| 1.902) -+.002/ .000| --.001/ .000] +.001
| (s)
Si 40117 005 0061) +.005| 4.006] -+.004| -—-.004] +.001] +.001
y=4 a
C., +.0182} +.0172! 1.016 0134 4.010] -+.006] -+-.005| +-.003] —.009] +-.001
| (¢c)
| Sis =10109| | ==.022 |) =0r'g2| sob e012 008) 002|) ==c0s lon
|

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 137

ML (¢) (s)
The quantities C,,, C;,,, etc., of the preceding tables have been divided by 2 to
save division after quadrature. To check the values of these coefficients we will take

the point corresponding to g = 22°.5, using the equation

+ S,sing + SS, sin 2g + ete,

noting that the tables give one-half of the values of these quantities.

Thus we have

Sal j= j=)! 6=FZ
(ce) WW 1 (e) 1]
2 Cio 453.571 120.046 So = 0185 — 0.558
a(c) : : (s)
1 + 1.013 moa Si, = + 1.306 ae Se
{s) ()
La — 094 = 2032 ne = te 020 He BH
(c) (s)
1,2 + .363 + .135 i = — 2a = 004
(s) (c)
1,2 aE iS 4, i io == LOD 4 Mi
(c) (s)
1,3 + 015 + .005 ig === 005 + .004
(s) (c)
13 ONT + .043 Si3 = 0 = 0m
(¢) (s)
1,4 0 1,4 —— 0
(s) (c)
1,4 0 — 0
17) // /} VT
> 55.126 +21.018 Se + O58 + 0.521
5 + 6.891 + 2.627 Ly = + 0.057 + 0.065
(c) (s)
1 + 6.893 + 2.629 ES OL05 i + 0.065

In this way we check the values of these quantities for all values of 2, in case of

both w(“), and wa(“).

Applying to the coefficients of the two preceding tables! the formula

($)" = 885(C, F S,,.) cos [(¢-F)g— 1B" | AabEX(Cl, + S,,) sin [(¢F2)g 6B]

2 3
noting that } has been applied, we have the values of u (G), a ) that follow :

A. P. S—VOL. XIX. R.

138

A NEW METHOD OF DETERMINING

GLa cos | sin cos sin
v) J} y) /}
0 @ +$[209.51455 | +1/364.6002 ]
1—0 0.25653 — (0.25027 +4.3500 —1.8014
2 — (i) * 0.00463 0.12279 —0.2566 +1.1803
a) 0.03070 0.05945 0.1118 0.5132
4—0 -L0.00037 +0.00055 0.0177 +-0.0182
|

—2—1 | +0.023 —().041 -+-0.1310 —().6464

== 0.427 —(.193 —().0112 SAT
0—1 —1].158 0.101 —3.2161 +-1.0496
ai | +53.571 0.735 +246.7810 +3.4388
2 — Il | 9 954. —0.144 19.4716 —1.2526
3 oo | 0.087 0.063 +0.1909 0.7842
4—] 0.016 0.041 0.0970 0.6740

—l1—2 0.099 — 0.287
0—2 --0.098 —0.129 —0.001 = 1283
1, —().891 0.029 —5.500 0.697
2—2 + 20.046 0.553 +147.068 +4.110
3— 2 1.656 —0().063 +13.246 —(.905
4— 2 0.093 0.0382 -+-0.590 -+0.483
0—3 0.00446 —0.01101 -.0750 —0.1753
1—3 0.04011 —0.07730 +.0591 —0.9741
Y—= 3 —0.56048 0.00322 —).0643 0.2912
3—3 +8.26978 --0.34414 32.9613 +3.4992
4—3 1.01947 —0.01936 10.8367 —().4375
> 0.07682 —-0.01603 -+-0.7185 +-0.2822
6—3 0.00879 0.01536 0.0868 +0.244]
lo dt +0.003 —0).004 0.053 —0(.098
Y— A --0.020 — (0.044 -- 0.082 —0.674
3—4 —().326 —0).005 —3.859 0.062
4—4 ---3.566 --0.199 +-45.338 2.562
5 — 4 0.585 —0.001 BE elales —0.149
6—4 -+-0.055 +-0.008 0.687 -+-0.1638
Pell 0.078 =-0.162
2} —— 5) 0.005 0.045 0.033 —0(0.049
35 — 5 -+-0.016 —0).025 0.088 —0(0.095
4—5 —().182 —0.007 —2.657 —(.041
5—5 1.576 --0.110 24.256 1-722
6 —5 +-0.325 --0.004 5.163 —().006
7—5) 0.081 0.004 +0.567 --0.436
4—6 --0.009 —0.008 --0.079 — 6.269
5 —6 —0.100 —0.006 1.717 —0.073
6 —6 +0.707 0.060 +12.781 +J1.095
16 +0.176 0.005 -+3.260 0.050
S==6 0.018 —0.005 0.426 0.057

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 139

5 a os [MUN oO 7
We have next to transform the expressions for «(—) and ua? (= ust given

into others in which both the angles involved are mean anomalies.

From

beginning with m = 5, we find the values of 7, for values of ¢’ from £ to e”’.
Then we find

Putting m= 4, we find the values of 1, as in the case of r;. Then we get ps from

1. = —
Dae

(0)

We proceed in this way until we finally have the values of p, Then we find J, . or

(Fi — 1) from

where J=h’¢,

(m)
and J je from
2

The details of the computation are as follows:

140

A NEW METHOD OF DETERMINING

Computation of the J functions.

iiss se! é ag 2e ag 3e a) de’
log. 1 8.38251 8.68354 8.85963 8.98457 9.08148 9.16066 9.22761 9.28560
log. 7; 2.31646 2.01543 1.83934 1.71440 1.61749 1.53831 1.47136 1.41337
log. p; 7.68354 7.98457 8.16066 8.28560 8.38251 8.46169 8.52864 8.58663
log. 7, 9.91955 1.91852 1.74243 1.61749 1.52058 1.44140 1.37445 1.31646
log. r,—log. p,|| 4.53601 3.93395 3.58177 3.33189 3.13807 2.97971 2.84581 2.72983
ech —1 == 5 = 12 20 — 31 — 45 — 62 = 81
221954 1.91847 1.74931 1.61729 1.52027 1.44095 1.37383 1.31585
log. p; 7.78046 8.08153 8.25769 8.38271 8.47973 8.55905 8.62617 8.68415
log. 73 2.09461 1.79358 1.61749 1.49255 1.89564 1.31646 . 1.24951 1.19152
Diff. 4.31415 3.71205 3.35980 3.10984 9.91591 9.75741 2.62334 2.50737
Zech —2 —9 —19 — 34 — 52 = (8 —=103 = 1155
2.09459 1.79349 1.61780 1.49221 1.39512 1.31570 1.94848 1.19017
log. ps 7.90541 8.20651 8.38270 8.50779 8.60488 8.68430 8.75152 8.80983
log. Tr, 1.91852 1.61749 1.44140 1.31646 1.21955 1.14037 1.07342 1.01543
Diff. 4.01311 3.41098 3.05870 2.80867 2.61467 2.45607 2.32190 2.90560
Zech —4 =i =38 =6F =105 = =208 —200
1.91848 1.61732 1.44102 1.31579 1.91850 1.13885 1.07186 1.01974
log. ps 8.08152 8.38268 8.55898 8.68421 8.78150 8.86115 8.92864 8.98726
log. 7; 1.61749 1.31646 1.14037 1.01543 0.91852 0.83934 0.77239 0.71440
Ditf. 3.53597 2.93378 9.58139 9.33122 9.13702 1.97819 1.84375 1.79714
Zech = 18 —ff —=ll4 —=%2 = 44 =618 207
1.61736 1.31595 1.13923 1.01341 0.91537 0.83480 0.76621 0.70633
log. p; 8.38264 8.68405 8.86077 8.98659 9.08463 9.16520 9.23379 9.29367
log. U 3.53004 4.73716 5.43852 5.93828 6.32592 6.64264 691044 7.14240
log. 2.92798 4.13210 4.83646 5.33622 5.72386 6.04058 6.30888 6.54034
— log. F 6.76502n 7.36708n 7.71926n 7.96914n 8.16296n 8.32132n 8.45522n 8.57120
Diff. 3.83704 3.23498 2.88280 2.63292 243910 2.28084 2.14684 2.03086
Zech =i 25 =i =10l =I =27 =899 409
loge ( + =) 6.76495n 7.36693n 7.71869n 17.96813n 8.16139n 8.31905n 8.45214n 8.56718n
3.23505 2.63307 9.98131 9.03187 1.83861 1.68095 1.54786 1.43282
Zech = She pp 0 GG es
log. J 9.99974 9.99899 9.99773 9.99599 9.99375 9.99104 9.98787 9.98495
log. p: 8.38264 8.68405 8.86077 8.98659 9.08463 9.16520 9.23379 9.29367
log. J™ 8.38238 8.68304 8.85850 8.98258 9.078388 9.15624 9.22166 9.27792
log. p» 8.08152 8.38268 8.55898 8.68421 8.78150 8.86115 8.92864 8.98726
log. J 6.46390 7.06572 7.41748 7.66679 7.85988 8.01739 8.15030 8.26518
log. ps 7.90541 8.20651 8.38270 8.50779 8.60488 8.68430 8.75152 8.80983
log. J®) 4.36931 5.27923 5.80018 6.17458 6.46476 6.70169 6.90182 1.07501
log. p 7.78046 8.08153 8.95769 8.38271 8.47973 8.55905 8.62617 8.68415
log. J 2.14977 3.35376 4.05787 4.55729 4.94449 5.26074 5.52799 5.75916

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 141

Noting that log. (J —1) = log. (— P+ oo A=, and t=W2', we form

bo| &

the following tables :

17 Te) ee) 1 2) 1 _@
he) Log. (Suv —1) Log. Fun Log. Jw Log. iin Log. 5 Fu
1 6.7649 8.38238 6.4639 4.3693 2.1498
2 7.0658 8.38201 6.7647 4.9712 3.0527
3 7.2415n 8.38138 6.9404 5.3231 3.5807
4 7.3661n 8.38052 7.0647 5.5725 3-JooL
5 7.4624n 8.37941 7.1610 5.7658 4.2456
6 7.5409n 8.37809 7.2392 5.9235 4.4826
7 7.6070n 8.37656 7.3052 6.0567 4.6828
8 7.6641n 8.37483 7.3621 6.1719 4.8562

» (Wi)

rr u
Value of i URN

1 —— fst se) WSs Was Wt WSs
1) 4.9712n 6.4639 6.76495n 8.38201 6.9404 5.5125 4.2455

2|3.3537n 4.6703n 8.68341n 7.366938n 8.68241 7.3657 6.0668 4.7835

3 6.9410 8.85913n 7.71869n 8.85764 7.6381 6.4006 5.1598

4 49714n 7.36675 8.98344n 17.96813n 8.98147 17.8413 6.6588 5.4583

5 5.6702n 7.6393 9.07949n 8.1614n 9.07706 8.0042 6.8709
6 6.1012n 7.8432 9.15756n 8.3190n 9.15471 8.1402
alpHlorsha—0» 6.4176n 8.0061 9.22320n 8.452In 9.21993
8 | we have 6.6689n 8.1423 9.27965n 8.5672n
9 8.38251n 6.87TTin 8.2594 9.32905n

In computing the values of the J functions, the lines headed Zech show that
addition or subtraction tables haye been used. For conyenience, (J“’—1) is em-

ployed instead of J\, its values being found in the line headed log. (— P rt).

A NEW METHOD OF DETERMINING

From the expression

(ni)

((¢, h’)) = So Tv (2, v),
h’ being the multiple of g’, and being constant, and 7’ being variable, we have

sim

ay A 1 are cos /,° y 2 Gee) Cos (7
((4, h )) = i Jy = (Ag) = 18)) =F i Tux Sn (ig — 2H’) + ete.

(+2
/

tee SD 9
Six sin (ag =F HH ) are Six

h hi’

)
98 (ig + 2H’) —ete.

Now for h’= + 1, we have, if we write the angle in place of the coefficient,

° {) (pe 2 2) cos (7° D
(ig —g')) =4dy S(ig—EB) +25. & (tg —2H’) + ete.

(2) (8)
— ty sn (ig + BA) — Fy Sn (tg + 2B") — ete. ;
and for h’ = —1, we have
‘ (2) (23) Bayes
((g + 9’)) = —tJ_y smn (19 — Ei") — 4 d_y sn (tg — 2.H") — ete.

(0) (1) Le
4B Lo 8 Gye 19) 2d 8 Gy Ee DB) 4 ate.

sin

Since

(=m) (m)

(—m) (m) (m) (m
J_ nh = Weo9

)
=(—1%h, de =(—IP Le ,

the last two expressions give
(0) (1)
(9g —9')) = Jy & (ig — B’) — 2, & (ig — 2B") + ete.

(2 (3)

)
— JS, SF (ig + BH’) — 2S, & (tg + #’) — ete.,
9) (3)
(ig +9) = —Iv 8 (ig — B) — Dy & (ig — 2B’) —ete.

(0 (1)
+ Jy % (ig + EB’) — dy 88 (ig + 2H) + ete.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 143

And for the particular case of ¢ = 1, we have

(1
(g— q)) = Iu ely — Bi) — OF, % (g — 2H) + 3J, © nin (q¢—3H’) Fete.

2 (4)
De ey 0 (g sp Jo" OW Sn (g 55 2H’) an dS, Sia (g aF 3H’) — Cte

(2) (3)
(9 +9')) = —Iv B(g— EB) —2Jy & (g — 28

(4)
Jy SS (g — 3H’) — ete.

(@)
+ Jy es (g-+ B’) 20, v (9 + 2H") + 8dy sm (9 + 3H’) ete.

(0) (0)
Instead of J, , we use (-/,, — 1), as has been noted.
If we put h’ = + 2, we have

0)
((ig —29")) = $ Jov 23 (ig —E") + Jy SE (ig — 2H") + BJy, 8 (ig — BE") + ete.

L Joy °° (ig +B’) —2 Joy % (ig + 2B") — ete.

a (h’—7’)
In the table giving the values of | tux , we have, under h’ = 2, which applies to

the equation just given,

1 : (3)
for = 1, log. 2 Ji, = 8.38201 log. (4 Jy) = 4.97120:

(0) (4)

for? =2, log.( ey —1) = 7.366938n — log. (— 3, ) = 3.85370;

(1)
ie 0 == 63 log. (— $ Jo ) = 8.859182 efel sete:
ete., ete. =a ete
(3) (4)
We find the values of — 3 -J,,, — 3, in the table under h’ = —2. We see that
1 (W—1)
these are the forms of the function we Vn whens anded.— bh and: 24 == 2)

In the expansion of the coefficient of (¢g —h’g’) indicated above by ((¢g —h’g’)),
we have coefficients of angles of the form (7g + 7H’). These can readily be put into
the form (— 7g — 7H’), but the form employed is convenient in the transformation.

144

Arranging the functions ju G); (ua (

A NEW METHOD OF DETERMINING

Zs
4

Bi laa
) in this form, we have

tos. o() Loe. 12)

gy) LH cos sin cos sin

—l1 0.06387n 9.0043 0.50740 0.0210
0 — 2 8.9912 9.1106n 7.0000n 0.10827
() —= 3} 7.6493 8.04187 8.8751 9.2437n
1+1 9.6304 9.2856 8.0493 0.1637
1—1 1.72893 9.8663 2.3923 0.5364
1—2 9.9499n 8.4624 0.7404n 9.8432
1—3 8.6032 8.8882n 8.7716 9.9886n
1—4 TATTL 7.6021 8.7243 8.9912n
2+ 1 8.3617 8.6128 QOS 9.8105
®% — Il 0.8530 9.1584 1.0959 0.0978n
2—2 1.30203 9.7427 2.1675 0.6138
% — 8 9.7486n 7.5079 0.70457 9.4642
i —— 8.3010 8.6435n 8.9138 9.8287n
Y — 5) 6.6990 7.6532
3—1 8.9395 8.7993 9.2808 9.8944
3 — 2 0.2191 8.7993n 1.1221 9.9566n
2 — ® 0.91750 9.5368 1.9189 0.5440.
3—4 9.5132 7.6990n 0.5865n 8.7924
3—5 8.2041 8.3979n 8.9445 8.97TIn
4—]1 8.2041 8.6128 8.9868 9.8287
4— 9 8.9685 8.5051 9.7709 9.6839
4h — 9 0.0082 8.2869 1.0348 9.6410
44 0.5522 9.2989 1.6565 0.4085
AL —— I 9.2601n 7.8451n - 0.4244n 8.6128n
4=— 6 7.9542 7.90938n 8.8976 9.4298n
j= 2 8.8855 8.2049 9.8564 9.4506
i —— A 9.7672 7.0000n 0.8905 0.1732n
5 — 5 0.1976 9.0414 1.3848 0.2360
5 — 8 9.0000n T.1782n 0.2347n 8.8633n
6— 3 7.9440 8.1864 8.9385 9.3876
6 —4 8.7404 7.9031 9.8370 9.2129
68 9.5119 7.6021 6.7129 7.7782
6 — 6 9.8494 8.7782 1.1066 0.0394
Cy 0.0224 8.84517
7—6 0.5132 8.6990
[—17 0.8222 9.8156
7—8 9.7973n 8.7924

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

We will now give examples to illustrate the application of the tables for trans-

forming from eccentric to mean anomaly, in case of the function u(5).

For the angle 39g — 3q’.

5 i! ae
«(3) ee
g Jaf cos sin (i; = 8) Log. Product.
3— 1 8.9395 8.7993 6.9404 5.8799 5.13897
3 — 2 0.2191 8.7993n 8.68241 8.9015 7.6817n
3— 3 0.91750 7.5368 7.718697 8.6362n 5.2555n
3—4 9.51327 7.69907 8.98344n 8.4966 6.6824
3 — 5 8.2041 8.3979n 7.6393 5.8434 6.0372n
For the angle g — og’.
: (Br == 0)
1—1 1.72893 9.8663 8.388251n O.11144n 8.2488n
1+1 9.6304 9.2856 8.58251n 8.0129n 7.6681n
For the angle g+q’.
(h’ = 1)
1—1 1.7289 9.8663 6.46392 8.19287 6.3302n

Jd5 ley S\N, BIDS (Se

Product.
+ .00008 + .00005
+ .07970 — .00308
— .04327 — .00180
+ .03189 + .00048
+ .00007 — .00011
+8.26978 +0.34414
+8.33775 +0.33973
—1.29259 — .01773
— .01030 — .00466
0.25653 —0.25027
—1.04636 —0.27266
” "
=. WG .000
+0427 £0,198
Qiu LOGE

146 A NEW METHOD OF DETERMINING
For the angle og — og.
W
D1 0.0637n ee 8.3825n 8.4462 .-. + 2029794
+104.75727

4104.78521

For the angles represented by (7g — g’), there may be cases when there are sensi-
ble terms arising from g + H’, g + 2H’, etc.; if so, we use the column for h’ = — 1,
and apply the proper numbers of this column to the coefficients of the angles named.
Likewise in the case of (¢g + g’), there may be terms arising from the product of the
numbers in the column h’ = 1 and the coefficients of the angles g + ZH’, ete. This

will be made clear by an inspection of the two expressions

©) a)
(@g—g))= dv SG — B) — 2S, sn. (1g — 2H") ete.

(2 (3)
—— Jy % (ig + E’) — 2S, & (tg — 2H’) — ete,

sin

2 > C OS y i) 8) COSIN (Ey U
(ig + J) = —dy & (ty — H') — 2Sy S&S (tg — 2H’) — ete.

(0) )
+ Jy % (ig + Bl) — Wy % (ig + 2B’) + ete.;

where ((ag — g’)), ((¢g + g’)) represent not the angles but their coefficients.
In retaining the form (¢g + 7H’) instead of the form (— 7g — 7H’) we can per-
form the operations indicated without any change of signin case of the sine terms.
Making the transformations as indicated above, we obtain the following expres-

a\3

sions for the functions ue) and ua?(“) §

THE GENERAL

PERTURBATIONS OF THE MINOR PLANETS.

147

g g cos sin | cos | sin
| ||
ah | | I
0—0 +104.78521 | " | +182.3777 "

ino 04636, | — 0.27266 | — 1,6046 —1.9194
O60 = OIO5 030 eI +0.12527 — 0.5606 +1,1949
Bei9 + 0.02860 | S005 GE | + 0.1067 | +0,4943
20 | | — 0.1974 | —0.6468
il ii SrOrat —0.193 | — 0.0830 | —1,4558
(ai hee 0.107 | = Bic So ANOR
a + 53.583 0.734 1246,9027 +3.4023
OA 12 1986 = oil | + 5.3656 —1.4496
Seas | + 0.014 0.066 i — 0.3758 0.8304
=> + 0.070 | —0.127 == O05 —1,242
V9 + 0.399 +0.053 | + 0.456 0.848
2 9 + 20.093 0.551 | +147.392 +4049
PD + 1.056 —0.086 + 7.914 eS
Jy + 0.027 +0.033 — 0.086 +0.537
v= 8 + 0.00815 —0.01707 + 0.0718 —0.2352
i=? + 0.04342 —0.07447 + 0.0041 —0.9231
8 + 0.40733 +0.03392 | + 9.0442 +0.5514
P= 3 aE) 81338 -+0.340 + §$3.537 +3,432
ba 8 SEONG) —0.036 | =5 61432 —0.659
eee + 0,028 0.010 f + 0.079 +0.449

7 |

- |
9—4 + 0.027 —0.043 1 + 0.050 EBT
3—4 +5) 0.275 -+-0.023 | 41° 9144 9.592
aed | + 3.628 +0197 | + 46.016 +9512
pee Ee O39 —0.013 4 4.898 | 0,223
6—4 + 0,021 0.008 | + 0,156 0.188
S's | + 0.020 — 0.023 ! + 0,080 | —0.074
4—5 | + 0.167 0.012 sao | +0,241
5 | + 1,623 0.109 | + 24.829 | +1.565
6—5 | + 0,224 —0.004 ] + 3.306 —0.148

| I

| |
4—6 EEO O1e —0.008 EEN OOT —0.250
FG + 0.092 0.007 | — 4,535 | 0.150
6 —6 Out 0.059 + 13.312 +1.085

148 A NEW METHOD OF DETERMINING

The transformation should be carefully checked by being done in duplicate, or
better by putting the angle 7g = 0, in all the divisions of the two functions, having
thus only the angles (0 — #’), (0— 2H’), (0 —3#’), ete., ete.; also (0 — g’), (0 —
2g’), ete. Adding the coefficients in each division of the functions before and after
transformation, and operating on the sums before transformation as on single members
of the sums, the results should agree with the sums of the divisions of the transfor-
mations given above.

The transformations of these functions were checked by being done in duplicate,
but we will give the check in case of another planet. We have for the logarithms of
the sums before transformation, and for the sums after transformation the following :

g Ja! cos sin Ge af COS sin
0—1 1.85407 1.62090n 0—1 + 10.548 — 40.188
0— 2 1.25778 1.51473n 0— 2 + 19.809 — 32.318
0— 3 9.7024n 1.26993n 0 = 8 + 0.906 — 19.852
0—4 0.71012 0.9147n Q0—4 — 4.540 — 9.268
C= 5) 066327) 0:3899n 0=h = Ary = B33
0 — 6 0.4387n 9.0934 0 — 6 — 3.059 — 0.330
0—1 0.1222n 9.8069 0—7 — 0.623 + 0.739
0—8 9.5965 9.8865 0—8 -—— 0.071 + 0.615
For the angle (0 —1), (0 — 2), OQ = 8.
1) ey Ww 1)
— 0.041 + 0.024 + 1.722 — 1.007 + .062 — .087
— 0.873 + 1.578 — 042 + .076 + 871 — 1.574
.000 — 0016 + .037 + 1.346 + .003 + .097
+ 71.462 — 41.774 — 012 — .O19 + 494 + (791
+ 70.548 — 40.188 + 18.104 — 32.714 — .020 — .O11
+ 70.573 — 40.196 + 19.809 — 32.318 — 504 — 18.618
fae + 19.811 — 82.319 + 0.906 -+ 19.852
Bene hs, + 0.902 - — 19.355

The numbers in the last line of each case are the sums of the divisions after con-

version when 7g is put = 0.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 149
To have close agreement it is necessary that all sensible terms in the expansion of
3
a of @ . . . .
u (“) and wo?(“) be retained. In the expressions for these functions given a large

number of terms and some groups of terms have been omitted as they produce no
terms in the final results of sufficient magnitude to be retained.

In transforming a series it will be convenient to have the values of the J functions
on a separate slip of paper, so that by folding the slip vertically we can form the pro-
ducts at once without writing the separate factors.

: ° a of @ 2 °
The numerical expressions for u(“) and wa?(“) being known, we need next to

have those designated by (#7) and (J), which represent the action of the disturbing
body on the Sun.
To find (#7) we use two methods to serve as checks. We have first

(HT) = s[hyiyy’ + Wd8] cos (g — 9) — 3 [ys + Uys] sin (g — 9)
+ alhyy — hoy) cos (—~g — 9) — aly. — ty,0;] sin (— 9 — 9’)
+ ly! cos(— yf) — Br sin (=)
+ 2[hyry! + 2'8:8:] cos (gy — 29) 9 — LM.’ + Uys8s] sin (gy — 29’)
4. 2[hyiy.! — W’8,5:] cos (—g — 29) — 2[ ldo’ — Uy,6,/] sin (— g — 29')
+ 2hyvy cos (— 29’) — 217d, sin (— 2¢’)
+ $[hyys! + W0,8.] cos (g — 3g) — $[ldys + U10,'] sin (g — 39’)
+ ete.

where
We Rk b= deh

(1) (3) () (3)
y2 = 4 [Jo — J, ] On = I [So oF J ]

(4)

by 2) e)
ye = 4[Ja —Jn] & = 44a + Ya I,

and similar expressions for 7,', 51’, 7’, 6.’, ete.; noting that y) = — de.

150 A NEW METHOD OF DETERMINING

The other expression for (7 ) is
(HI) = Shy’ — 8] cos (— Bg’) + My’ — V8] sin Bg)
+ Mhy! + Wy] cos (H— g) ~ — Sly’ + 18] ein (B— 9’)
— ehy, cos (— 9’) + el’/dy sin (— g’)
+ 2[ hy.’ — h’d,'] cos (— #— 29’) + 2[ ly.’ — U,'] sm (— # — 29’)
+ 2[hy.! + h’d:'] cos (H — 29’) -- 2[ty.’ + Vd,’] sin (H — 29’)
— 4ehy,! cos (— 29’) + 4el’d,’ sin (— 29’)

+ ete. -++ ete.

In both expressions for (#7) we have

h =" keos(1I1— KX)

> V
hi = — cos pcos q’ ky cos (Ml — KG) = Su —
a a

p. : > —_ 7, Osim |
1 =~-—coso¢k sin (ll— KX) = s4u—,
a a

pee aN, : > — 1. pecosP

U = cos @’ k, sin (11 — #) =

where as before

w= ,206264.”8 and a=“.
1-+m a

In the second expression the eccentric angle of the disturbed body appears and we
must transform the expression into one in which both angles are mean anomalies.
With the eccentricity, ¢, of the disturbed body we compute the J functions just as
we did in case of e’ of the disturbing body.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. Pk

We have in case of Althea

te € 3e 2e

(0)

Log. (J—1) = 17.20740n 7.808947 8.160251 8.408900
(0)

Log. J — 9.99930 9.99719 9.99368 9.98872
(1)

Log. J — 8.60344 8.90341 9.07774 9.20016
(2)

Log. J = 6.90632 7.5077 7.8587 8.1068
(3)

Log. J = 5.0329 5.9356 6.4630 6.8365
(4)

Log. J = F087 4.2384 4.9418 5.4403 '

- (h—7)

i : :
From these values we may form a table of —-/,, as was done for the disturbing

h =

body. The values of these quantities can be checked by means of the tables found
in ENGELMANN’s edition of BrssrL’s Werke, Band I, pp. 103-109.
Finding the numerical value of (£7) first by the second expression, we get

H g | cos sin
| /
ea | 448.154 40.651
== a0 | + 0,188 —0,102
Qa i | BEG: —0.044
1D SEegigad 40.062
ie + 0.018 —0.010
O29 — 0.344 —0.004
iene} + 0.37800 1-0,00510
a3 + 0.00141 —0.00081
=e — 0.03048 — 0.00036

To transform we change from (hH—7'q’) into (7’g’ —hH). Making the transfor-
mation, writing also the values found from the first expression for the sake of compari-
son, and the value of (J) which will next be determined, we have

152

A NEW METHOD OF DETERMINING

(1) (1)
Ge gy cos sin cos sin sin cos
” " 1 " " "

0—1 — 5.826 —(0).066 — 5,824 —0.066 +-4,799 2,043

0 — 2 — 0.560 —0.006 — 0.562 —0.006 +0.463 +0.197

0—3 | — 0:04566 —0.00057 — 0.04575 +0.038 0.016
== | + @149 —0.103 + 0.180 —0.103

t=—i-| Aone 10.650 148.079 0.650

1—2 sats 4.637 +0.062 + 4.605 + 0.062

i= 3 | | 0.387740 +0.00502 —_ 0.37738 -++-0.00510

2—1 Se iL Oi --0.026 + 1.927 0.030

panto | 4+ 0,186 0.002 | 0.186 10,002

Y= 3 - 0.011 0.000 + 0.015 0.000

To find the numerical value of (Z) needed in case of the function a” ( ab we have

(1) =

where

b = —“ cos¢’ sin 7 cos I’,
i

+ 460’, sin (— 2g’) + 407’, cos (— 29’)

+ 9 bo’, sin (— 39’) + 9 by’, cos (— 39’)

+ ete.

bs’; sin (— g’) +

+ ete.

Biy'scos (— 9)

j= sine acim
2

Having the values of « (4), war Gy. (77), and (J), we next find those of

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 1538

from
a =u (“)—(H)
UO TON Laer ee ei ON
oF dr Ie (5) E ~ @ a zit (‘) (i)
5 dQ 9 s ind Mig 7 , )
iP = jl (=) = % sin (f’ + Il’) + (7)
where
rn Le Wig ea) 7 A
a> 1+ 3@— 4J, cos g — 4d. cos 2g — 4d, cos 3g — ete.
sinI 7’ (A) (3) : ;
= jam (77+) =— Ge aye ¢, sing’ — s[doy + Sry | ¢ sin 2g’ — ete.

(0)

+ 3é¢ a oe Jecug—e oor — Shy 16 Cos Qari.

c, and c, being given by the equations

sin I
c¢, = —— cos f’ cos Il’

gma! _o
= sin IT’.

2 [.- = “|= = [9.5769400] — 2[8.38238] cos g’ — 2[6.46366 |cos 2g’— ete.
+ 2['7.99450] cosg + 2[6.29667] cos 2g +- ete.
—str x sin(/’+ II’) = [7.18046] + 2[8.39074] sing’ + 2 [6.77809] sin 2¢/

— 2[8.01941] cos g’ — 2 [6.40668] cos 29’
A. P. §.— VOL. XIX. T. °

154 A NEW METHOD OF DETERMINING

In multiplying two trigonometric series together, called by HansEN mechanical
multiplication,

let «a, the coefficients of the angles 2a in case of the sine,
2, those of the angles ux in case of the cosine,
y, those of the angles vy in case of the sine,

and 4, those of the angles py in case of the cosine.

The following cases then occur :

ad, Sin (Aw + py) + 30,0, sin (Ax— py)

a, sin A@ . 6, Cos py =

tol

6, cos ue. y, sin vy = 4 B,y, sin (ue + vy) — $6,y, sin (ue — vy)

GB, cos ua .d, cos py = & 3,0, cos (ux + py) + 3,6, Cos (ua — py)
a Sin Aw. y, Sin vY = — $ ~My, Cos (Aw + vy) + Fay, cos (Ax — vy).

In every term of the second members the factor } occurs. Hence before multiplying

we resolve the coefficients of one of the factors into two terms, one of which is 2.

3 : ozo d2 »6dQ
Performing the operations indicated, we have the values of aQ, ar As that

* follow :

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS.

Q (da
aQ, ar(“ ) a(“ =
dr dz
G Gf COs sin cos sin GOS
I Wy aA : /} dW
O10 +104.78521 we 116.5202 1T>: 0.2828
i= — 1.04636 —.27266 — 2.4398 — .6940 —2.6311
= 0 = (03031 Sosy || = song SEES — .059
3= 0 J) 20 ORG SE ay Sec = O17
a) aE ORi —.090 = 3 E355 .000
C= + 4.662 1.173 ; == iL168 4 481 es
ek + 5.504 1.084 eeis39 190 HENS
2 — il —= iil —.201 | = 1.652 —= n'a —1.596
Bye oie +" .066 | = O40 4+ 288 — .059
@—® a 2632 1 | —- 497 —= ul — 020
Wwe — 4206 —.009 | 9.136 + .200 — 9.474
ee 9 + 19.907 1.549 | 145.566 11.270 + .095
5) Seino sG —.086 | 1.642 == il — 922
Velo ee erp EOE Pes 1) Ses — .064
|
= 3 -_ (OD =o | 22 ms = 0G — 00
ees = $2306 =O) | = 245 = S208 — .045
9—8 J. S999 dL EBRD |) == PANGS — -L SES —1.494
= 8 +. 8.338 +.340 ee oTeoon 1,087 — .064
A} JL Gn —.036 ATINTOG — .269 = S19
53 4. 028 +.016 a 043 aL RK = 09
eae Op —.048 = 054 —= M0 = 048
4 JL ON 023 = a + .908 ae ed
ye 4 8.608 1.197 -+15.430 + .882 = 038
Heed JL) AG (O18 J 838 137 — OD
—— a ee oo 4.008 = Og + 063 =O
3 —5 a020 == 023 — 034 = O78 + .020
Nees 2 GY 1.012 = si + 044 = Amd
5 = 8 +L 1,028 +109 4+ 8.605 + 543 + 024
6—5 JL Oy —.004 JL OSI + .064 — 52
AG + 0.012 =008 — 0.075 —0.095
ieeeG Ap 1.007 — 9,995 + .026
6=—6 SE pol 1.059 + 4.559 e386

156 A NEW METHOD OF DETERMINING

3 : : 0 : dQ
Having a© we differentiate relative to g, and obtain a—.
ag

We then form the three products, A. a Bar =). Oxa: =. To this end

we find A, B, C, from

A = —3 = 22 = cos (7 — g)
+215 + €] eos (7 29)

— 2 [BS + 282] cos y

+ 2£ cos (y--39)
+ 2 cos (y —4g9)

+- ete.

B=—2— sein — 9)
—2 [$+ 3] sin (y—29)
— 2[5 + qee"] siny
— 236 sin (y — 3g)
— 2¢sin (y— 4g)

— ete.

C= 2[f—e] sn(y— 9)

+ 2 [$—75e'] sin (y — 29)
+ 2[—#e+ 46] sin y

+ 23

+ 2 te
+ ete.

sin (y — 39)
sin (y — 49)

The numerical values of A, B, C in case of Althza are

A=—38
+ 2 [0.802429] cos (y — g)
+ 2 [8.604489] cos (vy — 29)
. — 2 [9.304508] cos v
+ 2 [7.2076]

cos (y — 89)

B = —2 [0.001399] sin (y — g)
— 2 [8.604489] sin (y — 29)
— 2, [8.606234] sin y

— 2 [7.3836] sin (vy — 89)

C = + 2[9.697567] sin (y — g)

+ 2 [8.80066]

sin (y — 29)

— 2[8.77953] sin y

+ 2['7.08265]

sin (y — 39)

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 157

For the three products we then have

O O
Al. al =) if; ar(=) Ol ch)
dq dr ike I
g 9 sin cos | sin cos sin cos
VW VW yl VV dy ah
o=0 EROS 35 yeeO 537 Ian a = IetS4iee 0.6804 —1,3464 —3.0038
h—@ = He JL 565 Eon ee srs + 1987 + .2411
i =O — (9530 +1 .0439 — 32.9502 + 0549 = Ay == ASOD
2—0 = 9p 1 299 SE oisoee 1657 (9049 + 0998
20 4 2.079 = 507 |) ee LIBI® ) SL Gea | esis | Beoee
FO Je Gil 4.457 Z J068 2 B80 4+ 083 + 2404
| |

=O — I ++ 462 ae GH 4. £89 = 883 | = iG +E 948

Sees = AG = ON cess + 461 ase 14 454
Oi —10.992 SE 1158 —18.335 a. Sy ly as BR eGo
Ci ae GD ae ie a 4 (349 998 + 572
eae AL BRA | Se hop) 2S a0 Si aes
1a SGING) PS 0} Bon eG 41,906 —4.470
aT = Bho, aL darn 4+ 306 4 (216 4+ 1067 _ 098
oi pie 4 949 INGS58 = aS SECTS, | 859
Boi PSR) 1 — 929 4 1559 Se 5 ee nIn(G0
ey 033 aL 2G = 2 = HG

5,229 4.982 000 =— OEY — 09 2 iA
Oo 4 6.837 4+ 026 ST S00 nee 235 —1.230 = + 3.029
O— 9 ee se om = 00
i 5 —80.684 19.195 — 45.412 LIGA || a Gye eS Ba
129 eis 002 iy 132 4+ 406 = 18 dk 0
79 | acqgie Louies me omen 364 — (G7 LOL
22 SENGLES, | OL) Sony See ORs D798) 81083
3 5) 4 499 aL BiG 4 LS + .168 4 024 L 023
BaD —19.078 42.954 145.412 —1.264 = 053 > = 8
es a 3 0 — SG kB af OE
5 — 2 £08 4. 1955 He OO 1 = GR

G8 SE 15985) * —-s.1553 4. Gin = SOG |) as OE Ey
(ees = 7657 Tey | 9 =e IW 2b AGM | OS SET
13 = eel 9 Joanie So cin ora Eos! 20180
5) 8} —50.140 +1.905 —27.9994 11.0854 JL 8 eS tea
a8 aa) 60, Le ales = song | 2 oR; iG) EE SRG
36 = S30 | 2 ep — 2.8964 — 2201 1p a ae
28 nico 2 073 — fai == 0 | ae oe Sane
L—® 4 263 + 1190 See 2 010 4.005
dbede 3 49.676 +2.079 497.999 —1.083 to39 0 206
53 S95) e264 Eo oe eee em nO 4+ 534

158

A NEW METHOD OF DETERMINING

Ala) /55 ar(S) C. aS)
dg dr dz

Me Ol OF sin cos sin COs sin cos

ih dW ih II iA VW

1 Weed == J1G5 = ahd == 1038 JE ls

1 Oe 4 — 2,929 ae TY + 264 + 989 == 389 +1.029
= ee eee aL Oil +. Oli oe. rr 2008 + .014

ie eee —929.032 1.564 —15.481 + .915 L 022 33
—1 3—4 JL 058 ==, IR = 039) + 175 5 {pA + .051

1 4—4 — 1.0638 — 287 — 1.504 = (0S) — .140 — .3800
ied — 4 + 1,268 — 024 022 2938 L .390 —1.033
i he —28.751 +1.597 +15.479 = 5 | .033 9
=| G4! — 4,543 — .108 + 1.506 + .098

i B= 5 = 180 =5 156 L .002 + 088

ily ey ees — 1.654 a 132 ==" 1063 + .063 — .206 26 Bi)
78225 4. Oe + 014 = {001 — .008 _ 001 ae00S

i 235 —16.185 JL 08 == 8.66! + 1544 | 034 + .038
—l 4—65 + 015 — .148 — 045 36 8 — .035 + .004

il B= 6 == IOI = .153 = IL — .036 == (0 = n68
—l 5—5 —E 294 — .017 + 062 — .063 | 206 = G3
—l 6—5 —16.0388 +1,100 + 8.661 — .b44

i 86 = im = 08

Teer een — 1.088 aL 083 | + 9.052 + .088

1 5—6 = BOL aL 08 == Aol6 —- 387
= NT 226 == 8818 ae pul + 4.516 == 38K

Next from 2

we find the value of ——.

ndt

OW = 4. a(“) + B.ar (=)
nat dq Ta

Then we find W and —“. from
Cost

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 159

We first form a table giving the integrating factors. From log. n’ = 2.4758576,

log. n = 2.9823542, we have “ = 0.34954524,

° *, e °, nv ° * nv 1 ° ° * n’ ° *, nv 1
= =o = ary, L = og, = OF amar iy

4 vl a+r = Log. (‘4 a a Log-() CU) Cao Log (i+7 =) Log. (7)
—2 — 1| —2.34954 0.37098n 9.62902 3 — 3} +1.95136 0.29034 9.70966
—1 — 1} —1.34954 0.130187 9.869827 4— 3} +2.95136 0.47002 9.52998

Q — 1) — .34954 9.54350n 0.45650 5 — 3] 3.95136 0.5968 9.4032

1—1} + .65045 9.813217 0.186783 Th 2) SIE TESTL 9.60008 0.39992n

2—1} +1.65045 0.21760 9.78240 2—4| + .601819 9.77946 0.22054

3 — 1} +2.65045 0.4233 9.5767 3 — 4} +1.601819 0.20461 9.79539

4—1} +3.65045 0.5624 9.4376 4— 4) +2.601819 0.41528 9.58472
2) 1.69909 0.230217 9.76979n 5— 4) +3.601819 0.5565 9.4435

0 — 2} — .69909 9.84467 0.1554n 6 — 4; +4.601819 0.6630 9.3370

1 — 2) + .30091 9.478423 0.521577 2=—5) | 2522974 9.40187 0.59818

2 — 2} +1.30091 0.11425 9.88575 3 — 5| 1.259974 0.09770 9.90230

3 — 2} +2.30091 0.86190 9.63810 4— 5) --2.952274 0.385268 9.64737

4 2} +3.30091 0.5186 9.4814 5 = |) =3.2520714 0.5122 9.4878

5 — 2) +4.30091 0.6336 9.3664 }/6 —5| +4.252274 0.6286 9.3714

0 — 31 —1.04864 0.020627 9.97938n 138 — 6] + .902729 9.9556 0.0444

1 — 3) = .04863572 8.6869553n 1.3130447n ||4— 6] -+1.902729 0.2794 9.7206

2— 3) + .95136 9.97835 0.02165 5 — 6| +2.902729 0.4628 9.5372

In regard to this table we may add that the form of the angles is (¢g + vg’) =
(i 14 2) g= (7 4 ) nt. he differential relative to the time is (2 414 “) ndt.
g n 7

t
The preceding table is applied by subtracting the logarithms of the column headed
, 1
log. (i aE “), or by adding the Jogarithms of the column headed log. (aa):

We will now give the values of —— W, and ——, remarking that in the inte-
nat cost

grations the angle y is constant; after the integrations it changes into g.

160 A NEW METHOD OF DETERMINING

ae W =
ndt cosz
: |
vy OF of sin cos | cos sin | cos sin
7 ” an yy 7 Sale " ”

1 O=@ 2. BPRG 1S |) == Wei 26 BOR pi | SONS; — IAAL ia!

1 1—0 + .38901 + .9373. — .3901 ak 9857/33 == 1287 + .2411

1 1 0 32.6972 —- .0988 — 32.6972 + .0988 | SL BIST T — .4802°

1 Y— (() —~—s-—« 20738 = 4647 — 6.1036 + .23823 | + .0024 + .0114
=i 90 4 OAS =k OSL = A OS AE. LASS
—l 3 — 0 + .13850 + .0850 — .0450 + .0283 — .028 + 0801

Te) eal 4. O04 == W167 BSB aL Oy — .033 = ld

1—1—1 + 187 + .446 + .115 — .330 —0.62 — 1.60

nee, es = 307 sb B40 — 83.900 = oe +1.013 + 1.84
—l 0—1 — 015. + .530 — .045 — 1.516 — .652 — 1.64

1 1—1 + 4.609 —1.374 — 1.087 — 2.112 - +1.264 — 2.74

ie Seibel Lo Bn = AS) — 1.030 — 5D —1.370 —= R21

1 4 —— Il — 036) = 153 + 022 + .45 — .040 + .06
—l 2—1 aE OR) SE OGL — 4,263 + .038 | a KO § — 21

eee al = OG) Se LOOT JE 9B |
=I) Bi “LAB = O84 == 4) = Oy = 26 + .670

Teas al oT a als J OBL 4 029)

1—1— 2 — .03 — .ll

1 0—Q + 14.145 + .261 + 20.207 = oie —1.76 — 4.33

1 o¥ 12ALG ALBIN) +419.660 +11.503 59 — 128

he PBS) 116 + .408 + 2,380 + 1.356 + .46 + .96

1 2— 2 -—— 1.8387 —1.119 + 1.410 — .860 ++ .36 — .78
—Il 2—2 + 9116 — .475 — 17.008 — .365 — .95 — 2.34

la 8.2289 Ho AS — 204 42 = He Oil
<1 §—9 — 33.666 -+ .990 ++ 14.632 + .430 == 02 = 19

1 te59 = Ole = 1% SE D5 SE 033
Se vee) — 8150 = 1G + 9.469 — .035 — 4 4. 20
= 9 = PIO se 4099 e050 4 {pH

1 O08 J- 1009 = 4914 | - 10ie6 +. A501 == .05 Seals

1 ls 3} — 1.5475 -—- .3568 — 31.8180 — 1.335 —14.56 — 371,33
=I ils = (012 + 0893 | = O52 = Lear 4. D5 = BY

ae = 78 — 77.4394 +9.9904 | + 81.400 + 3.139 = 0 == 318

1 2 3 EL 2Ory Se ee — .3124 + 1631 + .06 + .14

Te 338 = 397G4 — jel | = ier9 = BB5 1 183 = 28
=—l 8—=8 | + 98708 = 235 | — 1916 = 192 = 28 = ol

1 Gea B | sk Avis se Bey = 050 a ealiity .00 .00
=| 48 = OA, +. $98 | — 74g JL SS = OI =
Sl 5-58 Seo 196 ee 047 a eo = 12 = 08 ae 8

ih Shas = Ge } = Ane = 008 = 2

i Q—4 — 1.96537 41.126 | + 3.265 + 1.871 ae 64 (eae ole

ae = Wiis LO | aL BILTOD + 1.548 = 0 =
=—1 8-4 = 81 = 12 SO = 00" 4 O15 + 08

1 he = W5G7 = £5 + 986 = 149 J BA =D)
=i 44 ae 002) == 7963 == Ba = Bi = 150 =

to esd = (PQ sb soy SAO G + .016
—l 5+4 = 1B Ae 832 + 3.686 + .190 = 00 —
=i G4 == ROBT — {00 + 660 = 2

" |

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 161

LW sae u
: W —.
ndt cost
Lh Bor eaY Ve RAE S : =
mG) of sin cos cos sin | COs sin
i ” " A Tima 1 E i wi
1 38—5 =— iovily —- 21195 se Tae — 156 + .165 46
—l 3—5 + 011 + 017 — .009 SE 04 — .001 el
1 4—5 — 24.846 +1.626 + 11.080 + 722 — 015 == 02
1 4 5 030 — .072 + .013 — .032 | —- .016 00
a 5 — 5 — 2473 — 194 + .160 — .060 | —— 025 — (05)
—l 5 — 5 -+ 306 — .080 — .110 — 0294 | G4: —= ,llis
1 6—5 — .089 + .160 - 021 + .038 |
=I 685 = IRir = §58 Sei aL 180 |
—l 5 + 1.413 + .036 == 2 + .007 |
| |
1 4—6 a 964 + .124 = py + 07 |
1 5 — 6 — 13.228 1.090 Se ei f5x5)5) + .38 |
—l 5 — 6 — 167 + .023 -- 057 + .06
1 6 — 6 — .946 — .002 —- 242 00
—l 6 — 6 — %.098 — .040 -- .038 — .01
1 7 6 3.302 —- .824 + 674 —— .09

The part of W independent of y arising from the factor, — 3, in the value of
A, has not yet been given. Its integral, or f= oa cr is the following:

dg
S = 8a)

GG! cos sin 1 Of GF Cos sin
1—@ | + 3.1399 aL ‘8181 lik tas 3 — 2.74 jullie
== 0) SRE! Sear Say es al — 08
3=0 | = (eas = aes
Qa | = 48
ee he 51 Ae OH | eA iy =13
L=—i | —39 = 0) A—f | S164 =P 91
Oey = 9.33 J Read | = 168 +.05
al = 204 — .22 || 6-4 | — .08 —.03
|
p= 9 +41,934 JE 0) | 3— 5 |= Aen a6
Qs 9 Teo —2s8 | 2=—5 | = 29 —.06
2 9 |) = 48 4k Bl Rf = YA) —.50
49) — 10 = 19 6—5 | = 86 ALD
1— 3 | 20,0020 =e] 2—6@ | — 07 +.05
Q— 2 | — IAB = Md 5— 3 = 68 —.04
3123 5 8846 —1.57 || 6—6 | — 3.35 2

A. P. S.— VOL. XIX. U.

162

A NEW METHOD OF DETERMINING

Having the values of the coefficients of (+ y + 7g + 7g’), both for W and —<,

we have next to find those of (vy + 7 + 7g’), and of (Oy + 7g + 7q) in the case

cosa

The expressions for this purpose

7? =
7) =—
vo) =
yO =

For Althzea we find

log. 7 = 8.60309 log

e. n® = 7.388368

are

aly _1 9
te — le — se
Bis ILS at
se 1T28@
le

— (ge + =4e + etc.)

lox. 1 = 9.081960

We multiply the coefficients of ( y + ig + 79’) by 7, and 7, respectively,

to find those of (+ 2y + tq + 79’),

(= 3y + 9 + 19’).

In case of (Oy + zg + vg’) in the expression for <= we add the coefficients of

(+ y + tg + 19’) to those of (— y + 2g + vg) and multiply the sum by 7.

We will give a few examples to show the formation of WW, and — 4—.

dy
With these two we give at once also their integrals, which are néz and v respec-
tively.
= 1W
Ww = i°
> ihe
(O—)
cos sin sin cos
ida W ” W
ail =O —=890909 =E.0988 +16.3486 +-.0494
— 2 2— 0 0190 —-.0017 + .0190 +.0017
—32.7162 +.0511
” wu
—32.7162 -.0511nt

THE GENERAL

W
”
—1 2—0 — .4i74
® T= 23766
2—1—0 —1.314
1 O—O —1.2175nt

st
(1 — 0)
” | ; uw
+ .042 } ++ .237
+ .818 | a
— 004 —1.314
— .6087nt

3.2376 nt

PERTURBATIONS OF THE

d W

+ .004
—1.6188n¢

ia)

MINOR PLANETS. a

” wW dA

1.351 —1.2175nt + .856 +3.2376nt

” wu

—1.07TT —.6087Tnt

Mu "

+ .025 —1.6188nt

“ UA W f? ” ” LA dh
+459 —1.2175nt —2.07 —3.2376nt —0.54 +.6087Tné —0.58 —1.6188nt
Cae pag)
” Ww | A "
1—2—1 + .883 + .070 +.191 —.035
= 1 0— 1 — .045 —1.516 + .022 —.158
—2 1—1 — .041 — .030 +.041 —.030
@) —= il —1l —..913 + .200 = —
—0.216 —1.246 | + .254 —.823
W A | W uM
SL G == oP AL --.61
(l= 1)
Ww " | Mp Wy
OD ores ee 99 = 2004 ! + 029 —.004
—1 Bil == 43} + .038 + 2.131 +-.019
0 1—1 — 25.390 — .390 — —
1 0—1 — 83.900 — .973 —41.950 -+-.486
—113.574 —1.329 | 39.798 +.501
MW uw | uw WwW
—174.61 +2.04 | +61.19 10.77
: : = W
In the integration we apply the proper factor to each term of VW, —3 ,and

obtain the values of ndz, », except in case of the terms (7g + 0g’).

Let us take the term (gy — 0g’) or (1 — 0), and let uw the integrating factor to

be applied.

Let c, a, d, b, represent the cos, sin, nt cos, nt sin terms respectively.

164 A NEW METHOD OF DETERMINING

Thus we have

€ d a b
V/ ih I // >
+1.351 —1.21T5nt --.856 +3.2376nt ;
and hence
(“e wb ud — Lea wd —ub
uw Wy} V1 1] Pet VT
-+1.351 3.2376 —1.2175nt —.856 —1.2175 —3.2376nt
or, since w is unity,
dt a VI 1]
+4.59 —1.2175nt —2.07 —3.2376.

In case of the term (2 — 0), u is E.

In the way indicated we derive the values of ndz, and ». In the case of sont

we have the values at once without another integration as was necessary for ndz and v.
In the value of W given above the arbitrary constants of integration have not

been applied.
We give these constants in the form

It) + hk, cos y + ky sin y + 7) k, cos 2y + yk, sin Zy + ete.

1dW

Then in case of —3 7, We have
r

$k, sin y — $k, cos y + 7) ky, sin 2y — 7 k, cos 2y 4 ete.

Having W from the integration of ae we form W from the value of W and

converting y into g.
We thus have from the equation

ait W+e() ,

Selah,
+(1”.351 + %,) cosg ~ + (0’.856 + k,) sin g
-— 1”.2175nt cos g + 3/.2376nt sin g
+ (—'.284 + 7 k,) cos 2g + (0.589 + 74 hk.) sin 2g
—'’.0488nt cos 2g + ’.1298nt sin 2g

+ ete. + ete.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 165

Tn the second integration the constants of ndz and » are designated by C and VV
respectively, and the complete forms are

C+knt + k sin g —k, cosg + $y, sin 2g — $n hk, cos 2g + ete.
N — 1k, cos g — 4k, sin g — 37 k, cos 2g — 37 k, sin 2g — ete.
In case of the latitude the constants of integration have the form
1, + l, sin g + 1, cos g.
We thus find
nz = O+[1+ & —82".7162]nt
+ [4.59 + k,] sin g + [—2’.07 —k,] cos g
— 1’.2175nt sin g — 3”.2376nt cos g
+ [—0”.11 + $y, &,] sin 2g + [—0”.31 — $7 k,] cos 27
— 0’.0244nt sin 2g — 0”.0649nt cos 2g
—+ ete. + ete.
vy = +0”.0511nt + NV
+ [—0".54 — h,] cos g + [— 0.58 — $h,] sin g
+ 0”.6087nt cos — 1’.6188né sin g¢
+ [07.05 — 37” k,] cos 2g + [— ”.24 — 37 &] sin 29

+ 0”.0244nt cos 2g — 07.0649nt sin 29
+ ete. + ete.
= = 10.3616 + 0.3623nt

Cos2
+ [1.52 +14] sin g + [—0”.68 + 1] cos 9
—1”.3464nt sing — 3’.0038nt cos g
+ 0.32 sin 2g — 0.16 cos 29
— 0’ .0539nt sin 2g — 0’.1204nt cos 2g

+ ete. + ete.

166

gg
0— 0
LW
2— 0
0—1
0— 2
0— 3
1—1
2— 2
3— 3
4—4
5—9d
6 — 6
1— 2
2—4
1—3
2—1
2— 3
3 — 2
3— 4
4—3
4—5
5 — 4
—l—1

The complete expressions for ndz,

A NEW METHOD OF DETERMINING

NOZ
sin cos
+k, nt
_39,7162nt
= Aah ook
— 1.2175nt — 3.2376nt
— 011 + 44k, — ‘31 — 37%k,
— 0.0244nt — “0649nt
+ 3.10 — 3.09
— 3.00 + 1.92
+ 0.23 — 1.76
—174.61 + 2.04
+263.97 — 7.21
+ 25.15 = 8
a” Ril — 025
an 164 = Oi
zee 49 = 0%
185.18 4. 9.1
— 41.10 — .71
+410.16 —87.44
— 5.25 + .87
— 31.94 + 8.03
+ 6.17 + .04
+ .90 — .86
i. + 04
ily — £8
= 34 Oil
+ .16 = WW

COS sin
+ N
+- ‘O51 1nt

0.54 — Hh, 58 — ih,
tL 0.608Tnt — 1.6188nt
aE 05 hey OA Ly Ke?
+ (0244nt — 0649nt
+ 2.12 — 1.54
— 1.30 — .95
1 HO 4 28
+ 61.19 eT
—156.21 — 4,24
— 18.30 — .56
— 4°68 — 29
— 1.45 — .09
— .50 — .04
— 43.97 + 07
-- 36 — .01
+ 14.64 + 3.15
+ 4,02 aL 52)
| 16.07 aL SG
= 208 — .Ol
— 1.05 — 300
=>. 369 4c 05
— .33 — .04
-— 38 .00
1 AO 4. Gil

v, —— in tabular form are the following :
Cost

U
COS 4

sin cos

ef 0.36

; a '3628nt

SEO sph ae h == 98 IL I,

— 1 3464nt _— 3.0038nt
a “39 — 16

— "05390 — 204nt
— 4.83 — 2.03
aes J Bil
— .87 + .25
+ 2.69 + 1.26
— 1.15 = OT
— 1.60 — .60
4c 08 ze 02
= 6:64 — 270
= Ail = 7
4 4,43 ss
— 1.98 == 299)
—38.24 —14.92
= py + 20
si 11.30 SRD
— 24 + .03
+ .28 + .10
— 1.62 — .63

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 167

The constants of integration are now to be so determined as to make the pertur-
bations zero for the Epoch. The following equations fulfill this condition :

C+ k,sing— k,cosg + 47% k, sin 2g — $y” kh, cos 2g + ete. + (ndz)y = Yo

us (ndz), = 0

k + k,cosg+ ksng-+ 7k, cos2g + 7°) k, sin 2g + ete ai

ole
NN — $k, cosg — 3k, sin g — 37 ky, cos 2g — dy k, sin 2g — ete. + (v)o =
Smale

+ hk, sin g — 3k, cosg + 7K, sin 2g — x7 ky cos 2g + ete

d+ %sing + 4 cosg + 7 1, sn 2g 4- 4%, cos 2g + ete. + (a) = 0.

Ll, cosg — l,sin g + 7” 1, cos 2g — 7° 1, sin 2g + ete. 4 ( = .)) = 0

To find %, and k, we have

k, [cos g — e + 7°) cos 2g + 7 cos 3g + ete.] + ky [sin g + 7 sin 2g + ete. |

— 84, + 6 (r)) + 4-5 (nbz) = 0

k, [sin g + 27° sin 2g + 37° sin 3g + ete.] — k,[cos g + 27” cos 2g + ete. ]
d aa
ar Dae (ro ==

where

N= —2k—ja4—44%4, 4 = — 82".7162,

6
ky being found from

hey = chy + 3%, — 3 (nd2)) — 6 (0)

We have also

1 = — él,

The symbols (ndz),, (v)o, etc., represent the values of néz, v, etc., at the Epoch.

168 A NEW METHOD OF DETERMINING

To find the values of the angles (2g + 7'g) at the Epoch we have

g = 332° 48’ 53.2
g = 63 5 48 6

The long period inequality, 5 Saturn — 2 Jupiter, is included in the value of g’.
From these values of g and g’ we find the various arguments of the perturbations.
Then forming the sine and cosine for each argument, we multiply the sine and cosine

coefficients of the perturbations by their appropriate sines and cosines.

In forming aan (ndz), ete., we can make use of the integrating factors, multiply-

ing by the numbers in the column (« a < )e Having their differential coefficients we
proceed as in the case of (ndz), ete.

We thus find

(néz)) = + 401”.7, (v)) = + 180”.6, (“) = —22”.6

Sy (te) = 89026, 2G) 7075) (——) = are.

“ndt ndt \eost

And from these we have

SLOOP i OD SOP, ih = OO
= — 45.2, iL =-+ 0".4, NG 28.3.
C= Be Wy 1a",

The new mean motion is found from (1 — 32’.7162 — 26”.21) nt, which gives
n = 855’.5196. With this value of n we find the only change is in the coefficients
of the argument (1— 3), having + 405’.29 instead of 410.16, and — 86”.30 instead
of — 87.44.

The constant C now has the value

C’ = 332° 44’ 16”.3.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 169
Introducing the values of the constants of integration into the expressions for

U
nz, v, and— _, we have
COS?

nz = 332° 44’ 16.3 + 855’.5196 ¢
+ 417.4 sing + 80.8 cosg
— 1'.2175sing — 387.2376 cosg

sip O72 sim2g ee o.0) | cos 2g;

0".0244 nt sin 2g — 0'.0649 nt cos 2g

+ ete. + ete.
i ==) 284.3 + 07.0511 né
— 206.9 cos g a 4079 sing

+ 0”.6087 ntcosg — 1.6188 ntsing
— 8'.2cos2g + 17.3 sin 2g

+ 0.0244 ntcos2g— 0.0649 nt sin 2¢

+ ete. + ete.
See 104 4+: 07.3623 nt
Cost

— 44’ 2sing — 0.7 cosg

— 1'.5464ntsing — 3.0038 nt cos g
— 1’ 5sin 2g — 0.2 cos 29

— 07.0539 nt sin2g— 0.1204 nt cos 29

From the expressions of the perturbations that have been given, and the elements
used in computing the perturbations, except that we use C’ in place of g) and the new
value of the mean motion, we will compute a position of the body for the date 1894,
Sept. 19, 10" 48™ 52%, for which we have an observed position. From a provisional
ephemeris we have an approximate value of the distance; its logarithm is 0.14878.

A. P. S.—VOL. XTX. V.

170 A NEW METHOD OF DETERMINING

Reducing the above date to Berlin Mean Time, and applying the aberration
time, we have, for the observed date, 1894, Sept. 19, 72800,

OG = sas? UB) sel, g = 60° 24’.1.
Forming the arguments of the perturbations with these, we find
noz = + 4 437.2, ps LBB, ee OH (Sy
To convert » into radius as unity and in parts of the logarithm of the radius
vector we multiply by the modulus whose logarithm is 9.63778, and divide by 206264’.8.

Thus we have from » = + 3”.6, the correction, + .000008, to be applied to the loga-
rithm of the radius vector. ;

U 4
In case of —— = — 2’.8, we have
cost
(2a — 28 S<alcos = 1 a)
Converting into radius as unity, we have éz’ = — .000055. 'The codrdinate 2’ is per-

pendicular to the plane of the orbit. As we will use coordinates referred to the
equator we have, to find the changes in a, y, z, due to a variation of 2’, which we have
designated by éz’, the following expressions :

da = (sin 7 sin 2) dz’
doy = (— sin 7 cos Q cos ¢ — cos? sin «) éz’

dz = (— sin 7 cos Q sin e + cos 7 cos «) 42

where ¢ is the obliquity of the ecliptic.
For 1894 we find

da = (— .0404) dz’, dy = (— .3123) de’, oz = (4.9491) dz’
And for the date we have

da = + .)00001 dy = + .000011 dz = — .000033

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 171

With 0 =5° 44 46, == PAB Oil 1” (5), = 7B Al MO tes

we compute the auxiliary constants for the equator from the formule

. tg
cote A = — tg 2 cos 7 iG dy = =
g g » WY ED = Coe
— ose cos (£, + «)
cote B= — ae (Zo ;
tg 2 cos E, cos ¢
cot (6) ee au : sin (Fo a ©)
tg 83 cos E, sin ¢
: ‘5 si OSé« 0 sin 2 sin «
sm @ = oes 2 sin 6 = SE ECORI sin G! — $3 § =
sin A sin B sin C

The values of sin a, sin 0, sin ¢ are always positive, and the angle E, is always
less than 180°.

As a check we have

sin } sin ¢ sin (C — B)

YU a ane cost
We find
A = 293° 45’ 29’’.3, B= 2022 59746795 C’ = 210° 45’ 55’.0
log sin a = 9.999645, log sin 6 = 9.977735, log sin ec = 9.498012

Applying néz = + 4’ 45’.2 to the value of g, we have

nz = 339° 24’ 215

By means of g or nz = E — e sin E we find
E = 337° 39’ 23.4

Then from

Jr, sin $v = J/a(1 +e) sin 3 E

Jr, cos} v = /a(l1—e)cos3 E

172 A NEW METHOD OF DETERMINING

we find
Sout? HUI, log r, = 0.878246

where v is the true anomaly.

Calling w the argument of the latitude we have
n= ® 2 a = @ = IB? bey 4Ol“{8.
Hence

AS ul omy B+ u= 346° 52’ 28.7, O+u = 354° 38’ 36.8.

And from
«= rsinasin (A -+ w)
y = rsinbsin (B+ w)
z=rsinesin(C+ uw),
where

log r = log 7, + 6 log r = log 7, + .000008,
we have
aw = + 2.331894, y = — .515433, z = — .070208.
The equatorial cobrdinates of the Sun for the date of the observation are
X = — 1.002563 Y= + .045198 Z= + .019611.
Applying the corrections da, dy, dz, we have

w+ da4+- X= + 1.329332, y+ dy+ Y= —.470224, z+ 62+ Z7= — .050630.

THE GENERAL PERTURBATIONS OF THE MINOR PLANETS. 173

Then from
rab } 7 : y er, i zg —-L Oz ae Z
tg a et nme Mere tg — 24% 47 ang = ! ea Sarr
7 a + 0% + X y +oz + Y z+ oa + X

ee ee Pape Sed

sin 6

’

we have, giving also the observed place for the purpose of comparison,

a, = 340° 31’ 11.4 6: = — 223’ 23" 1 log A = 0.149514.

a, = 340 33 49.1 d= —2 2 25.4

where the subscript c designates the computed, and the subscript 0 the observed place.

Both observed and computed places are already referred to the mean equinox of
1894.0. If the observed position were the apparent place we should have to reduce
the computed also to apparent place by means of the formule

Aa =f+gsin(@+a)tyd
MS = g cos (G + a),
the quantities f, y, and G' being taken from the ephemeris for the year and date.

If the observed position has not been corrected for parallax we refer it to the cen-
tre of the Harth by means of the formulee

Riess zpcosg’ sin (a — 4)
ip A “cos 0
ig 9’
oy = =
IY cos (4 — @)
Tae =p sin g’ sin (y — 9)
4 : sin 7

where

a is the right ascension, § the declination, A the distance of the planet from the
Earth, ~’ the geocentric latitude of the place of observation, 6 the siderial time of

174 A NEW METHOD OF DETERMINING THE GENERAL PERTURBATIONS, ETC.

observation, p the radius of the Harth, and =z the equatorial horizontal parallax of the
Sun.

For the difference between computed and observed place we have
O— 0 = — 2’ 37.7 in right ascension, and C’— 0 = — 57’7 in declination.

- By the method just given we have found the positions of the planet for several
dates and have compared with the observed places. The comparison shows outstand-
ing differences too large to be accounted for by the effects of the perturbations yet to
be determined, which are the perturbations of the second order, with respect to the
mass, produced by Jupiter, and the perturbations produced by the other planets that
have a sensible influence. We have therefore corrected the elements that have been
used in the computations thus far made, by means of differential equations formed for
this purpose, employing as the absolute terms in these equations the differences be-
tween computation and observation for the several dates. A solution of the equations
has given corrections to the elements that produce quite large effects on the computed
place. Thus recomputing the position of the planet for the date given above with the
corrected elements we find

a, = 340° 33’ 44.5 , 6, = — 2° 2 15”6.
And since
oy = 840° 33’ 49”.1 , 6) = — 2° 2 25” 4

we have, for the difference between computed and observed place,

C— 0 = — 4’.6 in right ascension, and C— O = + 9’.8 in declination.

ARTICLE II.

AN ESSAY ON THE DEVELOPMENT OF THE MOUTH PARTS OF
CERTAIN INSECTS.

BY JOHN B. SMITH, Sc.D.

Read before the American Philosophical Society, February 21, 1896.

Since the publication of my paper on the mouth parts of the Dzptera, printed in
the Transactions of the American Entomological Society for 1894, I have continued
gathering material, have examined the oral parts of a very large number of species of
all orders, and am more than ever convinced that in all essentials the conclusions
already published by me are correct —revolutionary as they seem at first sight. That
my ideas have not found unquestioned acceptance is not surprising; but no one has,
to my knowledge, published anything that disproves the points made by me. It has
been suggested, however, because I have not made continual reference to the works of
previous authors, that I was ignorant of the literature, and several papers have been
cited as contradicting my conclusions.

As a matter of fact I believe I am fully aware of all that has been written on the
subject, and have, in each case where my attention has been called to a paper, studied
it carefully, and found nearly always that the facts given bear me out, though the con-
clusions are adverse ; simply because no author has seriously questioned the univer-
sally accepted homology of the month parts in the various orders. My own studies
have been made on a basis so radically different from any heretofore accepted, that my
results must stand on them alone, and my conclusions, if valid, must stand on the facts
as they appear to me. I have used principally the dissecting needles in my work; but
have not neglected the section cutter. This latter instrument has been rather too
much used at the expense of the needles, and its results, though undoubtedly accurate
~as a record of facts, are easily misinterpreted if the basic homology which is assumed

176 AN ESSAY ON THE DEVELOPMENT

to exist is inaccurate. For the reasons just given no references to previous writers
will be made, except incidentally, and as I have in some respects modified my views
as to the homology of certain of the parts, I will go into the entire subject in such
detail as is necessary to prove my point; but without reprinting my first paper, which
should be herewith consulted.

I do not expect denial at this day, when I claim that no explanation of the homol-
ogies of the mouth parts of insects can be considered satisfactory which will not stand
the test of criticism by the theory of evolution. If we assume the origin of all insects
from one original type, we must, necessarily, assume that all the mouth structures are
derivatives of one type, and we must so study them as to be able to explain, step by
step, just what specializations have occurred. We may not be able to complete en-
tirely each link in the chain of evidence, but we can, at any rate, reach a result con-
sistent with all the facts known to us. Any explanation which satisfies all the require-
ments of a regular and natural development is to be preferred to one which demands
an unexplained specialization of any part, not in line with its function in other series.
It is therefore necessary to study carefully the make-up of every separate mouth
organ, and of every sclerite in each, to become thoroughly familiar with its uses and
to ascertain the lines in which it varies or develops.

It may be premised that the mouth parts of the Hemzptera in their present con-
dition are not included in the range of these studies. I have examined numerous
specimens and have devoted especial attention to Cicada and Thrips—the latter
classed as hemipterous for present purposes only—and I believed at one time that I
had made out the remnants of a mandibular sclerite, and so published it. Mr. C. L.
Marlatt questioned my conclusions and asserted that the mandibles are represented by
one pair of bristles. While I believe that I was wrong in my identification of the man-
dibular sclerite, 1 am yet convinced that I am correct in claiming that beak and sete
are all maxillary structures. I have concluded, however, after a careful review of all
my preparations and of what has been written, that the Hemzptera in the mouth struc-
ture are not descended from any well-developed mandibulate type, and that no trace of
true mandibular structure occurs in any present form.

In other words, the Hemiptera equal all the other orders combined in rank, for all
others are mandibulate or derivatives from a mandibulate type. The archetypal Thy-
sanuran with undeveloped mouth organs varied in two directions—toward the
haustellate type now perfected in our present Hemiptera, and to the mandibulate type:
and there has never since been any tendency toward a combination. The haustellate
type proved ill adapted for variation and there is, in consequence, a remarkable same-
ness throughout. This kind of stracture must be studied on an entirely new basis to

OF THE MOUTH PARTS OF CERTAIN INSECTs. al 77

get at the steps by which the present “beak” was developed, and my material is not
sufficient for that purpose. The mandibulate type, on the contrary, proved well
adapted for variation, and its differences and modifications are here traced.

For convenience, Kolbe’s figures of the mouth parts of a grasshopper are repro-
duced on Pl. III, Fig. 22, and may be referred to in connection with the following
explanation.

In a well-developed mandibulate mouth we have, forming an upper lip, the lab-
rum, often notched in front or toothed; but never a paired organ, never with appen-
dages, and never mechanical in function. It is articulated at base to the clypeus and
serves to shield or protect the mouth in front; as a matter of fact, not a functional
mouth structure at all. It is marked ldr in all figures.

More or less intimately associated with it on the inner side is the epipharynx, which
is compared in function with the palate of vertebrates, and is furnished with sensory
hairs, pegs or pittings. It may be so closely united with the labrum as to form, prac-
tically, a part of it, or may be entirely free. If free from the labrum, the epiphary nx
is more closely united with the other mouth parts, and in such cases its supports go to
the mentum or labial structures. Not infrequently it has attachments to both. In
form it may be a mere pointed process, or it may be a more or less divided, plate-like
organ; but its functions are gustatory or sensory in all cases—it never becomes a
functional mechanical structure, and I have never found it without a more or less de-
veloped labrum to shield it. It is lettered ep? in all figures.

Just below these covering and gustatory organs is a pair of mechanical structures
—the mandibles—set, one on each side of the head, and attached to the inferior margin
of the epicranium or an extension from it. These mandibles are never jointed, rarely
bear appendages, and never such as are functional, rarely have a movable tooth, and
are usually solid and highly chitinized. They are actually made up of a number of
sclerites, laterally united, but distinguishable in certain types like Copris, Pl. I, Fig. 8.
I have elsewhere named and homologized these sclerites; but as the matter is not in
dispute, and of no importance here, a simple reference to the figure in which they are
named is all that is necessary. The position of this pair of mouth structures is inva-
riable. They are completely disassociated from the maxillary or labial structures and
remain attached to the head when all the other parts are removed in a body. They
attach by socket joints to the epicranium and their tendons and muscles attach to
its inner surface. They never change in function, never become united with or
attached to the other mouth organs and never become internal structures. When not
needed for chewing or biting the tendency is to obsolescence: never toward a change
into a thrusting or piercing organ, so far as my observations extend.

A. P. S.—VOL. XIX. W.

178 AN ESSAY ON THE DEVELOPMENT

Below the mandibles are found a pair of maxillee, made up in all cases of a number
of sclerites, and nearly always supplied with palpi or jointed tactile organs. The
more particular consideration of these organs and their parts may be somewhat
deferred.

Forming the lower lip and closing the mouth inferiorly is the labium, also made
up of a number of sclerites and usually furnished with palpi. It is never entirely
paired in existing insects, but is assumed to be made up of two more or less united
structures, similar in essential character to the maxilla, as has been well stated by
Prof. J. H. Comstock. This labium is an exceedingly important structure and forms
the oral termination of the digestive tract or the mouth of the cesophacus.

Attached to the inner surface of the labium is the hypopharynx, a variably devel-
oped structure, which is supposed to be the remnant of another originally paired organ,
the endo-labium. I have never seen the genera in which it is said to be well devel-
oped, hence have no well-founded opinion to offer. I find it uniformly a single organ,
often highly developed and gustatory in function, sometimes a merely passive structure
more or less closely attached to the ligula, usually very near the opening into the
digestive tract.

Briefly recapitulated, the insect mouth, when most fully developed, consists of
two pairs of lateral jaws moving in a horizontal plane between an upper and a lower
lip, which are furnished with gustatory structures forming the roof and the floor of the
mouth respectively. This mouth is adapted for biting and chewing and varies to types
adapted to lapping, to sucking only, and to piercing and sucking. The problem before
me is to ascertain by what modifications these different changes in type have become
established.

If we examine the head of a well-developed mandibulate insect from the under
side—Copris carolina, P|. I, Fig. 7, may serve as type—we find, centrally, the gula or
throat, bounded laterally by the gene or cheeks, extending to the posterior margin of
the head and bearing anteriorly the labium. The labium when carefully dissected out
is found to consist of a broad basal plate, the submentum, more or less firmly articu-
lated to the gula and never, in existing insects, a paired organ. It bears anteriorly
another plate, the mentum, also a united organ, though sometimes traces of a division
are apparent.- It is usually smaller than the submentum, sometimes membranous,
often entirely separated and frequently so united with the latter part that the two are
not separable. Though the submentum is the most persistent and dominant structure
it has been customary to use the term mentum to apply to the united sclerites, and it
will become convenient for me to so use the term hereafter when no confusion or mis-
understanding can be occasioned. The structure is lettered m in all the figures.

OF THE MOUTH PARTS OF CERTAIN INSECTS. 179

Attached and articulated to the mentum anteriorly are the central ligula, a pair of
paraglossa bounding it, and a pair of palpigers, one at each outer edge, bearing the
labial palpi.

The ligula or glossa, marked gl in all the figures, is a paired organ only in the
more generalized orders, and is usually present as a single, central structure, which may
be either chitinous and rigid or membranous and flexible. It is the most persistent of
all the labial structures, is never attached except to the mentum, and always has asso-
ciated with it the hypopharynx where that is present. We always find at its base the
opening into the alimentary canal, or cesophagus, as this part of it is termed, and this
must eyer be the test of labial structures—that they are attached to the mentum and
have at their base the opening into the alimentary canal. The association is never
broken, and the base of the ligula, whatever its form or however it is modified, always
marks this point. On the other hand, by tracing the alimentary canal to its external
opening, we can always recognize the ligula by its position, however little it may re-
semble normal types.

The paraglossx are sometimes intimately united with the ligula, sometimes com-
pletely separated from it: they may be of the same or a different texture; but they
always arise from the mentum on each side of and close to the central structure. Their
tendency is to obsolescence, but they may become united and form a bed for the ligula
which remains the inner organ. Their range of variation is not great; they are never
jointed, and never become mechanical structures.

The palpi are tactile in function under all circumstances, though they may lose
this function in great part and may, by coalescence, form a sheathing to the ligula.
They are never, under any circumstances, attached anywhere except to the mentum,
directly or indirectly, and their location must be constantly the same. They cannot,
without losing their essential character, become disassociated from the mentum,
nor can they ever form an envelope or covering for it, or for the submentum, with-
out a change entirely at variance with any reasonable theory of development. ‘To
accomplish this they would first lose their character as labial appendages. In
brief, the labium is the external beginning of the alimentary canal, and none of the
parts ever lose this association. Whatever their modification, no labial structures
can eyer be joined to the sides of the head outside of mandibular or maxillary
structures.

As an illustration of the most generalized form of labium at present known to
me, the roach (Periplaneta orientalis, Pl. II, Fig. 16) may be selected. Here we find
the mentum with a well-defined impression resembling a suture, and bearing a broad
paired structure, from which arise the slender, two-jointed ligula, the broad, fleshy

180 AN ESSAY ON THE DEVELOPMENT

paraglosse, and the three-jointed labial palpi.. This generalized structure fixes the
relation of the parts, and from it we may pass to more specialized types.

In Harpalus caliginosus (P1. III, Fig. 7) we have a case where the ligula forms
a single, central organ, laterally bounded and on one side completely enveloped by the
softer paraglossze. The location of the palpi remains essentially the same. We have
here two cases showing the change of a two-jointed membranous paired organ into a
single, rigid, chitinous structure, and the identity of the parts is not questioned, nor
I believe, questionable.

If we carry our dissections one step further and from the fresh specimen remove
not only the highly chitinized parts, but also the softer attached structures, leaving
maxillze and mandibles undisturbed, we find in all cases the cesophagus in the cavity
below the mentum and submentum, and these sclerites afford attachments for neces-
sary muscles. They also form, by means of chitinous extensions and processes, a
chamber or cavity protecting the cesophagus and supplying muscular attachments
when a sucking or pumping structure is needed. Thus the mentum and submentum,
whether separated or united, are always inferior coverings to the cesophagus. To sup-
port this structure, processes sometimes extend almost or quite to the upper or anterior
surface of the head, and in many cases, where the epipharynx is separated from the
labium, it is connected by means of long processes with the mentum. This is true in
many Coleoptera, quite usual in the Hymenoptera, and occasionally found also in the
Diptera. In PI. I, Fig. 6, is a lateral view of the labium of Copris carolina when
completely dissected out, and the clubbed processes, loosely attached to the inferior
prolongation of the submentum, normally support the epipharynx. In PI. I, Fig. 9,
and Pl. II, Fig. 18, we note similar processes in Andrena vicina with part of the epi-
pharynx still attached, and in Polistes metricus, where the structures are complete.
Precisely the same structures occur in Simuliwm (P1. I, Fig. 1“), as will be more fully
noted hereafter. It may be stated that I have adopted the term “ fulcrum,” used by
Macloskie and others, to designate the structure formed by the mentum and submen-
tum and containing the beginning of the alimentary canal.

In Polistes metricus (P\. I, Fig. 18’) I show the labium completely dissected
out, with all its attachments, viewed laterally. It will be noted that here the mentum
and submentum are united, highly chitinized, and form a scoop-shaped structure,
bearing at one end the labial structures and enclosing normally the beginning of the
cesophagus. Attached by long chitinous rods to the posterior angles is the epiphar-
ynx, so that hypopharynx and epipharynx are borne on the same base, are closely op-
posed to each other and may be manipulated by muscles arising close together. The
origin of the palpi is shown from the mentum. On PI. II, Fig. 18%, are shown ligula

OF THE MOUTH PARTS OF CERTAIN INSECTS. 181

and paraglosse of this same Polistes. The structures are here membranous, some-
what bladder-like, and well adapted for lapping by means of flattened, bent processes,
set in series on the entire inner surface. The paraglossze are completely separated and
the mouth opening is shown at the base of the figure, as well as the chitinous ring
marking the beginning of the cesophagus.

In Andrena vicina (Pl. I, Fig. 9) we find a similar yet quite different structure,
v. é., the same parts, used for much the same purpose, yet considerably modified in de-
tail. The mentum is here much longer, more shallow, but similarly bears the epiphar-
ynx on chitinous rods. The ligula is more inflated and the paraglosse are much
reduced, but the palpi originate as before, and we have simply an illustration of the
variation in form found in this united mentum and submentum. It is important to
note here that in Polistes, Andrena, and indeed the Hymenoptera generally, the labial
structures are free from all lateral attachments to the head and may sometimes be pro-
jected forward quite a distance. The attachment to the head, indeed, is muscular and
membranous entirely, and there is no direct articulation to any point by chitinous or
rigid processes. There is nothing therefore to prevent the growth of the head sclerites
around the mentum, which would thus become an internal structure—as has actually
happened in the Diptera.

Another feature upon which Dr. Packard rightly places great stress is that a
salivary duct opens into the hypopharynx at the base of the ligula, which he thereby
identifies. As this ligula is always attached to the mentum, it follows that this struc-
ture may be identified in the same way, while no structures not originating from the
same point can be labial in character.

Before studying further the specializations of the labial structures, it may be well
to say that they sometimes tend to become useless or obsolete, or so much reduced that
they are difficult of recognition ; and, curiously enough, in such cases the palpi seem
to be the persistent organs, Thus in some species of Scolzide among the Hymenop-
tera the mentum bears only little, feebly developed palpi. A striking case is in the
Panorpide, where on Pl. III, Fig. 4’, the mouth structures of Bittacus strigosus are
shown. Here ligula and paragloss have disappeared entirely; but the palpi are dis-
tinct and the curiously developed hypopharynx marks the beginning of the opening
into the cesophagus.

A modification of this type is to be found in the Lepidoptera, where practically in
all cases the palpi alone, attached to a plate of variable size and shape, represent the
labial structures.

It seems a long jump from the reduced type in Panorpide to the fully developed
labium of the Apide ; yet, except for the fact that all the parts are much elongated,

182 AN ESSAY ON THE DEVELOPMENT

there is no difference from Andrena or Polistes, which have been already studied. I
have found no species which shows all the parts more fully developed than Xenoglossa
prumosa (Pl. II, Fig. 15). Here all the parts are equally developed and all are func-
tional; hence it makes a good starting point. The mentum is not shown in the figure
except at the point to which the other parts are attached, and surmounting it cen-
trally, we find the ligula; here a united, though extremely flexible organ. Lying cen-
trally upon it, so as to close a groove, is the hypopharynx, in this case not easily separ-
able from the ligula. Arising close to the central organ on each side are the para-
gloss; almost as long as the glossa itself, flexible, unjointed, flattened and a little
incurved at the margins so as to form, when closely applied to it, a partial shield for
the ligula. Outside of all, situated at the outer margins of the mentum, are the palpi.
These are four-jointed; but the basal joints are enormously elongated in proportion to
the terminal two, and they are also flattened out, broadened and infolded, so that when
at rest they cover and almost conceal the other labial parts, though not extending for-
ward as far as they. In this insect the structures just described are almost entirely
covered by the maxille, and a transverse section (PI. II, Fig. 15") is interesting and
instructive. It represents the structure at about the middle of the combined maxillee
and labium and illustrates the relative position of the parts.

The tendency in the bees is toward a loss of the paraglossz, which shorten grad-
ually until they disappear altogether, as represented in a species of Bombus figured in
Pl. II, Fig. 15. Every intergrade is represented in any good series of bee mouth
parts, and in their rudimentary condition, without function, they appear in Bombus sp.,
represented on Pl. III, Fig. 6. The palpi retain their unique development, and in
the figure just cited are seen to be as long as the ligula itself, the basal two joints en-
folding it almost completely, while the terminal joints are much reduced in size and
set near the tip of the second joint, on the outer side. in other species these terminal
joints are proportionately yet more reduced and are sometimes difficult to find. The
essential point to be noted is that at their best development the paraglossz are not
jointed and that they tend to complete obsolescence in the most highly specialized
types. The palpi in Bombus require a little further examination: Reference to the
figure last cited will show a short segment between the mentum and the first long
joint, and this is membranous in texture. The mouth parts in Bombus are folded
when at rest and the hinge is at the mentum; hence the necessity for some such pro-
vision to enable the palpi to bend safely.

Now let us assume that the ligula of this Bombus became rigid and chitinized,
and that the edges of the palpi enfolding it became united to form a complete cylinder ;
and then let us examine Hristalis tenax (Pl. ILI, Fig. 5) in the light of this assump-

OF THE MOUTH PARTS OF CERTAIN INSECTS. 183

tion. First let me say that I have already shown that a change from flexible to rigid
ligula is not uncommon, and the suggested union of the palpi is a much less violent
requirement than that imposed by the current explanation of the Dipterous mouth.
Referring for a moment to Pl. I, Fig. 3, we see the entire mouth structure of Hristalis
tenax. Above is the mentum and submentum, very like the structure already de-
scribed for Polistes and entirely homologous with it, and at its tip we find arising in a
group the structures further enlarged at Pl. IIT, Fig. 5. Centrally we find the now
rigid ligula, deeply grooved in the middle, the channel closed by a flattened, also rigid
and chitinized hypopharynx. Loosely enveloping this central ligula is a more mem-
branous cylinder, evidently made up of two lateral halves, two-jointed, and the ter-
minal joints separated or paired except at the base. As in Bombus the mouth of Hris-
talis is hinged, and the joint is also at the base of the ligula. The latter organ is so
articulated as to allow of the flexion; but in the palpi we find again the provision
already noted in Bombus—a flexible, membranous, pseudo-segment. Now if we sec-
tion the Bombus and Hristalis at the middle, we find the cuts alike, except that in
Eristalis the palpi are completely united over the hypopharynx and closely approxi-
mated at the opposite side. If we section near the tip, the cuts in both cases are
identical. That this united structure in Hristalis is the united labial palpi seems to
me beyond doubt. In the first place, the point of origin is normal, next to the ligula
and at the tip of the mentum; and, secondly, it is a jointed organ and therefore can-
not be paraglossa. It is in all points the structure of Bombus, with the terminal joints
lost and the two halves united for the greatest part of the distance. That the parts
named mentum and submentum are really such, is proved by the fact that the hypo-
pharynx, which is not in dispute, originates from and that the cesophagus originates
within it.

In Bombus fervidus the ligula is unusually developed and much longer than the
labial palpi, while the paraglosse are wanting. In PL. III, Fig. 12, is a camera lucida
sketch of the labial parts of a carefully mounted specimen. The structures here are
exactly as normally held when at rest, and only the mentum is a little crushed by the
cover glass on the shallow cell. Now chitinize this whole structure thoroughly, and
then compare with the drawing of Chrysops vittatus (Pl. III, Fig. 13) made in the
same way. The magnifications are different, of course, the Bombus being drawn at
short range with a four-inch lens while the Chrysops was drawn at long range under a
one-inch objective. The object was to get the two of approximately the same size for
convenience of comparison. In the Tabanids the mouth parts are rigid and not flexed,
and no sort of joint or hinge is required ; hence the structures are all rigidly united at
the base to the mentum. In Bombus fervidus the palpi are reinforced by a heavier

184. AN ESSAY ON THE DEVELOPMENT

chitinous rod a little to one side of the middle, and just this sort of structure we find
everywhere in the 'Tabanids, lying outside of the ligula at base, articulated to the
outer edge of the mentum. This, in fact, first led me to suspect the true nature of
the structure. If now we section Bombus and Tabanus near base, the cuts will be
alike, save that the palpi in the latter are united at one margin. If the cuts are made
toward the tip, the sections are alike—ligula and hypopharynx alone appearing in both
cases. We have then, in Chrysops also, a complete labium, save that the paraglossze
are absent and the palpi are united on one edge.

In the Simulide are many interesting species with generalized mouth structures,
and of these I have studied the “ Buffalo gnat,” from material kindly furnished by
Dr. Riley, an undetermined Simuliwm sent me in numbers by Prof. Aldrich, and an
undetermined little midge collected by me at Anglesea, N. J. The species are prac-
tically identical in the labial structures, and here again the mentum and submentum
strongly recall Polistes and other Hymenoptera. The hypopharynx is well developed
and the ligula are nearly divided; but I have no satisfactory sections of this insect
and the relations of the parts are not clear to me. At Pl. I, Fig. 1°, the labium of
the “ Buffalo gnat” is shown. In the species sent by Prof. Aldrich I succeeded in
getting a dissection illustrating the connection of the epipharynx with the mentum,
and this is illustrated at Pl. I, Fig. 1% This is really an exceedingly interesting speci-
men and it clears up the relation of the frontal prolongation of the mouth. That the
structure so labeled is really the epipharynx there is little room for doubt, and the
location of the little, chitinous, toothed processes, and their character, leaves no doubt
in my mind that they are mandibular rudiments—exactly as I claimed in my firet
paper. ‘That they can be dermal appendages, as has been claimed, does not seem rea-
sonable to me. They are too highly chitinized in comparison with their surroundings,
and why should they so completely resemble miniature mandibles? I do not know of
any case of dermal appendages of a similar character, and it is at least passing strange
that such should be developed exactly where, normally, mandibular rudiments might
be reasonably expected.

The tendency in the piercing Diptera is constantly in the direction of simplicity
of labial structures, and so we gradually note the loss of all trace of accessory labial
structures, leaving the ligula and hypopharynx as sole representatives. In the As-
ilide there are no other attachments to the mentum, as shown in PI. III, Fig. 1’.

These apparently single structures are sometimes interesting in section, as appears
in Stomoxys calcitrans, Pl. I, Fig. 11. Here the cut shows two crescent-shaped struc-
tures connected at one edge by the thinnest kind of a chitinous shell, and closed oppo-
site by a hypopharynx, which is almost tubular in structure.

OF THE MOUTH PARTS OF CERTAIN INSECTS. 185

Very interesting is the modification found in the Hmpzde, illustrating the extreme
in the loss of parts; for here the hypopharynx is also wanting, though the salivary
duct remains, opening into the grooved ligula, as shown in PI. III, Fig. 2%. In this
case the hypopharynx is replaced by an extension and peculiar modification of the
labrum. ‘This sclerite is elongated so as to extend to the tip of the labium, and is
very much dilated, somewhat bulb-like at its base. In PI. III, Fig. 2%, labrum and
ligula of Rhamphomyia longicauda are seen from the side, while in Pl. I], Fig. 13, are
shown the same structures in Hmpis spectabilis. The edges of the labrum are turned
under sufficiently to leave a central channel just large enough to receive the ligula,
with which it then forms a closed tube through which the food is taken.

In most of the Muscid flies we find a structure approximating Hristalis with the
labial palpi removed; and the parts may be longer, or shorter, or differently developed,
while adding nothing to what has been already shown; they are, essentially, reduced
piercing structures, no longer functional.

We have, however, in certain other species, where the mouth structures are short,
very poorly developed labial structures. So in Hermetia mucens (P1. II, Fig. 14) the
broad and large mentum bears only a short, scoop-like ligula. The specimen from
which the figure was made was somewhat distorted in mounting and the ligula is
turned just half round. Similar structures occur in the Bibionide, and Huparyphus
bellus (Pl. J, Fig. 12) is not essentially different.

Heretofore the hypopharynx has been referred to mainly in species in which it
was feebly developed and played but a passive part as a covering structure. It is
sometimes a highly specialized sensory structure, though it varies greatly, even when
functional.

A very curious type is found in Bittacus (Pl. ILI, Fig. 4’), where it takes the form
of a simple cylindrical process, set with spines, almost like an odd joint of some slen-
der palpus. In Copris carolina, Pl. I, Fig. 4, showing the epipharynx, may be
accepted as a fair representation of the hypopharynx as well, save that the latter is on
a much reduced scale. The opening of the salivary gland is in a dense mass of spe-
cialized spinous processes.

In the Libellula, among the dragon flies, we have an inflated, somewhat tongue-
like organ (PI. I, Fig. 10”), in which the salivary duet is plainly traceable to its open-
ing among a mass of crossed, specialized spines. The surface is richly supplied with
sensory pittings and tactile hairs. It is a great modification from a structure of this
kind to the simple, ribbon-like form of Bombus, or the flat, slender, chitinous form in
Tabanus ; but the intermediate stages are all present.

To recapitulate concerning the labial structures. The mentum and submentum
Ne TE Se VOL SIDS Oe

186 AN ESSAY ON THE DEVELOPMENT

cover the cesophagus. They may be united so as to form a single organ, and their
tendency is to become internal head structures. The ligula has at its base the opening
into the alimentary canal; it is rarely paired, may be rigid or flexible, and has closely
associated with it the hypopharynx, recognizable by the salivary duct which it shel-
ters. ‘The paraglossze arise on each side of the ligula or glossa, and may be chitinous
or membranous. They are never jointed, never developed for any specific mechanical
purpose, and their tendency is to become obsolete. The labial palpi are essentially
tactile and never become mechanical save as they may form a covering or sheath for
the ligula.

From the most generalized type found in the Glattide the modification is first
from a divided to a single ligula; next to a disappearance or obsolescence of the para-
glossze ; later the labial palpi also disappear, and finally the hypopharynx is also dis-
pensed with. There is no break, and nowhere is there any violent change of structure
or function.

We are now ready to take up the maxilla, which, though composed of a larger
number of sclerites, are usually more easily understood in the ordinary type of man-
dibulate insect. The organ is usually paired and never so completely united as the
labial structures. The two parts are always external to the labium, which it is their
tendency to enfold, and they never have any direct connection with the alimentary
canal. Though the maxillary structures tend to form a covering or sheath for the
labium and its appendages, there is never any intimate connection between them. No
part of the maxilla ever unites with any part of the labium or with any of its appen-
dages. The maxillz are essentially mechanical structures, and their range of variation is
sufficiently great to meet the most diverse possible demands made upon them. A dis-
tinct and fundamental characteristic is the fact that each set of sclerites has its own
peculiar possibilities and limitations, and once these are understood the most highly
specialized type becomes simply explicable.

On PI. III, Fig. 17, is a copy of Prof. Comstock’s figures of Hydrophilus, show-
ing the maxilla from both surfaces, and these may conveniently serve as a text to.
explain the sclerites composing it. At the base is the cardo or hinge, giving attach-
ment to muscles and tendons articulating it to the head. It is to be noted that there
is no firm or chitinous articulation to any head sclerite, and except by muscles or ten-
dons no direct attachment. This we found the case also in the labium in the more
specialized forms, and in the Hymenoptera, for instance, labium and maxille together
are easily dissected out without cutting any but muscular tissue, and without breaking
any chitinous connections or joints. This is in marked contrast with the mandibles
which, when functional, are always firmly articulated by chitinous joints to the external

OF THE MOUTH PARTS OF CERTAIN INSECTS. 187

head sclerites. Supported upon the cardo is the stipes or foot-stalk, deriving its mus-
cular attachments largely from the cardo; but to some extent from the head itself, and
this feature is a variable one. Surmounting the stipes is a palpifer or palpus-bearer,
to which is attached a palpus, varying in the number of its joints. This derives all
its muscles from the stipes in the typically developed maxilla. On the inner side of
the stipes is attached the subgalea, deriving its muscles from the head in large part ;
and this bears a two-jointed galea or hood. It is a matter of some importance to note
that this galea is never more than two-jointed under any circumstances, and that the
tendency is to maintain that number ; though in many instances it is reduced to one
only. It is the most persistent as well as the most variable of the maxillary struc-
tures, and is present when any of them exist at all. Inside of the subgalea, and
attached to it as arule, is the lacinia or blade, which may or may not bear a digitus or
finger. In the figures just cited we find what may be termed a normal or proportionate
development of all the parts, in which no one sclerite is unduly developed or special-
ized. Before attempting to study specializations it is important to note that, when
carefully examined, the sclerites are seen to be arranged in three parallel series. That
is to say three separable parts have grown together laterally, and this union bears with
it the possibility of future disunion or separation for special purposes. We have as the
inner series lacinia and digitus; as the middle, subgalea and galea; and as the outer
the cardo, stipes and palpifer with the attached palpus. Now if we examine some of
the Neuroptera, e. g., Stalis (Pl. II, Fig. 16), we find this lateral arrangement very
strongly marked, and it is easily understood that each of these parallel sets may have
their own peculiar limitations, and that each may be separately and independently
modified.

But lest this seem, after all, a far-fetched conclu<ion, let us examine the maxillee
of Bittacus strigosus (P1. III, Fig. 4”), and we find almost exactly the hypothetical
state of affairs actually existing! Lacinia, galea and palpifer all separated, of nearly
equal length, but of quite different appearance. ‘The appearance of a transverse sec-
tion made at about the middle is shown as Fig. 4”. Fora generalized type this form is
especially valuable, and we may fairly use it as a guide in our discussion of maxillary
possibilities.

There is no absolute rule in the matter, but usually the galea tends to become the
dominant maxillary organ. In many Neuroptera, and especially in their larval stages,
the laciniate structure is best marked, as illustrated in Pl. III, Fig. 9, representing
the maxilla of a Perlid larva Here the galea is reduced to a subordinate rank, and in
many predaceous Coleoptera it is truly palpiform.

In many Orthoptera the development of the galea justifies the name by forming

188 AN ESSAY ON THE DEVELOPMENT

an almost complete hood over the lacinia. This is well illustrated in the maxilla of the
oriental cockroach, Periplaneta orientalis, shown at Pl. ILI, Fig. 8. At this point a
comparison of the figure just cited with the galea of Simulium (PI. I, Fig. 1*) will
prove interesting and instructive.

In the Hymenoptera the galea dominate throughout ; no elongated palpifer is ever
developed, and indeed the maxillary palpi are sometimes almost rudimentary in the
Apide, as shown at Pl. III, Fig. 15.

In Polistes, illustrated at Pl. II, Fig. 18°, we find a common type of the Vespide,
where the lacinia forms a small, blade-like structure, free for almost its entire length,
and the maxille as a whole shelter a large part of the labium. In those cases in which
the “ maxille ” are elongated, the galea is usually the organ affected.

Thus in many Meloids among the Coleoptera we have the mouth parts elongated,
and a study of the maxilla of Memognatha (PI. III, Fig. 20) shows at once the scler-
ites concerned. Here the lacinia is much reduced, and if we remove it altogether we
have the normal Lepidopterous maxilla, which tends to a locking together to form a
complete tube. Recently it has been found that in certain Lepidoptera the lacinia are
actually present, and the figures which I have seen indicate a structure in all essentials
like that of Nemognatha.

While speaking of the Lepidoptera it may be well to cite Pronuba (PI. III, Fig.
21), in which the palpifer is elongated in the female and highly specialized into a sen-
sory and tactile structure, though unjointed. In a well-prepared specimen the point of
origin is perfectly clear, and it is entirely homologous with the structure seen in Bitta-
cus. In the male (Pl. III, Fig. 19) the “tentacle” is not developed, though the
palpifer is enlarged to some extent.

In the Apide, among the Hymenoptera, the lacinia disappear entirely in extreme
cases, or are at least greatly reduced, while as already stated the palpi are sometimes
scarcely visible. The galea, on the other hand, is very prominently developed, and
when at rest envelopes the ligula and paraglossze almost completely. In Pl. III, Fig.
15, is represented the usual appearance of all the parts separated, while at Pl. II, Fig.
15", the transverse section of the mouth structures of Xenoglossa pruinosa shows their
normal relation when at rest. It is seen that the galea actually overlap somewhat at
one margin, and a union along this line would be scarcely considered a violent stretch
of the range of variation. Assume such a union, eliminate the paraglosse which are
organs tending to obsolescence, and then compare with the transection of Hristalis
tenax (Pl. I, Fig. 38"). If the palpifer be eliminated from this latter figure the cuts are
practically identical.

Returning to our figure of Bombus (Pl. II, Fig. 15), we note at the outer edges

OF THE MOUTH PARTS OF CERTAIN INSECTS. 189

of the galea a series of ridges which, under a high power, look extremely suggestive
of the structures found in the labellz of Diptera, especially where, as for instance in
Bombylius, the pseudotrachea are imperfectly developed. These ridges vary much in
the species; but are particularly marked in a little Andrena near vicina, if not that
species itself. Here we see (PI. III, Fig. 3) the entire inner face clothed with a thin
membrane which is crossed by numerous closely set fine chitinous lines! I claim that
this structure is the homologue of the pseudotracheal structure in the Diptera, and that
in the latter order it is in the galea that the development occurs, as it does here in the
Hymenoptera. The relative differences in size are not of importance. As to the
particular use of this structure in Andrena I have no suggestion to make.

In the Proceedings Ent. Soc. Washington, Vol. III, Mr. Ashmead figures on
Pl. III, some very suggestive mouth structures of parasitic Hymenoptera, of which
that of a Pteromalid is reproduced on Pl. III, Fig. 18. The central labium with its
attached structures is much reduced in size, and the maxille, bearing the well-devel-
oped palpi, are reduced to a single structure, the galea, resting upon what may be con-
sidered the stipes. Now if we bring these two parts of the maxille a little more
closely together, we have almost the exact structure seen in 2bio (PI. ILI, Fig. 11°).
The basal ring, bearing the palpi, corresponds almost exactly to the basal ring of
Pteromalus except for size, while except that the surmounting galea are two-jointed,
the correspondence with the upper portion of the structure is equally marked. The
labium in Brbio is much like that figured in Pl. III, Fig. 14, for Hermetia, and in PI.
I, Fig. 12, for Huparyphus.

I am making no very risky statement when I assert that the sclerite to which the
maxillary palpi are attached must of necessity be maxillary; and further, it is equally
safe to say that no maxillary sclerite can bear a labial appendage: and certainly not a
labial palpus. It would be an absurdity, contrary to all the laws of a natural develop-
ment, for a modified labial palpus to become attached to the sclerite bearing also the
maxillary palpus; while if we consider it the two-jointed galea, its position is normal,
requires no assumption of change or character, and does not differ in any essential
points from the gale of the roach (Pl. III, Fig. 8). Yet these two joints in Bzbz0
will, with a ridged membrane thrown over them, represent the labellate tip of the
Muscid proboscis. That such a ridged membrane is well within the range of galear
variability we found in the Andrena near vicina (PI. III, Fig. 3).

The structure in Huparyphus bellus (Pl. I, Fig. 12) resembles Pteromalus yet
more closely, in that a single ring only surmounts the segment bearing the palpus. In
this instance the maxilla is reduced to exactly the same segments seen in the Hymen-
opteron, and logic demands that we recognize them as the same. In this case, how-

190 AN ESSAY ON THE DEVELOPMENT

ever, the lower ring is complete—7. ¢., the two halves of the stipes have become
united. That it must be stipes is shown by the fact that it bears the palpus, and
again the surmounting sclerite must be maxillary also.

There are other species allied to those already cited in which similar structures
occur; but I need for the present call attention to only one more; a species of Olfersia
(Pl. I, Fig. 19). Here the ring is complete in front, but broadly open behind, and
bears the chunky, single-jointed palpus. Surmounting is a single sclerite, very much
resembling in appearance that of Pteromalus, and undoubtedly homologous with it.
Of course Olfersia is parasitic in habit, and the mouth parts are specialized for blood-
sucking ; but the sclerites composing them are nevertheless derived from the same
source as in the “higher ” types.

I have several times referred incidentally to Simulium, and of this the galear
structures are figured (PI. I, Fig. 1"). Dissecting the parts out carefully we find an
almost complete ring at the base, the stipes, to which the palpus and palpifer are
attached. Surmounting this is a pair of sclerites, each almost a half cylinder, repre-
senting the subgalea, and bearing the two-jointed galea. Here again I claim that the
three joints just referred to must be maxillary because they are directly articulated to
the sclerite bearing the maxillary palpi, and the labial structures are all shown at
loiee, 1,

A step in the direction of union we find in the Anglesea gnat or midge—also a
Simulid, to which reference has been already made. Here we see (PI. I, Fig. 2") the
subgalea united most of their length at one side, while the galear joints are yet free.
The basal stipes is not figured because none of my specimens showed it clearly ; but
the palpifer, palpus and lacinia, as they are connected with it, are shown in the
specimen.

In the Aszlidew we find another suggestive structure, studied in the light of the
facts already set out. Here we see, as illustrated Pl. IL, Figs. 1* and 1’, the basal
stipes well developed, united posteriorly, but separated in front. The palpifer and its
attached palpus are situated at the sides, clearly articulated to the stipes, whose char-
acter is thus fixed. Attached to this stipes is a broad, infolded structure, united be-
hind but open in front; maxillary because of its attachment to the stipes, and sub-
galea from its location. It bears in orderly sequence the two-jointed galea of which
the terminal joints are free. The species of the Asiléde are large and easily dissected,
and the figures were drawn from a species of Laphria. The attachments are but
little different in the species, and as the figures illustrate the structure from both
front and rear, the position of the joints should be clear. These figures will be again
referred to in another connection.

OF THE MOUTH PARTS OF CERTAIN INSECTS. 191

Jn all the species heretofore cited the galear joints were more or less distinct and
the pseudotracheal system was little or not at all developed. As the face of the joints
becomes covered by a ridged membrane the texture of the entire structure changes.
It becomes less chitinized, and the chitine is not evenly distributed, causing sutures to
become indistinct and poorly marked. Yet, keeping in mind the general line of yaria-
tion, we can usually reach a correct conclusion.

In a Leptid, species unknown, we find the appearance shown in PI. II, Fig. 1.
Here there is a united basal plate, covered on one surface with a membrane, and from
the chitinous portion arises the palpifer with its attached palpus. Surmounting the
chitinous base are two joints, the galea, the chitinous parts of which only are shown
in outline, the balance of the space being covered by membrane. Here again the
attachment of the maxillary palpus to the basal sclerite determines the maxillary char-
acter of all the sclerites directly articulated to it.

In Hermetia mucens (PI. II, Fig. 17) the entire structure is much more membran-
ous, yet the basal chitinous plate is paired, and while the parts are shown in a dis-
torted position, the two galear joints and their relation to the basal, palpus-bearing
structure is yet perfectly obvious. The other maxillary structures have completely
disappeared, while what is left of the labium is seen at Pl. III, Fig. 14.

The mouth parts in some species of Zvpula are interesting, and a fair illustration
of one of the “snub-nosed”’ species is seen at Pl. I, Fig. 5. Here the origin of the
palpus at the immediate base of the chitinized part of the labella indicates its character,
and if we divest the chitine of the surrounding membrane we get the appearance shown
at Fig. 5" Practically we have a completely paired organ, the relations of which are
perfectly simple when the confusing and unimportant membrane is removed.

The peculiar relation of labrum and labium in the Hmpide has been already
noted, and this makes it easy to separate off all the other parts adhering to the margin
of the head, but not in any way connected with the labium. The relation of the parts
to each other in Hmpis spectabil’s is shown on Pl. I, Fig. 13, while on PI. III, Fig. 2’,
are shown the maxillary structures of Rhamphomyia longicauda. In this latter figure
we note that the parts, except palpifer, are entirely membranous. From the basal
sclerite the palpi arise so as to form only a continuation of the membrane itself with
an extremely slight attachment to the chitinous palpifer ; and to this very same mem-
brane there is articulated by a slightly thickened suture the subgalea, united poste-
riorly, but separated in front; and this bears in turn the indistinctly segmented galea.
This entire structure obviously belongs together and is one organ—necessarily the
maxilla.

A very similar structure is found in Chrysops (Pl. LI, Fig. 14) and in other species

192 AN ESSAY ON THE DEVELOPMENT

of the Tabanide. Now it will be remembered that in this genus I showed the con-
nection of all the labial parts with the mentum, where they normally belong; hence
all the other parts must be, of necessity, maxillary. So we find also in Pl. II, Fig. 14,
that the central labellate structure, two of the piercing structures and the maxillary
palpi all arise from a single united basal sclerite, the stipes.

In Hristalis tenax (Pl. I, Fig. 3) these labellate structures are shown, turned
aside to expose the labial structures. Here also I showed the presence of labial palpi
in close connection with the ligula and hypopharynx, normally attached to the men-
tum, and again it follows that the other structures must be maxillary. Again also
I must call attention to the fact that the palpi are mere continuations of the enveloping
membrane, and that this membrane continues without break to the tip of the labella.
Unless we are to believe that a continuous membrane may give rise to both the maxil-
lary and labial palpi, we cannot possibly consider the labella as labial structures.

I have now traced out what seems to me a continuous development of the modifi-
cations of the subgalea and galea, and have shown, I think, that from Pteromalus in
the Hymenoptera to Hristalis in the Diptera, a continuous chain may be constructed,
requiring nowhere any change of character, function or location. No disassociation
from other maxillary structures and no connection with labial structures.

In taking up the modifications of the palpifer I am confined almost entirely to
the Diptera, in which this sclerite is best developed. In Bettacus I showed its devel-
opment to an elongated structure of no particular type or function and of about the
same texture as the galea. In Pronuba I showed its development into a highly spe-
cialized “ tentacle,” tactile and sensory as well as mechanical in character. In the
Diptera it is quite usually present as an elongated, rigid, chitinous organ adapted for
piercing. It occurs in all the piercing types and is present as a rudiment in many
others. It undergoes a curious and interesting change in function as the Dipterous
mouth changes from the piercing to the scraping or lapping type, and as it becomes
flexed.

The simplest form occurs in those piercing Diptera in which the proboscis is not
flexed. Thus in the Buffalo gnat (PI. II, Fig. 9) it is a stout, semicylindrical piercing
organ, enlarged both at base and at tip, at which latter pomt it is also toothed. The
connection of the palpus with the subgalea was already shown on PI. I, Fig. 1%, and
this shows how the chitinous palpifer forms part of the combination. The palpifer
arises, normally, outside of the galea; yet at the tip it is found in connection with all
the other piercing structures inside of that organ. How it gets there is illustrated in
the Anglesea Simuliid (Pl. I, Fig. 2"), where all the maxillary parts are shown in
proper connection, and it is seen that the palpifer enters the galear envelope in the

OF THE MOUTH PARTS OF CERTAIN INSECTS. 193

incomplete articulation between galea and subgalea. By separating off the galear
structures, the relation of palpifer and lacinia in Simuliwm is illustrated (on PI. I,
Fig. 1°), and the convergence of the two at tip is not distortion, though perhaps a
little exaggerated by pressure. The result of this change of position is that a section
made near the base of the proboscis would show as illustrated on Pl. I, Fig. 2’, while
one made nearer the tip would show as in Fig. 1°. Incidentally it will prove interest-
ing to compare these sections with that of Bittacus strigosus (Pl. III, Fig. 4"), leaving
out of consideration the abnormal labium of the latter. The resemblance is perfect,
and the resemblance expresses fully the actual condition of the matter. A very simi-
lar state of affairs exists in the Astlidw (Pl. IU, Fig. 1°). Here the palpifer is the
only maxillary piercing organ, and the figure itself shows clearly how easily it would
swing inside the ample space left in the subgalea for its entrance. The curvature of
the organ is such, also, that when in place it meets the central ligula so as to form a
solid puncturing organ.

So in Chrysops (Pl. II, Fig. 14) the structure is seen to be similar to that in
Simulium ; but here, as almost everywhere else in the order, it is cylindrical or nearly
so, in marked contrast with the lacinia, which is always flattened.

As we get into types that have lost the piercing habit, the function of the palpifer
fails or changes. If the species have a short, nonflexed proboscis, it simply dwindles
from disuse. So in Stratiomyia and in Leptis (Pl. II, Figs. 1 and 2) it simply forms
a little chitinous appendage to the palpus—a mere remnant without function. If, on
the other hand, the species are able to flex the proboscis, another change takes place.
There is needed then some lever to which muscles for flexing can be attached, and no
structure seems to have been so easily adaptable as the palpifer. So we find in the
Eimpide, where only slight flexion is required, only a small basal extension, shown at
Pl. I, Figs. 4 and 3, for Hmpis spectabilis and Hulonchus tristis, and at Pl. IIT, Fig.
2’, for Rhamphomyta longicauda.

In the Bombyliide is a step forward. The insects are not predaceous, have the
habit of hovering over flowers and using the proboscis in feeding in that position.
This requires a much better control, and as a result the basal extension is much better
developed, as shown in Pl. II, Figs. 6 and 7, illustrating Bombylius and Anthraa.

As we get into types like Hristalis and other Syrphide, the basal extension be-
comes the most prominent and the piercing portion diminishes in size (Pl. H, Fig. 5),
and keeping step with this modification is a gradual separation of the palpus itself
from the palpifer. This is well illustrated both in Hristalis and Spherophoria, and
this tendency continues until in Duerllia (Pl. II, Fig. 10) the separation is complete,
though the piercing portion of the palpifer is yet distinguishable. In Callephora even

A. P. S—VOL, XIX. Y.

194 AN ESSAY ON THE DEVELOPMENT

this disappears and the chitinous rod is entirely disassociated from the palpus. Finally
in Stomoxys calcitrans (PI). Il, Fig. 12) there remains nothing to indicate the existence
of any relation between the slender chitinous rod and the distant maxillary palpus. It
is not in the least strange that guesses as to the character of this structure in Musca
domestica should have been so often wide of the mark; though with a proper series as
now shown, its origin is clear.

There remains to be accounted for the lacinia, and this in the Diptera is the flat,
blade-like structure generally identified as the mandible. It has been shown that
while the lacinia is often the dominant organ in many mandibulate insects, the tendency
is, on the whole, to a decrease in size, ending in the Hymenoptera in its entire elimina-
tion. In the Diptera it is present in the blood-sucking species only, and it may be
identified by its position and its relation to the other maxillary structures. It has
been several times referred to incidentally, and in the Anglesea Simuliid (Pl. I, Fig.
2") its relation to the other maxillary parts is shown. In PI. J, Fig. 1%, is illustrated
the connection between the palpifer and lacinia in the Simuliwm sent me by Mr.
Aldrich. This connection is not fanciful but actual, and no sclerite so intimately con-
nected with an admitted maxillate structure can be anything but maxillary.

Again in Chrysops (Pl. II, Fig. 14) I have illustrated the fact that all the struc-
tures which I consider maxillary have acommon origin. At Fig. 14* I show the lacinia
alone, and it is to be noted that at the base it is modified for attachment with reference
to the palpus. Now unless this is a maxillary sclerite, why should it be modified to
accommodate the maxillary palpus? Does it not seem rather absurd to believe that
this can be a mandible brought to originate from one point with the palpifer and modi-
fied to allow it to envelope at base the maxillary palpus ?

One of the most serious difficulties in the way of the proper understanding of the
mouth parts of haustellate insects has been the desire to provide for the mandibles on
the theory that they are among the permanent structures. Yet I cannot understand
why this should necessarily be the case. When functional, mandibles are essentially
chewing or biting organs, and when the insects do not require such structures, it seems
to me most natural that they should become obsolete: and that is exactly what has
occurred according to my reading of the facts. Their functional character never
changes; they simply dwindle from disuse and gradually disappear. So we find them
in the Lepidoptera as mere rudiments, connected with a highly specialized maxilla ;
and in the Rhynchophora they are sometimes mere remnants, occasionally reversed in
position—exactly as I pointed them out in Simulium. I think that in view of all the
evidence presented by me, none of the piercing organs of the Diptera can be consid-
ered mandibles, and I cannot even yet, after carefully weighing all that Dr. Packard

OF THE MOUTH PARTS OF CERTAIN INSECTS. 195

has written, see any reason why the rudimentary structures at the tip of the labral
extension in Semuliwm are not mandibles.

If we refer back again for an instant to the Panorpids we note (Pl. III, Fig. 4”)
that in Bettacus strigosus the origin of the mandibles form an extension of a lateral
head sclerite, with the labrum-epipharynx between them. In Panorpa the mouth
structures are much shorter, set on an immensely elongated stipes, and at the tip of
the frontal extension of the head we again have the mandibles, much reduced, with a
small, lappet-like labrum-epipharynx between them. Now the situation of the rudi-
ments in Simulium corresponds almost exactly with that of the undoubted mandibles
in Panorpa rufescens (Pl. III, Fig. 4°); but in the Hmpzide we find a yet more closely
allied structure. I have already called attention to the peculiar elongation of the front
of the head in this family, and now if we examine this at tip, in Hmpis spectabilis
(Pl. I, Fig. 13") its very close resemblance to Panorpa is at once evident. We find
a central lappet-like structure with a sensitive surface, which looks like and logically
should be the epipharynx, and moving below it is a pair of appendages which, in my
opinion, represent mandibles. They are membranous and probably not functional; but
this is no argument against their character. I believe that the similarity in the appear-
ance between PI. III, Fig. 4°, and Pl. II, Fig. 13%, is the expression of a true homol-
ogy, and that mandibles in the Diptera exist in no other form or situation. It is likely
that other species, showing them much more perfectly, will yet be discovered ; but so
indeed do I believe that labial palpi, properly connected with the mentum, will yet be
found, so distinct in character that, even if not functional, their homology cannot be
mistaken.

Labrum and epipharynx have been frequently referred to in the course of this
paper, and in the introduction the general relation of these two parts has been ex-
plained. Both structures occur in many families of the Diptera. As in the case of
the hypopharynx, the epipharynx has always connected with it a salivary duct. In its
intimate connection with the labrum it is shown on PI. I, Fig. 10°, illustrating the
epipharynx of Zzbellula. Here the chitinous tube giving passage to the duct is fully
shown. As an example of a highly developed structure, the epipharynx of Copris
carolina is shown (Pl. I, Fig. 4), and here the salivary duct opens among the dense
central mass of spinous processes. The epipharynx of Polistes was referred to in the
description of the labium, as was that of Andrena in the connection. In the Hemip-
tera the labrum and epipharynx are usually well developed and the salivary duct is in
many cases very well marked.

Among the Diptera some of the larger Syrphide have the labrum quite distinct,
and on the under surface is a sensitive surface into which an obvious duct, with chit-

196 AN ESSAY ON THE DEVELOPMENT

inous protecting margins, is led, as shown on PI. III, Fig. 10. A much better devel-
oped organ, strongly resembling that in some of the Hemiptera, we find in the Asilidw
(Pl. IH, Fig. 1”), and here also the salivary duct is obvious. The structure in Simu-
uum has been already referred to, as has that in the Hmpidw.

To recapitulate concerning the maxillee: The sclerites form three series, each of
which has its own possibilities of development. The lacinia never develope into any-
thing other than a chewing or piercing organ and always arises inside of the galea.
The galea varies in the direction of forming an enveloping organ for all the other
mouth parts, and the subgalea eventually unites along one margin for that purpose.
There is a tendency to develop a ridged membrane on the inner surface of the galear
joints which culminates in the pseudotrachea of the muscid labella. The palpifer has
a small range of development, from an unjointed, flexible, tactile organ, to a rigid,
piercing structure; and as this becomes useless, to a process for the attachment of
muscles used to flex the proboscis.

It remains only to acknowledge the assistance received from my entomological
friends. Dr. 5. W. Williston has from time to time sent me such specimens as I
thought might help me; Mr. C. W. Johnson has given me numerous species of fami-
lies selected because of apparent differences in the mouth structure; and to Mr. J. M.
Aldrich I owe many other species in some numbers, among them the Simuliid already
referred to. Mr. EH. P. Fell kindly sent me specimens of Panorpa and Bittacus, which
enabled me to make a much more complete study of these insects than would have
been otherwise possible. To all these gentlemen, as well as to the others who have in
any wise aided me, I desire to express my thanks.

Concerning the figures—most of them are camera lucida drawings. A few are
drawn from micro-photographs, assisted by the specimens themselves. The figures
of transections are largely made from actual preparations; some are redrawn from
other sources, while a few are ideal.

OF THE MOUTH PARTS OF CERTAIN INSECTS. 197

EXPLANATION OF THE PLATES.

The lettering of the parts, the same throughout, and the abbreviations, are as follows: ZLbr, labrum; epi,

epipharynx (the two sometimes combined as lbr-ept) ; md, mandible; car, cardo; st, stipes; pfr, palpifer; mp,

maxillary palpus; gal, galea; sg, subgalea; lac, lacinia; dig, digitus; sm, submentum; m, mentum; gl, ligula or

glossa ; par, paraglossa ; Ip, labial palpi; Ayp, hypopharynx.

ray

cw)

SOW ea Pw pp

Plate I.

Buffalo gnat. 1¢, galear structures with palpi attached ; 19, labial structures ; 1¢, lacinia and palpifer of

Simulium from Aldrich ; 14, labrum and labium of Simulium from Aldrich ; 1¢, transverse section through

middle of mouth of Buffalo gnat.

Simulium from Anglesea, N. J.

2, the maxillary structures in their actual relation to each other ; 2%,

transverse section of mouth parts toward the base of subgalea.

Mouth parts of Hristalis tenaz. 34, transverse section of same at the middle of subgalea.

Copris carolina, epipharynx.

Mouth structures of Tipula sp.; 5¢, the chitinous parts of the same.

Copris carolina ; labial structures dissected out and seen from side.

Copris carolina ; chitinous part of under side of head.

Copris carolina ; mandible with the sclerites named and homologized.

Andrena vicina ; labial structures, with part of epipharynx attached.

Libellula sp. a, the epipharynx ; b, the hypopharynx.

Stomoxys calcitrans ; transverse section through the middle of the ligula.
Mouth parts of Huparyphus bellus.

Maxillary structure of Leptis sp.
Palpifer of Stratiomyia.
Palpifer of Hulonchus tristis.
Palpifer of Hmpis spectabilis.

Plate I.

Palpifer of Spharophoria cylindrica.

Palpifer of Bombylius.
Palpifer of Anthrax.

Palpifer of Chrysops vittatus.
Palpifer of Simulium.
Palpifer of Lucillia.

Palpifer of Calléphora.
Palpifer of Stomozys.

Figs. 10 to 12 inclusive were accidentally reversed in making up the plate.

Mouth parts of Hmpis spectabilis.

134, elongated head structure at tip, showing mandibles and epipharynx ;

13, transverse section at middle of subgalea.
Mouth parts of Chrysops vittatus showing maxillary structures attached together. 144, the lacinia ; 14°, pal-

pifer and palpus ; 14¢, transverse section at middle of galea.

Labial structures of Xenoglossa pruinosa. 4, transverse section at about middle.

Labial structures of Periplaneta orientalis.

Maxillary structures of Hermetia mucens.
Mouth structures of Polistes metricus. 184, ligula, paraglossa and mouth opening ; 18°, labium as a whole,

with epipharynx attached ; 18¢, maxilla.
Maxilla of Olfersia. 194, seen from front; 19, seen from behind or below.

198 AN ESSAY ON THE DEVELOPMENT OF THE MOUTH PARTS OF CERTAIN INSECTS.

Fig.

Fig,

GU Gp Gl

Plate I.

Mouth structures of Asilide—Laphria sp. a, maxilla from front ; b, same from behind ; ¢, labium ; d, lab-
rum ; é, transverse section of mouth at junction of galea and subgalea.

Mouth structures of Ramphomyia longicauda. a, the labium ; 6, maxilla; c, extension of front of head ;
d, relation of this extension to the labium.

Galea of an Andrena allied to vicina.

Mouth parts of Bittacus strigosus. a, mandibles and labrum; 2, maxilla and labium; c, mandibles and
labrum—epipharynx of Panorpa rufescens. ;

Labial structures of Hristalis tenaz. 54, transverse section at about middle ; 5), same at about tip.

Labial structure of Bombus sp. 62, transection at about middle ; 6b, same made near tip.

Labium of Harpalus calignosus.

Maxilla of Pertplaneta orientalis.

Maxilla of Perlid larva.

Epipharynx of Hristalis tenaz.

Mouth parts of Bibiosp. a, maxilla from behind ; 6, same in front ; c, transection made near the base.

Labium of Bombus fervidus ; the transections are lined to the portions referred to.

Labium of Chrysops vittatus ; the transections are lined to the parts referred to.

Labium of Hermetia mucens.

Maxille and labium of Bombus, showing the relation of the parts to each other.

Maxilla of Stalis.

Maxilla of Hydrophilus from upper and Jower surface, redrawn from Comstock.

Maxilla and labium of Pteromalus, redrawn from Ashmead.

Maxilla of Pronuba, male.

Maxilla of Wemognatha.

Maxilla of Pronuba, female.

Mouth parts of Locusta from Kolbe. 7, labrum ; 7, mandibles ; 772, maxille ; ¢, labium,

NOTICE.

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Be TRANSACTIONS

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-_ VOLUME XIX.—NEW SERIES.

PART II.

John Van Denburgh, Ph.D.

as ARQICLE IV .— Results of Recent Researches on the Hvolution of the Stellar Systems.
A.M., Ph.D. (Berlin).

“Apnict V.—9n the Glossophayine. By Harrison Allen, M.D.
“Articte VI.—The Skull and Teeth of Hetophylla aiba. By Harrison Allen, M.D,

Hhiladelphia:
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bas et 5
A mene, Iil.—Some Hexperiments with the Saliva of the Gila Monster (Heloderma suspectum). By

By T. J. J. See,

JUN 16 1898

ARTICLE III.

SOME EXPERIMENTS WITH THE SALIVA OF THE GILA MONSTER
(IELODERMA SUSPECTUM).

BY JOHN VAN DENBURGH, Pu.D.,
CURATOR DEPARTMENT OF HERPETOLOGY, CALIFORNIA ACADEMY OF SCIENCES.

Read before the American Philosophical Society, September 38, 1897.

FE ENTRODUCION:

When, in 1651, Franciscus Hernandez published his Historie animalium et minera-
lum Nove Hispanie he gave to Europe the first account of a curious reptile native to
those far-western lands which the Spaniards had won beyond the sea. This was a large
lizard, said to grow three feet long, thick-set, heavy-jawed, protected by an armor of
wart-like bony plates, gaudily colored in orange and black—withal so repulsive that
Wiegmann, nearly two hundred years later, christened it Heloderma horridum.

For many years, this name was applied to these lizards wherever found, but in 1869
Prof. Cope discovered that those which had been caught within the borders of the United
States and Sonora differ in many details from their more southern relatives. He named
the smaller, northern species Heloderma suspectum. It is this species which, because
of its former abundance near the Gila river, in Arizona, has become popularly known
under the name Gila Monster.

The Indians and Mexicans claimed for these lizards power to inflict a bite even
more deadly than that of the rattlesnake, but, since they claimed like powers for other
reptiles known to be quite innocent of venom, their evidence was of little value. It
received some confirmation, however, when the herpetologists of Europe found that the
teeth of the Heloderma bear grooves similar to those which in some poisonous snakes
serve to introduce venom into the wound. Since this was discovered the question of the
poisonous nature of the bite of the Gila Monster has attracted considerable attention and
many opinions have been published.

A. P. S.—VOL. XIX. Z.

200 SOME EXPERIMENTS WITH THE

In 1857, Dr. J. E. Gray, of the British Museum, wrote :

“« This lizard is said to be noxious, but the fact has not been distinctly proved.’’

Seven years after this there appeared a popular account of the habits of the Mexican
species (HZ. horridum), in which M. Sumichrast, after dwelling at some length upon the
general habits of the animal, wrote:

“‘ In support of this pretended malignity, I have been told of a great number of cases in which ill effects
were produced by the bite of the animal, or by eating its flesh in mistake for that of the Iguana. I
wished to make some conclusive experiments on this point; but, unfortunately, all the specimens which I
could procure during my stay in the countries inhabited by it were so much injured that it was impossible to
do so. Without giving the least credit to the statements of the natives, I am not absolutely disinclined to
believe that the viscous saliva which flows from the mouth of the animal in moments of excitement may
be endowed with such acridity that, when introduced into the system, it might occasion inconveniences, the

gravity of which, no doubt, has been exaggerated.”’
Prof. Cope, in 1869, stated :

‘« That though the lizards of this genus could not be proven to inflict a poisonous bite, yet that the sali-
vary glands of the lower jaw were emptied by an efferent duct which issued at the basis of each tooth, and
in such a way that the saliva would be conveyed into the wound by the deep groove of the crown.”’

Six years later Dr. Yarrow said:
“« Tt is believed to be very poisonous, but such is not the case; for, although it will bite fiercely when
irritated, the wound is neither painful nor dangerous. . . . . The Pueblo Indians of this place said they

were quite common, and were regarded by the Mexicans as poisonous; the poison being communicated by
the breath as well as by the teeth. This has no foundation in fact.’’

The same year, M. Bocourt published some notes which he had received from M.
Sumichrast, who, having finally been able to make a few experiments, concludes :

““ Quoique ces expériences soient insuffisantes pour prouver que Ja morsure de |’ Héloderme est véritable-
p pour p q

—)

ment venimeuse, elles me paraissent assez concluantes pour faire admetire qu’elle ne laisse pas de causer

de trés-rapides et profonds désordres dans |’économie des animaux qui en sont |’objet. . . . .
““ Je ne doute pas que des expériences, faites avec des individus adultes et nouvellement pris, ne pro-
duisent des effets beaucoup plus terribles que ceux qu’oat pu occasionner Ja morsure d’un individu jeune et

affaibli par une captivité de prés de trois semaines.’’

In 1882, several opinions were published on each side of the question. A Helo-
derma, which had been received at the Zodlogical Gardens in London, bit some small
animals, and because these died several English writers—as Giinther, Boulenger, and
Fayrer—concluded that the Monster was poisonous, while some American authors haye
thought that death in these cases might have resulted from the mechanical injuries
received. The American Naturalist noted that “ Dr. Irwin, U. 8. A., experimented with
the H. suspectum in Arizona, fifteen years ago, and concluded that it was harmless.”

SALIVA OF THE GILA MONSTER. 201

Dr. R. W. Shufeldt had a personal encounter with an active Gila Monster, of which he
wrote :

“On the 18th inst., in the company of Prof. Gill of the [Smithsonian] Institution, I examined for the
first time Dr. Burr’s specimen, then in a cage in the herpetological room. It was in capital health, and at
fust I handled it with great care, holding it in my left hand examining special parts with my right. At
the close of this examination I was about to return the fellow to his temporary quarters, when my left
hand slipped slightly, and the now highly indignant and irritated Heloderma made a dart forward and
seized my right thumb in his mouth, inflicting a severe lacerated wound, sinking the teeth in his upper
maxilla to the very bone. He loosed his hold immediately and I replaced him in his cage, with far greater
haste, perhaps, than I removed him from it.

“« By suction with my mouth, I drew not a little blood from the wound, but the bleeding soon ceased
entirely, to be followed in a few moments by very severe shooting pains up my arm and down the corre-
sponding side. The severity of these pains was so unexpected that, added to the nervous shock already
experienced, no doubt, and a rapid swelling of the parts that now set in, caused me to become so faint as
to fall, and Dr. Gill’s study was reached with no little difficulty. The action of the skin was greatly
increased and the perspiration flowed profusely. A small quantity of whiskey was administered. This is
about a fair statement of the immediate symptoms; the same night the pain allowed of no rest, although the
hand was kept in ice and laudanum, but the swelling was confined to this member alone, not passing
beyond the wrist. Next morning this was considerably reduced, and further reduction was assisted by the
use of a lead-water wash.

“Tn a few days the wound healed kindly, and in all probability will leave no scar; all other symp-
toms subsided without treatment, beyond the wearing for about forty-eight hours so much of a kid glove as
covered the parts involved.

. ‘' Taking everything into consideration, we must believe the bite of Heloderma suspectum to
be a harmless one beyond the- ordinary symptoms that usually follow the bite of any irritated animal. I
have seen, as perhaps all surgeons have, the most serious consequences follow the bite inflicted by an
angry man, and several years ago the writer had his hand confined in a sling for many weeks from such a
wound administered by the teeth of a common cat, the even tenor of whose life had been suddenly
interrupted.”’

Only a few months had passed after the publication of Dr. Shufeldt’s article when
there appeared an account of the first carefully conducted series of experiments with the
saliva of the Heloderma. This was by Drs. 8. Weir Mitchell and Edward T. Reichert,
who conclude that :

“¢ The poison of Heloderma causes no local injury.

<< That it arrests the heart in diastole, and that the organ afterwards contracts slowly—possibly in
rapid rigor mortis.

«« That the cardiac muscle loses its irritability to stimuli at the time it ceases to beat.

‘That the other muscles and the nerves respond readily to irritants.

«¢ That the spinal cord has its power annihilated abruptly, and refuses to respond to the most powerful

electrical currents.

202 SOME EXPERIMENTS WITH THE

“This interesting and virulent heart poison contrasts strongly with the venoms of serpents, since
they give rise to local hemorrhages, and cause death chiefly through failure of the respiration, and not

by the heart, unless given in overwhelming doses.’’

For a time, it seemed that the experiments of Mitchell and Reichert had answered
the question of the poisonous power of the Heloderma once and for all. But five years
later, Dr. Yarrow, then Honorary Curator of the Department of Reptiles in the United
States National Museum, performed some equally careful experiments upon rabbits and
chickens. These, he says,

*“ Would seem to show that a large amount: of the Heloderma saliva can be inserted into the tissues
without producing any harm, and if, is still a mystery to the writer how Drs. Mitchell and Reichert and
himself obtained entirely different results. Were it not for the well-known accuracy and carefulness of
Dr. Mitchell, it might be supposed possibly that the hypodermic syringe used in his experiments contained

a certain amount of Crotalus, or cobra venom, but under the circumstances such a hypothesis is entirely
untenable.”’

Notwithstanding Yarrow’s results, Dr. Mitchell still held his original opinion in
1889.

The following year, Prof. Samuel Garman, of the Museum of Comparative Zoélogy
of Haryard University, published an account of experiments in which he caused an
active Gila Monster to bite the shayed legs of kittens without serious effect. He con-
cludes that

“The results of the experiments suggest danger for small animals, but little or none for larger ones.
Large angle worms and insects seemed to die much more quickly when bitten than when cut to pieces with
the scissors.”’

Thus while in England the Heloderma was unanimously held to be venomous, Dr.
Shufeldt, in 1891, summarized American opinion as follows:

“« Here in America the evidence would seem to be rapidly leading to the demonstration of the now
entertained theory that the saliva of this heretofore much-dreaded reptile is possibly entirely innocuous.”’

“« Thus the matter seems to stand at the present time—perhaps the vast majority of physicians who
followed Drs. Mitchell and Reichert in their experiments fully believe to-day that the bite of a ‘ Gila
Monster’ will very often prove fatal even in the case of man; while, on the other hand, naturalists
almost universally believe that the saliva of this saurian is hardly at all venomous, and then only under

certain conditions. ’’

Il. THE MOUTH FLUIDS.

In the winter of 1896-97 I began a series of experiments with the saliva of the Gila
Monster, the results of which are given in the subsequent pages. My object was to
answer the following questions :

(a) Is the bite of the Gila Monster poisonous ?

SALIVA OF THE GILA MONSTER. 203

(b) If poison is present what are its physiological effects ?

My Heloderma was the sole survivor of eight or ten brought from Arizona in
1892 and, although seemingly fat and healthy, was not very active. It was of moderate
size, being about eighteen inches long. The amount of saliva obtainable from it was so
small that it could be gathered satisfactorily only by causing the reptile to bite absorbent
paper wrapped around a piece of soft rubber and afterwards dissolving out the saliya in
water. For this purpose filter paper was used.

It would not do to let the Monster bite the pigeons, because if this were done and
the pigeons died the skeptics might justly claim that death was due to the mechanical
injury inflicted by the powerful jaws, with their long, curved fangs, rather than to any
poison haying been inserted. yen when the Heloderma’s saliva solution was injected
hypodermically and death could not have been occasioned by the severity of a wound
there might be some doubt as to the effect of a quantity of water suddenly placed under
the skin, or it might be claimed that some substance was present in the water or the paper
used quite poisonous enough to cause a pigeon’s death irrespective of any venom from the
Monster. So samples of all the materials used had to be subjected to careful tests to
show that they were harmless.*

Mucus.

A greater or less quantity of thick mucus is present in the back part of the mouth of

the Gila Monster. Some of this often adheres to the filter paper in stringy masses. It

is entirely without poisonous properties and need not be mentioned again.

THe Porsonous SALrva.

The water solution of saliva when extracted from the paper is a slightly yellow-
ish or opalescent liquid, often more or less stained with blood owing to injury to the
gums. It is faintly alkaline, and ordinarily possesses a pungent and highly characteristic
though not unpleasant odor. This odor becomes less and less noticeable when the Monster
is caused to bite every day, but its strength seems to be no indication of the lethal power
of the saliva. That the solution of saliva thus obtained contains a very powerful poison
is shown in the following experiments :

EXPERIMENT I.—Noy. 11, 1896. The Heloderma was caused to bite on paper three times. The

*In order to test my materials, and some other things as well, the following preliminary experiments were
performed, the first repeatedly :

ExPERIMENT,—A sample of filter paper was soaked in water, which was then injected subcutaneously in
front of the wing of a pigeon. During two hours there was no effect, and the next day the bird was still well.

EXPERIMENT,—Mixed human saliva with an equal quantity of water and injected about twenty minims in
wing of pigeon at 12.01 P.M. No effect. Next day well.

EXPERIMENT.—Mixed blood of horned toad (Phrynosoma frontale Van D,) with water and injected wing of
pigeon. No effect.

204 SOME EXPERIMENTS WITH THE

water solution—about twelve mimims—was then injected subcutaneously in front of the shoulder of a pigeon
at 3.18 P.M. In three minutes the pigeon was no longer able to stand, and fell over on its side with eyes
closed. At the end of the tenth minute the bird was unable to hold up its head when raised by its wings.
During the eleventh minute respiration was in gasps, and at the end of the eleventh minute the pigeon

was dead. [No local effects; heart beating regularly. |

ExPERIMENT II.—Nov. 12, 1896. Monster was caused to bite seven times during about as many
minutes. Saliva then dissolved in about seventy minims of water, of which ten minims were injected
under the skin in front of right shoulder of pigeon, at 11.24 A.M.

11.28. Pigeon barely able to walk.

11.29. Not able to walk.

11.30. Cannot stand; lies on side; eyes closed.

11.31. Head nods; respiration is forced.

11.32. Muscular straining; head drawn back between shoulders.

11.33-38. Respiration greatly forced; bill opens and shuts with each breath.

11.89. Violent contractions of caudal muscles.

11.40. Violent contractions of head and wings.

11.40}. Head falls forward onto table.

11.404. Death.

No local effects; ventricles empty, auricles full of clots; blood almost black.

Tf these experiments leaye any room to doubt that the bite of the Gila Monster is
poisonous it is entirely removed by the results of a large number of experiments which I
afterwards performed and in which death followed the injection of Heloderma saliva quite
as certainly and almost as quickly as when rattlesnake yenom is used.

It now became of interest to learn whether this powerful poison is affected by boiling
or decay, or the presence of alcohol, ete.

The Kyfect of Boiling.—Two experiments were performed which show that the
poisonous properties of the saliva are not injured by boiling. The solution becomes
opalescent and, if boiling be prolonged, loses its odor or gives off one similar to that of
boiled barley. .

EXPERIMENT IIT.—Noy. 12, 1896. The Heloderma was caused to bite seven times during about as
many minutes. Saliva then dissolved in about seventy minims of water. Ten minims of this solution,
having been boiled a few seconds, were injected under the skin of the right shoulder of a pigeon, at
2.21 P.M. The temperature of the pigeon before injection was 104° F.

2.22. Sits down, but is able to stand when frightened.

2.26. Sits down.

2.27. Sits down immediately after being caused to stand, seems dizzy.

2.29. Lies on side; temperature 100°.

2.54. Cannot stand; temperature 98°.

2.36. Violent respiration; temperature 96°.

SALIVA OF THE GILA MONSTER. 205

2.38. Violent respiration; temperature 98°.
2.39. Violent respiration; temperature 100°.
2.424. Violent respiration; temperature 1015°.
2.45. Violent respiration; temperature 100°.
2.48. Respirations about 108 per minute; temperature 99°.
2.50. Temperature 100°.
2.53. Respiration more labored; temperature 99°.
2.54. Temperature 98°.
2.55. ‘Yemperature 97°.
2.56-57. Temperature 93°; respiration short and forced, 39 per minute.
2.58. Wheezing; vomits.
2.584. No motion except quivering of wings; temperature 90°.
2.59. Wings and tail flapped twice.
3.00. Dead.
No local effect; small clot of blood in base of right Jung; ventricles full of black clots; auricles
beating; arteries empty; veins dilated with blood.

This experiment would seem to show that the action of the poison is slightly delayed
by boiling. Experiment IV shows that such is not the case.

Experiment [V.—Noy. 14, 1896. Ten minims of the solution used in experiments II and III
were boiled about five minutes on Noy. 12, and again Nov. 13 and 14, and then were injected under the
skin of a pigeon’s wing at 3.30 P.M.

3.34. Respirations 32 per minute.

3.37. Staggers about with peculiar circular motion.

3.3940. Respirations 48, becoming constantly more forced, so that at end of minute tail moves up
and down.

3.42. Cannot stand.

3.44-45. Respirations 49.

3.46. Falls on side.

3.47. Head nods; pupil seems slightly dilated.

3.52. Respirations 47, irregular.

3.53. Bill begins to open and shut.

3.54. Convulsive action of wings and head, heed drawn under to breast.

3.55. Death.

The Effect of Decay.—When a solution of saliva is allowed to stand for a few days
it soon begins to decay, and this process continues until a strong odor of putrescence is
given off and a muddy sediment appears at the bottom of the liquid. After this had
occurred, very large doses of the solution were injected into pigeons without producing the
slightest ill-effect. Decay, then, appears to destroy the lethal power of the saliva, but
my experiments are not absolutely conclusive because the solution was not tested while

fresh.

206 SOME EXPERIMENTS WITH THE

EXPERIMENT V.—Saliva of several bites was collected, November 14, and dissolved in about ten
minims of water per bite. November 16 there was a marked odor of decay. November 23 the odor of
putrescence was very strong and the liquid appeared muddy with a slight sediment. At 2.31 P.M., ten
minims were injected under the skin in axilla of pigeon whose temperature at 2.29 (when frightened )
was 106°.

2.35-36. Respirations 35.

2.40. Temperature 105°.

2.44-45, Respirations 32.

3.09. Temperature 104°.

3.10-11. Respirations 32.

3.28-29. Respirations 32.

3.31. Temperature 104°. Repeated injection.

3.33-34, Respirations 34.

3.55—06. Respirations 32.

4,21—22. Respirations 33.

November 24, ete. Still perfectly well.

EXPERIMENT VI.—December 1, 1896. Injected forty minims of solution used in experiment V
under skin of legs and wing of pigeon at 12.45 P.M.

4.30. Still no effect.

December 2. Well.

The Effect of Drying.—That drying does not affect the power of the venom was
shown by the following experiment, although the dose was too small to cause death.

EXPERIMENT VII.—December 1, 1896. A small quantity of the solution used in experiments II,
III and IV, having been dried, was redissolved in water and injected subcutaneously in a pigeon at
3.40 P.M.

4,10. Respiration slightly forced.

4.30. Cannot walk well.

4.45. Very ‘‘ tame;’’ respiration forced.

December 2. Pigeon recovered.

The Effect of Alcohol—When alcohol is added toa water solution of saliva, the solu-
tion becomes opalescent, as when boiled. This change in color is probably due to the
formation of a finely divided albuminous coagulate. It is not removed by filtration
through paper. Alcohol does not influence the action of the venom.

EXPERIMENT VIII.—About twenty minims of the solution used in experiments IJ, III, 1V and VII
was mixed with an equal quantity of ninety-five per cent. alcohol, November 14. About half of this had
evaporated when ten minims of the remainder were mixed with ten of water and thrown down the throat
of a pigeon at 11.25 A.M., November 18. ;

11.46. Seems well.

2.15 P.M. No effect.

SALIVA OF THE GILA MONSTER. 207

2.26. Injected the other ten minims in left axilla.
2.29. Shows uneasiness of left wing and cannot always contro] it.
2.294. Sits; cannot walk.

2.30. Pupils contracted; cannot stand.

2.31. Lies on side; respiration convulsive.

2.32. Respiration still more labored.

2.33. Seems unable to feel pinching of legs.
2.37. Rate of breathing very greatly increased.
2.38-39. Respirations 62.

2.40-41. Respirations 84.

2.43-44, Respirations 64.

2.45-46. Respirations 53.

2.46-47. No respiration; convulsions.

2.48. Death.

Auricles beating; ventricles stil]; blood black, clotted; auricles and veins full; ventricles and arteries

empty; slight extravasation in coat of small intestine near head of pancreas; no local effect.

Ninety-five per cent. alcohol when added to undiluted saliva does not injure its
poisonous properties, nor does the alcohol act. as a solvent of the venom, although its

solubility in water is unaffected.

ExprriMenr 1X.—November 23, 1896.

a. Filter paper containing saliva was washed in about one ounce of alcohol for about twenty hours.
The alcohol was then poured into an open dish. As soon as evaporation began a thin white scum
appeared on the surface of the alcohol, but did not increase much as evaporation proceeded to dryness. This
scum was not soluble in water, even after the addition of salt (NaCl). Placed under the skin of a pigeon,
it produced no effect.

6. The alcohol-washed paper was soaked during a few minutes in sixty minims of water. Twenty
minims of this water were injected under the skin of each wing of a pigeon at 3.25 P.M., November 24.
Half an hour later twenty minims were injected into the left leg.

4.07. Pigeon sits down.

4.12-15. Respirations 45.

4,15—21. Stands on right leg only.

4,22-23. Respirations 54.

4,23-24. Respirations 49.

4.25. Temperature stil] normal, 102°

4.35. Temperature 99°. :

4,39—-40. Respirations 48.

4,42. Temperature 98°.

4,44-46. Respirations 35 per minute.

4.47. Temperature 96°. Slides along on breast when trying to walk,
4.4748. Respirations 44, very weak.

Ac PB, WO, ADK, YA,

bo
S
oe)

SOME EXPERIMENTS WITH THE

4,52. Temperature 96°.

4.538-54. Respirations 44.

4.56-57. Respirations 31,

4.58. Temperature 96°.

5.00-OL. Respiration, wheezing pants.

5.01-02. Respirations, wheezing pants, 21.

5.02. Temperature 96°. Death without struggles.

The Kffect of Glycerine.—Glycerine seems to dissolve the poison and to partly destroy
its effectiveness, though this seeming injury may be due to the slowness with which the
glycerine is absorbed, preventing the poison from reaching the circulation rapidly enough
to result fatally.

EXPERIMENT X.—Paper containing saliva of four bites was placed in about forty minims of glycerine
and left for some hours. The glycerine, having been extracted, was injected in the breast muscles of a
pigeon at 12.10 P.M., December 4, 1896.

1.00. Still no effect.

5.15. Still no effect.

December 5. Well, but with yellowish-white swelling on breast.

December 17. Well, but breast muscles sloughing. Used in experiment XII.

EXPERIMENT XI.—December 4, 1896. Since it was quite possible that the poison had not been
dissolved by the glycerine, the paper used in the last experiment was well washed in alcohol to remove
glycerine, and then, after the alcohol had been removed by pressure and evaporation, was placed in water
(thirty minims). This water was injected into a pigeon at 3.15 P.M.

3.30. No signs of poison.

5.15. No effect yet.

December 5. Well.

December 8. Well.

Exprrment XII.—December 17. Saliva of the lower jaw from about three bites was collected
and divided into two parts, one slightly larger than the other. The larger part was then soaked in glycerine,
a little more than one-half of which was afterward injected in leg of pigeon used in experiment X.

4.35 P.M. Injected subcutaneously.

5.30. Seems slightly drowsy ; ‘otherwise well.

December 18. Found dead.*

ExpermMENT XIII.—December 17, 1896. To test the power of the saliva used in experiment XII
the smaller portion of the saliva-soaked paper was placed in a small quantity of water, and one-half of
the resulting solution injected in the breast muscles of a pigeon, December 18.

4.07. Injected.

4.30. Bird sitting; staggers when raised.

* Death may have been due to the rather extensive sloughing of the pectoral muscles, but that this was the

case does not seem probable.

SALIVA OF THE GILA MONSTER. 209

4.31-32. Respiration still normal, 7. ¢., 35.

4.35. Can still stand.

4.36-37. Respirations 30.

4.39-40. Respirations 31.

4.46-47. Respirations 29; sits with eyes closed.

4.53. Does not notice loud noises, as stamping on floor; cannot stand.
4.55-56. Respirations 31.

4.58. Head moves from side to side, slightly.

4.59-5.00. Respirations 30.

5.03—04, Respirations 34, slightly forced.

5.09-10. Respirations 34, slightly forced.

5.18-14. Respirations 43, a little more forced; head nodding.
5.15-16. Respirations 36, nearly normal.

5.18-19. Respirations 32, slightly forced.

5.2122, Respirations 50, much forced.

5.23-24. Respirations 32, convulsive.

5.24—25. Respirations 23, convulsive.
5.255. Raises tail and flaps wings.

5.26—27. Respirations 13, weak.

5.28. Heart still beating strongly and regularly.

5.30. Death.

Heart irritable and nerves of pectoral muscles, etc., likewise; blood very dark, semi-liquid, coagu-

lating quickly; no local effects.

THe HARMLESS SALIVA.

There is, then, in the saliva of the Gila Monster a yery powerful poison which may
be subjected to yery rough treatment without impairing its lethal vigor. This poison is
present in the saliva of one yaw only. If, when collecting the mouth fluids, the rubber be
properly placed between two layers of paper, the saliva from each jaw may be readily
obtained unmixed with that of the other. When thus obtained and dissolved in water,
the saliva of the upper jaw is a yellowish liquid, usually more or less tinted with blood,
slightly alkaline, without any odor, and absolutely harmless at the very time when the
lower jaw is flooded with deadly yenom. The quantity of saliva which may be collected
from the upper jaw at any one time is only a little less than is obtainable from the lower ;
but in one ease all of the saliva from the upper jaw was injected into a pigeon without
causing the slightest ill effect, while one-fifth of that obtained at the same time from the
lower jaw caused death in fifty-two minutes.

The following experiments are quite numerous enough to show beyond doubt the

difference in effect between the two kinds of saliva.

210 SOME EXPERIMENTS WITH THE

ExprRIMENT XIV.—November 24, 1896. Saliva of upper jaw from four bites was dissolved in
water one-half of which (ten minims) was injected into a pigeon at 11.40 A.M.

3.08. Still no effect; repeated injection.

5.40. Still no effect.

November 25. Well.

EXPERIMENT X V.—November 24, 1896. Same as last experiment, but with saliva of lower jaw in
another pigeon. ;

12.15 P.M. Temperature 104°.

12.17. Injected.

12.20-21. Respirations 31.

12.27-28. Respirations 31.

12.35. Temperature 100°.

12.36-37. Very ‘‘ tame.’’ Respirations 38.

12.38. Sways backward and forward.

12.39-40. Respirations 52.

12.42. Temperature 98°.

12.4748. Respirations 30.

12.50. Very drowsy. Temperature 97°.

12.54-55. Respirations 34, irregular.

1.03-04. Respirations 28, labored.

1.06. Temperature 95°. Can still stagger when placed on feet.

1.09-10. Respirations 38, very irregular.

1.11. Temperature 96°.

1.16. Temperature 95°.

1.17-18. Respirations 42, greatly labored.

1.28. Temperature 95°.

1.24-25. Respirations 46, bill opening and shutting. Can still walk slowly.

1.28-29. Respirations 55.

1.30. Temperature 96°.

1.33-34. Respirations 52. Can barely walk.

1.36. Temperature 96°.

1.37. Cannot walk.

1.37-38. Respirations 54.

1.46. Temperature 94°.

1.47-48. Respirations 49, head nods.

1.53. Temperature 94°.

1.54. No respiration.

1.55. Temperature 93°.

1.56. Death with convulsions.

Exprrmmenr XVI.—November 25, 1896. At 2.15 P.M., injected a pigeon with all of solution of
saliva of upper jaw from four bites. .

2.30. Still no effect.

from

2.40. Still no effect.
3.07. Still no effect.
5.05. Still no effect.
November 26. Well.

THE GILA MONSTER. Zh

EXPERIMENT XVII.—November 25, 1896. _ Injected one-half of the solution of lower-jaw saliva

same bites as last experiment.

3.02-03. Respirations 37; temperature 104°.

3.06. Injected as above stated.

3.14. Temperature 102°.
3.23-24, Respirations 38.

3.27. Very ‘‘ tame,’’ temperature 98°.

3.28. Cannot stand.

3.285-294. Respirations 53.

3.30. Temperature 98°.
3.32-33, Respirations 45.
3.33. Temperature 98°.
3.37. Temperature 98°.

3.38-39. Respirations 45.

3.40. Temperature 96°.
3.40-41. Respirations 45.
3.91, Temperature 94°.

.)38-54, Respirations 43.

wo 9

.56. Temperature 94°.

4.07. Temperature 93°.
4.15-16. Respirations 51.
4.21. Temperature 93°.
4,27-28. Respirations 26.
4,29. No respiration.
4.30. Death.

3.58-59. Respirations 45.

Heart (auricles and ventricles) beating strongly when exposed at 4.31 and until 4.36; blood in

veins; arteries and ventricles empty; no local effect.

Exprerment XVIII.—November 28, 1896.

pigeon, at 11.55. No effect.

EXPERIMENT XIX.—November 28, 1896.

bites as last experiment) in pigeon at 12.15 P.M.

Injected all of solution of saliva from upper jaw, in

Injected all of solution of saliva from lower jaw (same

12.19. Tips forward on legs, therefore cannot stand still.

12.20. Seems dizzy.
12.204. Sits.
12.22. Can walk well.

212 SOME EXPERIMENTS WITH THE

12.24. Very ‘‘tame;’’ hardly able to walk.

12.27. Can stagger with help of wings.

12.34. Respiration terribly labored, loud, wheezing pants, about 28 per minute.
12.39. Head drawn far back; still panting.

12.40. Still panting, but more slowly and weakly, 24 per minute.

12.41. Struggles, lies on side with head on floor.

12.42. Respiration practically stops.

12.421. Dead. ;

EXPERIMENT XX.—December 1, 1896. Injected solution of saliva of upper jaw from two bites, at
12.30 P.M.

1.30. Pigeon has shown no signs of poisoning.

3.30. Still no effect.

4,30. Still no effect.

5.00. Still no effect.

December 2. Well.

EXPERIMENT XXI. Injected solution of saliva of lower jaw from same two bites (experiment XX)
at 2.25 P.M., December 1, 1896.

3.25. Totters; lies down when set on feet.

4.00. Totters, leaning forward.

4.10. Can still totter.

4.20. Cannot rise or stagger.

4,30. Muscles all tense; bill opens and shuts.

4.30%. Respiration ceases.

4.31. Death.

EXPERIMENT X XIJ.—December 2, 1896. All of the solution of saliva of the upper jaw from three
bites was injected under the skin of the wing of a brown pigeon at 3.05 P.M. without any effect.

EXPERIMENT X XIII[.—December 2, 1896. T'wo-fifths of the solution of lower-jaw saliva from the
same three bites as last experiment were injected under the skin of wing of a pigeon at 3.15 P.M.

3.25. No effect yet.

3.28. Staggers slightly; sits immediately; respiration slightly forced.

3.32. Respiration very rapid—forced.

3.36. Respiration very slow but labored.

3.40. ‘‘ Skates ** on breast when trying to walk.

3.43. Convulsive quivering of wings.

3.44-45. Convulsive quivering of wings.

3.45. Lies stretched out on floor; convulsive respiration; wheezing with each breath.

3.48. No respiration.

3.484. Death.

SALIVA OF THE GILA MONSTER. 213

EXPERIMENT XXIV.—December 2, 1896. Two-fifths of the solution used in the last experiment
(XXIII) were injected in the breast muscles of a slate-colored pigeon at 3.16 P.M.
25. Barely able to walk.
6. Not able to stand; respiration forced.
8. Lies on side with head drawn back.

4. Respiration very rapid and convulsive, bill opening and shutting; head twisted on side.

94. Apparently dead.
0. Heart still beating.

9
oO.
9
oO.
3.
9
oO.
3.
9
oO.
9
oO.

2
2
3
39. Respiration ceases.
3
4

ExpERIMEeNtT XX V.—December 2, 1896. One-fifth of solution used in experiments X XIII and
XXIV was injected in a gray pigeon at 3.20 P.M.

3.25. Respiration deeper.

3.42-43. Respiration very rapid and shallow, 148 per minute.
3.51-52. Respirations 167; can still walk, but sits immediately.
3.58-59. Respirations 168.

4,02. Cannot stand.

4.04. Slight trembling.
-4.05-06. Respirations 149.

4.08. Head drawn back; bill opens and shuts.

4.09-10. Respirations 62.

4.10. Slight general contractions of muscles.

4.114-114. Respirations 4.

4.114-12. No respiration. —

4.12. Death.

EXPERIMENT XX VI.— December 8, 1896. Solution of upper-jaw saliva from one bite injected in
breast of a gray pigeon at 3.08 P.M without effect.

EXPERIMENT XX VII.—December 8, 1896. One-half of solution of lower-jaw saliva, same bite as
experiment X XVI, was injected in breast muscles of a gray pigeon at 3.16 P.M.

5.26. Pigeon very quiet.

4.00 Drowsy.

December 9. Well.

December 18. Well.

THE SouRCES OF SALIVA.

We have seen that two very different fluids are present in the mouth of the Helo-
derma; the one—from the lower jaw—capable of causing profound disorder when intro-

duced into the circulation of pigeons, the other—from the upper jaw—producing no more

effect than so much water. What are the sources of these fluids?

214 SOME EXPERIMENTS WITH THE

In Heloderma suspectum, there are two large glands, one on each side of the anterior
part of the lower jaw between the skin and the bone. When one of these glands has
been freed from its outer sheath it is found to be not a single gland but a series of three
or four glands, each perfectly distinct from the others and emptied by a separate duct.
These glands increase in size posteriorly, so that the last is very much larger than the
first. They vary in number because of the occasional union of the first and second
glands, or the presence, posteriorly, of a small, isolated, ductless portion. Their ducts
open between the lower lip and gum, as described by Stewart. It is shown later on that
these are the yenom-producing glands.

No glands have yet been described as existing in the upper jaw; indeed there seems
to be no room there for a well-developed gland. Nevertheless, paper which comes in
contact with the upper jaw during the bite collects almost as much fluid as is obtained
from the lower jaw. This, however, is true only when the paper is bitten a very few
times. The saliva of the upper jaw is exhausted much more quickly than that of the
lower. This fact, taken in connection with the absence of known glands, might lead one
to suspect that the upper jaw receives its saliva from the lower and holds it in the compli-
cated folds of its gums. This might perhaps be true if one or more segments of the sub-
labial glands secreted a harmless fluid, but the following experiments show that all are
specialized for the production of venom. I believe that the harmless saliva is secreted
by minute glands which lack of material has prevented me from finding—that it is in
fact the ordinary buccal liquid of lizards. That it is present in the lower jaw as well as
in the upper would seem to be shown by the fact that the fluids of both jaws are decidedly
alkaline, while a solution of the poison gland itself is quite neutral.

The following experiments were performed to show that each part of the sublabial
glands is deyoted to the production of yenom:

EXPERIMENT XX VIII.—January 5, 1897. Soaked the first portion of the right sublabial gland in
water and injected the resulting solution (three minims) into the breast muscles of a small finch, at
12.26 P.M.

12.28. Respiration forced; eyes closed.

12.29. Respiration greatly forced.

12.31. Flutters.

12.314. Convulsions and death.

12.35. Heart beating weakly; blood dark but lightens quickly.

EXPERIMENT X XIX.—January 5, 1897. Soaked the second portion of the right sublabial gland in
water and injected solution (four minims) into breast muscles of a small finch, at 12.00 M.

12.04. Eye nearly closed; respiration normal.

12.05. Respiration slightly forced.

—
Or

SALIVA OF THE GILA MONSTER. ale

12.054. Bill begins to open and shut.

12.07. Respiration greatly labored.

12.08. Convulsions followed by death.

12.10. Heart still beating; blood dark, lightens slowly.

EXPERIMENT XXX. Treated the third portion of right sublabial gland as the first and second were
treated in experiments XXVIII and X XIX, and injected four minims into a small finch at 11.34 A.M.

11.35. Wheezes; sitting down; eyes closed; tail moving up and down with each breath.

11.86. Same, but bill opening and shutting.

11.37. Does not open eyes when handled.

11.374. Respiration very short and jerky.

11.38. Respiration ceases, followed by convulsions and death.

11.41. Heart still beating, empty; blood dark brown, reddening very slowly.

EXPERIMENT XXXJ.—January 5, 1897. Injected four minims of solution of fourth portion of
right gland into a small finch, at 11.074 A.M.

11.084. Unable to stand erect; head drooping.

11.09. Respiration labored.

11.094. Respiration greatly labored.

11.10. Bill opens and shuts.

11.11. Bird falls on side.

11.124. Respiration in gasps.

11.13. Convulsions and death.

Heart responds to mechanical stimuli; blood black but becoming red on exposure.

EXPERIMENT XXXII. Injected five minims solution of first portion of left sublabial gland into a
small finch, at 2.41 P.M.

2.42. Hyes closed.

2.45. Respiration labored; bird leaning on side.

2.46. Almost unconscious; bill opening and shutting.

2.47. Convulsions.

2.474. Death.

Exprer(MenT XXXIII. Injected six minims of water into the breast muscles of a small finch

without effect.

Ill. THE PHYSIOLOGICAL ACTION OF HELODERMA POISON.

When a pigeon has received an injection of Gila Monster saliva it at first shows no
ill effects, and feeds or fights with its fellows as before. Soon, however, it begins to wink
very frequently, and ceases to show interest in anything about it. It stands thus for a
A. P. S.—VOL. XIX. 2B.

216 SOME EXPERIMENTS WITH THE

longer or shorter time and then sits down. If now it be frightened into attempting to
walk, it appears dizzy and staggers about, or, if unable to stand, slides along on its breast.
If not caused to arise, it never does so of its own accord, but becomes more and more
drowsy and sits with eyes closed. The rate of respiration now becomes yery rapid for a
time, but soon the breaths are shallower and then gradually fewer and fewer.* The legs
become more or less paralyzed, but the wings retain their power, although the codrdina-
tion of their motions sometimes is destroyed. The temperature falls as the respiration
becomes slower. The bird rolls over on its side. The head is drawn down over the back.
Respiration becomes nothing more than a series of wheezing gasps, with each of which
the bill opens and shuts. The head falls forward to the floor. The pigeon is unconscious.
Breathing ceases. There may be slight conyulsions followed by death, or death may
come quietly.

If the pigeon now be opened, it is found that the blood is very dark—often almost
black instead of red or blue. The heart either is beating or responds readily to mechan-
ical stimuli. The arteries and usually the ventricles of the heart are empty, while the
veins and auricles are full of blood which usually is more or less clotted. There is no
trace of discoloration about the point of injection, nor is the slightest extravasation of
blood to be found in any of the organs.

With all these facts in view, it is very evident that death is due to asphyxiation ; to
the failure of the blood to provide the various tissues of the body with the oxygen neces-
sary for their welfare. But, although. we may say that death is due to asphyxiation, we
have not really answered our question, for there are several ways in which this failure on
the part of the blood might be brought about :

1. If the poison acted upon the nerve centres which control the moyements of respi-
ration in such a way as to interfere with the action of the lungs, the blood would be
unable to procure its usual supply of air. We have seen that there is a very decided dis-
turbance of the respiratory function.; It may, perhaps, be due to direct nerve-poisoning ;
but I am inclined to believe that it is entirely a secondary phenomenon.

2. If the poison caused a breaking down of the capillaries of the lungs—such as
Martin{ claims to have found in certain cases of death from the venom of the Australian
black snake—the same effect would be produced, but there appears to be no such change.

3. Ifthe action of the heart became gradually weaker—as Mitchell and Reichert
have stated of their experiments—the flow of blood would be diminished and the tissues

*MThis is normally true, but respiration sometimes stops suddenly, even nearly at the time when it is

most rapid.

+ The table upon the opposite page shows the effect upon the number of respirations and the temperature.

+ Martin, Jour, and Proc, Royal Soc. N. 8. Wales, XXIX, 1895, 146-276.

MINUTE.

RESP. TEMP. |

RESP.

SS eee
imi Go G9 OD Go GY wD 29 t9
PS SRSA RSO ASE GS

Poems

108

39)

104

31
31
31

34.

SALIVA OF THE

TEMP.

104

100

Oman}

RESP. TEMP. || RESP.
\|

54
49

GILA MONSTER.

RESP. || RESP.

TEMP. | RESP. || RESP. RESP.
|| | |
| =| si || || ||
37 | 104 | 35 || || 34
32 |
| \| 36
| 102 | |
|| 48 | 1} ji) ie!
|| Nene za | |
| 49 84 | | |] 32
\] |
| 88 64 || || |
|| |
} 53m | 78
98 || Hl
|| 47 1D, || | || D.
i || 148
}) 953 98 ||
D. | 35 |!
|| 45 98 | I}
AS | |
| 30
98 |
| 167
45 31 ||
96 | |
45 || }
|
} |
| | 168
29° |
94
149
43
| || 31
| 94 || 62
‘ |
| | 4-0
| 45 I 80 D.
| i
|
84
\|
102 | |
93 | | 1]
| 34 |
| | 43 |
i] |
| || 36
99 51 | |
93 | 50
98 | | 32 |
| | steel
96 || 26 | | |
0 | D.
Dz D.
96 |
|
96 |
96 |
D.

218 SOME EXPERIMENTS WITH THE

would not receive their normal amount of oxygen. In all my experiments the heart con-
tinued to beat regularly long after respiration had ceased, so that this cannot have been
the cause of death.

4. If the poison acted upon the blood in such a way as to destroy its power to carry
oxygen—as Cunningham * says is true of cobra yenom—or,

5, if the poison caused the formation of clots in the veins, thus stopping the flow
of blood—as Martin tells us the yenom of the Australian black snake does—in either
case the effect would be the same as if the action of the lungs were to cease.

The sudden death of my Gila Monster prevented me from testing these possible
causes of asphyxiation from its poison, but I shall not be surprised if it be found that in
one or both of them exists the explanation of the phenomena exhibited.

But perhaps I should limit this statement somewhat, for Mitchell and Reichert state
very positively of their experiments that death was occasioned by the action of the
poison upon the heart. Here is an apparent contradiction of my results, and by the
highest American authority upon reptile poisons; but the seeming contradiction disap-
pears, perhaps, when we recall that Dr. Mitchell’s Gila Monster saliva was less dilute
than mine, and that it is known of some serpent poisons that “ with higher concentration
of yenom the heart is the more rapidly affected, but the continuous operation of the
poison in small concentration more quickly affects the respiratory ” system.

IV. SOME CAUSES OF DIVERSITY OF OPINION.

We have now reached our last question: Why has the bite of the Gila Monster so
often been considered harmless ?

Several reasons must, I think, already have suggested themselves. Dr. Shufeldt, it
will be remembered, was severely bitten on the thumb, and concluded that the bite of the
Gila Monster is no more poisonous than that of other angry animals; for example, a cat.
But Dr. Shufeldt expressly states that the wound was made by the upper teeth pene-
trating to the bone, and we have already seen that the saliva of the upper jaw is harmless
at all times, the venom being confined to the lower jaw. So it well may be that Dr.
Shufeldt owes his life to the circumstance that the injury to his thumb was inflicted by
the upper instead of the lower teeth of the Monster.

This same fact will account for the experiences of other authors who have thought
the bite of this reptile harmless, but there are other reasons for the occasional failure of
the Heloderma to inflict a deadly wound. The teeth, although sharp and long, are very
weakly fastened to the jaws, and often so many of them haye been broken out that the

*Cunningham, Sez. Wem. Med. Officers Army India, LX, 1895, pp. 1-54.
+ It would be interesting to know why the teeth of the upper jaw are grooved.

SALIVA OF THE GILA MONSTER. 219

Monster is unable to inflict a wound at all. Even if the teeth are in working order the
chances of the poison finding its way into the wound are very few, for the teeth are not
directly connected with the poison glands, and the latter are below the fangs instead of
above as in poisonous snakes. The poison simply flows out onto the gums below the
teeth, and, to be effective, has to be forced wp into the wound. Unless the flow of saliva
be abundant and the teeth all present and forced into the bitten flesh so deeply as to
press it down upon the poison ducts where they open between the lip and the gum, it is
difficult to see how even the smallest quantity of poison could enter the wound, eyen
though the teeth are grooved to afford it a passage. The strange thing, then, is not that
bitten animals should sometimes survive, but that they should sometimes die.

Nevertheless, small animals often do die from the bite of this, the only poisonous
lizard, and we must believe that a yenom which can kill a pigeon in seven minutes and a
rabbit in less than two might easily under favorable circumstances cause a wound to
prove fatal even to man—a belief which is rendered far from improbable by the extra-
ordinary virulence of the poison and the lizard’s habit of holding like a bulldog to what-
ever it bites.

V. BIBLIOGRAPHY:

1651. Hmrnanpez, F.—Historiz animalium et mineralium Nove Hispanie, p. 315.

1829. Wieamann, A. F.—Ueber das Acaltetepon oder Temacuilcahuya des Hernandez, eine neue Gattung der
Saurer, Heloderma. Isis, pp. 627-629.

1857. Gray, J. E.—On the Genus Necturus or Menobranchus, with an Account of ItsSkulland Teeth. Proc. Zool.
Soc. Lond., p. 62. "

1864. SumicHrast, F.—Note sur les Meeurs de quelques Reptiles du Mexique. Bibl Univers. et Revue Suisse (Ar-
chives des scien. phys. et nat.), XIX, pp. 45-61. [Reprint pp. 1-5.]

1864. SumicHrast, F.—-Notes on the Habits of Some Mexican Reptiles. Annals and Mag. Nat. Hist. (3), XIU,
pp. 497-500.

1869. Corn, E. D.—[Remarks.] Proc. Ac. Nat. Sci. Phila., p. 5.

1873. Gervais, P.—Structure des dents de l’Héloderme et des Ophidiens. Comptes Rendus Acad. des Sciences,
LXXYVII, pp. 1069-1071.

1875. Bocourt, F.—Observations sur les meurs de l’Heloderma horridum, Wiegmann, par M. F. Sumichrast.
Comptes Rendus Acad. des Sciences, LX XX, pp. 676-679.

1875. Yarrow, H. C.—Report upon the Collections of Batrachians and Reptiles made in Portions of Nevada,
Utah, California, Colorado, New Mexico and Arizona during the years 1871, 1872, 1873, and 1874. U.S.
Surv. W. 100th Merid., V, pp- 562, 563.

1878. Bocourt, F.—Mission Scientifique au Mexique et dans 1’Amerique Centrale, III, Reptiles, 5e livr., pp. 296-—
306, Pls. XX EH, Figs. 1-12, XX G, Figs. 1, 3, 6, 7, 8, 9, 10, 11.

1880. SumicuRast, F.—Bulletin de la Société Zoologique de France, p. 178.

1882. American NarurALIst.—[Note.] Am. Nat., XVI, p. 842.

1882. BounENncEerR, G. A.—[Remarks.] Proc. Zool. Soc. Lond., p. 631.

1882. Fayrer, J.—[Remarks.] Proc. Zool. Soc. Lond., p. 682.

1882. Fiscuer, J. G.—Anatomische Notizen tiber Heloderma horridum Wiegm. Verhandl. des Vereins Hamburg,
Bd. V, pp. 2-16, Pl. III.

SOME EXPERIMENTS WITH THE SALIVA OF THE GILA MONSTER.

SHUFELDT, R. W.—The Bite of the Gila Monster (Heloderma suspectum). American Naturalist, XVI, pp.
907, 908.

AMERICAN NATURALIST.—[Review of Mitchell’s and Reichert’s article.] XVII, 7, July, p. 800.

GARMAN, S.—Reptiles.and Batrachians of North America. Memoirs Mus. Comp. Zool. Cambr., VIL, 3,
p- Xi.

M , H. N.—The Physiological Action of Heloderma Poison. Science, I, 13, May 4, p. 372.

MircHe.t, S. W., and ReitcHEerRt, E. T.—A Partial Study of the Poison of Heloderma suspectum (Cope)—
the Gila Monster. Medical News, Phila., XLII, No. 8, Feb. 24, pp. 209-212.

BouLENGER, G. A.—Catalogue of the Lizards in the British Museum, Vol. II, pp. 300-302.

GuntueEr, A. C.—Biologia Centrali-Americana, Reptiles, pp. 43, 44, Pl. XXYVI.

BENDIRE, C. E.-Whip Scorpion and the Gila Monster. Porest and Stream, XXIX, 4, Aug. 18, pp. 64, 65.

SHUFELDT, R. W.--The Gila Monster. Forest and Stream, XXIX, 2, Aug. 4, p. 24.

Yarrow, H. C.—Bite of the Gila Monster. Forest and Stream, XXX, 21, June 14, pp. 212, 213.

Lussock, J.—[Letter from 8. A. Treadwell.] Proc. Zool. Soc. Lond., p. 266.

MircHeiy, §. W.—The Poison of Serpents. Century Magazine, XXXVIUII, 4, p. 505.

GARMAN, §.—On the ‘‘Gila Monster’’ (Heloderma suspectum). Bull. Essex Inst., X X11, pp. 60-69.

SHUFELDT, R. W.—Contributions to the Study of Heloderma suspectum. Proc. Zool. Soc. Lond., pp. 148-244,
Pls. XVI-XVIII.

BouLENGER, G. A.—Notes on the Osteology of Helodermu horridum and H. suspectum, with Remarks on
the Systematic Position of the Helodermatide and on the Vertebrie of the Lacertilia. Proc. Zool. Soc.
Lond., pp. 109-118.

BouLENGER, G. A.—The Anatomy of Heloderma. Nature, XLIV, p. 444.

SHUFELDT, R. W.—The Poison Apparatus of the Heloderma. Nutwre, XLII, p. 514.

SHUFELDT, R. W.—Further Notes on the Anatomy of the Heloderma. Nature, XLIY, p. 294.

SHUFELDT, R. W.—Some Opinions on the Bite of the ‘‘Gila Monster’’ (Heloderma suspectum). Nature's
Realm, Il, 4, April, pp. 125-129.

SHUFELDT, R. W.—Medical and Other Opinions Upon the Poisonous Nature of the Bite of the Heloderma.
New York Medical Journal, LIX, 21 (651), May 28, pp. 581-584.

STEWART, C.—On Some Points in the Anatomy of Heloderma. Proc. Zool. Soc, Lond., pp.119-121, Pl. XI.

ARTICLE IV.

RESULTS OF RECENT RESEARCHES ON THE EVOLUTION OF THE STELLAR
SYSTEMS.

(Plates LV and VY.)

BY T. J. J. SEE, A.M., Pa.D. (BrRuin),

ASTRONOMER AT THE LOWELL OBSERVATORY.

Read before the American Philosophical Society, January 7, 1898.

It is now two hundred and eleven years since Newton published the Principia,
embodying his grand generalization of the law of gravitation, and the proof of this law
for the most obvious and fundamental phenomena of the solar system. Geometers have
since been occupied with the development and extension of the principle discovered by
the illustrious Newton, and have finally explained with almost entire satisfaction the
motions and attractions of the planets, satellites, comets, and other bodies which revolve
about the sun. This great-development can hardly fail to excite the admiration of those
who contemplate the history of scientific progress, and must be accounted one of the most
noble and enduring monuments of the human mind. So sublime an achievement has
required the combined labors of a long series of men of transcendent mathematical and
mechanical genius, each building upon the foundation laid by his predecessors. Though
many distinguished geometers have borne an honorable part in this remarkable develop-
ment of Physical Astronomy, it will not be inappropriate to point out the great credit for
the perfection of the Newtonian theory due to Clairaut and Euler, Lagrange and La-
place, Gauss and Hansen, Adams and Leverrier. Among living investigators in mathe-
matical astronomy the names of Hill and Newcomb, Darwin and Poincaré occupy the
foremost place. These great men have brought the mechanics of the heavens to so high
a state of perfection that in almost every case we may now predict the heavenly motions
as accurately as we can observe them. In view of the rapid perfection of telescopes and
other instruments of precision, this achievement, from the intricacy of the analysis
required in the problem, and the abstruseness of the methods used in the reduction of

22 _ RESULTS OF RECENT RESEARCHES ON THE

observations, must be ranked as incomparably the most profound yet attained in any
branch of Physical Science.

Notwithstanding these splendid triumphs of the science of Celestial Mechanics, an
even greater and more recondite work remains to be done in a closely related field. This
is the investigation of the origin and cosmical history of the planetary and other systems
observed in the immensity of space. Even if some credit for pioneer work on this
problem be assigned to Kant, or, more remote still, to the Greeks of the pre-Socratic age,
it yet remains true that Laplace is the real discoverer to whom we are indebted for the
first ideas which proved fruitful for the advancement of science. About a century ago
this great geometer outlined for the solar system the celebrated Nebular Hypothesis, upon
which nearly all subsequent investigation has been based, and which has since been sub-
stantially confirmed, though but very little modified until within the last twenty-five
years. Passing over as irrelative in the present discussion the early work of Herschel
and Rosse, Helmholtz and Kelyin, Newcomb and Lane, we come down to the modifica-
tions introduced by Darwin about 1880.

In establishing the theory of gravitation, Newton assigned also the true cause of the
tides of the seas, though his explanation carried with it all the defects of the equilibrium
theory. More than a century passed before the dynamical character of the problem of
the oceanic tidal oscillations was clearly perceived, when Laplace developed and applied
the true theory with all the penetration characteristic of that great mathematician.
Yet in spite of the profundity which marks his treatment of the tides of the oceans, it
seems never to have occurred to him, or at least he made no record of the fact, that the
attraction of the moon necessarily produces tides in the body of, as well as in the aqueous
layers covering, the earth. We need not be surprised at this omission on the part of
Laplace and those who followed him, if we recall that for many years after the perfection
of Analytical Mechanics by D’Alembert and Lagrange, the subject was treated wholly
from the point of view of material particles, and the resulting system was what is now
called Rigid Dynamics. Little attention was bestowed upon the theory of fluid motion,
partly because of its intricacy, and. partly because there were no obvious applications of
the results except in the case of the tides, already treated by Laplace with great penetra-
tration and extreme generality. As mathematicians since the time of Newton had been
occupied chiefly with the development of the theory of planetary perturbations along the
line of rigid dynamics, it did not occur to them that they were building on a false
premise, that in reality the heavenly bodies so far as known are not solid, but fluid,
though Laplace with his usual sagacity had long foreseen that in the case of our planets
the nuclei are covered with fluid layers held in equilibrium by the pressure and attraction
of their parts. His grand treatment in the Mécanique Céleste recognizes the fluidity of

EVOLUTION OF THE STELLAR SYSTEMS. Deep)

the envelopes of the planets, and exhaustively examines the oscillations that will arise
therein. Nor did he fail to consider fully the deviations from spherical form and the
probable laws of density for the layers which compose the bodies of the planets.

The effect of so monumental a work as the Mécanique Céleste was twofold: on the
one hand it brought Physical Astronomy to an unexpected state of perfection, while on
the other it produced the impression on the less creative minds that there were no great
problems untouched by the master-mind of Laplace. His work had indeed well-nigh
exhausted the theory of Celestial Mechanics, so far as it could be built upon the assumptions
of rigid dynamics ; at least subsequent work has been for the most part little more than
refinement or perfection of the methods and processes given in the Mécanique Céleste. The
work of Laplace was designed for the solar system, and the idea that the universe is really
composed of fluid bodies, self-luminous stars and nebulz in space, seems never to have
occurred to him, or he would have foreseen that however adequate Rigid Dynamics may
be for effecting a first approximation, the true theories of ultimate Celestial Mechanics
must be founded upon the laws of viscous fluids in motion. So great is the influence of
tradition that it is difficult for us to realize fully that the stars and nebule are viscous
fluids, self-luminous liquid or gaseous masses, and that even in the solar system the
bodies are all fluids of various viscosities. This new point of view respecting the actual
facts of the universe has brought about an important modification in the nebular hypothesis
and in the ultimate theories of Celestial Mechanics, of which we shall now give some
account.

About 1875, G. H. Darwin, who had qualified himself for the Law and been called to
the Bar, on account of ill-health, abandoned his profession to undertake for Lord Kelvin
some scientific work, which among other things included the reduction of a great mass of
Indian tide observations with a view of throwing light upon the problem of the rigidity of
the earth. This work, besides leading Lord Kelvin to the celebrated conclusion that the
earth as a whole is “ probably more rigid than steel, but not quite so rigid as glass,” was
the oceasion* of the younger Darwin developing the theory of bodily tides, or the
theory of the tides which would arise in the earth on supposition that it is not rigid as at
present, but a viscous fluid, as it must have been, according to Laplace, at some past age.
While some allusions to bodily tides can be found in scientific literature as far back as
Kant, and especially in the papers of Delaunay on the secular acceleration of the moon’s

*In the Atlantic Monthly. for April, 1898, Prof. Darwin remarks: “It was very natural that Mr. See should
find in certain tidal investigations which I undertook for Lord Kelvin the source of my papers, but as a fact the
subject was brought before me in a somewhat different manner. Some unpublished experiments on the viscosity of
pitch induced me to extend Lord Kelvin’s beautiful investigation of the strain of an elastic sphere to the tidal dis-
tortion of a viscous planet. This naturally led to the consideration of tbe tides of an ocean lying on such a planet,
which forms the subject of certain paragraphs now incorporated in Thomson and Tait’s Watwral Philosophy.

A. P. S—VOL. xix. 2c,

QA RESULTS OF RECENT RESEARCHES ON THE

mean motion, it is yet indisputable that Darwin was the first writer to treat the problem
in a systematic, thorough-going and original way. Recognizing that at some epoch in
the past, the earth was probably a mass of viscous fluid, he set for himself this problem :
To determine the bodily tidal distortion of the earth, and the effects of this alteration of
figure upon the orbital motion of the moon, and upon the earth’s rotation. His papers
were communicated to the Royal Society between 1878 and 1882, and are celebrated con-
tributions to the general theory of tides. In these papers he has traced the moon back
to close proximity to the earth, when the two, at the breaking off of the moon, were most
probably revolving in about 2h. 41m. The moon has since receded from the earth under
the action of tidal friction, while the rotation of the earth has been slowed up in correspond-
ing degree. It was rendered certain that in the origin of the Lunar-Terrestrial System,
the action of tidal friction had played a prominent, if not a paramount part, and the
question naturally arose whether it had not been equally potent in the development of other
parts of the solar system. When, however, Prof. Darwin came to apply the results to
other satellite systems and to the solar system as a whole, it was found that here the
effects had been much less considerable than in the case of the earth and moon, owing
chiefly to the small masses of the attendant bodies. Thus the major axes of the orbits
had perhaps been very slightly increased, and the rotations correspondingly exhausted,
but no radical change had taken place. Under these circumstances it was natural that
Darwin should drop the subject without further search for extension of the principle he
had developed.

About November 1, 1888, while I was still an undergraduate at the Missouri State
University I became much interested in the origin of the double stars. The immediate
cause of my taking up the subject was the Missouri Astronomical Medal, occasionally
awarded by the University to a graduate of highest standing in the Mathematical and
Physical Sciences. Having been informed by Prof. W. B. Smith that I was eligible to
write for the medal, by virtue of my standing in the Physical Sciences, our conversa-
tion drifted on to the probable subject of the Thesis, and in this way he was led to
suggest a criticism of Darwin’s work on the origin of the moon. He remarked: “ You
may find this only a pocket, already worked out, and not a continuous vein of rich ore,
but it seems to me worth thinking of. At any rate I would not advise you to write on
the orthodox Laplacean Nebular Hypothesis, for that subject is worn threadbare.”

The suggestion of a critique of Darwin’s work did not quite meet my approval, for I
feared the subject was already exhausted and would leaye no field for future progress.
As I had been observing various double stars for the past two years, and had seen no
suggestion regarding their mode of development, it occurred to me that perhaps the tidal

theory might find application among the stars. When I had collected such orbits as were

EVOLUTION OF THE STELLAR SYSTEMS. 225

available in the books at my disposal (Humboldt’s Cosmos, Herschel’s Outlines, ete.), I
discovered to my surprise that unlike the orbits of the planets and satellites, they are
very eccentric, though not so eccentric as those of the periodic comets. It was at once
evident that it would be hopeless to attempt to explain the origin of the stellar systems,
if we could not explain the cause of the high eccentricities of the orbits. The next day
I called on Prof. Smith and told him of the discovery that the orbits are very eccentric,
and asked whether he thought [ might explain this peculiarity on the tidal theory ; rub-
bing his head for a moment in quiet reflection, he replied: “Oh! I see what you mean ;
you think the dragging of the tides in the bodies of the stars has produced the elongation
you find in the orbits. Such an idea can hardly be discussed off-hand, but it is at least
worth examining; it may prove fruitful.’ “That is exactly what I mean,” said I,
“and you have correctly interpreted my line of thought.” After this conversation, which
is here reported exactly as it occurred,* there was nothing else before my mind for
several days, as I was wholly occupied with finding out whether the problem undertaken
was soluble, and, if so, whether it would result in any important Physical Truth. Having
established the fact of high eccentricity as thoroughly as the published orbiis at my dis-
posal would admit, I set about that same day the problem of explaining the cause of
the eccentricities ; and as I worked the impression continued to grow on the mind that
since the stars are not solid, but self-luminous fluid bodies like our sun, and the two
members of a system comparable in mass, the action of each body would produce tides in
the other, and the lagging of-the tides in the two stars would gradually expand and
elongate the orbits as now observed in space. And before I had obtained access to the
learned papers which Darwin had communicated to the Royal Society, or even to his

article ‘‘ Tides” in the Hneyclopedia Britannica, I proved by an elementary process that
when the bodies rotate more rapidly than they revolve, the eccentricity of the orbit would
gradually increase. Here then was a result confirmatory of the happy intuition, and
for the past nine years my energies have been largely devoted to the extension and
generalization of the theory of bodily tides in relation to cosmical evolution.

After concluding my undergraduate studies at the University of Missouri, I con-
tinued the work at the University of Berlin. It is particularly of that work and the
extension which I have since made of it that I shall speak to-night. The theory of tidal
friction developed in the Inaugural Dissertation presented to the Faculty of the Uni-
yersity of Berlin is essentially a special treatment of the general theory as it occurs in
nature, while that previously developed by Darwin in connection with the moon and
planets is restricted by the condition that the perturbing body is very small. I shall
therefore discuss the general case as presented in my own researches.

_ *As the occasion of my beginning this work has never been published, I trust it will not be thought inappropriate
for me to recall it in this paper to the American Philosophical Society.

226 RESULTS OF RECENT RESEARCHES ON THE

Suppose we denote an element of the mass of a spheroid by m, and its distance from
the axis of rotation by d; then the moment of inertia is

IL = Sa

If the spheroid be rotating with an angular velocity y, then Jy will be the moment
of momentum of the body about its axis. For a second body whose moment of inertia is
I’, and angular velocity z, the moment of momentum is /’z.

Foilowing the analogy of Darwin’s procedure, we choose a system of units designed
to simplify the resulting equations. Let us take as the unit of mass

UM.

MM”
and as the unit of length a space 1 such that the moment of inertia of the spheroid about
its axis of rotation shall be equal to the moment of inertia of the two spheroids treated as
material points, about their common centre of inertia when distant apart [. Then
we have

wr)? Mr)?
M | M+ a | + MW ea } = /Lor

I (M+ W))3
ae a |

Let the unit of time be the interval in which one spheroid describes 57°.3 in its
orbital motion about the other when distant I’. In this case, 1 is the orbital angular
velocity of the body. The generalization of Kepler’s law gives

G7 i= Gh-— i). and

9 = {Pore my:
ine pe (ILM)?
Now suppose the two stars to revolve about their common centre of inertia in a cir-
cular orbit, with an angular velocity ©, when the radius vector is p. Then the orbital

moment of momentum is

a eal OM, 7) 2 rs (rsa) pO

In a circular orbit the law of Kepler gives 0’p? = uw (M + M); and Qo’
= w (M+ IM)? p; andon inserting for Op’ its value, we have u? MMW (M+ M’)~? ¢',

EVOLUTION OF THE STELLAR SYSTEMS. 227

which in special units is p?. Now the total moment of momentum of the system is con-
stant, and is given by

JEL == Joys JPe se flu (UE TEL) seepenpepaecoee coon (1)
The kinetic energy of orbital motion is

7 (, GP LP | oO aa crops ne

The kinetic energ ey of rotation is

i 9

The potential energy of the system is

‘ MM’
p

By adding all these energies together we get the total energy of the system :

E rime oe inl ae
where £ is twice the whole energy.

In the system of special units, J, wJZM/, are equal to unity. If we put = = we
shall get

H=y+k?

Let ¢ = ©, and then O* = 63, x = 9}, and we have finally

If we suppose the two stars to turn on their axes in the same time in which they
revolve in their orbits, so that they show always one face to each other, the motion of the
system will be as if the masses were rigidly connected. ‘This condition is given by

OF =)

l|
&
o
Lard

Oke th == 2 = ee OL

ys Moa a aeialitns is. Beste sb hed mea ehk ees, (3)

228 RESULTS OF RECENT RESEARCHES ON THE
Accordingly we haye the system of fundamental equations :

HH = y => ke => 2, plane of momentum, |

1 : !
Pye pe at AA ee
= y¥ + kz — —, surface of energy, r Me ed (4).
9 9 a . 878 1
oy = laws = il, quae OF mechiny. J

These equations represent all possible interactions of the system, but in their present
form are very difficult to interpret. The general problem to which they give rise
seems to be insoluble, but we can solve and interpret them fully for one particular
case which is in close accord with the conditions existing in nature; and it is possible to
show by analogy that all other cases will be essentially similar to the one of which we
shall treat.

By taking the case of two equal stars rotating in the same direction with equal
angular velocities, or substituting (3) of (4) in (1) of (4), we reduce the plaiie of momentum
to a particular line of that plane :

ae — He + (14 k) = a — Ae’ + 2 = 0, since & = 1.

The equation of the energy surface passes into the form
fo)

see al
Go

The curve of rigidity becomes

n = 24, wheren = Vy 4 #2

Every point in the plane of momentum represents one configuration of the system, 2. é.,
one distance apart, one velocity of axial rotation, one moment of momentum of orbital
motion. This point therefore determines the dynamic condition of the system, and by the
motion of this point we may discover the changes which are taking place in any case that
may be imagined. As we have restricted the plane of momentum to one line, the guiding
point representing the configuration of the system will simply glide back and forth along
this line. In the same manner the surface of energy is now restricted to a curve formed
by cutting that surface by a certain plane; the guiding point that would slide along
the energy surface is thus restricted to one line of the surface given by the transformed
equation. [The reader who may desire to examine this question exhaustively must be
referred to my Inaugural Dissertation, Die Hntwickelung des Doppelsternsysteme, Berlin,
1893, R. Friedliinder & Sohn. |

As the tides raised in the stars are subjected to frictional resistance, energy 1s

EVOLUTION OF THE STELLAR SYSTEMS. 229

thereby converted into heat, and lost by radiation into surrounding space; thus the total
energy of the system must decrease with the time. Hence it follows that, however the
system be started, the guiding point representing the configuration of the system must
slide down a slope of the energy curve. In the accompanying illustration the curves are
drawn for the value of /Z = 4.

If the guiding point is set at @ it may move either of two ways: it may slide down
the slope ac, im which case the stars fall together ; or it may slide down the long slope
ab, in which ease the stars recede from each other under the influence of tidal friction.
This latter case is the one of chief interest in respect to systems actually existing in
space, and the several other ideal cases need not be discussed in this paper. The con-
dition at ais dynamically unstable, and corresponds to that of the system at the instant
when the stars are first separated. At this juncture they rotate as a rigid system, but as
each is losing energy by radiation, the axial velocities will soon surpass the velocity of

orbital motion, and then the tides will begin to lag, and the mutual reaction of the stars
will drive them asunder. Thus the guiding point in general slides down the slope ad.
This means that as the stars recede from each other, the period of revolution for a long
time surpasses that of axial rotation, but that in time the two periods again become
synchronous when the guiding point has reached the minimum of energy at 4, where the
bodies once more revolve as if rigidly connected.

The question now arises with respect to the changes of the eccentricity. The
differential equation for the change of the eccentricity is shown to be

ff IT

; = HB
8a np =
(@ @) ge Ese

uU—
— arcan ——

ical 1a) sone \ aidan debates (5)
J

’ ih eee —$—__——__—
= i eapea
Roe

where B is an arbitrary constant; a, 0, a + (, are the roots of the biquadratic equation.
a’ — He + 2=0. Equation (5) is illustrated in the lower part of the preceding
figure, the origin being shifted downward to 0’ to prevent confusion of too many curves.
in one diagram. Now as the guiding point on the energy curve slides down the slope
ab, the eccentricity at first very slightly decreases, then increases slowly, finally much
more rapidly, until a high maximum is reached, after which it again diminishes, owing
to the libratory motion in the system. Thus it is clear that as the stars recede from

each other, the orbit becomes highly eccentric, but will ultimately become circular when

230 RESULTS OF RECENT RESEARCHES ON THE

the system revolves as a rigid body. This last condition cannot come about while the
stars are still contracting and shining by their own light, and hence all visible systems
are characterized by highly eccentric orbits.

To leave no doubt that tidal friction is a sufficient cause to account for the elongation
of the orbits of the double stars, I applied the theory to a special case, in which the
masses, distances and velocities are known. Taking two spheroidal fluid masses each
three times as large as the sun, expanded to fill the orbit of Jupiter, and set revolving in
an orbit of 0.1 eccentricity at a mean distance of 30 astronomical units, I find that by
tidal friction the major axis of the orbit will be increased to 48 astronomical units, while
the eccentricity will rise to 0.57. In this problem the masses are set rotating at such a
rate as will produce an oblateness of about 2, so that the equilibrium is stable. Different
conditions will produce different results, but it is easy to see by this numerical example
that tidal friction is a sufficient cause to account for the observed elongation of the
orbits of double stars.

Though it may be supposed that there could be little doubt of the generality
of the law of the eccentricity which I inferred in 1888, yet the importance of this
fundamental fact of the universe is so great that I did not feel satisfied till all the obser-
vations of double stars had been examined anew and this conclusion touching the
eccentricity established upon the most unshakable foundation. At length I have been
enabled to show by the most exhaustive investigation of stellar orbits ever attempted, that
the most probable eccentricity is 0.48; while on the other hand extremely eccentric and
extremely circular orbits are equally rare, and must be referred to some unusual cireum-
stances. Thus of the 40 orbits now well-known, it turns out that none lie between the
eccentricities 0.0 and 0.1; two between 0.1 and 0.2; four between 0.2 and 0.3; eight
between 0.5 and 0.4; nine between 0.4 and 0.5; nine between 0.5 and 0.6; two between
0.6 and 0.7; four between 0.7 and 0.8; two between 0.8 and 0.9, and none between 0.9
and 1.0. It follows therefore that by whatever process the stars developed, their orbits
assumed a form which is about a mean between the nearly circular orbits of the planets
and the extremely elongated orbits of the periodic comets.

Now a double star can originate by but one of two processes: either such a system
is the outgrowth of the breaking up of a common nebula, or it is made up of separate
stars brought together in a manner analogous to that involved in the capture of a
comet. That these systems are not the outgrowth of accidental approach of separate
stars we may at once affirm; for if we suppose them to be so produced, there being
no third disturbing body which acts like the sun in the capture of comets, the
captured star would recede to a distance equal to that from whence it came. In that

eyent we should observe stars moving in paths of very immense extent, and consequently

EVOLUTION OF THE STELLAR SYSTEMS. 231

revolving at the quickest in some hundreds of thousands of years. If the paths be
elliptical, the major axes of these ellipses would be of the same order of magnitude as
the distance which separates us from a Centauri; while if the paths be parabolic or
hyperbolic, the two objects would pass and then separate forever. On the other hand we
can conceive of nothing which could diminish the dimensions of a very long ellipse,
unless it be something analogous to a resisting medium. Such a medium to be effective
in reducing the size of the orbits would have to act for a great period of time, and
besides would probably be visible in space as diffused nebulosity. No nebulosity is
observed about revolving double stars, nor is there any evidence of a sensible resisting
medium either among the stars or in our own solar system. We may therefore reject
the idea that the dimensions of the orbits were originally very large, and have since been
diminished. As the orbits are now of the size of those of our greater planets, and there-
fore comparatively small, it follows that the stellar systems have originated by some
process other than by the union of separate stars.

As a nebula is a very rare and expanded mass, and is yet held in equilibrium by
the pressure and attraction of its parts, it necessarily rotates very slowly ; and hence
when it divides into two parts under the acceleration of rotation due to secular condensa-
tion, the orbit pursued by the detached mass must be of small eccentricity. For even if
the forces producing separation could be exerted suddenly to produce a violent rupture,
the detached mass in pursuing its eccentric orbit would again come to periastron, where it
would encounter resistance in its orbital motion, and the result of the grazing collision
would be a diminution of the size of the orbit, and consequently an exaggeration of the
resistance at the next periastron passage; in this way the system would very soon
degenerate into one mass. On the other hand were the initial eccentricity small, the
newly-divided masses would pass freely, and when the orbit eventually became highly
eccentric the secular contraction in the size of the masses would prevent disturbance at
periastron. Subsequent collision could not possibly occur, because the periastron distance
would steadily though perhaps only slowly increase as the stars are pushed asunder and
the orbit is rendered constantly more and more eccentric.

It follows therefore that in the beginning the orbits are only slightly eccentric, and
that the eccentricity is developed gradually as the result of secular tidal friction working
through immense ages. Accordingly in the elongation of the orbits now observed we
see the trace of a cause which has been working for millions of years. The existence of
this cause and its effects on stellar cosmogony could probably never be inferred except
in the manner by which I approached the problem. On the one hand it appears that
we have inferred the true cause of the expansion and elongation of the stellar orbits,
while on the other the trace left by this cause has enabled us to detect the existence of

A, BsS:—-VOW. XUX. 2D:

232 RESULTS OF RECENT RESEARCHES ON THE

unseen tides in every part of the heavens. In a fluid universe tides necessarily result
from gravitation, and are as universal as this great law of nature. In my later researches
I have therefore been much concerned to show from the discussion of reliable observations
that gravitation is really universal* and consequently that the tides we have assumed
actually exist in the bodies of the stars. It is thus made certain that the foundation upon
which our cosmogonie speculation rests is as enduring as the Newtonian theory itself.

We now come to the second part of the problem: By what process did the stars
separate ? In college lectures I had heard the annular theory of Laplace expounded for
the solar system, and yet I failed to see how this theory could account for the separation
of equal or comparable masses, such as we observe among the stars. Realizing that
the double stars are in fact made up of two bodies of comparable mass, I reached the
conclusion while still at the Missouri University that there must exist some process by
which a nebula divides into equal or comparable parts, in a manner analogous to that
of fission among the protozoa. About November, 1889, very soon after I entered upon
my studies at the University of Berlin, I found that Darwin had recently published an
important mathematical paper on the figures of equilibrium of rotating masses of fluid,
and had referred therein to the profound work of Poincaré. published about a year
before. When I beheld the figures of equilibrium which these mathematicians had com-
puted, I recognized at once the cosmical process I had already assumed to exist; it
was indeed a great satisfaction to see a demonstration that under gravitational contrac-
tion homogeneous incompressible fluid masses may divide into equal or comparable
parts. The next question was: Are there nebulee of this form in the actual universe?
In searching over the paper of Sir John Herschel in the Philosophical Transactions
for 1833, I found some drawings of double nebule almost exactly like the figures
mathematically determined by Darwin and Poincaré. It was no longer possible to doubt
that the real process of double-star genesis had been discovered. Further investigation
and reflection haye confirmed this inference, and I believe we may now accept with
entire confidence the result reached at Berlin in November, 1889.

In the first investigation Poincaré begins with the Jacobian ellipsoid of three unequal
axes, and imagines it shrinking in such a way as to remain homogeneous, and yet gain
constantly in velocity of axial rotation. When the oblateness has become about 2 he
finds that the equilibrium in this form becomes unstable, and the mass tends to become
a dumb-bell with unequal bulbs—an unsymmetrical pear-shaped figure which I have
called the Anioid. As the contraction continues the whole evidently ruptures into two
comparable masses, and the smaller will then revolve orbitally about the larger. If

* RESEARCHES ON THE EVOLUTION OF THE STELLAR SystmMs, Vol. I: On the Universality of the Law of Grav-
tution and on the Orbits and General Characteristics of Binary Stars (Tue Nichols Press, Lynn, Mass., 1896).

EVOLUTION OF THE STELLAR SYSTEMS. LUBY

we suppose either mass to contract still further, it is evident that the rotation will begin
to exceed the orbital motion ; and the tides raised in either mass by the attraction of the
other will lag, and tidal friction will henceforth play just the part we have already
described.

Starting from a different point of view, Darwin was already at work on essentially
the same problem when Poincaré’s paper appeared, and he held his results back for
nearly a year longer, hoping to make application of the principle Poincaré had
announced. In this second method of treatment two masses of homogeneous fluid were
brought so close together that the tidal distortions of their figures caused them to coalesce
into one mass; set in motion asa rigid system, the problem was to find the resulting
figure of equilibrium. It turned out to be a dumb-bell with equal or unequal bulbs
according to the relations of the primitive masses. Thus we see it proved from two

The Apoid of Poincaré, showing how a rotating mass of

fluid separates into two unequal parts.

independent points of view that a division such as I assumed in 1888 can theoretically
take place ; and among actual nebule of space such division seems to be a general law.
During the years of 1896 and 1897, I have examined a number of such objects in the
southern hemisphere, and find them substantially as drawn by Herschel many years ago.
Burnham and Barnard had previously assured me that the interpretation of the figures
of double nebulse based on the drawings of Herschel was in accord with the phenomena
of nature, but the studies more recently made with the great Lowell telescope supple-
ments their large experience in a very happy manner, and may be said to remove the
last doubt that could attach to the division of nebule by the process of fission.

Before concluding these remarks it ought to be pointed out that in space we have
to deal with masses which are not homogeneous, nor are the nebule by any means

incompressible ; yet many considerations lead us to believe that in most cases the density of

23: RESULTS OF RECENT RESEARCHES ON THE

a nebula is not very heterogeneous, and hence in general the foregoing conclusions would
not be greatly modified. In this reasoning I have assumed nothing but that the nebule
are figures of equilibrium under the action of gravitation. That these masses are fluid
is certain, for the bright lines of their spectra indicate that they are self-luminous gas ; on
the other hand the same force which controls the motions of the stars must operate
among the particles of the nebule, and thus determine the figures of the masses in accord-
ance with the laws of mechanics.

As the conditions here assumed certainly exist in the heavens, we need only add
that when the masses separate they are probably revolving as a rigid system. When
they contract under the influence of gravitation, they must by a well-known mechanical
law gain in velocity of axial rotation, and tidal friction then begins expanding and
elongating the orbits; in the course of some millions of years we have a double star like
a Centauri or 70 Ophiuchi.

The stellar cosmogony here suggested may be regarded as a very general theory.
Our solar system is so remarkable, that it is uncertain whether a theory which explains
the formation of double stars could assign also the cosmogonic processes which have given
birth to the planets and satellites. The masses of the planets are very small compared to
that of the sun, and the masses of the satellites are equally insignificant compared to
those of the planets about which they revolve. Moreover the orbits are very circular,
and these various circumstances make our system absolutely unique in the known crea-
tion. Yet so far as our researches on the double stars may illuminate the problem of
planetary cosmogony, they indicate that the separation took place in the form of lumpy
or globular masses—not in rings or broad zones of vapor such as Laplace supposed.

From the survey thus hastily made of a very large subject, it appears that we have
taken a step in the generalization of the theory of tides and of tidal friction, and have
indicated the probable mode of formation of the stellar systems. Little or nothing 1s
known of the development or even of the mechanism of star clusters; the problem of
explaining the more complicated systems must ultimately occupy the attention of
astronomers if we are ever to trace the development of the visible universe. As a step
in the direction of accounting for the origin of multiple systems, it may be said that
observations on triple and quadruple stars have shown that they, too, developed by repeti-
tion of the fission process. One or both components of a binary have again subdivided,
just as I inferred was the case when still at the Missouri State University in 1888. While
the views here expressed are the results at which I have arrived after a partial investiga-
tion of the theory of tides and of the figures of equilibrium of rotating masses of fluid
and a comparison of these theories with the phenomena observed in the heavens, I

reserve the right to modify any opinion or conclusion which future research may show

EVOLUTION OF THE STELLAR SYSTEMS. 235

to be unsound or incomplete. That tidal oscillations which were first noticed by the
navigators of our seas are at length seen to be but special phenomena of a general law
operating throughout the universe is alike honorable and gratifying to the human mind.
It is equally inspiring to recall that by the known laws of these phenomena we are
enabled to trace existing systems through immeasurable time, and thus disclose cosmical
history which mortal eye could never witness. In our time it is no longer sufficient to
maintain the traditions of the past, to trace the planets, satellites and comets through
centuries, and explain observed anomalies in their figures, attractions and orbital motions
by the law of gravitation. We-must essay to discover the cosmical processes by which
the existing order of things has come about. Though it seems probable that a fair begin-
ning on this problem has already been made, a much greater work remains to be done
during this and the coming century.

What is needed is a more thorough exploration of the face of the heavens, by
astronomers who are familiar with the laws of mechanics; and a far-reaching investiga-
tion of the general theory of tides in viscous liquid and gaseous masses such as the stars
and nebule of remote space. Even if the full extent of the hopes here expressed can be
realized only after the lapse of several centuries, I venture to believe that the achievement
will not be unworthy of the past history of Physical Astronomy.

ARTICLE V.

ON THE GLOSSOPHAGIN

(Plates VI-XV_)
BY HARRISON ALLEN, M.D.

Read before the American Philosophical Society, January 21, 1898.

Having an impression that the genera of bats are best defined by minute characters
in the skull, teeth and wing membranes, I am led to review the Glossophaginee—a sub-
family of the Phyllostomidide, concerning which unsatisfactory accounts exist both as
to structure and relationship.

The bats embraced in the group are characterized by a slender protrusile tongue, an
elongated jaw and a deeply cleft lower lip.* The temporal impression is faintly marked
and the sagitta is absent or confined to the frontal bone. The thumb and forearm are
‘long. The olecranon lies on the upper side of the wing membrane. The canine teeth
are long and the upper molars without hypocone. ‘The incisors are so diminutive as to
permit the tongue to be freely projected without wide separation of the jaws.

According to P. Osborne (Proc. Zodl. Soc., 1865, 82) the thumb aids in the seizure
of small fruits, the teeth tear through the skin and the long tongue extracts the semi-fluid
contents. As in the Edentata, the elongation of the jaws and tongue has led to the sim-
plification of the teeth. But reduction in number of the teeth has gone on scarcely at
all; indeed, the most highly specialized forms are those having the largest number of
teeth.

The genera are arranged in three alliances—the glossophagine, the chcernycterine and
the phyllonyeterine. The first is composed of G'lossophaga, Leptonycteris and probably
Monophyllus ; they certainly relate closely to the Vampyri. The second of the highly
specialized and more doubtfully placed group of Chernycteris, Lonchoglossa and Anura,

oe asin as are indebted to Prof. W. Peters (If. B. Akad., Berlin, 1868), for a revision of the group of the glos-
sophagine bats. The diagnoses are unfortunately sometimes inadequate and without critical analyses of synonymy. The
confusion arising from the circumstance last named is to be acknowledged ; as a result, the task of identification when
not aided by inspection of type specimens is difficult. Dobson in his well-known catalogue of the Chiroptera in the

British Museum, 1878, follows Peters closely—often indeed merely translating or paraphrasing his language—and on the
whole shows less acumen than characterizes his admirable work elsewhere.

238 ON THE GLOSSOPHAGIN &.

is probably also of Vampyrine origin. The third division contains but a single genus,
viz., Phyllonycteris. It is so near Brachyphylla that it would be easy to effect the
transition and remove the genus to the alliance expressed by the term brachyphylline.
It is akin, therefore, if not annectant, to the subfamily Stenodermine.*

The material available for the study just completed was not large, and two genera,
namely, Monophyllus and Glossonycteris, 1 have not seen. I have concluded from the
published descriptions of G'lossonycteris that doubts can be frankly expressed concerning
the validity of this genus. Perhaps not enough stress has been laid upon the effects
of age in attempting to separate it from Anura.

Reliable characters are found in the lower molars. The extension forward of the
ridge (anterior commissure) between the protoconid and the paraconid is more marked
than in any other group, and is in consonance with the compression of the crowns. The
ridge is not spinose, and is scarcely raised. In Glossophaga the ridge is constantly as in
the Vampyri, but in the other genera it is an extension forward from the protoconid.
No trace of hypocone is seen in the upper molars.

The row of glands lying to the outer side of the nostril is discernible in all genera
except Phyllonycteris. Minute distinctions are found in the degree of development of
these glands. They are best developed in the glossophagine group, and least so in the
cheernycterine. In Phyllonycteris the ecto-nareal gland-row is occupied by a flattened
fold of skin which becomes incorporated with the nose leaf.+

The proportions of the width of the third and fourth digital interspaces taken at the
distal ends of the metacarpal bones when the wing is extended is found to be as valuable
an aid in determining affinities as elsewhere in the order. In like manner the shapes of
the terminal cartilages of the fourth and fifth digits, the arrangements of muscles and
nerve markings of the wing membrane are noted as furnishing excellent characters.

The following scheme of interdigital diameters is given :

Second Third Fourth Second Third Fourth

Interspace. Tnterspace. Interspace. Interspace. Interspace. Interspace.
Glossophaga soricina .....- Q 12 17 = Lonchoglossa.. cog 16 23
Glossophaga truei........-.. 2 11 15 JAGR acconscqcon0nc0ss9008002 3 15 30
Eeptony Ctr is ...-.eseeeeeee 3 15 25 Phyllonycterts ....+++...... 3 13 25
Cher ony cteris........-000+ 2 11 20

Enough can be gleaned in the way of inductions from the shapes of the anterior

* In a paper by myself, entitled ‘‘On Ametrida minor” (Proc. Bost. N. Hist. Soc., 1892), I used inadvertently the
term Stenodermatide for this subfamily.

+ The genera of the remote megaderminine genera are in like manner distinguished by characters in rows of glands
as contrasted to folds of skin, though the structures are here not ectonareal, but infranareal. In Megaderma the glands
are distinct, while in Lyroderma and Lavia they ave supplanted by a skin-fold which becomes an integral part of the
nose leaf,

ON THE GLOSSOPHAGIN &. 209

extremities and the details in the phalanges and terminal cartilages to warrant the intro-
duction at this place of a few remarks on the subject of flight.

Leptonycteris. The greatest restriction in the movements of the digits is found in
Leptonycteris. The sharp flexure of the second row of the phalanges on the first impede
rapidity of flight, while the axially disposed, terete terminal cartilages show absence of
strain. The second and third metacarpals always maintain an acute angle to the forearm.

Glossophaga and Chernycteris. These genera resemble Leptonycteris, differing
therefrom in degree only in the greater degree of interphalangeal flexure and in the
angulation of the second and third digits to the forearm.

Anura shows scarcely any tendency to flexure or angulation of the parts above
named while the terminal cartilages of the third and fourth digits are markedly deyiated
from the axial positions and thus appear to correlate with increase of wing strain.

Lonchoglossa is intermediate between Anwra and the preceding group.

Phyllonycteris shows an isolated position from the foregoing group as a whole, on
account of the terminal cartilage of the fifth digit being entirely embraced by the wing
membrane. It is a curious circumstance that the remote Leptonycteris exhibits a similar
peculiarity.

It cannot escape notice in studying the group that the extraction of soft pulp from
a fruit is not unlike the lapping of blood. Acquirements apparently so diverse as
fruit-eating and blood-taking are not so improbable as they might appear to be at
first sight. Geoffroy, who established Glossophaga, yet who had no knowledge of the
habits of the species, concluded from the structure of the tongue that the animal was a
blood-sucker.* In adapting the head so as to create a blood-lapping from a pulp-
extracting form the greatly elongated jaws are shortened, the face flattened, and the
teeth become knife-like. In this manner we may trace the transitions which have taken
place in the Vampyri in creating on one hand the Glossophagine and on the other hand
the Desmodine.

In Glossophaga the Flexor carpi radialis passes along the upper border of the radius
as far as the distal third, at which point it crosses the curved radius to reach the carpus.
In Chernycteris and Lonchoglossa the tendon of this muscle lies to the lower border of
the nearly straight radius.

The Flexor sublima digitorum has the weakest development in Chernycteris,
which form it supplies the first and fourth digits only. In Phyllonycteris it omits only
the second, while in Lonchoglossa and Glossophaga it supplies all the digits.

* The stomach in the Glossophaga villosa Rengger (Naturgesch. der Sdugcthiere von Paraguay, Basel, 1830, 80) was

found to contain blood with remains of insects. Jt is not known what forms would now be included under this title.
See remarks on Ania.

Ay B &=VOm, SUK, YD.

240, ON THE GLOSSOPHAGINE.

The origin of the Glossophagine is easily traceable to the group denominated by
Peters the Vampyri. But the division between the genera composing the Vampyri is
of a character to suggest two groupings at least, and the term Vampyri is best used in a
restricted sense. Indeed, it is a small cluster of four genera only ( Vampyrus, Macrotus,
Schizostoma and the aberrant Hemiderma), which possess a large, triangular, first upper
premolar and an inflated, weak periotic region. .

Of the second group (Phyllostomi), of which Phyllostoma is the type, I have imper-

fect knowledge—haying studied besides this form the genera Lonchorhina and Lophostoma.

But they agree in having the first upper premolar small and acicular, a peculiarity I find
figured in Geryais (Hxp. du Sud.) as characteristic of Tylostoma and Monophyllum
(Dolichophyllum). 1 infer that Trachyops, Phylloderma and Mimon are members of this
group from Dobson’s statement (Br. Cat. Chir.) that they resemble Phyllostoma.. I have
no satisfactory knowledge of the periotic region in this group, but can say that it is boldly
defined, concave, and not inflated in Phyllostoma, Lonchorhina and Lophostoma.

Now it has been seen that the Glossophagine yield two groups—that of the Glosso-
phagi and that of the Lonchoglossi. In my judgment these do not haye a common origin.
The Glossophagi agree with the Vampyri as above restricted in the shape of the first
upper premolar and the inflated periotic region, while the Lonchoglossi are much nearer
the Phyllostomi. Chenycteris possesses a triangular premolar (with large denticles)
and a moderately truncate concaye periotic region, but its other characters, taken as a
whole, connect the form intimately with the Glossophagi.

The taxonomic value of the terminal cartilage can be determined only by the
examination of extended series. At first I had inferred that the shapes of the cartilages
of the fourth and fifth digits were of considerable value. But inspection of the largest
number of individuals of the most common species—namely, Glossophaga soricina—gave
me an impression that they were really variable structures ; thus in one individual from
Costa Rica they were both spatulate ; in another from Bahama Islands they were both
aciculate ; and yet in a third specimen from the last-named locality the fourth digit was
spatulate and the fifth aciculate. Nevertheless the variability itself is of interest and I
have, therefore, figured the cartilages, believing that after extended observation they may
assist In more firmly defining the minor groups of species than is now the case.

GLOSSOPHAGA.

Upper incisors in a continuous row. Length of forearm not exceeding 56 mm.; thumb,
8 mm.; calear present; the tail is short with free tip on the dorsum of the interfemoral
membrane. Proencephalon creates an eminence on brain case ; fronto-maxillary inflation

conspicuous ; mastoid process small.

Dental formula: 1. +— ¢. +— prm. 4—m., $= 21.

ON THE GLOSSOPHAGIN®. ZAI

The Mexor profundus digitorum supplies second and third digits oniy. The
Semimembranosus and Biceps femoris are absent. The tendons of the Gracilis and Semi-
tendinosis closely approximate and give the appearance of being fused, but by gentle
traction they can be shown to be distinct.

Pallas first described Glossophaga soricina as haying no tail (Mise. Zodlog., 1766,
48), the type beimg a female. He subsequently described and measured a second speci-
men (Spicil. Zool, III, 1767, 24), a male, which he dissected. He now noted the
presence of a short tail and figured the skeleton in which the tail is plainly
seen. Geoffroy accepted the first description as final, and proposed a separate name
(G. amplexicaudata) for the assumed new species possessing a tail. Gray (Ann. and Mag.,
N.58., 1858, IT, 490) acting on these erroneous premises proposed the name Phyllophora
for Glossophaga amplexicaudata. Geryais (Expn. Amerique du Sud., 1855, 11, mem., 40)
sustains Gray’s position without comment. Peters set the matter to rights in 1868, over

~a hundred years after Pallas’ first simple error of observation.

Of the elaborate measurements of Pallas those taken of the male are the most accu-
rate and include those of the skeleton as well. The figure of the head by Geoffroy also
conforths in vertical measurement. The width of the basal part of the nose leaf is less
than in our figure. Pallas, Geoffroy and Spix all accurately figure the interfemoral
membrane as approaching the ankle, certainly reaching a point below the level of the
middle of the tibia, which is the distance given by Dobson.

The fact that the two forms of Glossophaga differ so widely makes it desirable that
the characters of the first recorded species be carefully noted. A review of the original
description of Pallas is of restricted value, other than the anatomy of the soft parts,
notwithstanding the praise Geoffroy and Dobson award it. Geoffroy states he had dis-
sected an alcoholic specimen and confirmed Pallas’ observations. But Pallas did not note
so conspicuous a fact that in the first digit the metacarpal bone is much shorter than the
combined lengths of the phalanges. The cranial and dental outlines are worthless ;* but
one cannot gainsay the value of the figure of the fimbriated and elongated tongue.

evokes tats @
Synoptical Table of Genera.

Palatal portion of premaxilla forming a rostrum in advance of median incisive foramen;

gland mass confined to sides of nose leaf; occipito-squamosal suture without foramen;

tympanic bulla separated from postglenoid process by a conspicuous interval; ethmoid

;
} bone convex in brain case; no ectopterygoid lamina; in third to fifth digits first
| phalanx smaller than second; fimbriz: not confined to tip, but extending well back
L

along the tongue.

| Glossophagina vera.

* Gervais (1. c.) believes the form is not Glossophaga at all, but Hemidernu.

242 ON THE GLOSSOPHAGIN®.

a. Median upper incisors larger than lateral; premolars 2; crown of lower canine
with base lying inside position of lateral incisor; median incisor foramen
barely in advance of paired foramina; upper incisors inclined; pit over
proximal third of face vertex.

b. Upper incisors in continuous row; molars 3; thumb one-fourth the

length of forearm (31-34 mM. )....<.- 2.0... .2..cee-seescusecucccnescensscneneeans Glossophaga.
b. Upper incisors with wide interval between centrals; molars 2; thumb
one-sixth the length of forearm (45 mm.)..-.--+-+sseeeeeeeeeeeees Leptonycteris.

£; crown of lower canine

a’. Median upper incisors smaller than lateral; premolars
with base not lying inside position of lateral incisor; median incisor fora-
men well in advance of paired foramina ; upper incisors vertical.
e. Lower canine compressed, with cingulum; metacarpal bone of
thumb exceeds length of phalanges.
d. No phalanx to second digit of manus; premolars #; tail
present; thumb one-seventh the length of forearm
(AD aa, )) coscocmos ooocoasenocotosacoSeconUncoodISCoROASEnoSoACoSADCSOSS Charnycteris.
c’. Lower canine rotund, no cingulum; metacarpal of thumb equal
length of phalanges.
d'. Phalanx to second digit of manus; tail present; thumb
one-eighth the length of forearm (38 mm.).......-....-...-+5 Lonchoglossa.
d''. No phalanx to second digit of manus; no tail; thumb

one-sixth the length of forearm....-..--.-+.-.:01.eseeeeseeseen Armura.

Palatal portion of premaxilla not rostrum-like; gland mass crosses muzzle back of nose leat ;

tympanic bulla almost touches postglenoid process; occipito-squamosal suture with large
Tul. foramen; ethmoid bone not convex in brain case; an ectopterygoid lamina. In
third to fifth manal digits first and second phalanges equal; premolars 3; molars }; fim-

brize of tongue at tip only.

|
|
4
|
L

Glossophagina aberrantia.

Tail present; exceeding short interfemoral membrane;
thumb one-fourth the length of forearm (45 mm.)..Phyllonycteris.

Glossophaga soricina Pallas.

Auricle emarginate at upper half of the outer border ; internal basal lobe free from
head and indications of basal ridge. Lappet in side of the external basal lobe stout,
pointed. Wing membrane from ankle. Terminal cartilage, fourth digit spatulate.
Rudiment of an ascending process from the zygoma.

Auricle subrounded, internal basal lobe with suggestion of vertical ridge, outer
margin of auricle sinuate ; external basal lobe large, obtuse, retroverted, internal lappet
a mere projecting nodule. Tragus straight on inner, convex or obscurely serrate on outer,
margin. The nose leaf hairy and small, midrib confined to the pedicle. The leaf proper
projecting nearly one-half its length above the conspicuous gland mass. The upper lip
as well as the borders of the groove in the upper lip furnished with four to nine minute
warts. Above, the fur is dark, sooty gray, at the tip the remainder of the hair being
lighter but nowhere white. Beneath paler, unicolored. Interfemoral membrane almost

ON THE GLOSSOPHAGIN”. 243

as long as tibia. The calear is one-half the length of the tibia. The interfemoral mem-
brane is often incised rather than semicircular.* The tip of the tail projects from the
free margin of the interfemoral membrane. Tongue on dorsum free from retrose papill.

The first phalanx of the first digit is as long as the metacarpal. Entire digit one-
fourth or nearly one-fourth the length of the forearm (10 to 40, or 8 to 36). The first
phalanx of the second digit is one-thirtieth the length of the metacarpal; the entire
digit is not as long as the third metacarpal. The first phalanx of the third digit is
smaller than the second; the third is flexible; the separation from cartilage tip is
indeterminate. Metatarsi equal. The row of first phalanges of toes equal.

The Skull—The brain case papyraceous; the position of the body and hemispheres
of the cerebellum—the mesencephalon and prosencephalon—hbeing clearly outlined on the
periphery. Pretemporal crests scarcely defined and not continuous with the orbital
margin; mesotemporal not seen ; posttemporal not distinct from the occipital.

The face vertex is flat with shallow median depression over the ethmoid bone. The
convex nasal bones are outlined by grooves, of which the median is the widest and
deepest. Each nasal bone is incised on its free margin at the anterior nasal aperture.
The sides of the face are convex, with a conspicuous, though small fronto-maxillary
inflation. The infraorbital foramen answers in position to the junction of the premolars.
The lateral border of the anterior nasal aperture is produced ; between it and the promi-
nence over the canine tooth a groove is defined. The height of the alveolus is one-third
the width of the neck of the canine, and one-seyenth the vertical diameter of the anterior
nasal aperture. The posterior border of the hard palate near the zygomatic root is
spinose. The palatal notch at the mesopterygoid fossa is acutely incised, carried back to
a line answering to the glenoid notch and is without median spine. It reaches a point
opposite the posterior third of the zygomatic arch. The tip of the pterygoid process lies
opposite the oval foramen. The ascending process of the zygoma is inconspicuous and
rounded. Base of cranium with prominent, median, vomerine ridge. The lateral depres-
sions on the basioccipital are conspicuous, the mastoid process is ebtuse. The tympanic
bone is separated from the postglenoid process by an interval. The coronoid process of
the lower jaw is carried above the level of the condyle and is subacuminate. The angle
is hamular and deflected outward with a notch between it and the lower border of the
masseteric impression and projects backwards slightly beyond the condyloid process.
Symphysis not carinate. The junction of the ethmoid and sphenoid bones in brain case
convex.

The Teeth—TVhe teeth of Glossophaga are the best defined of any of the group.
The cusps are sharp, the incisors and premolars are adapted for cutting, and the molars

* Geoffroy expressed it thus, ‘‘ coupée en angle rentrant,’’ but this shape is often absent.

2A4 ON THE GLOSSOPHAGINA.

for grinding. In the upper jaw, with the exception of an interval on either side of the
canine, all the teeth are contiguous.* In the lower jaw there is no interval on either
side of the canine, for the lateral incisor and the first premolar are in contact with it.
The upper incisors are arranged in a small are, which is smaller than the space between
the canines.

The central incisor is hatchet-shaped, the outer margin concave. The lateral incisor
is smaller than central, with inner border twice the length of the outer. The canine is
coneave on the palatal surface. The premolars are triangular subequal, yet the heel of
the second tooth is twice the size of the first. The cingules are scarcely discernible.
The first molar is subtriangular with W-shaped crown reduced, the fiuting on the para-
conid, rudimental ; the metacone is united to protocone by a ridge. The second molar is
subquadrate, W-pattern scarcely reduced; the fluting on the paracone marked; the
ridge from the metacone not reaching the protocone, but a distinet though narrow valley
intervening. The third molar is one-half the size of the second, the second V being
rudimental. The longitudinal axis of both second and third molar is oblique to axis of
the alyeolar processes. The third molar slightly oyerlaps the second at the buccal
border.

The lower incisors are proyided with flat smooth edges to the crowns and are
adapted to crushing rather than to cutting food. The canine is directed slightly back-
ward and is provided with a small heel. The premolars are triangular, equal, the bases
increasing in thickness from before backward. The molars exhibit marked commissural
extension in advance of protoconid and paraconid. The hypoconid is cuspidate and as
high as metaconid ; all the teeth are much alike, but become progressively smaller and
narrower from the first to the third, while the extension in front of the paraconid and
protoconid become Jess and less marked. The third tooth is not more than two-thirds
the length of the first.

In a skull of an embryo which measured 8mm. long, the lower jaw projected well in
front of the upper and. bore the deciduous canines. The shapes of the incisors and pre-
molars could be discerned, while the upper jaw was edentulous.

Tn an adult which retained the right upper lateral incisor only and the molars were
much worn, the only teeth in the upper jaw that were in contact were the second and
third molars. In the lower jaw the third molar was separated from the tooth both the
first and third. The lower incisors were much worn and placed slightly in adyance of
the lateral teeth. I am inclined to believe these are variations due to advanced age.

* The upper incisors as represented by Leche (Studier ofver Mjolkdentionen och Tindernas Homologier hos Chiroptera,
1876, Tab. If, VIL) do not touch.

ON THE GLOSSOPHAGIN &. 245
Glossophaga true, n. s.

In the Proc. U.S. Nat. Mus., XVIII, No. 1100, 1896, 779, I described a new species of
Glossophaga under the name G. villosa. Since Rengger (J/. ¢., p. 80) described in 1830
a species under this name I have concluded to rename the form, notwithstanding that
the species is quite different from the genus Glossophaga as now restricted. See
remarks under Anura. I take pleasure in dedicating this species to the accomplished
Curator of Mammals of the National Museum, Mr. F. W. True. I herewith reproduce
the description, which now has the advantage of appearing with appropriate figures of
the head, skull and teeth.

It is a remarkable circumstance that the genus Glossophaga, while the most common
of any of the forms embraced in the group of Glossophagi, and has been collected from
he widest range of any of its race, should haye presented degrees of variations so low as
neyer to have permitted the recognition of more than a single species. The complicated
synonymy successfully unraveled by Peters, it is true, contains a number of names of
species, but these were proposed through misapprehension of assumed generic values and
bear no relation to questions of specific distinction.

A eareful study of two specimens (Nos. 9522 and 9525) belonging to the United

coo)

States National Museum has conyinced me of the necessity of recognizing two species of

Glossophaga—namely, Glossophaga soricina and the one which I here name

Glossophaga truei.

Auricle entire on outer border or slightly emarginate. Internal basal lobe bound
down to head without -trace of ridge. Excepting in length of head and trunk eyery-
where smaller than G@. soricina. The ascending process of the zygoma twice the size of
the same part in that species. Wing membrane from distal fourth of tibia. The termi-
nal cartilage of the fourth digit terete.

The auricle is without ridge at base of the internal: basal lobe, which is scarcely
defined and closely bound down to head ; outer margin almost entire; external basal lobe
and nodule inconspicuous. Tragus with trace of serration on outer margin, basal lobe
large, quadrate.

The nose leaf, hairy, without midrib at internarial pedicle, projecting scarcely at all
above the simple gland mass of the upper lip, which it almost entirely occupies. Thumb
one-fourth the length of the forearm

namely, nine to thirty-two. The tail had
evidently occupied a position similar to that seen in G. soricina, It had been remoyed
in preparing the skin,

246 - ON THE GLOSSOPHAGIN A.

Based on skins of two adults: No. 9525, U.S. N. M., La Guayra, Venezuela ;* and
No. 9522, U.S. N. M., co-types.

No. 9525, U.S. N. M., fur soft, shrew-lke; dull ash at basal two-thirds, sooty at
apical third; it extends along the entire length of the dorsifacial region. No. 9522,
U.S. N. M., quite the same, but is dark brown instead of sooty.

The skull + closely resembles that of G. soricina, but is smaller and thinner walled.
The ascending process of the-zygoma is longer and more pointed than in the species just
named; the palatal notch is less acute. The fronto-maxillary inflation is conspicuous.
The symphysis menti is carinate. The angle of the lower jaw projects backward slightly
beyond the line of the condyloid process. The brain case is 12 mm. and the face 7 mm.
long.

The upper central incisors broad with slightly concave cutting edges; the lateral
incisors are narrow with oblique cutting edges. The premolars are slightly separated
from one another and the second premolar from the first molar; they are compressed,
subequal, and triangular ; the second premolar is thickened posteriorly. The other teeth
closely resemble those of G. soricina. ‘The first upper molar is longer than the second
and the second longer than the third; there are no ridges extending from the paracone
to the metacone. The third upper molar does not overlap the second molar at the buccal
border.

The muscle fascicles and nerve markings of the endopatagium disposed as in
G. soricina. This system is the weakest of any of the group of the Glossophagi. The
terminal cartilages are throughout terete. —

On the whole the descriptions of Pallas and of Geoffroy agree well with Glossophaga
soricina of Peters’ revision, and exclude those specimens here embraced under G. true.
In Geoffroy’s figure { the measurements of the nose leaf agree with those of G’. soricina,
but the shape of the tragus and internal basal lobe of the auricle are like those of the
form under consideration. But the figure is evidently based upon a dried specimen.

The isolation of the premolars in G. true: answer fairly well to the arrangement of
the teeth in an old example of G. soricina. This is an interesting fact, inasmuch as it
suggests that senile characters in one species may be the same as those found in young
adult life of another.

The following proportions are noteworthy: The first phalanx of the third digit is
longer than the second. The third metacarpal bone is as long as the forearm. The

*Tt is not certain that the locality here given is the correct one. The record in the National Museum catalogue is
imperfect.

+ In addition to the skull in the type specimens, I possess a skull from Brazil presented by the late Mr. Harte,

which answers to the above description.
{ Ann. du Mus., 1810, XV, Pl, XI.

ON THE GLOSSOPHAGIN ®. 247

forearm is 1.15 mm., the smallest in the group. The calcar is one-third the length of the
tibia. The first phalanx of the first toe extends slightly beyond the first phalangeal joint
of the second toe. The first row of phalanges decreases progressively from the second to

the fifth toe.
Type.—No. 9522, U.S. N. M.*

Measurements of G'lossophaga truet.

Millimeters.

Head and body (from crown of head to base of tail) ..........::cccceessessseeeeeeeeeseeee seseeeaasenenes 45
Eecadvand et rears sense sees sects eere sete eeneae sce ence sae nase Son oe senor ce wate sce soos cocoueee Soaoa0ic09000 32
First digit :

Wengthyof firstimetacarpall DONes...sacce0stsseteoceeecncvssosseee-rccesecncnoceecececeseceecesasessecece 4

en op hvonehastap all axeasme erect eemtcentensrscostreenscecccetessseeeeseeeaceseresceeeceteeeeneeeteres 4
Second digit :

Length of second metacarpal bone coo SS

ILGNYE AN Oi Tah 79) 11 ENGb-<: ceaponosadccoscoooScooccaqdboSHo4Ro5o0e saps sbngcIDEsaee0HoNGekeRD0IINNINITEbOOIHHSOOS 2
Third digit :

Length of third metacarpal DOne.............000.scsse-seseccesescceeccceecccesesceeeteetesecseteneesens 30

Length of first phalanx........ccccsccceecccseseeeeeesceseeceeseenseeee coo» dal

Length of second phalanx 14

JURA ERIN Mie NTE: TAR ETE < ococnconbopcesqoodsasacnoaconoqecddacoasq¢aRcocOsaondanpLoondGUaSUDDOBUOHGoOHObOND 6
Fourth digit :

Length of fourth metacarpal bone 27

Heng thnotsirstgphalanxeeeeteccsemonsecracscasasencserscosen so seeites ect eeteeeteceecsceeceeteiisetenas 9

eng thotsecond yphalanxgeseesnsceeccesoseseassseecsedeecesatee cen oeeeeeeeeecee etree eseerstntes 9
Fifth digit :

idens thy of fiithtmetacarpall ones--c-casn-casccesencacsesccorseecasesscesesmccecceseersseteeereecereee Q7

Length of first phalanx ....... 50060600000000000000 6000000) -BondoNOdan9uDS BeSoDSqSUEHESodeosanNDSSEHAGCBGOD 8

ILEMEHT Ot Ses ~MnG! TATE <onqosccnG BoocaSdonDsoHedodonahnccogsedasonGscadocenccoccadusosdccorASooaCosEO0000 8
Length of head - 21
TEIGHEAING OE GAT oson0n5 coc cenpcoccoDN6o nag dooADDoNsDDODNDOBSoEnOKHODIDESORH SoARtODEaROBOOdOCODHHODdoODDESdDEAaDOODDRGOoS 11
IBIGTGNH Ot TERETE codec000002c0cccog oqo asa coacadobsocoOSobONCdD HB. oDNDSnEsoSSaceoaNaLaoDONSSONCOEDDSUodERDDHoGOIEGGOO0 3
JUGINSIAT, Ci GHDYE pene ooo conaacedégacaebocgoanqnabaccoqDdosddccosHaocaCGocHAOcOSocHSd=cbopoocadssenesnocssna pcocoaHoCoRSeSE 11
Length of £006 ......2......sceceesesceeneeeees mee 8
Length of interfemoral membrane...............ccccecsccneececceececneeeeacecesenscecsseesesseeseenseccauecsess 9

MonopHyYLuws.

Upper incisors not in a continuous row. The first and second upper molars with hypo-
cone. Length of forearm, 37 mm.; length of thumb, 10 mm. The tail projects from the
margin of the short interfemoral membrane. The proencephalon does not create an

eminence on the brain case. No vertical line is found on any of the interdigital spaces.

Dental formula: i. + — ce. +— prm. 2 —m. 3 = 21.

osloo

* The measurements of No. 9523, U. S. N. M., are the same asin No, 9522, U. S. N. M., excepting in the second
phalanx of the third manal digit, which is but 12mm. long.

A. P. S—VOL. XIX. 2F.

248 ON THE GLOSSOPHAGINE.

The single specimen of J/onophyllus which was available was that of a skin of an
adult (No. 83347, 9°, U. 8. N. M.) obtained by exchange from the Berlin Museum. The
genus is in close alliance with Glossophaga—closer, indeed, than any two genera of the
group. The retention of the hypocone in the first and second upper molars, the presence
of a keel on the symphysis of the lower jaw and absence of the vertical line in the inter-
digital spaces, separate the two forms. Other characters if they existed unassisted by
those just named would be those of relation and proportion. The presence or absence

ot the calear could not be determined.

Monophyllus redmani Leach.

Auricle with blunt tip, scarcely emarginate on outer border. Wing membrane from
basal third of the tibia: terminal cartilage of the fourth digit, spatulate. Marked rudi-
ment of ascending process from the zygoma. Nose leaf, upper lip and membrane much as
in Glossophaga truer.

The auricle resembles G. truer nearer than G’. soricina. It is blunt at tip, scarcely
at all concave on the outer margin. <A faint emargination is noted on the inner margin
which may be exaggerated in the dried skin. The external basal lobe was everted by
the method used in preparing the specimen. The parts do not differ from those studied in
Glossophaga. The tragus is blunt, presenting two coarse sinuations at the outer side and
two denticulations at the base. The nose leaf, upper lip and mentum almost precisely
the same as in G. truei. No warts are anywhere present.

Fur above is dark brown; the head, neck and shoulders a lighter shade than the
back of thorax and lom. Examined with a lens, the fur has an admixture of fine gray
hairs, which are more numerous on head, neck and shoulders than elsewhere. The fur
beneath is gray and brown, about equally admixed. Both above and below the hair is
unicolored. Sparse gray hairs extend below on arm to elbow and slightly over the endo-
patagium. The legs are naked.

There is no vertical line on the membrane of any of the interdigital spaces. The
endopatagium exhibits a few coarse vertical lines. The fourth interdigital space is
obscurely areolate.

The skull was mutilated at occiput and posterior third of the base. It closely resem-
bles Glossophaga. The fronto-temporal crest is more defined, while the fronto-maxillary
inflation is less defined than in that genus. The posterior palatine notch, narrow. Seen
from above, the posterior border of the infraorbital foramen appears as a blunt spine. A
narrow but well-defined groove extends the entire length of the face, beginning at a
foramen near the pretemporal ridge. The ascending process from the zygoma is

greatly in excess of the same character in G'lossophaga. The external auditory opening

ON THE GLOSSOPHAGIN &#. 249

is smaller than in the genus just named. The thick skull does not admit of the divisions
of the brain being discerned. The lower jaw is more robust—the depression in advance
of the angle most marked of any genus in the group; the angle is raised high above the
level of the lower border of the high ramus as in the Lobostomina; the symphysis is pro-
vided with a large keel.

On the whole the skull is more robust in texture and is of a larger animal than
Glossophaga, but the face structures more extended, and presumably from the symphysal
modifications, a longer and more prehensile tongue.

The Upper Teeth.—The incisors are not arranged in a continuous row or in pairs, but
intervals* are found between the teeth.

The space between the central incisors is wider than that between these teeth and
the laterals. The central incisors are obscurely hatchet-shaped, while the laterals are
conical. Wide intervals also exist between the canine and the first premolar and between
the first and second premolars. The other upper teeth are contiguous. The premolars
are aciculate, compressed, with prominent base conules. The first and second molars are
quadrate with conspicuous hypocone. The third molar is more triangular and resembles
the first and second molars of Gilossophaga.

The Lower Teeth.—TVhe incisors are reduced to tubercles, arranged in pairs, which are
widely separated both from the symphysis and the canine tooth, though nearer the latter
than the former. The central incisor is larger than the lateral. All the other teeth are
contiguous, except the second and third premolars, which are separated by an interval
equaling that in the upper series. The first premolar is distinctive. It closely resembles
the homologous tooth in Glossophaga and anteriorly overlies the base of the canine. The
second and third premolars are similar to those in the upper jaw. The molars are of the
same type as in Glossophaga, but elongated and compressed in advance of the protocone
- and paracone as in Leptonycteris.

The comparison of the skull and lower jaw seen from in front with Glossophaga is
instructive in the differences in the shapes and relations of the shapes of the teeth already
noted. ‘The upper canines are observed to be longer and more trenchant in Jonophyllus
than in Glossophaga.

Rugee ten in number, the anterior five undivided and the posterior five divided.

Measurements of Monophyllus redmani.

Millimeters.
Head and body (from crown of head to base of tail)...

Length of arm

Length of forearm. ............cccsccee seeeeeseeeecanesseceneeees sadocdosncna9sHs8 2060 sosNanSEcOSHONERaGAASHESETESTOUeC 37

*According to Dobson’s text, the upper incisors are in a continuous row, but they are figured with an interval

‘between the central incisors. In the table of genera all the upper incisors are said to be arranged in pairs.

250. ON THE GLOSSOPHAGIN#.

First digit : Millimeters.
Heng thy offirsimetacanpalupones-nss-0.--e0ssseeseaeereeereee easter cer secer enero reece emt eeeceesene tees 4
ensthiob first, plalantxcdcncsmcc- ses daecens-eectcacceseeseteecter Peaenaeaareeaeeee secon ence oeeceeoeree 6

Second digit:

Length of second metacarpal bone

IUGR OL TESTS TOO Ray <e00300000000000000000000060031050 nAdoacanDSEcneaDEDEoHosacGaNce BODDODABNIECNsFOSsARRDSAC 2
Third digit:
Wenethvor third metacarpal POuesee--emsceseeeecressossesees ese sche -eceeasseeeeieassceessteeeseeeee 38
Meng thvorsinstyplaalankcesmscsechessescaseseseceeeeseceet eter cess eeeeeeen =oqon000ndanaans0000000000 13
Meng thronsecondgphalanxererecrerchen-teescmertcceecstesesteetersseeacereesseeeerseeeeeeeee eee eeeeeseners 19
Weng thvoimihindsphalansceccesrecreoeasteeseetea cease ceeseceen eee esesseeeeete sees eee eeeee eee ease eee 9
Fourth digit:
enethiotstourthymetacanpalybonescse--eceastmereeeeee ase sees seeee er eeeeeseee esac eeeee eee eecere 35
IUGR Oe ISG TOMATO apsoconc00809 cooanonoobconnsDqDanBGONoNDHSHs5 AcHBaAHSE SHOAGHanODASSOBOEEAS 3000000000 8
Mmenothvorsecondephalanixeamereseessermere teeter sascha tects corer erect eee sseech ese eeeeeenteeertsrens 12
Fifth digit: :
Length of fifth metacarpal Done.........-:.csseccseececeeeseeseeseeeeseercesersasecneseneesecessscneserass 30
ILE Or THESE 7) OTT ocosc000G000200 non9s0n.0nG0 So CocaDDapDNFDODOnODAnDCoGEONDanODANSSHONSBNECOKODANDSAADDS 9
engthiofssecondsphalanxe.n.cccreccrsccstasdsescesenches seivenc ce osetieenissneestiseen res shes = seaececennenenss 10
ILEIRAD OF WSC l-ocoooganoqdcososconncoandonsooc0sq5c0bn99 DEC otaDDSOUaTS>NRIoOSE9ONNDoOSOOEDOSARdRaEDSECCHHODATSbONEsCS od 25
TSIEN DE EBPs sc0s093c9 0090009005 a09n0 qoNoNDaDED ANON O saDDESOOAODACAIDIANEHOSEDaCHONSHOAaDDSHOSDDERISSSRSoBEoRDSADORODER 10
Te (Gr Ne OE HEV FS)590080000005 900065000d90n04000000000000000000 2ooa0 00D ShDSaGRASSHeDGoNUsaDbOadboDERDAAGORAG peteseeeseeee 3
TLS OP MIEN, cooosac00s0900 000008 c00300000 soDdDDODN cOSODDBUDSSSECONDOCHODEHDSCSHODOBRADAADOEGOSE aoocnoDODOIRODORCOONN 11
IL SN GE THlOMocoscoocsa0e aaponno sods 980709903 0cdoNsCDID0009N000 ganoano9nOHONGE DoDOSeDHDScDSEODE. CUasHOURESSSSNBNOEBESCC 15
TURIN CL? TO Foon occ onoss ono ann conDab0Roo nDOSTEODOREDOGOCACHOSSD ConSOoDSODOSSEAbOSizaNSdOBON pabODOnAREAEOOHOBSsEDDeCRN! 11
Length of interfemoral Membrane. .........2..22cceceecccseeceeeceecseeseseccnessnesccecsaneceseeeecseeceeneeeees 4

Length of tail

LEPTONYCTERIS.

Upper central incisors separated by wide interval. Proencephalon not forming an
eminence on the brain case. No spine at upper margin of the anterior nasal aperture
caused by union of the free margins of the nasal bones. Tail none. Second phalanges
of third, fourth and fifth digits sharply flexed on the first.

Dental formula: i. 4—c. +— prm. 3—m. ?= 18.

Leptonycteris nivalis Saussure.

Auricle small, nearly one-half the length of the face, slightly emarginate at basal
half outer border. Internal basal lobe scarcely free; external basal lobe convex, inner
lappet crescentic. Tragus straight on inner, convex on outer side; basal lobe conspicuous.
Nose leaf projects far beyond non-ribbed pedicle. The latter forms a wart-like contour
inferiorly. The upper lip is narrow and provided with two inconspicuous nodules. Car-
tilages at the end of digits are as in Glossophaga. Calcar rudimental, scarcely one-fifth
the length of the tibia.

Tongue furnished on sides and dorsum with minute, hair-like papille. The side of

ON THE GLOSSOPHAGIN#. 251

the mental groove furnished with an obscure row of minute warts and the chin beyond
the groove thickened with gland clumps.

Fur short, villose, longer on neck, above deep ash verging to gray, base white, below
paler. On neck, basal part tawny, but abdomen almost unicolored. The hair is slightly

whiter at pubis. Distal half of humerus (above and below) hairy—the rest of the limbs,

except the base of thumb, second digit and all of dorsum of foot, covered with a sparse
growth of short hair.

The muscle fascicles on wing membrane are much the same as in Phyllonycteris.
They are wide apart generally, but do not extend over so large a field. The reticulated
arrangement of fibres near the forearm is conspicuous. The longitudinal lines in the
third and fourth interspaces distinct. The nerve markings are characteristic. Both arise
from the digits far above the joint, the anterior being at distal third of the fourth meta-
carpal bones.

The terminal cartilage of the fourth digit scarcely spatulate ; that of the fifth digit is
terete and not free. In this respect Leptonycteris resembles the remote Phyllonycteris.
The skin in the second interspace is not pigmented.

The Skull—Skull not papyraceous ; proscencephalon not defined. The pretemporal
crests subtrenchant and form a short, faint conjoined line with its fellow at the sagitta ;
the scarcely discernible mesotemporal depressed, not reaching sagitta; pasttemporal reaching
occipital crest. Face vertex with depression over ethmoid, but the nasal bones are scarcely
defined in median line and not separated at all laterally from the concave sides of the
face. Fronto-maxillary inflation barely discernible and crossed by the orbital ridge.
Alveolar process in height equals one-seventh the width of the neck of the upper canine
and one-twenty-second the vertical diameter of the anterior nasal aperture. The depression
between the lateral margin of the anterior nasal aperture and the root of the canine tooth
much deeper than in Glossophaga soricina. Ascending process of zygoma rudimentary.
The premaxilla weak in advance of the large incisive foramina ; posterior border near the
zygoma root not spinose. The rounded notch at the mesopterygoid fossa midway between
zygoma root and glenoid cavity. Scarcely any difference observed between the level of the
basioccipital and the basisphenoid. The mastoid process acuminate. The tip of the
pterygoid process in advance of the oval foramen. ‘The nasals are incised at the anterior
nasal aperture. The angle of the lower jaw acute, not hamular; it is on the same plane
with the masseteric impression, not separated therefrom inferiorly by a notch, and projects
backward beyond the condyloid process. Symphysis not carinate. The lower border of
the masseteric impression carried in a semi-circular line beyond the horizontal ramus.

The Teeth—Teeth crowded for the most part. Upper incisors as in G'lossophaga
soricina ; the central hatchet-shaped, separated by an interval. The lateral incisors as

252 ON THE GLOSSOPHAGINA.

large or larger than centrals. Canine concave on palatal surface. The first premolar with-
out basal cusp and separated from the canine and the second premolar. The second pre-
molar with basal cusp and in contact with the first premolar. The first molar much larger
than the second, the paracone subtriangular, the outer surface of the paracone and mesacone
are scarcely at all fluted, hence the W-pattern not evident. The second molar without
fluting on the rudimental mesocone, hence the posterior limb of the second V is absent.

The single lower incisor which is seen in the two examples lies in close contact. with
the canine. The canines are large and divergent, projecting to the inner side of the lateral
incisor. The three premolars are triangular with conspicuous cingules; lingual aspect of
the first premolar concave and in contact with the canine; the second free from the first
and the third premolar. The protoconid with a long anterior extension which has the
value of a second functionalized cusp. The paraconid is small and placed slightly back
of the protoconid. The mesoconid is higher than either of the other elements, and
together with. the hypoconid form a low, broad heel. Molars slightly overlapping at
buccal borders ; the metaconid and hypoconid are of great size with wide yalley.

Metatarsi equal ; first row of phalanges decrease progressively from the second to
the fifth.

The measurements of Dobson do not agree in some respects with the three specimens
examined. The thumb is smaller, while the first phalanx of the third finger is much
larger. He states the “tail none or exceedingly short.”

In the cheernycterine alliance the genera Chernycteris, Lonchoglossa and Anura

are placed. They have in common three premolars and three molars in each jaw.*

CH@RNYCTERIS.

Naked skin fold defining nostril laterally. Pterygoid process in contact with tym-
panic bone. No phalanx to second digit. Length of forearm, 42 mm.; thumb, 7 mm.

Dental formula: 1. 4 — c. + — prm. 3 — m. 3 = 22.
Chernycteris mexicana Tschudi.

Auricle subelliptical, emarginate on posterior border; internal basal lobe large,
entirely free from the head and hairy; external basal lobe small, acute; internal lappet
conspicuous. Tragus elliptical; basal lobe simple, deflected backward.+

Interfemoral membrane longer than tibia, semicircular. Calcar half the length of the

* The only other forms possessing the same armament are the remote genera Vespertilio, Cerivoula, Natalus and
Thyroptera.

+ In one specimen the tragus exhibited near the tip two papille seen on both the anterior and posterior borders and
an additional cluster of three on the posterior surface.

ON THE GLOSSOPHAGIN®. 253

tibia; the tip projects slightly beyond the interfemoral membrane ; wing membrane
attached at a point midway on metatarsus. Nose leaf acuminate, sparsely hairy. Inter-
nareal pedicle with midrib ; below two warts at median line in the short lip; outer flange
at the nostril broad, tumid and gland-bearing. The gland mass proper well defined, but
not across the face back of the nose leaf.

Tail two-thirds the length of the femur and appearing free above the interfemoral
membrane. Vibrissee on muzzle very long. Fur everywhere silky. Above, tips dark
brown, the remainder of hair lighter brown. Beneath, lighter in shade, light brown,
unicolored. No. 399, Acad. Nat. Sci., is smaller than the specimen named. The
length of forearm is 33 mm. (about 1/50), and shorter than that assigned Chernyc-
teris minor Peters. ‘The calcaneum, however, is not as long as the foot. The central
incisors are absent in the upper jaw. In other respects the specimen resembles C. mezi-
cana. I do not identify this specimen with C. minor, but regard it as a variation of
C. mexicana.

The Skull.—Skull papyraceous ; the divisions of the cerebellum and cerebrum discern-
ible through the periphery. Temporal ridge almost n7/, not forming union at any part of
the sagitta. Fronto-maxillary inflation absent, but the inner wall of the orbit and the
fronto-nasal depression unite to form a ridge which bears a foramen. Face vertex without
median fronto-nasal pit, but in its place a flat surface which bears a median ridge. No
groove indicating positions of the nasal bones, but the outlines are seen through the
translucent periphery. The sides of the face uniformly convex. The upper border of the
anterior nasal aperture incised. The lateral margins of the anterior nasal aperture
scarcely produced; the groove between them and the eminence over the canine teeth rudi-
mental. The simple infraorbital foramen over the first premolar tooth.

Alveolar process in height one-thirty-first the width of the neck of the canine and
one-thirteenth the vertical diameter of the anterior nasal aperture. Six inconspicuous
ruge. Zygoma incomplete. The infraorbital foramen on same vertical line between the
second and third premolars. Hard palate acutely arched in molar range. The posterior
border near root of zygoma with slightly convex margin; oval foramen well in advance of
the pterygoid free tip which reaches the tympanic bone. The tympanic bone not
reaching the postglenoid process. The palatal bone extends to the anterior lacerated
foramen before forming the large subacuminate notch. Pterygoid process convex out-
ward, forming bulla-like recesses. The mesopterygoid fossa with a faint vomerine ridge
which is continuous with the conspicuous basioccipital ridge. The coracoid process
acute, deflected outward, the angle produced beyond the condyloid process, and con-
tinuous with the depressed lower border of the masseteric impression. Symphysis with
pronounced carination. Brain case, 16 mm, long; face, 14 mm. long; or the face almost

as long as the brain case, ©

254 ON THE GLOSSOPHAGINA.

The Teeth—Wide interval between upper incisors. The central as described by
Dobson, is smaller than the lateral. But in two specimens examined by me the centrals
were larger than the laterals. Both teeth are inconspicuous and scarcely raised above
the gum line. The palatal surface of the slender canine flat. Of the two premolars
present, the first possesses both anterior and posterior cingules and without increase of
width back of the cusp. The second is without posterior cingule, but is widened
back of the cusp. The first molar with paracone extending the entire length of
the tooth, but sloping from before backward. Protocone and’ mesocone without buccal
fluting or palatal ridges. The second molar as the first, but the protocone ends at the
beginning of the mesocone. The third molar as the second much smaller and all parts
rudimental.

The lower incisors deciduous. The slender canine with rudimental lingual cingule
which does not extend beyond the level of the lateral incisor. The first premolar close
to canine with cingule subequal to the cusp. The second and third premolars with cusp
much larger than the prominent cingules. The first molar with protocone and paracone
almost coalesced ; the protocone well advanced. The posterior border of the tooth is
furnished with a prominent cingule apparently developed from the hypocone. The first
molar is separate from the third premolar and the second and third from one another.

Chernycteris exhibits vertical muscle fibres in the endopatagium, the nerve markings
of the interdigital spaces and the shapes of the terminal cartilage of the fourth digit in
a manner quite the same as in G'lossophaga, though the structure last named is less spatu-
late than in that genus.

Measurements.—The first phalanx of the first digit shorter than the metacarpal; no
phalanx is present in the second digit. The metatarsi and the first row of phalanges
equal.

Tongue attached to floor of mouth at the level of the space between the second and
the third molars, or 12 mm. from the symphysis. Penis not pendulous.

ANURA.

Interfemoral membrane hairy ; tail absent; wing membrane attached to midtarsus ;
calcar absent ; no phalanx to second digit; two warts on upper lip; groove in lower lip

wide with many warts. First premolar large remote from canine.
Dental formula: i. + — ¢. + — prm. 3 — m. 3 = 22.

Resemblance to Lonchoglossa very close. The general appearance the same even to
the shape of the terminal cartilages of the phalanges. Skull and number of the teeth
the same. But it is held that the tail, calear and phalanx to the second digit all being

absent, separate Anura from the genus just named,

ON THE GLOSSOPHAGIN®. P55

The first lower premolar possesses a small, anterior, basal cusp and is, therefore, almost
as large as the other premolars. The main cusp throughout scarcely higher than the

basal cusp.

Anura wiedii Peters.

Auricle much the same as in Lonchoglossa. The tip of the tragus is pointed. Nose
leaf simple, acuminate, no depression above nostrils. The gland mass at the side of the
nostril continuous with that extending up to the side of the nose leaf. Upper lip with
two equidistant warts. Fur everywhere long and silky. Above, apical third dark brown,
basal two-thirds Isabella brown. Below, apical third Isabella brown ; basal two-thirds dark
gray. Thus the arrangement of color is boldly contrasted with that of other forms in the
group. Fleshy mass of forearm, the interfemoral membrane, the thigh and the feet
covered with short hair. On the ventral aspect the forearm is covered with fur which
extends thence a short distance on the interfemoral membrane.

The proportions of the wing of Anuwra are those of a larger animal than Loncho-
glossa, though the thumb is of the same size. The lower extremities are almost identi-
cally the same in size, the calcar alone being larger in Lonchoglossa. The absence of the
phalanx has already been noted in Chernycteris. Alliance with this genus is suggested
in the great width of the cleft in the lower lip and in the possession of warts on the
upper lip.

The muscle fascicles and membrane markings are as in Glossophaga, but the
terminal cartilages of the fourth digital interspace while spatulate exhibit the limb on
the somad side greatly prolonged. This character is not seen elsewhere in the group.
The cartilage of the fifth digit while terete is also greatly prolonged on the free margin of
the endopatagium. These characters indicate that there is more strain on the wing
during flight than in any other genus. .

The Skull—Vhe skull is almost identical with that of Lonchoglossa. The alveolar
height is one-third the width of the neck of the canine and one-seventh the vertical dia-
meter of the anterior nasal aperture. The zygoma by careful maceration is shown to be
cartilaginous. A specimen of Lonchoglossa shows the same structure. The skull is
24 mm. long. The brain case is 60 mm. long, and the face 40 mm. The lower border
of the masseteric impression is not produced. Dobson’s figure, Pl. XX VII, Fig. 4, does
not agree in all respects with our example.

In 1830, Rengger (Naturgesch. der Sdugeth. von Paraguay, 80) described a species
of bat under the name Glossophaga villosa. Since Wagner (Suppl. Schreb. Siugeth.)
assigns this forma place under Chernycteris, it is well to state that while G. villosa
Rengger retains three premolars in both jaws, that the tail is absent, the interfemoral]

A. P. S— VOL. XIX. 2G

256 ON THE GLOSSOPHAGIN®.

membrane is but half an inch deep at the rump, and the lateral upper incisors are
smaller than the centrals. The interfemoral membrane is hairy. This species is nearer

Anura in most of its characters than any other genus in the group.

LoNncHOGLOSSA.

Tail short ; wing membrane attached to ankle; calcar present but small, about one-
third the length of the tibia; a phalanx to second digit; groove in lower lip narrow with
a few inconspicuous warts; no warts on upper lip; basal part of nose leaf rudimental ;

apical third of tongue filamentose ; interfemoral membrane not hairy.

Dental formula: 1. 4#— ec. +— p. 3 — m. 3 = 22.
The first lower premolar small and without anterior, basal cusp; the main cusps of
the entire series twice the height of the basal cusps.
The presence of the tail and a phalanx to the second digit are sufficient grounds to

separate Lonchoglossa from Anura.

Lonchoglossa caudifera Geoff.

Auricle pointed, internal basal lobe bound down to head. External border faintly
sinuate scarcely ; any external basal lobe; the inner lappet large. Tragus blunt at tip.
Nose leaf simple, without pedicle ; lateral gland mass of base rudimental ; wpper lip short,
without warts.

Large numerous vibrissee from face, especially from mentum. Filaments on tongue
large, not meeting in middle line of dorsum. Wing membrane reaches to calcar. Seven
rugee on the hard palate, the last two alone divided. The tail not quite as long as the
short interfemoral membrane, the tip not free.

The hair of the dorsum exhibits apical third brown, basal two-thirds pallid. Beneath
paler, prevailing hue brown (but with scarcely a contrasted shade toward base), tending
to become grayer, almost unicolored on loin. Limbs naked.

The wing markings both in the nerves and muscle fascicles are as in G'lossophaga, but
the terminal cartilage of the fourth digit is terete, and that of the fifth digit is small and
scarcely deflected.

The Skull—The bones yery.thin, permitting the subdivisions both of cerebellum
and cerebrum to be seen through the periphery. The pretemporal ridge unites with its
fellow at the anterior fourth to form a faint, linear crest; the mesotemporal and _post-
temporal ridges not separately defined, scarcely discernible. Fronto-maxillary inflation
small. Face vertex without pit at the fronto-nasal region ; outlines of nasal bones not
defined. Side of face conyex. The lateral borders of the anterior nasal aperture mod-

erately produced, The foramina between the two premaxille near the incisor margin large,

ON THE GLOSSOPHAGIN A. 207

The alveolar process so slender that it cannot be measured. The parts as viewed
from in front embrace the floor of the nasal chambers at the premaxillary part and permit
the median foramen to be seen. The zygoma without a trace of ascending process. The
posterior palatal margin near the root of zygoma spinose ; the posterior palatal notch with
conspicuous spines. Pterygoid process almost reaching tympanic bone and extends
beyond the oval foramen. Mastoid process aciculate. Mesopterygoid fossa with incon-
spicuous yomerine spine. Basioccipital depressions shallow. |The coronoid process
scarcely raised above the level of the condyloid process. The deflected hamular angle
projects in a marked degree beyond the condyloid. The lower border of the masseteric
impression is produced conspicuously beyond the border of the ramus. Symphysis with
large keel. One skull 21 mm. long; face 8 mm. long; brain case 15 mm. long.

Upper Teeth_—The small central incisors separated by wide interval, and each tooth in
close contact with the large lateral. The central incisor with ovoid crown scarcely wider
than neck ; the lateral incisor projecting below the level of the central with crown wider
than neck and conspicuously oblique outer border. The interval between lateral incisor
and the canine no greater than in other genera. Canine with inner surface flat. First
premolar one-half the size of the others; separated from the canine and the second pre-
molar, but nearer the last-named tooth. The second and third premolar triangular, with
large basal cingules.

The W-pattern of the molars discernible. In one specimen the long, sloping proto-
cone with suggestion of hypocone, recalling the parts as in Macrotus ; in the second the
teeth were without hypocone. Canine with rudimental heel. First premolar separate
from the canine and second premolar. Second premolar separate from the first and third ;
third premolar separate from the second, but contiguous to the first molar. First molar
with cingule of the protocone extended forward, scarcely deflected inward and overlap-
ping third premolar; protocone and paracone approximate, united at base.

Lower Teeth.—First lower premolar without anterior basal cusp, and is, therefore,
much smaller than the other premolars. In the entire series of premolars the main cusp
is twice as high as the height of the basal cusps. The first and second molars of
the same plan with the foregoing, the third being slightly the smaller.

The lower teeth with jaw are figured by Leche (/.¢., Taf. I, Fig. 8). The first pre-
molar is represented as being exactly like others of the series. This character would
prevent the Lonchoglossa of Leche’s identification being received under Lonchoglossa
caudifera of this essay. ;

Variations.—The above description is based on two specimens, which were subject
to some variation. In one the pretemporal crests did not unite. In one the cusps of the
teeth were much worn.

258 ON THE GLOSSOPHAGINZ.

Notes on the Skeleton.—Ribs thirteen ; first costal cartilage not wider than the rib.
Humerus with pectoral crest relatively high, one-half the diameter of distal end of bone.
The sternal crest after careful removal of the pectorals is very high and apparently with-
out notch, but the greater part of the interpectoral septum is membranous. The phalanx
of the second digit about as in Vespertilio. The metatarsi and first row of phalanges of
toes equal. .

Measurements.—Forearm, 36 mm.; foot and thumb of same length, viz., 8 mm.; fore-
arm, 1.35 mm.

BRACHYPHYLLINA.

I propose to establish the Brachyphyllina to include the genera Brachyphylla,
and Phyllonycteris,* forms which have hitherto been assigned separate groups in the Phyl-

lostomidie, the first named to the Stenodermata and the second to the Glossophagina.

Brachyphyllina.

Leaf-nosed bats with tip of tongue retaining clump of papille extending across
dorsum. In the Glossophagina the papille are arranged not only at the tip but the
sides for great lengths. The minute first upper premolar wedged in between the
canine and large second premolar; coronoid process acute, raised high above the level
of the condyloid process. Mesopterygoid fossa deep, apex answers to the junction of the
anterior and middle third of the zygoma. Nasal bones high, arched, defining a depres-
sion between them and the maxilla. Sagitta entire with well-defined pretemporal crests.
The glands of muzzle continuous behind nose leaf. Thumb large, one-fourth the length
of the forearm, nearly. Auricle narrow, oval with pointed tip. Tragus coarsely serrate
entire length of outer border. Upper lip hairy, without warts. Lower lp with shallow
median groove, margined with large warts. Lips not fringed internally.

BrAacHYPHYLLA.

Upper central incisors very much larger than the laterals. Length of forearm, 65
mm.; that of thumb, 16 mm., this being about one-fourth the length of the forearm as in
Phyllonycteris. Grinding surfaces of molars with numerous large mammillations, cuspi-
dation distinct. Angle of lower jaw quadrate, massive ; nostril entire, the wide outer
margin and the side of the rudimental nose leaf continuous. Tragus entire on inner
border. The tail rudimental, one-fourth the length of tibia, and concealed im the inter-
femoral membrane.

Dental formula: i. ¢— ce. +— prm. #— m. 3 = 20.
* T have not studied Riinophylla, but the conclusions arrived at after reading the accounts of Peters and Dobson

induce me to place the genus in the same alliance with genera just named. But in the absence of material I am com-

pelled to confine my comparisons to Grachyphylla and Phyllonycteris.

ON THE GLOSSOPHAGIN #. 259

Brachyphylla cavernarum Gray.

The auricle lanceolate with shghtly convea margins, basal lobes rudimental. The
tragus pointed, one-half the length of the inner margin of the auricle ; convex on thickened
inner, and coarsely serrate on outer, margin.

Nose leaf with entire nostrils and wide ectonareal flange ; erect portion of nose leaf
rudimental—concaye and often minutely crenulate on midmargin. Supranarial margin

concaye on either side of an obscure median ridge. Infranarial margin wide, continuous

with upper lip and faintly incised. The basal gland-clump continuous across face—vertex
back of nose leaf. The upper and outer parts are thick and bear a few coarse bristles,
while the lower are thin and lost on the upper lip. ‘Twelve warts are arranged in pairs
on the side of a mental V-shaped group, the median groove being shallow. Two median
warts may be said to haye slight morphological significance.

The fur above is yellowish white except the tip, which is brown. Below the tints
are the same, but the shaft is more tawny and the tips much lighter. The distal third of
the arm above and below is covered with hair. The distal half of the thigh is similarly
covered. A sparse growth of hair is limited to the upper half of the dorsal surface of the
interfemoral membrane.

The calear is rudimental. The terminal cartilages of the fourth and fifth digits are
uniform, elongated and scarcely wider at free margin than on the sides. The second
interdigital space is almost devoid of pigment. The third space retains a vertical line for
nearly its entire length, while the fourth exhibits one for about an inch near the free
margin, the rest of the space being areolated. The endopatagium is furnished with
numerous thick muscle fascicles ; near the tibia it is thick and leathery.

Second interspace, Third interspace, — Fourth interspace,
Pteral formula :

3 mm. 19 mm. 35 min.

The Skull—vThe walls of the skull are thin and permit the divisions of the brain to
be discerned. The sagittal, pretemporal and occipital crests are well defined and tren-
chant. The fronto-maxillary inflation is conspicuous and bears the pretemporal crest.
The inner orbital wall is moderately conyex, and is marked by a conspicuous foramen.
The infraorbital foramen is placed well in adyance of the orbit in line of the second
premolar. The zygoma with a rudimental ascending process at the posterior third, but
none anteriorly to contribute to the limitation of the orbit.

Lower Teeth.—The incisors are stout, in continuous row. The palatal basal cusp is
on level with the crown, which thus presents a broad, quadrate surface, marked in the
middle from before backward by a ridge. Canine without conspicuous basal cusp. Pre-

molars subequal, the first the smaller and triangular, the second with large basal cusp.

260 ON THE GLOSSOPHAGINA.

First and second molars with quadritubercular cusps well defined, a large mammillation
on the anterior commissure of the second molar; the third molar triangular, tri-
tubercular.

Upper Teeth—The central incisors are very large, triangular, nearly fillmg the
interval between the canines. The lateral incisors are minute, not over one-fourth the
size of the centrals. The anterior surface is concave; the crown is blunt and quadrate,
with basal cusp and cutting edge equal. The canine with anterior and posterior denticles,
the posterior of the two being enormous and presenting the aspect of being an outshoot
from the side of the crown. The first premolar minute and of the same form as the
lateral incisor. The second premolar large, triangular and projecting beyond the molars.
The basal cusp (denterocone) conspicuous. Molars tritubercular, without W-shaped
pattern. Several mammillations are present on the grinding surfaces. Third molar is

one-half the size of the second.

Measurements of Brachyphylla cavernarum.

Millimeters.
Head and body (from crown of head to base Of tail)......ss0.cssseeeecseeeeeeeeseeeeeeeeeneeeetee essere eeees 66
Thength Of Arm -...22....cccccesense-eeccserecanscenescessnrsssserecnnesnsens pedsso95ES00000900989 oAODIOCeH d500800 0 eo0300 40
Length of forearm, ...-.0.....ccesessseesssnececcoreceansecsocteccanesscceonevsretestnacerenssecnnsosseacsscoateccneess 65
First digit:
Length of first metacarpal DOnE.......-.:1:esssceceeceeeeensscceereseceeceesseseereceesrsscscceeenansense 4
Thength Of phalanges.......00..--0.sesesserssensersrcenesnccnesenenseoessuoersressvseecesnrecreuscennaesesrnes 12

Second digit:

Length of second metacarpal bone..

Length of first phalanx.......::scccccsesceeececneenseseccueneeeeeseceeessececsceeccssrsaenscsseseanecererens
Third digit:

Length of third phalanx
Fourth digit:

Length of fourth metacarpal bone.........-.:seccecceeeeteerenneteenceseceeteeses rescue sasteecsueeoers 51
Length of first phalanx...:.....0....0ccssscssecssecccnssossaacseusscesccocrereroness secneersascemuercersnnce 15
Length of second phalanXx..........::sc:ossccecssessscssecencteecceterceenercnerscsunceesene sodoenonpgaoo5000 17
Fifth digit:
Length of fifth metacarpal DOme..........sccseecsecerneeceseneetsencenneceeseseenesesereessueeeaecuserens 55
Length Of first phalamx......-01seccssesssscssrnnscescnasennrsiecacnneseserreesecccusetrressscecsseranevesers 15
Length of second phalanx.......0..ssssssssecersecsasccsescecececcenneasnesesecerevenccesesseeueccssteeeeners 14
Length of head
TCI SG) OF (CAaL:conssrcnccnvclsvons-seqnocesncerisscsiusnacseancsseecm=ran= ns -écQnan0oDGNSo3soNnGEDoSONNDOTHCOOGBOSLOD +212
Height of tragus.........csese-.seeee en0ece000019990000005R000 so0D00BDoBNaRaRDqHOoNS, EoGAGRdAHGEDs9Hdudacaaq905056000R3 9

Length of thigh

Length of tibia.....
Length Of £00b...c.0sccccccecceecenecreetsccnecseeraeeanscensscesasanecuseuseesarseceoascaussssoscesesenessesareceassses

Length of interfemoral membrane
Length of tail

ON THE GLOSSOPHAGIN®. 261

PHYLLONYCTERIS.

Upper incisors separated from the laterals by wide intervals; naked skin-fold
defining nostrils laterally ; nose leaf not reaching aboye the level of approximate club-
shaped gland masses. Thumb the largest in the group nearly one-fourth the length of
the forearm. Length of forearm, 45 mm. Teeth with cusps nearly obliterated, no W-
pattern on molars. Large vacuity between occipital bone and pars-squamosal of the
temporal. Fimbriz not arranged in rows, but form a uniform coyering to the tip of the
tongue. The first and fifth metatarsal bones longest. The first row of phalanges of third
to fifth digit of manus, same length as the second row. Calcar wanting. Zygomatic

arches fibro-cartilaginous.

Dental formula: i. 4 — ¢. + — prm. 3? — m.3 = 21

Phyllonycteris was described by Gundlach, but published under the care of Peters,
who does not appear to have known the form. Gundlach correctly compares the genus
to Brachyphylla. Dobson follows Gundlach closely, his description being little more
than a translation of the original article. When he departs from the text he makes
statements which do not agree with the specimen on which the present essay is based.
Thus he says, “the incisors are as in Glossophaga; the molars like those of Carollia

(Hemiderma), but the W-shaped cusps scarcely developed ;” whereas the upper lateral
incisor is twice the size of the central and the zygoma may be complete. With the

exception of the skulls, Dobson did not study Phyllonycteris at first hand.
Phyllonycteris sezecorm Gundl.

Auricle simple, ovate, with rounded pointed tip. External outline without subdivision
or inner lappet near the base. Internal basal lobe scarcely free. Tragus convex on inner
side, straight on outer. Both sides marked by three, coarse, teeth-like processes. Basal
point scarcely longer.

Nose leat simple, obtuse with internarial pedicle. The perinarial flange is lamillar
and distinct from gland mass. The structure last named well defined, apparently
crossing muzzle back of the nose leat, but two club-shaped masses are nearly approximate.
Upper lip high without warts. Interfemoral membrane deeply incised, extending from
distal third of the tail to the caleaneum. The tail is short, scarcely projecting beyond the
interfemoral membrane. The fur long and silky above, light gray tipped, subtip sooty,
the rest of the hair pale verging to white. Beneath much paler, nearly uniform gray.
The tip of hair tawny, the rest of the hair of a somewhat lighter shade.

Almost the entire field of the endopatagium filled with widely separated nearly

equidistant vertical muscle fascicles, There is no reticulated arrangement of fibres, The

262 ON THE GLOSSOPHAGINA.

nerve markings in the fourth interspace as in Glossophaga except that from the fourth
digit there are three instead of one nerye. The terminal cartilage of the fourth digit is
obscurely spatulate.

The Skull—The skull not papyraceous, the division of the cerebellum, but not of
the cerebrum, discernible on periphery. The pretemporal crest distinct. It begins over
the moderate fronto-maxillary inflation to form a delicate crest by union with the fellow
of the opposite side at the anterior third of the sagitta. _Mesotemporal and posttemporal
crests not discerned. The orbital ridge is rudimental, but the frontonasal pit conspicuous
at proximal end of the slightly convex nasal bones. The large infraorbital foramen
lies over interval between second premolar and first molar and is thatched by a ridge.
The alveolus (7. e., the distance from the central incisor to the anterior nasal aperture)
equals in height one-fifth of the base of the upper canine and one-eighteenth of the ver-
tical diameter of the large, anterior, nasal aperture. The zygoma often complete.* The
maxilla at root of zygoma with a very small ascending process. The premaxilla at the
side of the anterior nasal aperture salient. Neither the grooye between the nasal bones or
the depression on the maxilla at the side of the nasal bones are conspicuous. The depres-
sion between the aperture last named and the eminence over the canine is shallow. The
hard palate just back of the last molar is sharply defined by a double crescentic trans-
verse ridge; the palatal notch is acute and deep, the apex reaching the level of the -
anterior third of the zygomatic arch, the pterygoid process corresponding in position to
the oval foramen. The tympanic bone touches the postglenoid process. The junction of
the ethmoid and sphenoid bones in the brain case not convex. A vacuity is found in the
line of junction of occipital and squamosal bones.

The basioccipital bone with scarcely any pit-like depressions ; the vomerine ridge
scarcely discernible in the mesopterygoid fossa. The mastoid process small, conical.
The proportion of the face to the brain case is as 9 to 15 mm.

Lower Jaw.—Coronoid process acuminate. The hamular angle not deflected or pro-
jected beyond the condyloid process ; lower border of the masseteric impression not dis-
tinguished from the corresponding border of the horizontal ramus. Back of the molars
and at base of coronoid process a tubercle for insertion of temporal muscle is seen.
Symphysis-menti broad, non-carinate, the surface near the incisors marked by coarse
venous foramina.

The Teeth—The upper central incisors hatchet-shaped, contiguous ; laterals much
smaller, not half the size of centrals and separate therefrom. The incisors not entirely

occupying space between the canines. Canine broad at base, robust, convex entire length

* Dobson ( Cat. Chirop. Br. Mus.) in text states that they are incomplete, but acknowledges the fibro-cartilagium

arch in a footnote,

ON THE GLOSSOPHAGIN#. 263

of palatal surface. First premolar very small, nodular, about one-fourth the size of the
second and not much larger than the lateral incisor. Second premolar triangular, with-
out basal cusp; posterior half of palatal surface concave. Molars without well-defined
cusps and decrease in size gradually from before backward. The third molar one-half
the size of the second. ‘The protocone, paracone and metacone scarcely indicated ; no W-
shaped pattern.*

Lower lateral incisors twice the size of the centrals; all are non-contiguous and
nodular. Canine with conspicuous concave heel; all other parts convex; cingulum
extends inward so as to lie back of the lateral incisor. The premolars thick and robust,
subequal ; the first smaller. The molars decreasing in size from before backward without
details.

Of the measurements it is noted that the first phalanx of the first digit is scarcely
longer than the metacarpal bone. In the second digit the single phalanx is one-tenth
the length of the corresponding metacarpal bone. The entire second digit is as long as
the third metacarpal bone. In the third digit the first and second phalanges are equal—
the third phalanx is nearly one-half the length of the second. The terminal cartilage of
the fourth digit is moderately spatulate, and that of the fifth digit is deflected toward the
body. The wing membrane attached to the tibia at the distal seventh or to the ankle.
Interfemoral membrane attached to tip of the small calcaneum.

The Skeleton —The sternum is boldly keeled over the presternum and metasternum.
The ribs are twelve in number. The first costal cartilage is discoidal. The humeral
pectoral crest is relatively low and not half the diameter of the proximal end of the bone.
The fifth metatarsal bone is much the largest of the series. Palatal rug eight, last three
to four interrupted in centre. The first and fifth metatarsals are longer than the others.
The bones of the first row of phalanges of the toes are equal.

* Peters and writers following him give all glossophagine genera W-shaped pattern of molars. I have had no oppor-

tunity of examining the type of Phyllonycteris in the Berlin Museum, but I have received through the kind offices of Mr.

Paul Matschie a photograph of the skull which I find conforms to the account above given.

A. P! S.— VOL, Xix. 24,

264 ON THE GLOSSOPHAGIN®.

Table of Measurements (in millimeters).

&: | & 3 | fmee Wey es =a 5 2
S | 8 aes Saf ree lush
Head and body (from crown of head to base of tail) 45 45 57 | 55 40 49 | 32
TL SIAN GP PE ease os cnocesanoecocas onnaosonnecoosnoce aoe oon sb ansstoaoansoGN: ssbarcossoaee 19 2 | 20 20 20 | 25
Teng ih Of fOrea tM accass--ceseeseresseneraaanne=aneeeece ner oeceo= = neanencccssce=acennean iar 36 39 50 | 42 | 35 38 45
First digit : } |
Length of first metacarpal bone 4 4 4 3 5
Length of first phalanx .-------.----<-2-22----0--2-2c-ceeeenennneneesnanennee one 4 4 4 3 3 3 7
Second digit : :
Length of second metacarpal bone......---.+--:+++----s0eeeesseeeeeeeee sees 30 25 40 40 29--) 33 33
Length of first phalanx.......- Toaoencose 1 2 3 0 yet ONC 3
Third digit : a
Length of third metacarpal bone........-..---+:+s++0eeeeeeeeereeeeeeseeeeeees 34 30 | 47 45 37 38 | 38
TeeMStH Oe Tash [PRA on sec conse taco nosceoscnacaceesspeosessoneoscesccccoccees 13 sal 14 17 42 | 13 | 44
Length of second phalanx....00:.0...::22seeceeseeeeece see ce ese c es neeeeeeeeeeees 16 12 | 923 21 18) |) 21 S14
Length of third phalanx......--..--.+0:cseseececsseeeeeesceeeeeeeeeeeeneeeseeees 7 Gj 9 Quslestt eels

Fourth digit : | |
Length of fourth metacarpal bone

Length of first phalanx ...---.+--2:::ceeseseseeeees eee eneneeesceesessseeeeeeees 10 9 11 12 | 10 | 13
Length of second phalanx...-----.---:2:::sseeseseeeessseeeeeeesecceeeseeeeeees eat) 9 16 15 12 13 11
Fifth digit : .
Length of fifth metacarpal bone 30 20 40 35 30 30 35
Length of first phalanx........-..... pasagia swt ae eetectiewcet wos seescnesteen weet oee 9 8 10 10 7 8 1d
Length of second phalanxX..--.-++..:ssssssscceceeeeceeseneesssseeeeceeeeeseeenees 9 8 10 13 To es | 10
Length of head.......e..ceeeeeceeceeeeeeeceecesseeseseeeecneeeererenteeceeeereeececeseasanes 23 21 o7 32 25 29 25
Height of ear 14 ) 11 12 13 13 14 11
TS erTey SNE re ea Scene celareecrner ees Ree Beason a eeeaR occas eceeocec acto 6 ASS |e} 4 5 eae 4] 5
Length of thigh......-....--:s:e++- PR ee eae abe Hees soa oaSe 10 2 15 A. jel Sella STS
Tareas reer eae ee es MR ne SE, 14 [ee top |e Penta te etree
Length of foot 8 8 12 10 vw 7 13
Length of interfemoral membrane in median line........---+--++111s12++eeeeeeees hal 9 20 | 4 6 7
5 2

Length of tail....----..:::eceeeeeeeeeeeeeecee cee eeeeee cece eneeeseeeaaee cesses eeeeeeseneaeees

Norre.—The Secretaries deem it proper to state that this, as well as the succeeding paper, was presented to
the Society after the author’s death, which lamented event occurred on November 14, 1897, and that, therefore, it

has not had the benefit of his revision in its passage through the press. !

Glossophaga soricina.
Glossophaga soricina.
Glossophaga soricina.
Glossophaga soricina.
Glossophaga soricina.
Glossophaga soricina.
Glossophaga soricina.

Glossophaga soricina.

Glossophaga truet.
Glossophaga truet.
Glossophaga truet.
Glossophaga truet.
Glossophaga truet.
Glossophaga truet.
Glossophaga truet.

Monophyllus redmani.
Monophyllus redmani.
Monophytlus redmani.

Monophyllus redmani.

Monophyllus redmani.
Monophyllus redmani.

Brachyphylla cavernarum.
Brachyphylla cawernarum.
Brachyphylla cavernarum.
Brachyphylla cavernarum.

Brachyphylla cavernarum.

Brachyphylla cavernarum.
to 39. Brachyphylla cavernarum.

Leptonycteris nivalis.
Leptonycteris nivalis.
Leptonycteris nivalis.
Leptonycteris nivalis.
Leptonycteris nivalis.
Leptonycteris nivalis.

ON THE GLOSSOPHAGIN ji.

EXPLANATION OF THE PLATES.

PLATE VI.

Head seen from in front.
Skull vertex. xX 3.
Skull profile. xX 3.
Skull base. X 3.
Jaws with incisors and canines seen from in front.
x 10.

Lower teeth seen from above.

Xx 2.

Upper teeth.
x 10.
Left lower molars seen in profile from lingual aspect.

Puate VIL.
Head seen from in front. X 2.
Skull vertex. xX 3.
Skull profile. X 3.
Skull base. xX 3.

Upper teeth. xX 8.
Lower teeth seen from above.

x 8.

Left lower molars seen in profile from lingual aspect.

PLATE VIII.

View of head from in front, showing ear and nose leaf.

Skull of same. Norma verticalis. x 3.
Skull of same. Norma lateralis. x 3.
Skull of same. Norma basilaris. X 3.

Upper and lower jaws seen from in front. xX &

Teeth of the same as seen from the surfaces of crowns.

PLATE IX.

View of head showing ears and nose leaf.

Skull of same. Norma verticalis. x 3.
Skull of same. Norma lateralis. x 3.
Skull of same. Norma basilaris. x 3.

Upper and lower jaws seen from infront. X 8.
PLATE X.

Teeth of same seen from the surfaces of crowns. X

PLATE XI.
Head seen from in front. xX 2.
Skull vertex. xX 3.
Skull profile. xX 3.
Skull base. xX 3.

Jaws with incisors and canines seen from in front.
x 8.

x 8.
Upper teeth.

IGD

Xx 8.

The first molar is to the

X 2.

oS (eb

Terminal cartilages of the fourth and fifth digits.

266 ON THE GLOSSOPHAGINE.

Fig. 46. Leptonycteris nivalis. Lower teeth. X 8.
Fig. 47. Leptonycteris nivalis. Left lower molars seen in profile from lingual aspect. The first molar is to the

right. X 10.
PLATE XII.

Fig. 48. Chernycteris mexicana. Head seen from in front. X 2.

Fig. 49. Charnycteris mexicana. Skull vertex. X 3.

Fig. 50. Charnycteris mexicana. Skull profile. X 3.

Fig. 51. Charnycteris mexicana. Skullbase. X 3.

Fig. 52. Chernycteris mexicana. Jaws with incisors and canines seen from in front. X 5.

Fig. 53. Chernycteris mexicana. Upper teeth. X 10.

Fig. 54 Chernycteris mexicana. Lower teeth. X 10.

Fig. 55, Chernycteris mexicana. Left lower molars seen in profile from lingual aspect. The first molar is to the

right. 10.
PLATE XIII.

Fig. 56. Lonchoglossa caudifera. Head seen from in front. X 2.

Fig. 57. Lonchoglossa caudifera. Skull vertex. X 3.

Fig. 58. Lonchoglossa caudifera. Skull profile. x 3.

Fig. 59. Lonchoglossu caudiféra. Skull base. XX 3.

Fig. 60. Lonchoglossa caudifera. Jaws with incisors and canines seen from in front. X 8.

Fig. 61. Lonchoglossa caudifera. Upper teeth. X 8.

Fig. 62. Lonchoglossa caudifera. Lower teeth. X 8.

Fig. 63. Lonchoglossa caudifera. First and second right lower molars seen from lingual aspect. The first tooth
is to the right. x 10.

PLATE XIV.

Fig. 64. Anuwra wiedii. Head seen from in front. X 2.

Fig. 65. Anura wiedii. Skull vertex. X 3.

Fig. 66. Anura wiedii. Skull profile. X 3.

Fig. 67. Anurawiedti. Skull base. X 3.

Fig. 68. Anwra wiedii. Jaws seen from in front showing incisors and canines. > 8.

Fig. 69. Anura wiedit. Upper teeth. X 8.

Fig. 70. Anwra wiedii. Lower teeth. X 8.

Fig. 71. Anuwra wiedii. Left lower molars seen from lingual aspect. The first tooth is to the right. X 10.

PLATE XY.

Fig. 72. Phyllonycteris sezecorni. Head from in front. X 2.

Fig. 73. Phyllonycteris sezecorni. Skull vertex. X 3.

Fig. 74. Phyllonycteris sezecornt. Skull profile. X 3.

Fig. 75. Phyllonycteris sezecornt. Skull base. X 3.

Fig. 76. Phyllonycteris sczecorni. Upper teeth. X 10.

Fig. 77. Phyllonycteris sezecorni. Lower teeth. X 10.

Fig. 78. Phyllonycteris sezecornt. Jaws seen from in front showing incisors and canines. X 8.

Fig. 79. Phyllonycteris sezecorni. Left lower molars seen from lingual aspect. The first tooth is to the right. xX 10.

ARTICLE VI.

THE SKULL AND TEETH OF ECTOPHYLLA ALBA.
(Plate XVI.)

BY HARRISON ALLEN, M.D.

Read before the American Philosophical Society, January 21, 1898.

In 1892 (Proc. U. S. Nat. Mus., 1892, No. 913, 441), I described a bat from
Honduras under the name of Lctophylla alba. The single specimen was without skull.
I have been permitted through the courtesy of Mr. Oldfield Thomas, of the British
Museum, to inspect a second example of the genus. The material consisted of a dried
skin and a skull of a male individual which was mutilated by shot in the ptery-
goid and orbital regions. The specimen was collected at San Emilio, Lake Nic-Nae,
Nicaragua.*

The norma verticals shows faint fronto-temporal lines which barely approximate near
the bregma, but recede from that point posteriorly so that no trace of a temporal crest
exists. The fronto-maxillary inflation is conspicuous and makes a swollen border for the
upper and anterior orbital margins. The nasal bones are sharply elevated above the
plane of the maxilla. Sufficient of the norma dasilaris remains intact to show that the
hard palate is elongated and the palatal bones are produced, thus separating the genus
sharply from Stenoderma and its allies and allying it to Vampyrops (see Synoptical
Key). The basioccipital bone is deeply pitted for muscular impressions. In this respect
it presents a marked contrast with Vampyrops, in which this bone is nearly fiat. The
tympanic bone is small, leaying the greater part of the cochlea exposed. The norma
occipitalis shows a weak occipital ridge. The junction of the ectopetrosal + surface of the
pars-petrosa with the occipital bone is complete, while in Vampyrops a vacuity exists.

The lower jaw retains a curved aciculate angle relatively twice the size of the same

* The skin was badly mutilated by shot and the nose leaf and chin plates so distorted that no attempt is made to
compare the parts with the original description. The second interdigital space is without pigment, head and neck both
above and below are pure white. The lower third of the body both on dorsum and ventre is tipped with ash-gray.

+ I propose naming that part of the pars-petrosa lying in the brain case the endopetrosal, and that lying exposed

back of the pars-squamosa the ectopetrosal part (Journ. Acad. Nat. Sci., 1896, Philadelphia).

268 THE SKULL AND TEETH OF ECTOPHYLLA ALBA.

part in Vampyrops. ~ The masseteric muscle extends to the lower margin of the ascending
ramus. The coronoid process is one-third smaller than in the genus last named.

Dental formula: 1. 2 — c. + — prm. 2 — m. 2 & 2 = 28.

The Teeth—Upper incisors conical; the centrals larger than the laterals with rela-
tively broader bases. The centrals are separated from each other by a smaller interval
than exists between these teeth and the laterals, or between the teeth last named and the
canines. The canines are slender and slightly longer than the second premolar. The
first premolar is pointed, root much exposed and is about one-third the size of the second.
The first upper molar is quadrate with trenchant marginal cusps in position of proto-
cone, paracone and metacone; the crown defined by these elements is concaye. ‘The
second molar is pyriform, the base being toward the palate. A pointed marginal cusp is
seen in the position of the paracone and a second in that of the metacone. The crown is
concave and simple, save for a longitudinal ridge. The premolars and molars are separate
from one another ; the greatest interval being between the premolars.

The lower incisors are blunt cones, contiguous, filling space between canines; the
teeth last named are deeply excavate posteriorly. Premolars are aciculate, the first tooth
almost touching the canine and is smaller than second. The second tooth is deeply con-
cave posteriorly with a conspicuous heel and cusp. The molars are subequal, without W-
pattern. The first molar is obscurely quadrate, slightly narrowed in front with enormous
sharply pointed paraconid; other cusps are absent; the lingual border is not raised.
The second molar is subrounded, no trace of cusps being present other than a longitudinal
ridge in the middle of the deeply excavate crown. The front and lingual borders of the
tooth are greatly elevated, the former furnished with two sharp processes, the latter
crenulate. The teeth are all separated from one another beyond the canine, the smallest
interval being that between the canine and the first premolar and the widest between the
premolars. :

Letophylla is in alliance with Vampyrops. It resembles this genus in the upper
incisors and first upper premolar being conical and in the prolongation of the palatal
bones. The shape of the lower first molar possesses a large paraconid, but is without
protoconid. In the dental characters last named ctophylla is like all other Steno-
dermine, excepting Brachyphylla, Artibeus, Dermanura and Sturnira.

The forms exhibiting the stunted, first, lower molar are again divided into two groups
by the palate andthe lower jaw. In Chiroderma, Vampyrops and Ectophylla the palate
is oblong ; the palate bone extends to a point answering to the anterior root of the zygoma,
or eyen the posterior third of the arch, and the lower jaw has a well-defined posterior
border to the ascending ramus, with no deflected angle. In Pygoderma, Stenoderma and

THE SKULL AND TEETH OF ECTOPHYLLA ALBA. 269

Trichocorys, the palate is rounded, as a rule excavated and rarely reaches a point
answering to the anterior root of the zygoma; the lower jaw has no well-defined posterior
border, the boldly deflected angle almost reaching the condyloid process.

The position of Eetophylla in the Stenodermine is shown in the synoptical natural
key. Brachyphylla is an annectant genus to the Glossophagina through Phyllonycteris
Artibeus, Dermanura and Sturnira apparently relate to the Vampyri, but while the
structure of the molars is essentially that of this group, no annectant form is known.
Sturnira in the simplicity of the tooth structure recalls Hemiderma. The relation
between the remaining genera of the table is intimate. The Stenodermine constitute,
with the exception of the Heamatophilhia, the most aberrant group of the Phyllostomidide.

I recognize, therefore, the following natural arrangement of the genera :

Subfamily STENODERMATIN#.

eae linam ny llinie ne Seen aes. cste ha hreiad. see Brachyphylla.
( Artibeus.
P Nor tillye lines eis eee eh re Mice Bare | Crake
Dermanura.

| Sturnira.
( Chiroderma.
*, | 4
Ghimocleriminiiee eee ehh ae = ca haan ee : Vampyrops.
| Eetophylla.
( Stenoderma.
| Pygoderma.
aie 1 Centurio.
Stemod ermine cy eet kaa ks saan :
Trichocorys.
Ametrida.
y .
| Spheronycteris.

A Natural Synoptical Key of the Stenodermide, Based on Characters Derived from
the Skull and Teeth.

I. First lower molar elongate with paraconid distinct.

[ a. Angle of lower jaw broad, scarcely pointed, concave above, not deflected, ascending
\ ramus defined. Hard palate oblong, palatal bones produced. Upper incisors coni-
| cal, molars $ ; crowns coarsely ridged ; all cusps of the first lower molar subequal...
l Brachyphylla.

Group Brachyphyllini....

* Ohiroderma is not as near Vampyrops and Ectophylla as the members of other groups are to each other,

270 THE SKULL AND TEETH OF ECTOPHYLLA ALBA.

( a. Angle of lower jaw narrow, aciculate, not deflected ; posterior border of ascending
ramus defined ; hard palate oblong ; palate produced.

b. Palatal bones extend to point answering to the middle of zygoma. Upper incisors

flat ; first upper premolar broadly lanceolate ; crowns of molars rugose ; proto-

conid and paraconid of first lower molar prominent, subequal, the others rudi-

ee mental.
CEO pe OUCI NS eres Gy WIDE #roccsccecocsscontcooscs0ssdoonnoos.osnao2esenscasnseosoososcaerocnBNNONOADNODSONN Artibeus.
Gl. WIGS 2 -cococonencoso op eanbacoadapo Scand cnoSesndoorocosDosHONENDOAboSSHHoooSSANNIG Dermanura.

b’. Palatal bones extend to point answering to the anterior third of the zygoma.
Upper incisors conical, contiguous ; first premolar narrow lanceolate ; crowns
of molars smooth ; all cusps of first lower molar subequal, anterior commissure

@msjaneeine 8 NOMEN 2 oocossocoasos ope cocogasQnecdencencocosoeponHSSconBeGREoSioconnCoD4 Sturnira.
Il. First lower molar subquadrate without paraconid.

if d. Hard palate oblong, palatal bones produced. Upper incisors conical.

e. Angle of lower jaw quadrate, not deflected, posterior border defined.
Nasal bones absent in adult ; palate bones produced nearly to
the line of glenoid cavity. First upper premolar acicular ; first —

ae lower molar with protoconid and mesaconid subequal. Molars 2...

Group Vampyropini....-.- Cee

e/. Angle of lower jaw acuminate, not deflected. Protoconid of first
lower molar aciculate, enormous.
Jf. Hypoconid first lower molar rudimental ; molars 2... Vampyrops.
jf’. Hypoconid first lower molar none ; molars 3.........-.. Ectophylla.
ad’, Hard palate round, palatal bones scarcely, if at all,* produced.

| e/’. Angle of lower jaw rounded, deflected, posterior border ascending

ramus not defined.

g. Frontal bone in orbit greatly inflated ; palatal bones extend
to a point answering to the anterior root of the zygoma ;
| pterygoids produced, inflated and nearly teaching the

panic bones; upper incisors conical ; protoconid of
first lower molar searcely larger than other cusps ; hypo-
| conid of the same tooth marginal, rudimental molars 2...

| Pygoderma.

Cremicrenodencie | g'. Frontal bone in orbit not inflated ; palate bone produced
to anterior third of zygoma; upper incisors conical ;
protoconid first lower molar enormous; hypoconid of

same tooth marginal ; molars }...-.....-.---..-.-+ Ametrida.

g'’. Frontal bone in orbit scarcely inflated ; hard palate with
posterior margin excised; pterygoids not produced.

h. Palate excised to first molar ; hypoconid of first lower
| molar inside contour. Molars .........-. Stenoderma.
| h’'. Palate excised to middle of first molar ; hypoconid of
|

first lower molar marginal. Molars 2 ... Trichocorys.

* Mr. O. Thomas (Ann. and Mag. Nat. Hist., 1889, p. 70) first employed this character to separate this group from
the foregoing.

THE SKULL AND TEETH OF ECTOPHYLLA ALBA. 271

Measurements of Ectophylla alba (in millimeters).

Head and body (from crown of head to base of tail)... 36 36
Men ot hpoisanmirenn cerca terrane scar mere atcctcine seameeiooecisciie ceccacioe tition caste celecs vce vena shade svcetestersevsiics 17
IL@IGIN Ot TOREAI TT qeaosadsoanseda0.an9odbsedb0adg6n0s8094Geqshdd90s00N60d0K000 000 Jodo bco0desHSsodDnECHE AG DoS He SEnBBEDEAOOSOEe 25 26
First digit :

ent hvolsnirshame tacan pala On eseerspheecsenteecsanceimasce ses sscsteneeresreeetimesaes see seceeeessesseemececees : 3 3

Length of first phalanx...........-- 3 3
Second digit :

Wencthvotsecondentetacanpalliib one ys eeeerteeer creer steerer cece enlaces eee sesereseiseereseneaseeeteeceere 21 20

TLS ON THRE, THAT TAS oc soog3n50000s900030N= INS cao SoocONbonSSaaTEG2Goqsabsoco9qsaROveS>9aHNGNDBAD9S9GBe0000 HOBHaDAea 3
Third digit :

Length of third metacarpal bone.........- 25 25

ILE GE aS TONE FATED 0053090000000 ssaDDaDoADDOAGOCODBEHEDHODSEdoONdoHSHODoSENcHOADScROBOeSHASEAaEHAGIS 9

ILemaHn @ii S@COMEl TRAN, soocec cov e3c000n0009s900099c3000R00080800 neagdaNDSSoODDoGONSOD oADeooAoeSAANGACES HoaHSo90a 12 13

ILEWSUN Oi WantREl TNA eva CeccosoHoaconsdoooono5coaeINaESDDOGNbSONoROnOSCASDDKDDN|AGOGG soNEEHoSDHEOLOdesADoaEeDeGouAS 6 6
Fourth digit :

Length of fourth metacarpal bone 25 25

Meno thyotuirstaphalamxerscyeccades veces ose de eeicishae cinch anced ene mare ueeseunaec ce acca sene ae soecee? sant 7h 8

ILEMAWA Oi Seeded! [MAMI .coccnacsooocoGseDoGASEABOOBEOCHESoDUA SoBNSooogonnScoRaSDoDNEgOIBAALONATeREDEOgOSUDNeS 8 7
Fifth digit : |

‘Length of fifth TEASE EN! INO, paovoboodcongkosossoass0nGenbon 7OSG9nONEGERO anEDosONH/-aagodaDRoOEsCORONanDOGaeOD 25 ?

Length of first phalanx.. 6 6

ILemaWn OF SecouGl Tp ogee, coosopon20096590Racoa9Hoas nonDobsbopadeooosnHocHosHoo;endDoRSopSsOGASHADBORSoEHOODE Anae 7 ri
ILE HD OF VEAL ceosenodasccsen600399c00905000 099 D500N00090000d0s NoNDOS NSESEES OED gBOsOSAOIcCoGNOoOOoRGNOSHOS PoaDbosaneHooAeaDAG 14 14
JEIGREING @E © JProsccsaqnedancoconesecoq das pss bos0Hbo0000¢bac0000000 00500 aod ObUDDUsHAIGHE teEoRAODODDAESHOSeOUSbOARsODHsoGovOHONONS 10 10
TEIGTEINE OF WREGWE .000900900060 on00000c0 0900900990000d000000000000950020000-,eaqD0D60 DonooDOINOTODOAGENNSNNOROHO SoncvasoROseDABHeE 5 2
Length of t. igh.... 8 i
ILEMAIN OP HDI, coocococcecas oscosonon donc coseaa0cnqqCnObESURCBSCONDABROSaqOCaD0 EqDNSIDDboaENcHDLAGOHDOOOD ScHoCODONDOTAGUOnEOTE » 10 10
Length of foot ........ poo soaseso99eecbapcaDoDUaR ADDED DODHOUDEESORCO EDU DOONoDEANOCOORD.SoDAcHOsedSedonSEDonaaEEuDaEEeoSRaCRENcaS 8 8
Length of interfemoral membrane........... SCORE GUE SNHSGABEAdoOSGnED. cosdacHaonbacoonaDoode Load bHebsaenEoooecanereones 4 4

In concluding the account of this interesting specimen, I will call attention to the
molar teeth of Cephalotes, a member of the remote group of the Pteropodidee. The two
genera, however, resemble one another in being frugivorous, in retaining few or no
tubercles to the molars and, probably on this account, in exhibiting elongated crests in the
centre of deeply excavate crowns. A tenable hypothesis for the origin of this central cusp
may be expressed as follows. The grinding away of the crowns has gone on to a degree
that brings the enamel cap down near to the division in the alveolus, between the sockets
for the roots of the teeth, so that this ridge acts as a point of resistance to further wear
and leads to a reassertion of the principle of cuspidation at this point,

A. PB. S.——Vow. XIX. 21.

DD THE SKULL AND TEETH OF ECTOPHYLLA ALBA.

One of the most marked characteristics of the teeth of fruit-eating bats is the dis-
position for the loss of cusps in the molar teeth. This takes place without intermediate

grades so far as is known. In two of the three subdivisions of the Phyllostomide it

occurs as exceptions to the rule—Hemiderma in the Vampyri and Phyllonycteris in the
Glossophaginee, but is the rule rather than the exception in the Stenodermine. In the
Pteropodide the tendency to the loss of cuspidation is the rule, the genus Pteralopex
being the only exception. Such abrupt variation within the limits of small groups indicates
that the tendency to external specialization has weakened the type and exposes it under
the influence of environment, ordinarily acknowledged as active in modifying forms, to

gross modification always on the side of deterioration.

EXPLANATION OF PLATE XVI.

. Eclophylla alba—norma verticalis.
. Eetophylla atba—norma lateralis.
. Ectophylla alba—upper and lower teeth.

=
(>)
mw wm ee

. Ectophytla alba—lower molar (profile).
. Ectophylla alba—ramus of lower jaw.

. Cephalotes peroni—first right upper molar.

leo)
=!
asI OD

. Cephalotes peroni—first and second right lower molars.

7
-

£4,
ae
Lig

NOTIGE.

A few complete sets can be obtained of the ‘Transactions, New ates Ve
I—XVII. Price, ninety dollars.

Acdress, THE LIBRARIAN.

OCT 4@ lave

fens ACTIONS

OF THE

AMERICAN PHILOSOPHICAL SOCIETY,

HELD AT PHILADELPHIA,

FOR PROMOTING USEFUL KNOWLEDGE.

VOLUME XIX.—NEW SERIES.

PART TTT.

. ARTICLE VII.—The Osteology of Elotherium. By W. B. Scott.

Articue VIIT.—Notes on the Canide of the White River Oligocene. By W. B. Scott.

ARTICLE IX—Contributions to a Revision of the North American Beavers, Otters and Fishers. By
Samuel N. Rhoads.

a, Philadelphia:
4 PUBLISHED BY THE SOCIETY,
¥ AND FOR SALE BY
ae: Tue American PuirosopHicat Society, PHILaDELPHta,

N. TRUBNER & CO., 57 and 59 LUDGATE HILL, LONDON.

: 1898,

OCT 4& 1898

ARTICLE VII.
(Plates XVII and XVIII.)
THE OSTEOLOGY OF ELOTHERIUM.

BY W. B. SCOTT.

(INVESTIGATION MADE UNDER A GRANT FROM THE ELIZABETH THOMPSON FUND OF THE A. A. A. Ss.)

Read before the American Philosophical Society, February 4, 1898.

Elotherium is one of the many genera of fossil mammals concerning which the
growth of our knowledge has been exceedingly slow, and only of late has it become prac-
ticable to give a complete account of its bony structure. The genus was named in 1847
by Pomel (47 a, 6) and shortly afterward renamed Entelodon by Aymard (’48) from a
better specimen, but for several years only the dentition was known and that imperfectly.
In 1850, Leidy (50, p. 90) described the first American species, but, not suspecting its
generic identity with the European forms, he at first referred it to a new genus, Archio-
therium. Leidy’s material enabled him to give a fairly complete account of the skull.
Kowaleysky, in 1876, described an imperfect skull found in France and he further
showed that the feet were didactyl, a very unexpected fact in view of the pig-lke char-
acter of the dentition. In this country Profs. Marsh and Cope have added materially to
our knowledge of this remarkable animal (Marsh, ’73, 793, 94; Cope, ’79) and the
former has published a restoration of one of the species. In spite, however, of this list
of workers who have, from time to time, occupied themselves with the study of Hlothe-
rium, much still remains to be learned regarding its structure, and its phylogenetic rela-
tionships are even more obscure. ;

In the summer of 1894, Mr. H. F. Wells discovered in the White River Bad Lands
of South Dakota certain bones, which, with the expenditure of infinite pains and skill,
were excayated from the rock by Mr. J. B. Hatcher, and which proved to be a most
remarkably complete skeleton of EHlotherium. This beautiful specimen (Princeton Mu-
seum, No. 10885,) formed the subject of a preliminary communication which I made to
the third International Zodlogical Congress, at Leyden (Scott, ’96), and will be more fully

described in the following pages. Except for a single thoracic vertebra (and perhaps a

QA THE OSTEOLOGY OF ELOTHERIUM.

few caudals) and part of the hyoid apparatus, the skeleton is complete; it is represented
in Pl. XVII, which will enable the reader to judge of its unusual state of preservation.
Additional material, belonging to several species, will also be made use of for purposes of
of comparison, but the description will deal almost exclusively with the White River
forms.

The Artiodactyla may almost be designated as the despair of the morphologist. So
manifold are the forms which this puzzling group has assumed, and so variously are the
characteristics of its minor groups combined, that the confusion seems hopeless. ‘The
only way in which this tangled skein can be unrayeled and its many threads separated
and made straight, is by the slow but sure method of tracing. the phylogenetic develop-
ment of each family step by step from its incipient stages. Many years must pass before
sufficient paleeontological material has been gathered to make this possible, but already
some progress has been made in the work. Each successive form in a series, as soon as it
is recovered, should be fully described and illustrated for the benefit of other workers, a
necessity which must excuse the minuteness of detail into which the following deserip-
tion enters. For the sake of conyenience the entire bony structure of the animal will be
described, including those parts which are already well known, in order that the reader
may be spared the trouble of searching through many scattered papers, written in several
languages.

I. Tue Deyririon.

The teeth of Hlotherium are already familiarly known and require but a brief account

here. The dental formula is I 3, C 4, P 4, M 3.

A. Upper Jaw.—The incisors, three in number, increase regularly in size from the

first to the third, the latter being much the largest of the series; it has a conical or some-
what trihedral crown and resembles a canine in shape and appearance. In some individ-
uals the crown of this tooth is worn in a peculiar manner, a deep groove or notch being
formed on its posterior side, in a place where it cannot haye been made by the attrition
of any of the lower teeth. The other incisors have spatulate crowns, with blunted tips,
the attrition of use wearing down the apices as well as the posterior faces of these teeth.
This description applies more particularly to the larger White River species, such as
Lf. ingens and E. imperator ; in E. mortoni the upper incisors are of more nearly equal
size and more conical shape. In all, the median incisors are separated from each other
by a considerable notch, and the whole series is much more extended antero-posteriorly
than transversely, the external incisor standing behind the second one. I 3 is separated
by a short diastema from the canine and at this point the premaxillary border is quite
deeply notched to receive the lower canine.

The canine is a very large and powerful tusk, with a swollen, gibbous fang; the
d c=) I ? co) fo)

THE OSTEOLOGY OF ELOTHERIUM. 275

crown is long, massive, recurved, and bluntly pointed; it is oval in section, and has a
prominent posterior ridge.

The premolars are very simple in construction. The first three are well spaced
apart and have compressed, but thick, conical crowns, without accessory cusps of any
kind, and each is implanted by two fangs. In size, they increase posteriorly and p ® has
a decidedly higher crown than any other premolar. P 4 is smaller than p 2 in every
dimension except the transyerse, this diameter being increased by the addition of a large
internal cusp (the deuterocone) and the crown is carried upon three fangs. In the
smaller species of the genus, such as H. mortoni, p 2 and p 4 are placed close together,
while in the larger forms these teeth are separated by a short space, and the diastemata
between the other premolars and between p + and the canine are relatively somewhat
greater, the enlargement of these teeth hardly keeping pace with the elongation of the
muzzle. In the European species, #. magnum, the arrangement of the premolars is
somewhat different, p 2, ® and 4 forming a continuous series, while p + and 2 are quite
widely separated.

The molars are relatively quite small; m 2 is the largest and m ? the smallest of the
series. The crowns are low and bunodont, bearing six tubercles arranged in two trans-
verse rows. ‘The hypocone, though functionally important, is decidedly smaller than the
protocone, and structurally is still a part of the cingulum. Schlosser is, however, mis-
taken in supposing that there is any important difference between the American and the
European species of Hlotherium with regard to the position of the protocone. In m 3,
which has a more oval crown than the other molars, the sexitubercular pattern is
obscured by the development of numerous small tubercles upon the hinder half of the
tooth. The cingulum of the molars is quite strongly marked, especially upon the ante-
rior and posterior faces.

B. Lower Jaw.—The incisors resemble those of the upper jaw, except that they are
of more nearly equal size and somewhat more spatulate shape ; 1 y is little enlarged and
is much smaller than the corresponding tooth in the upper jaw.

The canine is a very large, recurved tusk, like the upper one in size and shape; it
bites between the upper canine and enlarged external incisor, the three teeth together
making up a very formidable lacerating apparatus. An interesting hint as to the habits
of this animal is given by a peculiar mode of wear of the lower canine which occurs in
some well-preserved specimens. In these we find a deep groove on the posterior face of
the tooth, beneath the enamel cap and close to the level of the gum. No other tooth can
reach this point to cause such a mode of attrition, and the groove is doubtless due to the
habit of digging up roots with the lower tusks; the pull of the roots, especially when
covered with sand or other gritty material, would naturally wear such a groove.* The

* This ingenious and highly probable explanation of a somewhat puzzling fact was suggested to me by my
colleague, Prof. C. F. Brackett.

276 THE OSTEOLOGY OF ELOTHERIUM.

same explanation applies to the curious notches sometimes worn in the external upper
incisor. ‘The numerous specimens examined do not indicate that there was any difference
between the males and the females in the size of the canines, the tusks being invariably
large and powerful. If, as here suggested, the canines served other purposes than those
of weapons, the lack of any such sexual difference would be intelligible enough.

The premolars are very simple and quite like those of the upper series in shape;
their crowns are massive, compressed cones, without additional cusps. The cingulum is
usually prominent, but varies in the different species. P is much the highest of the
series, especially in #. imperator, where it rises to the full height of the canine, and gives
a very characteristic appearance to the lower dentition. Pz has its posterior face flat-
tened, forming an incipient fossa with a number of small tubercles in it. P sand ; stand
quite close together, and p ; is separated by a short space from the canine, while p; is
isolated by considerable diastemata both in front of and behind it.

The lower molars are small in proportion to the size of the jaw and to the space
occupied by the premolar series. In size they increase posteriorly, and they have a
simple, quadritubercular pattern, the crowns surrounded by a strong cingulum. There
is much variation in the development of the fifth or posterior unpaired cusp (hypoconu-
lid); it is frequently absent and represented only by a strong cingulum, though some-
times it is present as a distinct cusp on m > or mz. It is less commonly found on m 3
and only in the very large /. leidyanum is it well developed.

The Milk Dentition—The temporary canines and incisors differ from the permanent
ones only in size. It is uncertain whether the first premolar, in either jaw, has a prede-
cessor in the deciduous series, none of the specimens distinctly showing such a predecessor.
In one individual, howeyer, the tip of p 1 is just visible in the centre of a large alveolus,
from which a milk-tooth has apparently been shed. If this change does actually occur, it
must take place at an early stage, and, on the whole, it seems probable that, at least in the
upper jaw, the number of deciduous premolars is four. Dp 2 has a compressed, elongate,
conical crown, without accessory cusps of any kind; it is carried on two widely separated
fangs, and is isolated by diastemata both in front of and behind it. Dp 2 consists of
three principal cusps. The antero-external cusp (protocone) is an acutely pointed pyra-
mid, while the postero-external cusp (tritocone) is lower and smaller. The internal cusp
(tetartocone) is posterior in position and placed on the same transverse line as the trito-
cone, while between the two is a small conule. The cingulum is distinct on the front and
hind faces, obscure on the outer and absent from the inner face of the crown. Dp ¢ is
molariform, but differs somewhat from the molar pattern in the fact that the postero-

internal cusp is even more distinctly an elevation of the cingulum and that the posterior
conule is double.

THE OSTEOLOGY OF ELOTHERIUM. 277

The lower milk-premolars are even simpler than the upper; dp 5 and y are com-
pressed and conical, without accessory cusps, but with serrate edges and sharply-pointed
summit. Each of these teeth is supported upon two fangs. Dp < is of the usual artio-
dactyl type, consisting of three transverse pairs of cusps, of which the median pair is the
largest, and the anterior pair the smallest. A small talon is formed by the elevation of
the cingulum in the median line, behind the posterior pair of cusps.

This account of the milk dentition applies only to £. mortoni; I have not seen these
teeth in the larger species.

Measurements.

| No. 11156 | No. 10885 | No. 11009 No. 11440

|
Upper dentition, length I1to M3...... Ramee ve te pho nar acti Seti | | 20.270
Ce 2 TON Games, IOMGWISc coc onevcdoos Goce benodo cose oeuE 118 104 | .064 .065
(C joReMOlayr SOmies, NeMADN. -s2corcccsoa0cancacnu0ds Ste kiacsens .238* 175 .124 .113
CRMTOE, HIN DOM, GHANA, .16 congeadosaocosssedscaud | .048* .046 | .082
GG ee Mans Versen diam ecleneeemeniceia te emcee nee .03885* .042 .022
Come Pale mlen thie sanss acne ease tron oe aac er Toes ian Gaee> .080* | 024 .019
CEE TED De WRG File OUR elo oA Seo te i | 088% | .088 02 een e O23
EID Be lye KBs Meee ee RS Me ee mR AL Rtgs | 041 028 | .028
CMe AR pmeaL ier haar teniara® ME Se Ra ae RAS Mae | . .085* | 081 0195 .018
cS > TST TL iepayetitn d ea 0 ire i a ge emecOS52. le 038 | - .020
E09 OOD SATIN pe nee ee eer aA Gai oe alee eget | .086 | .019
05. INL Bh TTC dle Soe RR RR ea | 04 | 085 025 | .028
cy 9 "007 SRR g Oar | 039 0235 | 024
6G) NEG) Tan iatllie gas tison Raab O Une suaap Ore cone et SE eaneeree ne OR 2 ORE I ae a ies
ces) 6 0G" RENT ten rN rascal ae ga eI | 088 022 | «0215
Lower dentition, length IT1toM3............................ | -432* | | 261
GG AOA SOMES, IEMA cc copcocs oss Goeoooe Guns Sosscnngen 121% | -108 | | 070
Re == premolar series: lengthy. o.% s. cnc os 0s se0ees tanec one } Bile? |) SRD 126
a 1B il Nena noes deino.ce ceeds Coen CD ODS Oe aE eaGEn coon. | .028* | .026 | 017
s cc INGA! Oi, GiOneas opbecbocossEnesbes sooo LeRsBEEGSS -026* | -019
CRaPEPEOM enh sa sais oe eres teas te SNE Sie BME lipeeeO3tcans | Giar.0asit ste] .020
Si, Bae STy@TCANIP Seas Senet ee See eer .038* | .023
“i 1D B, IGMEHI -oocoousonas oo pedoegaesouedbeeadoDanL 00000 .046* .043 027
RRC Mee pli beet me Tae Ade teh et Pee o61* | 031
Gs “TBA. Thana de Se pe Uae Sle dese ao Spec Soa Meee eb bras 046 | 037 | 025
3) 1G Teele Nice sane seer ce Reena rare Maen, 0445 | 020
Gl TAT Te Ten geoceecs ook acu OI eee enn 037 | 081 0215
65-7) (GS SIN Ha oe pes o> (geen eR REE MEE RSE ERIE EBC aese 029% 027 | 018
CINE DBT Miggooneee dosece aed: Ae aNROOU Ce Oar oR esEee .0895* | .035 | 0225
Ob. 08 ORT St Hea a: cob bE ROBO REAR eRe Re meee aee .036* .030 > neorG
ci) SUL S) Deming ope’ canbe Otte: Eee nae mm nat .043* 039 | 0245
a Scam UL eecya ee a te ore eA weioe sis lemons 037% | 028 | .016
|

*No. 11161.

278 THE OSTEOLOGY OF ELOTHERIUM.

Il. Tue SKULL.

The skull of Hlotherium is one of the most remarkable features of this very curious
animal. It is characterized by great length and slenderness, with the supraoccipital and
nasal bones lying in the same horizontal plane. The muzzle is exceedingly long and
narrow, and tapers somewhat anteriorly, though expanded by the sockets of the great
tusks; the orbit has been shifted far back, its anterior border being, in some species, over
m 2, and in others above m 8. The cranium is short and of absurdly small capacity,
which, with the great temporal openings, gives an almost reptilian appearance to the
skull when viewed from above or below. The sagittal crest is very high and thin, and
the zygomatic arches, though rather short, are enormously developed. One of the most
peculiar features of the skull is the great, compressed plate which is given off from the
ventral surface of the jugal and descends below the level of the lower jaw, and this gro-
tesque appearance is further increased by two pairs of knob-like processes on the ventral
borders of the mandible. The occiput (Pl. X VIII, Figs. 1,2) is high and very broad at -
the base, but narrowing rapidly to the summit; above the foramen magnum it forms a
broad, flat projection of almost uniform breadth, with a very deep fossa on each side of it.

The basioccipital is stout and rather short, keeled in the median ventral line and
slightly contracted to receive the auditory bulle ; at its junction with the basisphenoid it
forms a pair of small, roughened tubercles. The exoccipitals are yery large bones, espe-
cially in the transverse direction along the base of the occiput, dorsally they narrow fast.
Above the foramen magnum they form the very broad, prominent and nearly square pro-
jection which has already been mentioned ; this is thick and is filled with cancellous bone,
the fossa for the vermis of the cerebellum making but a slight depression upon its internal
face. On each side of the projection is a large and deep triangular fossa, which, how-
ever, is not confined to the exoccipital, the periotic and squamosal both being concerned in
its formation. The inferior part of the exoccipital extends widely outward, reaching to
the line of the glenoid cavity, and ending in the large, prominent and massive, but not
elongate paroccipital process. In this region the exoccipital is brought very close to the
zygoma, but, ventrally at least, does not quite touch it, a narrow band of the tympanic inter-
vening between them. The foramen magnum is strikingly small and of a transversely oval
shape. The occipital condyles are relatively rather small, especially in the vertical dimen-
sion, laterally they are well extended, and they are widely separated both aboye and below.
In the very large /. imperator the external angles of the condyles are abruptly truncated
in a curious way, and bear flat articular surfaces, though in some individuals this trunca-
tion is found only on one side; while in the smaller species the condyles are of the usual
form. The supraoccipital is a large bone, widest at the base (7. ¢., the suture with the

exoccipitals) and narrowing dorsally. Superiorly it is drawn out into two posterior wing-

THE OSTEOLOGY OF ELOTHERIUM. DATES)

like processes, such as are found in Oveodon and other White River ungulates. Between
these wings the hinder face of the bone is concaye and at the bottom of this concavity are
two small, but profound pits. The supraoccipital is continued over upon the roof of the
cranium and forms a part of the sagittal crest.

A considerable part of the periotic is exposed on the surface of the skull, at the bot-
tom of the lateral occipital fossa, where it is enclosed between the exoccipital and the
squamosal ; it does not give rise to any distinct mastoid process.

The oceiput of the European species, Z. magnum, as figured by Kowalevsky (76,
Taf. XVII, Fig. 5), is different in many details from that which characterizes the Amer-
ican species. It has more of an hour-glass shape, not so wide at the base, more contracted
in the middle and more expanded at the top, but with much less conspicuous wing-like
processes, and it has no such projection above the foramen magnum, nor such deep lateral
fosse. The condyles are larger and of an entirely different shape, haying their principal
diameter vertical, instead of transverse. The paroccipital processes are longer, more com-
pressed and not so widely extended laterally. -The foramen magnum is large and of more
nearly circular outline.

The basisphenoid is narrower than the basioccipital and is not keeled on the ventral
surface, but is otherwise like that bone. So much of its course is concealed by the union
of the palatines and pterygoids along the median line that its length cannot be deter-
mined, while the presphenoid is nowhere exposed to view.

The tympanic is very extensively developed (Pl. X VIII, Fig. 1). Part of it is inflated
into an oyal, somewhat flattened and rather small auditory bulla, which differs from that
of Hippopotamus and of all existing suillines in being hollow and not filled up with
spongy tissue. On the outer side of the bulla the tympanic is extended as a narrow strip,
which broadens considerably between the squamosal and the exoccipital, with both of
which it articulates suturally, as well as with the alisphenoid in front. The bulla itself
terminates anteriorly in a blunt spine. ;

The alisphenoid is small and forms yery little of the side of the cranium. It is most
elongate antero-posteriorly along the ventral line, but has hardly any distinctly developed
pterygoid process. At the line of the sphenoidal fissure, which notches but does not per-
forate the bone, the alisphenoid is narrowed, to expand again at its suture with the parie-
tal and frontal. The orbitosphenoid is relatively rather large, but is low in the vertical
dimension, and does not extend upward into the orbit proper. Two sharp ridges on the
external face of the bone enclose a V-shaped grooye, in which lie the optic foramen and
foramen lacerum anterius.

The parietals are very large proportionately to the size of the cranium, but quite
small as compared with the entire length of the skull; they roof in most of the cerebral

Ne By VO, WiIDK, DY A

280 THE OSTEOLOGY OF ELOTHERIUM.

chamber, but toward the yentral side they rapidly contract, forming narrow strips
between the squamosal and frontal. Throughout their length the parietals unite to form
the very high, thin and plate-like sagittal crest, which is one of the most characteristic
features of the skull. In the European species, #. magnum, this crest has a remarkably
straight and horizontal course, but in the known American species it is gently arched
from before backward. Large sinuses are developed in the parietals, so that the cerebral
chamber is eyen smaller than it appears to be, when viewed from the outer side. These
sinuses extend over the entire roof of the cerebral fossa, even invading the supraoccipital ;
they appear to be traversed by numerous small trabeculee, the ends of which are seen, in
the sagittal section, embedded in the matrix which fills the sinuses.

The frontals are much larger than the parietals. In the postorbital region they are —
very narrow, in conformity with the very small size of the brain, but at the orbits they
expand widely to form the broad, lozenge-shaped forehead, which is convex from side to
side, though slightly depressed, or “dished” in the middle; the supraciliary ridges are
very inconspicuous. Anteriorly the frontals diverge to receive the nasals between them,
sending forward long, pointed nasal processes, which, owing to the great elongation of the
muzzle, are widely separated from the premaxillaries. The orbit is large and projects
prominently outward ; it is completely encircled by bone, the long and massive postorbital
process of the frontal uniting suturally with the shorter process of the jugal. The orbits
do not rise above the leyel of the forehead, as they do in Hippopotamus, and present
more anteriorly, less directly outward, than in that animal. Mention has already been
made of a groove on the orbitosphenoid, which terminates below and behind in the fora-
men lacerum anterius; this groove is continued upward and forward upon the frontal,
steadily widening as it advances. The postero-superior ridge bounding the groove is the
more prominent ; it extends almost to the postorbital process, from which it is separated
by a distinct notch, while the antero-inferior ridge dies away within the orbit. In most
of the American species the forehead rises yery gradually and gently behind to the sag-
ittal crest, but in F. imgens the rise is much more sudden and steep. The frontal sinuses
are large, giving the conyex shape to the forehead which has been described; these
sinuses appear to communicate with those formed in the parietals.

Except posteriorly, the sywamosal forms but little of the side-wall of the cranium, its
suture with the parietal curving abruptly downward and forward; its compressed and
prominent hinder margin forms nearly the whole of the lambdoidal crest, though a con-
tinuation of it extends upward upon the supraoccipital, ending in the wing-like processes
of that bone. The zygomatie process is enormously developed ; it extends widely out-
ward from the side of the skull as a massive, vertical plate, which is shaped much as in

Hippopotamus, and is not continued forward as a broad, horizontal shelf, such as is found

THE OSTEOLOGY OF ELOTHERIUM. i 281

in Sus. The superior border curves. upward into a great, hook-shaped process, which
resembles that seen in Merycochawrus, and gives a highly characteristic appearance to this
region of the skull. That portion of the zygomatic process which is directed anteriorly
is short and, though massive, is much less so than that which extends out laterally ; in
front it is received into a notch of the jugal. The glenoid cavity is large, transversely
directed and quite deeply concave, though the postglenoid process is not strongly deyel-
oped and is hardly more conspicuous than the preglenoid ridge. This disposition is
unusual among the ungulates, but it occurs also in the Eocene genus Achanodon and in
the modern Dicotyles. The glenoid cavities of the two sides are very widely separated,
their inner margins lying external to the line of the paroccipital processes. The posttym-
panic process of the squamosal is small, and is closely applied to the paroccipital process.
The shape of the zygomatic arches, together with the extreme narrowness of the cranium
proper, causes the temporal openings to be very large and to appear widely open when
the skull is viewed from above. ‘These openings are, however, less extended transversely
and more antero-posteriorly than in Hippopotamus, while in Sus they are hardly visible
from above.

The jugal isa very remarkable bone and constitutes one of the most extraordinary
features of the Hlotheriwm skull. Posteriorly it is notched to receive the zygoma, and
sends out a process along the ventral face of that bone, extending to the preglenoid ridge.
The jugal forms the inferior half of the nearly circular orbit, and for this purpose its
dorsal border is made deeply concave, giving off a stout postorbital process to meet that
of the frontal, while anteriorly it is moderately expanded upon the face in front of the
orbit, where it is wedged in between the lachrymal and the maxillary. The most pecu-
har feature of the jugal, however, is the immensely developed vertical plate, which
descends from beneath the orbit downward and outward to below the level of the ven-
tral border of the mandible, recalling the similar, but much Jess massive processes found
in certain edentates, e. g., Megatherium. ‘These plates are laterally compressed, but quite
thick, and when the skull is viewed from the front, they are seen to diverge quite
strongly downward ; their shape varies in the different species. In the very large forms
from the Protoceras beds, such as #. imperator, the process retains its plate-like form
throughout, its free end being only moderately thickened. This appears to be true also
of £. mortoni, though my material is not sufficient to allow me to make this statement
positively, but in the large species from the Titanotherium and Oreodon beds (£. ingens)
it forms a club-like thickening at the tip, which in /. ingens is coarsely crenulate on the
posterior border (see Pl. XVII). These processes are, so far as is yet known, quite unique
among the hoofed mammals, and it is difficult to form eyen a conjecture as to what their

functional significance may have been. Some misunderstanding has arisen as to the spe-

282 THE OSTEOLOGY OF ELOTHERIUM.

cies in which these jugal plates are found. Nothing is known concerning their presence
or absence in the European representatives of the genus. Leidy’s material gave him no
reason to suspect their occurrence in the species described by him, and he consequently
restored the zygomatic arches without them (769, Pl. XVI). Marsh first discovered the
processes in a skull of the species named by him /. crasswm, and it has sometimes been
assumed that they were more particularly characteristic of that form. As a matter of
fact, they have been observed in all of the American species of which well-preserved
skulls are known, viz., 2. mortoni, EL. ingens, and E. imperator, and, in all probability,
all the American forms, at least, possessed them.

The achrymal is a rather large bone and forms nearly half of the anterior boundary
of the orbit. On the face it is expanded into quite a large plate, which articulates below
with the jugal, in front with the maxillary, and above with the frontal, the long anterior
process of which prevents any contact between the lachrymal and nasal. In Aippopota-
mus the very short, broad frontal has no anterior process, and so the nasal and lachrymal
are connected, as they are also in Sus. Within the orbit the lachrymal is but little
extended ; the foramen is single, very small, and placed inside the orbital margin. The
lachrymal spine is very low.

The nasals are narrow, slender and very much elongated. Their greatest width is
at the anterior end of the nasal processes of the frontal, and here is also their greatest
transverse convexity ; ‘from this point they narrow and flatten, both in front and
behind. Anteriorly they contract very gradually and terminate in sharp points, with
their free ends quite deeply notched. In /. wmgens the nasals appear to be relatively
shorter than in the other species. In /Hippopotamus these bones have much the same
shape as in Hlotherium, but they narrow more abruptly behind the poit of greatest
width, and their free ends are not notched. In Sus the nasals are truncated posteriorly
and in front their free tips project far beyond the borders of the premaxillaries.

The premaxillaries ave very large and heavy bones, the horizontal or alveolar portion
especially so. Posteriorly, this portion is constricted, forming a groove for the reception
of the lower canine, expanding again in front to carry the large incisors. The palatine
processes are not much developed, the very large incisive foramina leaving but little
space for them; the spines are long and slender, extending behind the canine alveolus.
The ascending ramus of the premaxillary is low and rises gradually behind, and though
broad at first, it rapidly becomes very slender, terminating behind in a fine point.
Though these bones in Hlotheriwm haye a very different appearance from the immensely
enlarged premaxillaries of Hippopotamus, yet both may have been formed by divergent
modifications of a common plan.

The maxillary is greatly extended antero-posteriorly, in correspondence with the

THE OSTEOLOGY OF ELOTHERIUM. 283

elongation of the whole muzzle ; its facial portion is low, gradually diminishing in height
forward, where its suture with the premaxillary forms a very gentle, sweeping curve.
The longest suture of the maxillary is that with the nasal, the connection with the frontal
being very short, owing to the. extension of the lachrymal. Posteriorly, this bone pro-
jects but little beneath the orbit, which has an imperfectly developed floor, and the pro-
jection which it sends out to the jugal is much less massive than in Hippopotamus. The
face gradually narrows forward, until it reaches the infraorbital foramen, expanding
again in front of the foramen and swelling out into the prominent canine alveolus. The
palatine processes of the maxillaries are long and narrow, and as the molar-premolar
series of the two sides form almost straight and parallel lines, the bony palate is of nearly
uniform width, slightly concave transversely, but almost plane antero-posteriorly. In
front, these palatine processes are deeply emarginated by the large incisive foramina, and
in the median line are still further notched to receive the long premaxillary spines.

The palatines make up but very little of the bony palate, forming only a narrow
strip in front of the posterior nares, and narrow bands along the sides. The palatal
notches are small and shallow. The pterygoids are elongate, but quite low; there are no
hamular processes or pterygoid fossee; the two bones meet suturally along the median
dorsal line, completely concealing the presphenoid from view. The posterior nares are
long, narrow and low, extending forward to the middle of m 2; the opening gradually
contracts posteriorly, where it becomes very narrow, while the side-walls slope upward
and die away upon the alisphenoids. Anteriorly the nares are divided by the very large
vomer, Which is distinctly visible, and which at its hinder termination expands into a
transverse plate, articulating with the palatines. The meeting of the two pterygoids
forms a small canal, which appears to overlie the whole length of the posterior nares and
to open forward into the nasal chamber on each side of the vomer. This is a very excep-
tional arrangement, and I am unable to suggest what its functional meaning may be
(see Pl. XVIII, Fig. 1, c).

The cranial foramina are, in some respects, quite peculiar. The condylar foramen is
large and conspicuous, being placed well in front of the condyle ; it is, however, smaller
than in the specimen of /. magnum which Kowaleysky has figured. The close approxi-
mation of the paroccipital and stylomastoid processes, and the outward extension of the
tympanic between them, have given a somewhat unusual position to the postglenoid and
stylomastoid foramina; they are crowded close together at the postero-external angle of
the auditory bulla, and both of them perforate the enlarged tympanic bone. The fora-
men lacerum posterius forms a long, narrow and curved slit at the postero-internal angle
of the bulla, while the foramen lacerum medium and the opening of the eustachian canal
occupy their ordinary position at the front end of the bulla. No distinct carotid canal is
visible externally.

284 THE OSTEOLOGY OF ELOTHERIUM.

Kowaleysky inferred from the study of his specimen that the foramen ovale “ nicht
als selbstiindiges Foramen existirte, wie z. B. bei den Ruminanten, sondern mit dem For.
lac. med. verschmolzen war, wie bei den heutigen Suiden und bei Hippopotamus” (’76,
p- 483). This is probably a mistake; at all events, it is not true of the American
species, in which the foramen ovale is a long, conspicuous opening, of oval shape, perfo-
rating the alisphenoid. As in the ungulates generally, there is no separate foramen rotun-
dum, that opening being fused with the foramen lacerum anterius. The latter is a large
and somewhat irregular opening, which notches the anterior border of the alisphenoid,
passing between that bone and the orbitosphenoid. The optic foramen is small and well
separated from the foramen lacerum anterius, lying in front of and at a slightly higher
level than the sphenoidal fissure ; it does not open so far forward as in L. magnum, and,
in consequence, it does not form such a remarkably elongated canal as in the European
species (see Kowalevsky, 76, Taf. XVI, Figs. 1 and 3, dd), but, on the other hand, it is
far from being a simple perforation of the orbitosphenoid, such as occurs in the recent
ungulates. This elongation of the optic canal should probably be correlated with the
very small size of the brain, which would seem to have been relatively smaller than in
the ancestors of the genus. Though the orbits are far behind their primitive position,
the backward shifting of the optic tract would seem to have kept pace with the change in
the position of the orbits.

The posterior palatine foramina are large and conspicuous openings, placed at the
maxillo-palatine suture, and separating the two bones at these points; the palatine plates
of the maxillaries are deeply grooved for some distance in front of the foramina. The
incisive foramina are likewise large, invading both the maxillaries and the premaxilla-
ries; indeed, their size preyents the development of any considerable palatine processes
on the latter bones. These foramina are in very marked contrast to those of ippopota-
mus, in which the enormously expanded and massive premaxillaries are perforated by
two small and widely separated openings; in Sus also the incisive foramina are propor-
tionately much smaller than in Hlotheriwm. The infraorbital foramen is large and is
separated from the orbit by a considerable interval, opening above the anterior border of
p 2. In front of the foramen a deep groove channels the outer face of the maxillary for
a short distance. The canal itself is much elongated, in correspondence with the great
length of the jaws, and its posterior orifice, within the orbit, is very large. The lachry-
mal foramen, which is single, is quite small and is placed inside of the orbit.

The supraorbital foramen is subject to some variation in the different species. In
E. ingens, from the Titanotherium beds, these openings are of good size, are placed quite
near to the median line, and have well-marked vascular channels running forward from

them. In specimens of #, mortoni from the Oreodon beds, and in the very large species

THE OSTEOLOGY OF ELOTHERIUM. 285

(£. imperator) from the Protoceras beds, the openings haye become minute; they are
shifted laterally and haye no anterior grooves leading from them.

The mandible is not the least curious part of this remarkable skull. The horizontal
ramus is extremely long and nearly straight, with an almost horizontal inferior border.
The depth and thickness of the ramus yary considerably; even in skulls of the same
length the mandible is decidedly more slender in some specimens than in others. The
materials are, however, not yet sufficient to determine whether this difference is of a spe-
cific, sexual, or merely individual character. -A remarkable knob-like process is given
off from the ventral border of the mandible, beneath p z, which is subject to much -yari-
ation in shape and elongation, in accordance with the age and size of the animal. In
young indiyiduals still retaining the milk-dentition, the process is a mere rugose eleva-
tion, and in the adults of the smaller species it is hardly more than a knob, while in the
large forms it becomes greatly elongated and club-shaped. No marked difference in this
regard is observable between the species from the upper and those from the lower hori-
zons of the White River formation, the process being relatively quite as long and promi-
nent in Z. ingens from the Titanotherium beds, as in /. imperator from the Protoceras
beds, but in the huge John Day species it has become particularly long and heavy.

The symphysis is quite long and very thick and massive; the two rami are indis-
tinguishably fused together and laterally expanded, so as to somewhat resemble the sym-
physis of Hippopotamus, though not attaining any such extreme degree of massiveness as
in the modern genus. The chin is abruptly truncated and flattened, and rises very
steeply from below; on each side, beneath or a little behind the canine alveolus, there
arises from the ventral border a second club-shaped process, similar to, but much heavier
and more prominent than the posterior process already described. These two pairs of
knobs give to the jaw a highly peculiar and characteristic appearance ; they form another
of the enigmatical features of the Elotherium skull, for it is difficult to imagine what part
they can have played in the economy of the animal.

The two inferior dental series pursue a nearly parallel course, diverging backward
but little, but behind the molars the two rami turn outward and diverge rapidly, so that
posteriorly they are very widely separated, in correspondence with the great interval
between the glenoid cavities of the two squamosals. The angle of the mandible is prom-
inent and descends below the ventral border of the horizontal ramus, much as in Hippo-
potamus, though not to the same extent. The ascending ramus is not high, but of con-
siderable antero-posterior extent. The masseteric fossa is quite small, but very deeply
impressed, and is situated quite high upon the side of the jaw. The condyle is relatively
little raised above the level of the molar teeth, and it is sessile, hence inconspicuous,

though it is large, transversely expanded, and strongly convex. The coronoid process

286 THE OSTEOLOGY OF ELOTHERIUM.

is strikingly low and small; it is of triangular shape, erect and not at all recurved, and
is separated from the condyle by a yery wide sigmoid notch. The mental foramen is
small, single, and placed below p 3.

Several of the hyoid elements are preserved in connection with the skeleton of L.
ingens which forms the principal subject of this description. The stylohyal is quite long
and slender ; its proximal portion is laterally compressed and yery thin, but moderately
broadened in the fore and aft direction. For the distal two-thirds of its length the bone
is thicker and of a compressed oval section, expanding into a club-shaped thickening at
the lower end, which is excavated for the connecting cartilage. The ceratohyal is con-
siderably shorter than the stylohyal, but of quite similar shape; its proximal end bears
a cup-shaped expansion, beneath which it becomes yery thin and much compressed, but
broadened antero-posteriorly ; the inferior part of the shaft is slender and oyal in section,
with another cup-shaped expansion at the distal end. The epihyal and basihyal haye
not been preserved. The thyrohyal is of remarkable length and slenderness, and obyi-
ously was not codssified with the basihyal; the bone is of subcylindrical shape, with
expansions at the proximal and distal ends.

This hyoid apparatus does not resemble that of any artiodactyl with which I have
been able to compare it. The elements of the anterior arch somewhat resemble those of
Fippopotamus, but are more slender and elongate. In the modern genus, on the other
hand, the thyrohyals are very short, and are ankylosed with the basihyal, a totally differ-
ent arrangement from that which characterizes Hlotherium.

From the foregoing description and accompanying figures it will be obyious that the
skull of Hlotherium is an extremely peculiar one. Among recent animals that of Hippo-
potamus approximates it most closely, and displays, with many striking differences, sey-
eral decided and, it may be, significant resemblances. Some of these resemblances, such
as the straight cranio-facial axis and the long sagittal crest, are of no particular import-
ance, because they occur so yery generally among the primitive ungulates of all groups.
Other similarities, again, are not of this nature. The proportions of the cranial and
facial regions, the degree of backward shifting of the orbits, the relations of the zygo-
matic and paroccipital processes, the broadening of the muzzle, and the general plan of
skull construction, are all similar in the two genera. On the other hand, each genus has
certain peculiarities correlated with its manner of life. Thus, the elevation of the orbits
and the backward displacement of the posterior nares in Hippopotamus are adaptations
to its aquatic habits. Doubtless the extraordinary peculiarities of Hlotherium, such as
the dependent processes of the jugals and the great knobs on the mandible, are of a sim-
ilar nature, though, in the absence of the soft parts, it is difficult even to conjecture what

their use may have been.

THE OSTEOLOGY OF ELOTHERIUM. 287
Measurements.

No, 11156. No. 10885. No. 11009. No. 11440.

Skull, extreme length on basal line........................... 0.803 20,648 20.460
«« width across zygomatic arches (behind jugal process).. . 2.500 443 2297 -264
66 AINA 1) 546 po sgeodocamounalds Goodeabecad elsjelelejetelalele.ls 133 140 :089 -082
Cranium, length to anterior border of orbit.................... .282 -288 | .198 193
Face, length to anterior border of orbit.-..................... | .518 2.378 270
Occiput breadthwote base reer ee terre eee | 281 252 .160 158
Fe on AGHA TNE Uke eoeodd b GOSAEOE ies oS aU ate eee OR Ia Oe San | .158 .120
Bony palate, length in median line................-.......... 2.376 | 247
AP SOMANE OREM, WOMB, > o choco peoces cob oso ons acosudEeogo0NH 279 Sarat | 146 146
Descending process of jugal, length................-......06- | 830 256 126
Mandible wlenothencyacea eters tele csh cits aerate ever | .659* 608
Es heightaticoronoidsprocessies-eee sees bieas | .253* 171 107
co GI NUN 1) sos omc a ssaustcosdos soc laed anne aoamiacetre tc .133* 091 052

* No. 11161.
Ill. Tue Bra.

Attention has been repeatedly called, in the foregoing description of the skull, to
the extraordinarily small size of the brain-cavity. Even on viewing the skull externally,
this smallness of the cranium proper strikes the observer immediately, and, in connection
with the long, slender muzzle, gives the skull something of a reptilian aspect. When
the cranium is sawn open in longitudinal section, it becomes apparent that the brain is
even smaller than would be inferred from the external view alone, much of the space
being, so to speak, wasted in the great frontal and parietal sinuses which overlie the
whole cerebral chamber. In a large, full-grown skull this chamber will hardly contain
an ordinary human fist.

The olfactory lobes are very large and are connected with the cerebrum by short
thick olfactory tracts. The lobes are not at all overlapped by the hemispheres, but are
entirely exposed for their whole length.

The cerebral hemispheres are relatively small, though they are, of course, much
larger than the other segments of the brain; so short are they that they do not extend
over the olfactory lobes in front, or the cerebellum behind. In shape, they are low and
wide, narrowing gradually forward, but with blunt anterior termination. The frontal
lobe is yery small, for the frontals take but little share in the roof of the cerebral chamber.
The parietal lobe, on the other hand, is relatively large and forms the greater part of the
hemisphere, for there is, properly speaking, no occipital lobe, the occipital bones not tak-
ing any part in the formation of the cerebral fossa. The temporo-sphenoidal lobe is also
quite large and prominent, but is short antero-posteriorly. The brain-cast shows that the

As 1 Se AVOWs BIDS 1

988 THE OSTEOLOGY OF ELOTHERIUM.

hemispheres were conyoluted, but the conyolutions are so feebly marked that they are
hardly worth description. It is obvious, howeyer, that the gyri were fewer and simpler
than in any of the modern ungulates.

The cerebellum is rather small, though the cerebellar fossa has a vertical diameter
not much less than that of the cerebral fossa. Antero-posteriorly the former is quite
short and its transverse breadth is not great. This breadth is still further reduced by the

relatively very large size of the periotic bones which extend freely into the fossa.
IV. THe VERTEBRAL CoLumy.

The vertebral formula is: C 7, Th ? 138, L. 6,S 2, Cd 15 -+

The atlas (Pl. X VIII, Fig. 3) is very wide transversely, and at the same time it is of
considerable antero-posterior extent, a shape which recalls that of Anoplotherium, rather
than that of the recent ruminants or suillines. The anterior cavities for the occipital
condyles are deep and wide, but low and depressed. Dorsally, these cotyles are widely
separated by a broad, but not very deep emargination of the neural arch, nor do they
approximate each other very closely on the ventral side, a notch of considerable width
intervening between them at this point. The neural arch is thick and heavy, but short
from before backward and quite narrow transversely ; it is also low, not arching strongly
toward the dorsal side, and nearly smooth, being free from any but the most obscurely
marked ridges. The foramina perforating the arch for the first pair of spinal nerves are
unusually large. The neural spine is rudimentary and forms only an inconspicuous
tubercle. The neural canal is low and broad, forming a transversely directed ellipse.
The inferior arch is considerably more elongated antero-posteriorly than the neural, and
has but little transverse curvature, except laterally, where it rises to form the sides of
the neural canal. The hypapophysis is represented by a small, backwardly directed
tubercle, which arises from the hinder margin of the ventral arch, and occupies the same
position as in the pigs, but is much less strongly developed. The articular surfaces for
the axis are low and broad, and haye a very oblique position, presenting inward toward
the median line, almost as much as backward; they have also a slight dorsal presenta-
tion. In shape, they are yery slightly concave and are surrounded by prominent
borders. The facet for the odontoid is wide, and deeply conecaye in the transverse direc-
tion, but quite short antero-posteriorly. This facet is connected at the sides with those
for the centrum of the axis, but distinct ridges are formed along the line of junction.

The transverse processes of the atlas extend out widely from the sides of the arch,
attaining their greatest transverse breadth along the posterior line; they are also very
long in the fore-and-aft direction, reaching far behind the surfaces for the axis. For

most of their course the transverse processes have thin borders, but posteriorly the

THE OSTEOLOGY OF ELOTHERIUM. 289

margin becomes much thicker and more rugose. The vertebrarterial canal, which is
notably small, occupies much the same position as in Sus, opening posteriorly upon the
dorsal side of the hinder border. The anterior extension of the transverse processes
has conyerted into foramina (atlanteo-diapophysial) the notches for the inferior branches
of the first pair of spinal nerves. On the ventral face of each process is a large fossa,
enclosed between the side of the inferior arch and the greatly thickened posterior border
of the process. The resemblance in shape to the atlas of Anoplotherium, to which atten-
tion has already been called, affects more particularly the form of the transverse processes
but they are more extended transversely than in that genus and are not so pointed at the
postero-external angles.

The avis (Pl. XVIII, Fig. 4) is a short, but very massively constructed bone,
which in general shape and appearance resembles that of Lippopotamus. The centrum is
short, anteriorly yery broad and depressed, but thickening posteriorly, and with a nearly
circular and slightly concave hinder face. A strong and prominent keel runs along the
ventral face of the centrum, enlarging backward, and terminating behind in a trifid
hypapophysis. The odontoid process is short, heavy and conical, with no tendency what-
ever to assume the depressed and flattened shape which occurs in so many White River
ungulates. The yentral articular surface of the odontoid seems like something super-
added to the process itself, for it is clearly demarcated by a groove running all around it,
and projects slightly in front of the body of the process. On-the dorsal side of the
centrum a broad and well-defined ridge runs backward from the odontoid along the floor
of the neural canal. The atlanteal articular surfaces are yery broad and low, not rising
so as to enclose any part of the neural canal. They are very oblique with reference to
the median line of the centrum, with which they form angles of about 45°. These
surfaces are slightly convex in both directions, and ventrally they project much below
the level of the centrum.

The transverse processes are short, thin and compressed, much less massive and
widely extended than in Hippopotamus ; they are perforated by very large foramina
for the vertebral arteries. The pedicels of the neural arch are low and short, but very
heavy ; they are not pierced for the passage of the second pair of spinal nerves, as they
are in Hippopotamus and in some of the pigs. The neural canal is decidedly small,
especially its anterior opening; behind, it enlarges somewhat, particularly in the dorso-
ventral dimension, the posterior opening being high and narrow, while in Hippopotamus
it is low and broad. The neural spine is a large plate which is very thin in front, but
becomes thick and massive behind, ending in a broad rugosity. This spine resembles
that of Hippopotamus, but is not produced so far backward and does not overhang ‘the

third cervical, The pestzygapophyses are large, slightly concaye, and present obliquely

290 THE OSTEOLOGY OF ELOTHERIUM.

outward, as well as downward; their bases are separated by a broad and deep grooye,
which is continued upward upon the posterior side of the neural spine.

The third cervical vertebra also bears a considerable resemblance to that of Aippopo-
famus, differing only in some points of detail. The centrum is short, heavy and moder-
ately opisthoccelous, depressed, but increasing posteriorly in vertical thickness. It bears
-a strong yentral keel, which terminates behind, as in the axis, in a trifid hypapophysis.
The pedicels of the neural arch are not, as in the pigs, pierced by foramina for the
spinal neryes; they are low and short, but very thick, and the neural canal is strikingly
small. The dorsal side of the arch is short, broad and nearly flat. The neural spine is
remarkably well-developed (when the anterior position of the vertebra is taken into
account), rising as high as that of the axis. It is rather thin and compressed, although its
base occupies the whole fore-and-aft length of the arch. From the base, however, it rapidly
tapers upward and terminates ina small, rough tubercle. In Hippopotamus the third
cervical has an even better developed neural spine, not higher, but broader and less
tapering than in Llotherium. The prezygapophyses are large, oblique and somewhat
conyex ; they are placed very low, so that their inferior margins are separated from the
centrum only by narrow notches. The posterior zygapophyses are much larger and
more prominent than the anterior pair; they are also less oblique in position and are
raised higher above the centrum, corresponding to the posterior elevation of the neural
arch. The transverse. process is a compressed plate, which has no great vertical height,
but is well extended from before backward, exceeding the centrum in length; the pos-
terior portion of the process is thickened and recurved, ending in a rugose hook. The
absence of any distinctly marked diapophysial element distinguishes this vertebra from
the corresponding one of Hippopotamus and Sus, and in the latter genus the inferior
lamella is more slender and rod-like, while the spinal nerves make their exit through
foramina in the pedicels of the neural arch.

The fourth cervical vertebra is different, in many respects, from the third. The
centrum is somewhat shorter and is less distinctly carinate on the ventral side, but is more
decidedly opisthoccelous. The neural arch is remarkably short in the antero-posterior
dimension, so that the articular faces of the postzygapophyses actually extend forward
beneath those of the anterior pair, which gives to the pedicel of the neural arch, when
seen from the side, a curiously notched appearance. The neural spine is higher, but
more slender and recurved than that of the third cervical. The transverse process is
altogether different in shape from that of the latter. It has, in the first place, a very
prominent diapophysial element, which projects outward as a heavy, depressed bar,
thickened, rugose, and slightly upcurved at the distal end. In the second place, the
inferior lamella is much higher vertically, but decidedly shorter from before backward,

THE OSTEOLOGY OF ELOTHERIUM. 291

In Hippopotamus and in Sus this vertebra is very similar to that of Elotherium, but the
neural spine is notably heavier.

The fifth cervical vertebra las an even shorter neural arch than the fourth-and a
much higher neural spine. ‘The spine tapers rapidly from the base upward and becomes
very slender, but it is nearly straight and only slightly recurved. The neural canal
is somewhat larger than in the fourth vertebra, but, as in all the cervicals, it is strikingly
small as compared with the size of the vertebra as a whole. The diapophysis is strong
and prominent, but more slender than on the preceding yertebra, while the inferior
lamella, though relatively short from before backward, has attained great vertical height
and is strongly everted. In Elotheriwm the fifth vertebra is of the same type as the sixth,
whereas in Aippopotamus it more nearly resembles the fourth.

The sixth cervical is very like the fifth, but displays certain obyious differences.
Thus, the neural arch is even shorter antero-posteriorly, and the neural spine is higher,
heavier and much more strongly recurved. The postzygapophyses are decidedly smaller
and are very characteristic in their markedly oblique position, for they rise steeply back-
ward in a way that occurs in none of the other yertebree. The diapophysis is shorter but
heavier than that of the fifth, while the inferior lamella is of similar shape, but larger,
higher and with the free margin more thickened. In Hippopotamus this vertebra has
much the same construction as in Lotherium, but the spine is shorter and more massive
and the inferior lamella is much larger. In Sus the sixth cervical bears considerable
resemblance to that of the White River genus.

The seventh cervical is characterized by the height and thickness of the spine, which
in these respects much exceeds that of the sixth. This spine tapers superiorly, but
expands again at the tip into a rough tubercle. The posterior zygapophyses stand at a
higher level than the anterior pair and are unusually concave. The peculiarities seen in
the postzygapophyses of the sixth and seyenth vertebree are to provide for the curvature
of the neck, which changes its direction at this point. From the occiput to the sixth
cervical the neck is nearly straight and inclines downward and backward, while the
seventh vertebra begins the rise which culminates in the anterior thoracic region. This
change in direction requires greater freedom of motion, which is supplied by the modifi-
cation of the zygapophyses upon the vertebree mentioned. The transverse process is, as
usual, not perforated by the vertebrarterial canal; it is rather short, but heavy and much
expanded at the distal end. On the posterior face of the centrum are large facets for the
heads of the first pair of ribs. In Hippopotamus the neural spine of the seventh cervical
is relatively much longer and heavier than in Hlotheriwm or in Sus.

As a whole, the neck of Hlotherium is short and massive, with very strongly
developed processes for muscular and ligamentous attachments, as are indeed necessitated

292 THE OSTEOLOGY OF ELOTHERIUM.

by the immense weight and length of the head. Among recent artiodactyls Hippopotamus
has cervical vertebrae most like those of Hlotheriwm, though there are many differences
in the details of construction. The most apparent of these differences lies in the greater
and more uniform height and thickness of the neural spines in the modern genus.
Doubtless the even more exaggerated massiyeness of the skull in the latter is the occasion
of this increased development of the ceryical spines. In Sus the perforation of the neural
arches for the passage of the spinal nerves constitutes an important difference from
Hlothervum.

The thoracic vertebra would appear to haye numbered thirteen, though this point
cannot, as yet, be determined with entire certainty, and while the thoraco-lumbar vertebree
were, in all probability, nineteen in number, as is well-nigh universal among the artio-
dactyls, yet there were doubtless variations in the number of ribs, as is very frequently the
case among existing animals.

The first thoracic has a rather small centrum, with decidedly convex anterior and
nearly flat posterior face; the facets for the rib-heads are very large and deeply
concave. ‘The transverse process is rather short, but very large, heavy and rugose, and
bears an unusually large, concave facet for the tubercle of the first rib. The prezyga-
pophyses are of the cervical type, but present more obliquely inward than in the vertebree
of the neck, while the postzygapophyses are, as in the other thoracics, placed upon the
ventral side of the neural arch. The neural canal is high and narrow and its anterior
opening has assumed a cordate outline. The neural spine is inclined strongly backward,
much more so than that of the seventh ceryical, and though laterally compressed it is
extremely high, broad and massive, greatly exceeding in all its dimensions that of the
last neck vertebra.

The anterior six thoracic yertebree (see Pl. XVIII, Fig. 5) are very much alike in
appearance. The first three have broader and more depressed centra, which in the others
become deeper vertically and more trihedral in section. The transverse processes are
very large and prominent and carry large, deeply concave facets for the rib tubercles.
The neural spines are very high, thick and heavy, and are strongly inclined backward,
with club-shaped thickenings at the tips. At the seventh thoracic begins a rapid reduc-
tion in the length and weight of the spines, a process which reaches its culmination on
the eleventh vertebra, which has a remarkably short, weak and slender spine. This
arrangement results in a great hump at the shoulders, somewhat as in Zitanotherium,
though in a less exaggerated form. In both genera, the length of the anterior thoracic
spines should be correlated with the great elongation and weight of the skull which
requires immense muscular strength in the neck and shoulders. Hippopotamus has no

such hump, but this is probably explaimed by its largely aquatic habits,

THE OSTEOLOGY OF ELOTHERIUM. 293

A change in the character of the facets for the rib tubercles occurs simultaneously
with the shortening of the neural spines ; they suddenly become much reduced in size
and are plane instead of concave. The transverse processes, however, remain very large
and prominent as far back as the eleventh thoracic. In no case are these processes per-
forated by vertical canals, such as occur in Sus. The twelfth thoracic is the anticlinal
vertebra and has a nearly erect spine of lumbar type, though somewhat more slender
than in the true lumbars. On the thirteenth the spine is quite lke that of the lumbars
and inclines slightly forward. Transverse processes are absent from the last two thoracic
yertebree, which display the feature, very unusual in an ungulate, of large and conspicuous
anapophyses.

As far back as the eleyenth vertebra the zygapophyses are of the ordinary thoracic
type; they are small, oval facets, the anterior pair on the front of the neural arch and
presenting upward, the posterior pair on the hinder part of the arch and presenting
downward. On the eleventh thoracic a change takes place ; the anterior zygapophyses
are as before, but the posterior processes are flat and present obliquely outward, rather
than downward, the two together forming a prominent, wedge-shaped mass. The
prezygapophyses of the twelfth vertebra are correspondingly modified; they present
obliquely inward and together constitute a cayity which receives the wedge-like projec-
tion from the eleyenth. Prominent metapophyses also make. their appearance on the
twelfth thoracic. The posterior zygapophyses of the latter and both pairs of the thirteenth
are of the cylindrical, interlocking type characteristic of the lumbars. These processes
are remarkably complex and in a fashion that does not occur in Hippopotamus, but is
found in Sus and many of the Pecora. The complexity is occasioned by the development
of large episphenial processes, which give an additional articular surface above the
zy gapophyses proper ; in section these processes have an S-like outline, and they constitute
a joint of great strength.

The lumbar vertebra (Pl. XVIII, Fig. 6), almost certainly six in number, have
rather short, but massive centra. In the anterior part of the region the centra are some-
what cylindrical in shape, but they become more and more depressed and flattened as we
approach the sacrum. The neural canal is broad and very low, especially in the pos-
terior part of the region. The neural spines are inclined forward and are of moderate
height ; they are broad antero-posteriorly, but thin and laterally compressed, except at
the tips, where they are thickened. The spine of the last lumbar is a little different
from the others in being more erect and slender. Episphenial processes are present on
the first, second and sixth vertebre, but not on the third, fourth or fifth. These
processes are apt to be somewhat asymmetrical and better developed on one side than on

the other, and it is probable that more extensive material would show them to be subject

294 THE OSTEOLOGY OF ELOTHERIUM.

to much individual variation. Metapophyses are prominent only on the first and second
lumbars, rudimentary on the third and absent from the others. The transverse processes
are very feebly developed in proportion to the size of the vertebrae. On the first lumbar
they are short and straight, and gradually increase in length up to the fifth, but in all
they are strikingly thin and slender. The last lumbar has transverse processes of unusual
length, space for them being obtained by the sudden eyersion of the anterior ends of the
ilia, but even here they are weak.

The trunk-vertebre of Hippopotamus are much more massively constructed than
those of Hlotheriwm, the decrease in length of the thoracic spines posteriorly is more
gradual, while the neural spines and transverse processes of the lumbars are much longer
and in every way heavier. The thoraco-lumbar series of Sus bears considerable resem-
blance to that of Hlotheriwm, but in the former the transverse processes of the thoracic
vertebree are perforated by vertical canals, and those of the lumbars are much longer and
stouter.

The sacrum consists of two vertebree only. The first has a broad, depressed centrum
and yery large pleurapophyses, which carry most of the weight of the ila, though the
second sacral has also a limited contact with the pelvis. On the first vertebra the
prezygapophyses are very well-developed and haye large episphenial processes to receive
those of the last lumbar. The two neural spines are co(ssified into a high but short
ridge. The second sacral has a yery much smaller and especially a narrower centrum
than the first, and retains moderately complete postzygapophyses.

In Hippopotamus and in Sus the sacrum is relatively much larger than in Hlotheriwm,
and consists of at least four vertebrae, sometimes eyen as many as six. Hyven in aged
individuals of the White River genus I haye not seen more than two vertebre in the
sacrum. ; ;

The caudal vertebre (Pl. XVIII, Figs. 7, 8, 9), of which fifteen are preserved in
association with one individual, indicate a tail of only moderate length, and present a
number of peculiarities. The first caudal has somewhat the appearance of a miniature
lumbar ; its centrum is short, broad and depressed, with quite strongly conyex faces; the
neural canal is relatively large and a distinct, though small, neural spine is present.
The zygapophyses, especially the anterior pair, are large and prominent and project
much in front of and behind the centrum. The transyerse processes are quite long and
heavy, and are directed outward and backward. A pair of tubercles on the ventral side
of the centrum represent rudimentary hemapophyses.

The succeeding caudal vertebrae resemble the first in a general way, but passing
backward, the centra become more and more slender and elongate, while the neural canal

diminishes in size, and the various processes are reduced. The hemapophyses, on the

THE OSTEOLOGY OF ELOTHERIUM. 295

other hand, increase in size and on the (?) fifth vertebra they curye toward each
other, almost meeting and enclosing a canal, which continues as far back as the (?) eighth
vertebra, behind which the hemapophyses are again reduced. The middle portion of
the tail is composed of very long, cylindrical vertebree, which in shape strikingly
resemble those of the great cats, and which are proportionately much longer, though
apparently less numerous than those of Anoplotherium. At the anterior end of each
vertebra are six prominent, nodular processes, the zygapophyses, transverse processes and
hemapophyses respectively. Posteriorly the centra become more and more slender, but
are not much diminished in length, for what appears to be the penultimate vertebra is
nearly as long as those in the middle region. The various processes are, however,
reduced to yery insignificant proportions. The last vertebra has its anterior portion
shaped like that of its predecessor, but it rapidly tapers behind to a smooth, slender,
compressed and subeylindrical rod, with a club-shaped thickening at the end. As I have
seen but a single specimen of this curious vertebra, I cannot feel quite confident that its
shape is a normal one and not due to some injury or morbid process.

The tail of Hippopotamus is of about the same relative length as that of H/othe-
rium, but the individual vertebree are very different, being all shorter and heavier, and
diminishing in size more gradually to the end. In Sus the caudal vertebrae are somewhat
more like those of Hlotherium, but none of them have such slender elongate centra. Little
is known concerning the caudals of Anthracotherium. Kowaleysky says of them: ‘“ Von
den Schwanzwirbeln liegt mir nur ein einziges yor. Obwohl seine Erhaltung sehr
mangelhaft erscheint, kann man doch aus diesem kleinen Sttick den Schluss ziehen, dass
der Schwanz bei den Anthracotherien kurz war und somit gar keine Aehnlichkeit mit
dem sonderbar langen Schwanze der Anoplotherien hatte” (73, p. 333; Taf. x, Fig. 36).
The vertebra described by Kowaleysky is an anterior caudal and is much smaller and in
every way more reduced than the corresponding ones of Elotherium. Among existing
artiodactyls, it is the giraffe which most resembles the White River genus in the peculiar
character of its caudal vertebre.

Measurements.

ENE Sp TORE, copsascs coocconan soxsco Spo euESgONDNSSESOSOCOOIS OOSCOIUSOHODOSOND SenocasoosnosecONSeDcCeN CONOR boxeCes 0.160
JNTES, ERGRTIESE TAGLUTR. .. cononeoconscecoenabaosbsanSodosesuSeCCESoONdAN=dos poSSoRHNSsHeanccOOSSCEGHSOAENSSUECHSAOOH J 270
Axis, length GT GST TIT aces AOR Gn ECBO CCD NEC OOOH ERE EEE Boa nicsaaa2 Saccon OO pao Dodo SEER TSC IaA EAC ICRU eee Se 085
ANSE), TGTIRUE ©? OOM OTT ococaconsosscospesgoSoSHESORCoceTRESeoO NcbSoOSHNSSSKOeOOIES SoEeoSSoecanOAbTSIoDIEONS! GOSH. 6 026
Ata Sm anite nl OmMbread uhisascessonnesessncdnecct sec natacacsncscisercanceusenterte se netes seneaeoesemareesnecacaceeaaes -109
JASSS,, POOSHSHIOW | TENN hens concd ccqsonaoadosSuaNsscn5 005095 coscDONSOnoHOSOSURsaSno SondES SsancosEpSRaNSccODONDSSoOOS 054
Third cervical, length

Seventinice nyaea lee en pstihveseetssesecenta reese neenanedee ca elene a eeeenertereneee eee nominee anne ea ter eenenee eens .056
TRY TIRGREO KE, LIGLEVE Nace coccescosocteoncdecbe BO ACBOSBECEDABREDSECOATIASE C90: SOSCREEIE HONE EIRE TOSI COSEADOQUOOSES! & 051

KD SSO, cade Wve

296 THE OSTEOLOGY OF ELOTHERIUM.-

Measurements.

Fifth thorecic, height of neural spine. ..-.....-ecesecseeeeeegeceeteceaeeetscecersceseesensessecssescreenesses esi
First lumbar, IGG MoosocsosconesanscosoracznencoABAS.EeESENNCERADNa HoDOUDBONNE> sonSECOOOBDOONDOSEEEDEEODODDSEOSOD .050
SIDR REM ORNR, NE Ixa (EO jccocsocsoxdc000c0s03 29000000 90009n200299R00 D9Sdos09NDC0N9r joNHSaDEDGEgHDODOCHONBeOUE9INSIcGGN 048
Sixth lumbar, breadth across tramsverse PYOCESSES.. .+--.+02cseceecesecs ete ecn sec seeeceseeeeseceserseees 176
fSHYoTUITAS, IETDVR A Blo ccpcqoa0oomcoodocooan00se0cbon.cdoooboSdoAaSant non HDSEdoReOydOBDUDSEDCHaLCaSDOGe gaqNOKOOTaCeUAEADODOOdES c 098

Furstisacrals-waidthoots cent iimissceccsesccsscasc ccssclscGarecivencssrestincsn apices stirasleetns cascelomeensicecensne .068

Second sacral, width of centrum.....

ANTEETTOR CAC, HET 29000 cosG0000Kc0 000 cbnan00009NqS00;300 p>noNIOoASHoDOnGNOAOHAEDOCIGHUHDSRDOIHCKOGBARHOEO 3 032
Wicca Gamal, WER ccopsscoaccoonnnponannnoanoscoonocH.CHDaNNSAN}Guo9qo4 AHo.oAHonDOoHOSOnUONDONEY HoBooDNNAHOOET .063

VY. THe Riss and STERNUM.

The ribs of Elotherium are decidedly smaller and lighter and indicate a less capacious
thorax than we should expect to find in such a large animal, a fact which adds to the
apparent height of the skeleton, because of the long interval between the thorax and the
ground.

The first rib is short, subeylindrical proximally, but broadening considerably at the
distal end; it has only a slight lateral curvature, appearing nearly straight when viewed
from the front, but it arches moderately backward. The head is large and compressed,
and is separated by a deep and narrow notch from the yery large and conspicuous
tubercle, which is also compressed laterally. The ribs increase gradually in length up to
the seventh or eighth of the series, and the posterior five, though successively shortening,
retain a considerable relative length throughout. The first five or six ribs are laterally
compressed and of moderate breadth, but the posterior part of the thorax is composed of
very slender and subcylindrical ribs, very different from those which we find in most
ungulates, except in the more primitive groups. The tubercle reaches its maximum of
size and prominence on the third rib, behind which it gradually diminishes in size and
becomes more and more widely separated from the head, and more sessile in position.
On the twelfth and thirteenth pairs the tubercles are absent, corresponding to the lack of
transverse processes on the twelfth and thirteenth thoracic vertebre.

In Hippopotamus the ribs are relatively yery much longer, broader and heayier than
those of Mlotherium, and grow broader toward the hinder end of the thorax, where the
great bony slabs are in the sharpest possible contrast to the slender and subcylindrical
rods of the extinct genus. In Sus the ribs are more like those of Hlotheriwm, but they
have not such a regular and symmetrical curvature as in the latter.

The sternum of Elotherium is a yery remarkable structure, and although it is of
distinctly suilline type, it is, nevertheless, not altogether like the sternum of any known
genus, recent or fossil. The presternum, or manubrium, forms a very large, thin, com-
pressed and keel-shaped plate, which is especially remarkable for its great vertical depth,

THE OSTEOLOGY OF ELOTHERIUM. 297

this dimension exceeding the antero-posterior length, and is proportionately much greater
than in Hippopotamus or the modern suillines. The body of this segment is extremely
thin, but the anterior border, and to some extent the ventral border also, is thickened and
rugose. ‘The facets for the first pair of sternal ribs form prominences, which are situated
near together and close to the postero-superior angles of the segment, so that nearly the
entire length of the latter projects in front of the first pair of ribs.

Of the mesosternum four segments and a part of the fifth are preserved. The first
segment somewhat resembles the presternum in shape, being short, narrow and yery deep ;
the dorsal border is much thicker and wider than any other part of the segment, and the
ventral border is also thickened, though in a less marked degree. Posteriorly, this
element becomes somewhat wider and shallower. The second segment of the mesosternum
is decidedly broader and shallower than the first, but still retains a very unusual
degree of vertical depth. Both the dorsal and ventral surfaces are much broadened,
while the body of the bone is a thin, vertical plate, which connects the horizontally
directed dorsal and ventral borders, giving a cross-section somewhat like that of an
I-beam. In the third segment these progressive changes are carried still farther, and the
bone becomes very distinctly broader and lower than the second segment. The dorsal
and yentral borders still project much beyond the vertical connecting plate ; this plate,
however, is much thicker transversely than in the preceding segment. The ventral
surface is rendered quite strongly concave by- the elevation of its lateral borders. In
part, this concavity may be due to the pressure which has somewhat distorted the entire
sternum, but the ventral groove is so symmetrical that it can hardly be altogether due to
distortion. The fourth and fifth segments exhibit similar changes, each one being
broader and lower than the one in front of it; the vertical plate becomes very much
thicker and the ventral groove more widely open. Though the specimen is of an animal
past maturity, yet the last three segments distinctly show the median. suture, along which
their lateral halves united.

In Hippopotamus the breast-bone is quite like that of H/otherium, but the presternum
is longer and not of such exaggerated depth, and the rib-facets are placed much nearer to
the anterior end, while the mesosternum consists of fewer, broader and shallower
segments. In Sus the sternum is still more like that of Hlotheriwm, but has a decidedly

longer and lower presternum.
VI. Tue Fore Limes.

The fore limb of Elotheriwm is quite elongate and, in connection with the shallow
thorax, and very long neural spines of the anterior thoracic yertebre, it gives to the

skeleton a somewhat stilted appearance.

298 THE OSTEOLOGY OF ELOTHERIUM.

The scapula is remarkably high, narrow and slender, at least in the White River
species, while in the John Day forms there is reason to believe that its proportions are
quite different. The glenoid cayity forms a narrow, elongate oval, with its long axis
directed antero-posteriorly, and is not very deeply concave. The coracoid is a large, but
not very conspicuous rugosity, which sends off from its inner side a compressed, hook-lhke
process ; when the shoulder-blade is seen from the external side, this process is concealed
from yiew. ‘The neck of the scapula is broad and rather thick, and there is no distinct
coraco-scapular notch. The coracoid border in its upward course inclines forward but
little, and for the upper one-third of its height curves gently backward, to join the
suprascapular border, which is exceedingly short. The glenoid border is more oblique,
and inclines backward and upward at a moderate angle. The spine is shifted far forward,
dividing the blade very unequally, so that the prescapular fossa is very much smaller
than the postscapular. Indeed, the distal one-third of the shoulder-blade can hardly be
said to have any prescapular fossa at all. The spine itself is rather low, and for much of
its course its free border is curved backward and thickened to form a massive meta-
cromion. ‘The acromion is very short and inconspicuous, ending considerably above the
level of the glenoid cavity. .

The scapula associated with the large species of Elotheriwm from the John Day
beds, which Cope has described under the name of Bodcherus (79, p. 59), is very
different in shape from that of /. ingens from the White River, to which the description
in the preceding paragraph more particularly applies. The blade is very much broader,
both fossee widening rapidly toward the dorsal end; these fossee are of nearly equal width
and the spine is placed almost in the middle of the blade. There can be little doubt that
this scapula is properly referred to the incomplete skeleton with which it was found
associated. Aside from its similarity in color and texture to the rest of the skeleton,
there is no other amimal known from the John Day horizon to which so large a scapula
could belong.

The shoulder-blade of Hippopotamus is much broader, in proportion to its height,
than that of /. ingens ; the coracoid is more prominent and the coraco-scapular notch is
distinctly marked; the postscapular fossa is somewhat larger than the prescapular, but
the difference is much less extreme than in the White River species, the spine occupying
a more median position ; the acromion is much the same in the two forms, but the meta-
cromion is larger in the fossil. In Sus also the scapula is relatively broader than in
E. ingens, and, in particular, it has a wider prescapular fossa, but is without any distinct
coraco-scapular notch. The spine rises from the suprascapular border yery steeply to
the high (but much smaller) metacromion, and then descends gradually to the neck,
without forming an acromion. In spite of these differences, the resemblance in the

character of the scapula between Sus and Hlotherium is unmistakable.

THE OSTEOLOGY OF ELOTHERIUM. 299

The humerus is relatively long, but is, at the same time, a massively constructed
bone. The head is large and very strongly convex, especially from above downward,
although it is not set upon a very distinct neck, nor does it project far behind the plane
of the shaft. The external tuberosity is very large, forming a massive and roughened
ridge, which runs across the whole anterior face of the head and rises toward the internal
side, where it terminates in a high, thick and recuryed hook, overhanging the bicipital
groove. The internal tuberosity is very much smaller, but is, nevertheless, quite promi-
nent; it likewise projects over the bicipital groove, which is very broad and deeply
incised into the bone. The great transverse breadth of the external tuberosity displaces
the groove far toward the internal side of the humerus. The shaft is long and heavy ;
its proximal portion has a great antero-posterior diameter, and its transverse thickness,
though less, is still very considerable. The fore-and-aft diameter gradually diminishes
downward, until the shaft assumes an almost cylindrical shape, below which point it
begins to expand transversely. The deltoid ridge is rugose and prominent, and runs far
down upon the shaft, but forms no deltoid hook. The distal end of the shaft is very
heavy, being both broad and thick. The supratrochlear fossa is low, wide and shallow,
while the anconeal fossa is very high, narrow and deep, its depth being much increased
by the great production of the posterior angles of the distal end. The supinator ridge is
rough, heayy and prominent. The trochlea, which is very completely modernized, in
correspondence with the advanced differentiation of the ulna and radius, is somewhat
obliquely placed with reference to the long axis of the shaft, descending toward the ulnar
side. The trochlea differs very markedly from that of such primitive artiodactyls as
Oreodon and Anoplotherium ; it is high, full and rounded and is divided into two unequal
radial facets, of which the inner one is decidedly the larger. The intercondylar ridge,
which, in most primitive artiodactyls, forms a broad and rounded protuberance, is, in
Klotherium, compressed into a sharp and prominent ridge, and shifted well toward the
external side. The internal epicondyle, which is so largely developed in Oreodon and
other early artiodactyls, has practically disappeared.

The humerus of Hippopotamus is relatively much shorter and more massive than
that of Hlotherium ; the external tuberosity is not extended so far across the anterior
face of the bone and the bicipital groove is, in consequence, not shifted so far toward the
inner side; the deltoid ridge is much better developed and gives rise to a prominent
deltoid hook. In the existing species of Hippopotamus the intercondylar ridge is
narrower and less conspicuous, but in a Pliocene species from the Val d’Arno it has
quite the same appearance as in H/otherium (see de Blainyille, Ostéographie, Hippopot-
amus, Pl. V). The epicondyles are much more prominent than in the latter, and
the postero-internal border of the anconeal fossa projects much more than does the

300 THE OSTEOLOGY OF ELOTHERIUM. :

external border, while in Klotherium this difference is decidedly less marked. In Sus
the humerus resembles that of the White River genus in form, but is proportionately
very much shorter; the deltoid ridge is shorter and less prominent, while the supinator
ridge and the epicondyles are more so.

The radius and ulna (Pl. XVIII, Fig. 10) are firmly coéssified in all the known
species of Klotherium, though the suture between them is clearly marked, eyen in old
animals. ‘The radius is relatively very long, but rather slender; the head is quite thick,
but of only moderate breadth, projecting most toward the external side. The humeral
surface is composed of three connected facets, of which the internal one is much the
largest and bears an elevated ridge for the corresponding depression on the humeral
trochlea. The groove for the intercondylar ridge of the latter is quite broad and notches
the anterior border of the radius. The shaft is rather narrow transversely, but quite
thick and heavy, and arches forward but moderately ; the distal portion is broadened and
thickened and bears upon its dorsal face a deep tendinal sulcus, bounded by very promi-
nent ridges. The distal face is quite broad, but without much dorso-palmar extension,
and carries two well-distinguished carpal facets, which pursue an oblique course, from
before backward and inward. The scaphoidal facet, which is the smaller of the two, is
concave in front, saddle-shaped behind, and is reflected up upon the posterior face of the
bone. The facet for the lunar is much larger than that for the scaphoid, and has a
somewhat similar shape, but the anterior concayity is not so deep, and the articular
surface is carried much farther up upon the palmar side of the radius. The radius has
no contact with the pyramidal.

In Hippopotamus the forearm bones are ankylosed, though somewhat less intimately
than in Elotherium. The radius is very short, broad and thick, and is almost straight.
The external facet for the humerus is larger and more concaye and the carpal facets are of
more nearly equal size, while that for the lunar rises much more steeply toward the ulnar
side. In Sus the two bones are separate, and the radius is short, very heavy and arched
forward ; its distal end is much more thickened than in Elotherium, the facet for the
scaphoid is relatively larger, while that for the lunar is smaller and is extensively
reflected upon the palmar face of the radius. In Dicotyles the ulna and radius have
coalesced even more completely than in Elotheriwm.

The w/na has a very long, thick and prominent olecranon, which projects far behind
the plane of the shaft. The process is conyex on the outer side and concave on the inner,
thickened and club-shaped at the free end, which displays a broad, shallow sulcus for the
extensor tendons. The sigmoid notch is deep and the coronoid process prominent, as is
required by the great depth of the anconeal fossa on the humerus. ‘The articulation of

the ulna with the latter is confined to the posterior and superior aspects of the humeral

THE OSTEOLOGY OF ELOTHERIUM. 301

trochlea, no part of the articular surface on the ulna presenting proximally, for the radius
occupies the entire distal aspect of the humerus. Only the proximal portion of the facet
for the humerus extends across the entire breadth of the ulna; for the rest of its course
this facet is confined to the inner side. The shaft of the ulna is somewhat reduced, but
is not interrupted at any point and, indeed, it is quite stout for its entire length ; its prin-
cipal diameter is the transverse, the antero-posterior thickness being decidedly dimin-
ished. Below the head it narrows and then expands to its maximum breadth, from
which point it narrows gradually to the distal end. On its external side the shaft is
quite deeply channeled. The distal end is small and bears a saddle-shaped facet for the
pyramidal, which is concave transversely and conyex in the dorso-palmar direction ; its
external border is compressed and extends as a sharp edge behind the body of the bone,
forming a concavity on the palmar face. The pisiform facet is continuous with that for
the pyramidal. The ulna extends distally below the level of the radius and thus arises the
very exceptional condition of an articulation between the ulna and the lunar. The facet
for this carpal element is small and is entirely confined to the radial side of the ulna, the
distal end of the latter not extending at all upon the proximal face of the lunar. In most
artiodactyls in which the functional digits haye been reduced to two, the radius tends to
encroach more or less extensively upon the proximal face of the pyramidal, for which
extension the diminution of the ulna makes a way. - In Elotherium the arrangement is
different, the ulna occupying the entire proximal surface of the pyramidal, and by
extending below the level of the radius securing a lateral contact with the lunar. Indeed,
this arrangement quite precludes the attamment of the more usual radial-pyramidal
articulation. ;

The ulna of Hippopotamus is proportionately much shorter and in eyery way more
massive than that of Hotherium ; it also has a very much larger and more prominent
olecranon, as would naturally follow from the immensely greater weight of body which
requires support upon the limbs. There appears to be a slight disto-lateral contact
between the ulna and the lunar; at all events, the radius does not extend over upon the
pyramidal. In Sus the ulna is free throughout and its shaft is relatively much shorter
and heavier than in Elotherium ; the ulna and lunar do not come into contact. The
ulna of Dicotyles is more reduced than that of the White River genus and the connections
of the carpals with one another and with the metacarpus are upon quite a different plan.

Measurements.

Scapula, height. ...-...--.seeesceeceeseeeeeesnereneeneecesesesceaceces, cteeecseccrecscesnececesssennseeccensceessnsees 0.430
Scapula, greatest Widthh...........-.c0s--sssececsneceeceeteceeeeecncecerenee cuneecccueseccceccccnseseceeererscerene 245
Scapula, breadth Of NECK -..--+.-..-s0scescnceeeeccececcnaeevteeouanasssscancavestceccccattsencteacncesessccccrese= 065

Scapula, glenoid cavity, ant.-post. Ciameter........:1.cesecseeeeeseceeessseeeseecceceneereesssesaneeereeees .068

302 THE OSTEOLOGY OF ELOTHERIUM.

Measurem ents,

Scapula, glenoid cavity, transverse CiaMeter..........ccsccccesseceseeneceecceeecanccessececeesecssesseses -050
Pimerus, Lem eth <-ee-- eee em nine nen ennnieccaneisse ste cieciens\asieseerisiosensriss>ssaseass-er/ssensenseaseee saan. rssnns .405
Humerus, width of proximal end,.........06. consoscresssereerssecssecseeeseeeecssssevsreeseesteveresanresers 132
Humerus, thickness of proximal end....-...-:.-secsesseeeeeeeeeeeeeeeecee ee .128*
Etumeruss wiadtihvordistaltemdccs-ceateeessccrsecceesscesciccceneasessecheeskerneeeerenteseee cee rerceeeeesecee .095
TREGITS, JIGME H Nooo ocoons00an sno noHSoDSADNoTOMADSGOdNODD soqDoOONoECHSgOFODSEC nocoD DHS SACHCOoDONboBHOSHOSODSHODHSONNSCO .300
DEH USS yy Le fy pox ena ea Cl ep ta ett ee ele ele tel le le ele lee leet see se alee eee eee eee 074
Radaus; widbhuot Gistalliendsaneccscscscssnc sce sece casement siseseseiteaeiec seateneecbenilsshiveasisndseessce=ncteieecce .062
UME, TSE Mescrscae ceeednses2c902009d000 BooNoD oaaSoaDdsSCEDDTONOGNA DLioBeOSOUNGoOSNOSOSHoDOOCUS OCS SBOoOASoooRSOBOCRON 443
Ulna, length of olecranon fr. CoronOid ProCeSs....-.--.--.eseceeceeseccnecceccereeentsasscecenseeceessseners 103
Wika, spadlidn Ge Chie iElscc6on: sonecnacond copoonoGaBEC DOG ADCoOATAgSDoCoOHOOONUNEHSosaDEDSeONS SoeeencSoEEeNOoIOCNCON .037

VII. Tse Manus (Pl. XVII, Fig. 11).

co}

The principal facts of the structure of the fore foot have already been determined by
Kowalevsky, but the material now at command permits a more complete account to be
given. Certain differences also which obtain between the European and American repre-
sentatives of the genus should not be passed over without mention.

The carpus of Elotheriwm is a curious one in many ways, and while modified to suit
the didactyl condition of the foot, by the reduction of the lateral and enlargement of
the median elements, it has yet retained many of its primitive characteristics.

The scaphoid is high and thick in the dorso-palmar direction, but very narrow trans-
versely. The dorsal and internal (7. ¢., radial) surfaces of the bone are very rugose, and
on the palmar border, which is the narrowest part of the scaphoid, is a blunt and massive
mammillary process. The articuiar surface for the radius is of unusual shape. It is
divided into two parts, an antero-external and a postero-internal ; the latter is much the
larger and is saddle-shaped, conyex transversely and concave in the dorso-palmar
direction, while the former is convex and descends steeply toward the ulnar side. These
two parts of the articular surface are continuous, but they meet at nearly a right angle,
and their junction forms a ridge, which is the highest point of the scaphoid. On the
ulnar side are three facets for the lunar ; the largest one is proximal and dorsal, and is
continuous with the surface for the radius, which it meets at almost a right angle ; this
facet is very oblique and presents distally as well as laterally, the scaphoid here forming
a projection which extends over the lunar. The second lunar facet is dorsal and distal in
position ; it is small, nearly plane, and not very distinctly separated from the facet for
the magnum. The third lunar facet is distal and palmar, and is placed upon the ulnar side
of the mammillary process already mentioned ; it is of oval shape and nearly flat. The
contact between the scaphoid and the lunar is confined to these three points, and as the

* Somewhat reduced by crushing,

THE OSTEOLOGY OF ELOTHERIUM. 303

facets on both bones are more or less prominent, they are elsewhere separated by con-
siderable interspaces. The distal side of the scaphoid is much narrower than the
proximal and is occupied by facets for the trapezoid and magnum, no articular surface
for the trapezium being apparent. The trapezoidal facet is considerably the smaller of
the two, and is simply concave. The magnum facet is in two parts, a very slightly
concave distal portion, and a somewhat smaller lateral portion on the ulnar face of the
scaphoid.

In the European species figured by Kowalevsky (’76, Taf. XX VI) the scaphoid is
somewhat broader than in the American forms, In both groups a remarkable resem-
blance to the scaphoid of Anthracotherium is observable, which extends to even the details
of structure (see Kowalevsky, ’75, Taf. XI, Fig. 38). As Anthracotherium is, however, a
tetradactyl form, the scaphoid is somewhat broader in proportion to its height than that
of Elotherium, though hardly so much so as would be expected. In Hippopotamus and
Sus the scaphoid is of quite a different shape from that of the fossils, being distinctly
shorter and wider.

The /unar is a very large and complex carpal, which exceeds the scaphoid in all of
its dimensions, and especially in breadth. The radial facet is in two parts, continuing
those which occur on the scaphoid ; the anterior or dorsal part extends across the width
of the bone and is very convex antero-posteriorly, while the palmar portion is very much
larger and is concave in the same direction. The dorsal border rises steeply toward the
ulnar side, where the lunar is drawn out into a blunt, projecting, hook-like process, which
extends over the pyramidal, as the scaphoid does over'the lunar. On the radial side are
three facets for the scaphoid, corresponding to those on the latter, which have already
been described. The palmar face is greatly extended transversely, and, though lower, is
much broader than the dorsal surface. On the ulnar side are two facets for the
pyramidal, which constitute an interlocking joint of unusual firmness and strength. One
of these facets is proximal and dorsal and overlaps the pyramidal; the second, which is
very much larger, is palmar and distal in position, and has a saddle-like shape ; it interlocks
closely with a similar facet upon the pyramidal. When seen from the front, the contact
between the lunar and the magnum appears to be entirely lateral, but as it passes toward
the palmar side, the magnum facet broadens, becomes very concave, and assumes a distal
position. The unciform facet is aiso oblique and the beak between the two is not in the
median, but shifted far toward the radial side. Dorsally the unciform facet is consider-
ably wider than that for the magnum, but on the palmar side these proportions are
reversed.

The lunar of L. magnum figured by Kowalevsky resembles that of EZ. ingens, except
that its proximal surface does not rise so steeply toward the ulnar side and does not

A, P, S.—VOL. XIX. 2M,

304 THE OSTEOLOGY OF ELOTHERIUM.

project over the pyramidal. The lunar of Anthracotherium (see Kowalevsky, ’73, Taf.
XI, Fig. 37) is like that of Hlotheriwm, but is narrower, especially its palmar face, and
much thicker, and the distal beak is more nearly in the median line. In Hippopotamus
the lunar is broad and rests almost equally upon the magnum and the unciform, as it
does also in Sus.

The pyramidal quite resembles the scaphoid in shape, but is much broader, not so
thick antero-posteriorly, and generally of a more rugose and massive appearance. In
view of the reduced lateral digits and the codssified radius and ulna, the relatively large
size of the pyramidal is somewhat surprising. The proximal end is occupied by the
ulnar facet, which is convex transversely and deeply concave antero-posteriorly. On the
palmar side is a narrow, plane facet for the pisiform, which is very oblique in position.
This facet is carried upon a compressed and slightly recurved, hook-like ridge, which
runs for nearly the full vertical height of the bone, though not quite reaching to the
distal end. On the radial side are two facets for the lunar, separated by a wide and deep
suleus; the palmo-distal one is larger than the corresponding surface on the lunar, and
its curvatures are, of course, in opposite directions to those of the latter, being concaye in
the vertical, and conyex in the dorso-palmar diameter. The distal end of the pyramidal
is taken up by a large, but shehtly concave facet for the unciform.

In the material described by Kowalevsky the pyramidal of Hlotherium is not repre-
sented, while that of Anthracotherium is so badly preserved and of such uncertain
reference, that any comparison founded upon it would be valueless. The pyramidal
of Hippopotamus is broad, square and heavy, as is also that of Sus, on a smaller scale.

The pisiform is quite small and slender, though of considerable length ; it is strongly
recurved toward the median side of the carpus, presenting the conyexity externally ; the
distal end is thickened and club-shaped, though but little expanded in the vertical
dimension. The pyramidal facet is nearly plane and oblique in position, broadest exter-
nally and narrowing to a point on the radial side. The ulnar facet is very much smaller
and somewhat concave; the two meet at almost a right angle.

The pisiform of H. magnum (Kowalevsky, ’76, Taf. XX VI, Fig. 27) is not unlike
that of H. ingens, but is of a more irregular shape, which looks as though it might be
due to disease, that of Anthracotherium (XKowalevsky, ’73, Tat. XI, Fig. 58) is of quite
similar shape, though much larger. In Sus the pisiform is of an entirely different shape
from that of either of the extinct genera, being much deeper vertically, more compressed
and plate-like, and less strongly recurved. That of Hippopotamus is more like that of
the fossil forms.

The trapezium is not associated with any of the specimens which I have seen, nor is
any facet for it distinctly visible on either the scaphoid or the trapezoid. If present at

THE OSTEOLOGY OF ELOTHERIUM. 305

all, it must have been in a very reduced and rudimentary condition, haying lost all
functional importance.

The trapezoid is high, narrow and thin ; it is closely interlocked with the magnum,
lying in a depression on the radial side of that bone. The facet for the scaphoid is
simple and strongly convex. ‘Three facets for the magnum occur on the ulnar side, one
proximal and two distal; the former is much the largest of the three, but is confined to
the dorsal part of the ulnar side. Of the two distal facets, one is dorsal and one palmar ;
they are separated by a narrow space and are situated in different planes, almost at right
angles to each other. On the radial side, near the distal end, is a shallow depression,
which may haye lodged a rudimentary trapezium, though there is no facet for such a bone.
The distal side of the trapezoid bears a small, plane facet, of triangular shape, for the
rudimentary second metacarpal.

The trapezoid is not yet known in connection with the European species of /lothe-
rium, or with Anthracotherium. In Hippopotamus it is lower and broader and of more
functional importance than in Elotherium, as it also is in Sus, and in the latter, differing
from all of the other genera mentioned, it articulates extensively with the third meta-
carpal.

The magnum is a relatively large and massive bone, the three diameters of which are
nearly equal, though the dorso-palmar dimension somewhat exceeds the other two. The
dorsal moiety of the bone is the lower, quite a prominent head rising proximally from
the palmar portion. The palmar hook is represented by a short, but broad, rough and
massive ridge. The proximal end is unequally divided between the facets for the
seaphoid and lunar ; dorsally the former is much the wider and occupies almost the entire
breadth of the bone, but it does not extend so far posteriorly and on the head is con-
fined to the antero-internal aspect of that elevation. The lunar facet is very narrow on
the dorsal side, and lateral rather than proximal in position, but posteriorly it widens and
coyers nearly the entire head. When yiewed from the ulnar side, the lunar facet
appears to be of a horseshoe-shape, narrow arms extending far down upon the dorsal
and palmar borders, and separated below by a very large sulcus. These two arms of
the lunar facet are obscurely demarcated from the two small facets for the unciform, in
which they may be said to terminate distally. The distal end of the magnum is covered
by the large, saddle-shaped surface for the third metacarpal, which is convex transversely
and concave antero-posteriorly ; and proximal to this, on the radial side, is a small facet
for the second metacarpal. On the radial side also is a depression, running almost the
full vertical height of the magnum, for the reception of the trapezoid. The depression
contains a larger proximal and two smaller distal facets for the trapezoid, corresponding

to those already described on the latter.

306 THE OSTEOLOGY OF ELOTHERIUM.

The magnum figured by Kowaleysky (76, Taf. XX VI, Figs. 21, 32) is of the same
general type as in the American species, but with some differences of detail. Thus, the
bone is of relatively greater antero-posterior thickness; the palmar face is narrower and
the palmar hook very much more prominent; the sulcus which, on the ulnar side,
separates the two arms of the lunar facet is much narrower, and, in consequence, the
arms themselyes are broader; the head of the magnum rises less abruptly toward the
palmar side. The magnum of Anthracotherium is not sufficiently well known for com-
parison. That of Hippopotamus is low and broad, and differs from the magnum of
Llotherium in that the dorsal portion of the lunar facet is proximal in position. In Sus
also the magnum is low and wide; its lunar facet is relatively larger than in Hippopota-
mus, and it has no articulation with the second metacarpal, from which it is excluded by
the contact of the third metacarpal with the trapezoid; the head is low.

The wnerform is the largest and most massive bone of the carpus; in shape it is low,
broad and thick, with its principal diameter directed transversely, and has on the palmar
side a hook-shaped process, which is not very prominent, but broad and heavy. The
proximal end is occupied by the facets for the lunar and pyramidal, of which the latter
is much the wider; the junction of the two forms a prominent ridge which curyes across
the proximal end, from the dorsal to the palmar side. These two facets are both slightly
concave transversely, but very strongly convex antero-posteriorly, being reflected far
down upon the palmar face. On the radial side are two vertical articular bands,
separated by a wide and deep sulcus. The dorsal band, which is much the wider of the
two, is composed of two very obscurely separated facets, a minute proximal one for the
magnum anda very large distal one for the unciform process of the third metacarpal.
The palmar band is a high and narrow facet for the magnum only, and is much more
extended vertically than the corresponding surface on that bone. The distal end carries
a large facet for the head of the fourth metacarpal, and on the ulnar side is a minute facet
for the rudimentary fifth metacarpal.

The unciform of Kowaleysky’s specimen does not differ in any significant way from
that of the American species. In Anthracotherium this bone is much wider and lower
than in Klotheriwm and the facet for the fifth metacarpal is more distal than lateral. In
Hippopotamus the unciform is exceedingly large, and its dorsal face is of a low, wide,
rectangular outline, and its great breadth corresponds to the large size and functional
importance of the fifth metacarpal. The proximal end is divided almost equally between
the lunar and pyramidal facets, and the absence of a distal beak on the lunar allows a
larger contact between the unciform and magnum. In Sus, which has much reduced
lateral digits, the unciform is narrower than in Hippopotamus, but broader than in
Hlotherium, and the facet for the fifth metacarpal is not so completely displaced toward
the ulnar side as in the latter.

THE OSTEOLOGY OF ELOTHERIUM. 307

The metacarpus consists of four members, two functional, the third and fourth, and
two mere rudimentary nodules, the second and fifth.

Metacarpal IT is not preserved in any of the specimens which I have seen, though it
is figured by Marsh (93, Pl. VIII, Fig. 4), but the facets on the neighboring bones show
that it was carried by the trapezoid and retained a lateral connection with the magnum,
excluding me. i from any contact with the trapezoid. The manus of Elotheriwm is thus
-a typical example of what Kowaleysky has called the “inadaptive mode” of digital
reduction.

Metacarpal IT is long and massive. The head is heavy, enlarged in both dimensions,
and has a stout prominence upon the palmar side; it bears a broad, saddle-shaped surface
for the magnum. On the radial side is a depression for me. ii, at the proximal end of
which are two small facets for that bone. The unciform process is very large, prominent
and heavy, and projects far over the head of me. iy, but is, as usual, confined to the
dorsal half of the head. On the distal side of this process and on the ulnar side of the
shaft is a continuous, concave facet for the head of me. iv. A second facet for the same
metacarpal is borne upon the palmar projection from the head. The shaft of mc. ii is
broad, but much compressed and flattened antero-posteriorly ; both width and thickness
are nearly uniform throughout, but increase slightly toward the distal end. The distal
trochlea is broad and rather low, but is reflected well up upon the palmar face; on the
dorsal side it is demarcated from the shaft only by an obscure ridge, with no deep
depression aboye it. The carina is yery prominent, but is confined entirely to the palmar
face. The lateral pit on the ulnar side is large and deep, but that on the radial side is
faintly marked. 7

In Kowalevsky’s specimen (’76, Taf. XX VI, Fig. 21) the third metacarpal does not
differ in any important way from that of the American species, though the magnum
facet is somewhat more concave transversely and the shaft is rather more slender. In
Anthracotherium (Kowalevsky, ’73, Taf. XIII, Fig. 80) me. iii is very similar to that of
Elotherium, but is relatively heavier; at the proximal end the tubercle for the insertion
of the extensor carpi radialis muscle is more conspicuous, and the palmar projection of
the head more prominent.

Metacarpal IV is a little shorter and narrower than me. iii, with which it articulates
by two large facets, separated by a wide and deep groove ; of these facets the dorsal one,
which is overlapped by the unciform process of me. iii, is strongly convex, while the
palmar facet is flat and borne upon the palmar projection. The ulnar side has a shallow
groove, in which lies the nodular me. vy; the articulation with the latter is by means
of a single, small, triangular facet. The shaft is somewhat narrower transversely than

that of me. ili, but is otherwise like it, as is also the distal trochlea.

308 THE OSTEOLOGY OF ELOTHERIUM.

In £. magnum, Kowaleysky’s figure shows a somewhat differently shaped proximal
end (76, Taf. XX VI, Figs. 21, 24), the head is somewhat more extended transversely,
especially toward the ulnar side, while the palmar projection is narrower and_ less
prominent. In Anthracotheriwm the head of me. iii has no such transyerse extension.

Metacarpal V is an almond-shaped nodule, almost exactly like the specimen figured
by Kowalevsky (Taf. XX VI, Fig. 25), though of a rather more regular outline. Proxi-
mally the nodule has quite a large, subquadrate, and slightly concave facet for the unci-
form, which presents more laterally than superiorly, and forming a very obtuse angle with
this surface, is a smaller, triangular facet for me. iy.

The metacarpus of Hippopotamus has four functional members, though the median
pair are longer and stouter than the lateral. Compared with those of /lotheriwm they
are relatively shorter and much heayier. In Sus there are also four metacarpals, but
the laterals are much reduced, while the median pair, which carry most of the weight,
are yery short and thick, and the distal carina surrounds the entire trochlea, dorsal as
well as palmar. The mode of articulation between the carpals and metacarpals is
quite different from that found in either Hlotherium or Hippopotamus, the head of
me. iii being much broadened and articulating extensively with the trapezoid, so that
me. ii is cut off from any contact with the magnum. This is what Kowaleysky has
called the “adaptive method” of digital reduction, and it is in decided contrast to the
inadaptive method exemplified in L/otheriwm.

The phalanges, which are quite short, as compared with the length of the meta-
earpals, are developed only in the median pair of digits. The proximal phalanx of
digit iii is relatively elongate, straight, broad and depressed ; its proximal end is both
wide and thick, and carries a concaye facet for the metacarpal trochlea, which is deeply
notched on the palmar border for the carina. Toward the distal end the phalanx
narrows but little, though diminishing much in the dorso-palmar diameter; the distal
trochlea is low, wide, depressed and only slightly notched in the median line. The
second phalanx is short, broad and thick, and of quite asymmetrical shape ; its proximal
trochlea is obscurely divided into two facets, of which that on the radial side is the
larger and extends more in the palmar direction, while the median dorsal beak is not
prominently developed. The distal trochlea is much thicker than that of the first
phalanx, is reflected much farther upon the dorsal face, and is more distinctly notched
in the median line. The course of this surface is oblique, so that it faces somewhat to
the ulnar side. The ungual phalanx is curiously small and nodular in shape, and is
short, but quite broad and thick; the proximal trochlea is imperfectly divided into two
slightly coneaye facets. The palmar surface is nearly plane, except for its rugosities,

while the dorsal margin descends abruptly to the blunt distal end.

THE OSTEOLOGY OF ELOTHERIUM. 509

In Anthracotherium (Kowalevsky, ’73, Taf. XI, Figs. 53, 54) the phalanges are of
the same general type as in Hlotherium, but are proportionately much shorter and
stouter. In Hippopotamus they are short, broad and yery heavy, while the unguals are
reduced and of nodular form. In Sus the three phalanges of a digit are together con-
siderably longer than the metacarpal, which is far from being the case in Elotherium ;
they are also of quite a different shape from those of the latter. The proximal phalanx
is much thicker in proportion to its length, and its proximal trochlea is deeply grooved
across 1ts whole face for the metacarpal carina. The ungual phalanx is longer, broader

and more depressed and pointed.

Measurements.

(CETDES INGIGIING cos00000 nod on 0sg nesses anandnanason0sdHSaD;BoseEouDaqdeBd00qbq0200e-0RKonddD oAaoDDaDAGGNSAUEDELOOOLODE 0.072
CHIROTIS, TWAlaltHNscscoonsca9asc0800 s¢scqoDo000 000000000 He0oaandad BOOTH. GdoDNCHOdOODUSGDOADaddd BddobOHODGODASHHSOGuRGOGND 077
Scaphoid, height...

Scaphoid, breadth ......

Se NNONGL, TAMONEES io. conccaes stoooe0don002 09000000 nbossscoOsEaboRoCOBUDCbbROsScoOONSHODEREONNAE snonscaodonbo90n05000 047
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310 THE OSTEOLOGY OF ELOTHERIUM.

VIII. THe Hinp Line.

The pelvis is remarkable in many ways. As a whole, it is curiously long and
narrow, except anteriorly, where the sudden and strong eversion of both ilia gives it con-
siderable breadth. The ilium is elongate, and has a long, heavy, trihedral peduncle,
which expands quite abruptly into the broad anterior plate. This plate is very strongly
eyerted in its antero-inferior portion, and in shape is not at all like that of Sus, or of most
existing artiodactyls, but rather resembles that of such ancient perissodactyls as
Paleosyops. The plate rises high above the sacrum and conceals much of that bone from
view, when the pelvis is seen from the side; the gluteal surface is concave and the sacral
surface strongly convex; the suprailiac border is quite thin for most of its course, but
becomes very thick and rugose at its inferior angle. The ihac surface is relatively wide
and may be traced through the whole length of the bone, the pubic border being very
distinctly marked throughout. The ischial border is, for the most part, thick and
rounded, but becomes sharp and compressed above the acetabulum. The pectineal process
is a very prominent and rough tuberosity, and a second rugosity lies above and behind
it. The acetabulum is rather small, but deep, and is of almost circular form; its
articular surface is but little reduced by the deep and narrow sulcus for the round
ligament.

The ischium is likewise elongate, though much shorter than the ilium; above the
acetabulum its dorsal border arches upward into a high, thin and roughened crest, the
ischial spine, very much like that seen in Sus, behind which is a distinct ischiadic noteh |
a difference from the true pigs, which have no such notch. For most of its length, the
ischium is laterally compressed, but expands posteriorly into a large, thick plate, with
eyerted hinder border and yery massive tuberosity. The pubis is short, heavy and
depressed. The symphysis, in which both the pubes and the ischia take part, is very
long, the posterior notch between the two ischia being shallow. Consequently, the
obturator foramen is much elongated antero-posteriorly, and of oyal shape. This region
of the pelvis is entirely different from that of Sus, in which the ischia are widely
separated behind, the symphysis is short, and the obturator foramen is nearly circular in
outline. In Hippopotamus the pelvis is more like that of Hlotherium, but is much larger
and more massive in every way; the peduncle of the ilium is not so elongate or so
slender, the spine of the ischium is very much less prominent, and the posterior expansion
of the ischium is yery much larger and heayier. Unfortunately, the pelvis is not
sufficiently well known in Ancodus or Anthracotherium for comparison with that of
Hlothervum.

The femur is a long and proportionately rather slender bone, The proximal end is

THE OSTEOLOGY OF ELOTHERIUM. 311

quite widely expanded in the transyerse direction; and in shape recalls that seen in the
camels and llamas. The head is almost hemispherical in form and has a small, deep pit
for the round ligament; it is set upon a very distinct neck, which is connected by a long,
narrow bridge of bone with the great trochanter. The latter is very large and massive,
especially in the antero-posterior direction, but does not rise above the level of the head,
and hence is not very conspicuous, when the femur is seen from the front. The digital
fossa is deep and widely open, which is due to the great thickness of the trochanter, but
is not much extended in the vertical direction. The second trochanter is also large and
very rugose, but not very prominent; it projects almost entirely backward, so that the
trochanter is hardly visible, when the bone is viewed from the anterior side. There is no
plainly marked intertrochanteric ridge, connecting the great and second trochanters, but
from the latter a ridge runs proximally and almost reaches to the head.

The shaft of the femur, which in its proximal portion is much expanded transversely
and compressed antero-posteriorly, rapidly narrows downward, and below the second
trochanter becomes quite slender and subeylindrical in shape. Toward the distal end
the shaft widens considerably, though increasing little in thickness. Above the external
condyle is a long, narrow pit, with rugose margins, which serves for the origin of the
plantaris muscle. The rotular groove is very broad, but quite shallow; its inner border
is much thicker and more prominent than the outer, and ascends higher proximally,
where it terminates in a short, overhanging hook, while the external border dies away
more gradually. The condyles are relatively small; they present directly backward,
though not projecting very strongly behind the plane of the shaft, and are of almost
equal size, the external one but slightly exceeding the internal in height and breadth.
The intercondylar fossa is broad and deep and has nearly straight borders.

The proportionately small antero-posterior diameter of the distal part of the femur
in Elotherium is in decided contrast to the thickness of this region in Ancodus. The
femur of Anthracotherium is much like that of Hlotherium, but it is even more slender
in proportion to its length, and the condyles are smaller. Sws has a femur of quite a
different type; the proximal end is not so wide, the head is more sessile and has a much
larger pit for the round ligament; the bridge connecting the head with the great
trochanter is shorter and much thicker, and the trochanter itself is more prominent ; the
shaft is relatively less elongate, the rotular grooye has borders of nearly equal height,
and the condyles are more prominent. The femur of Hippopotamus, though extremely
massive, has yet a certain resemblance to that of Hlotheriwm, as may be seen in the
transverse expansion of the proximal end and in the obliquity and asymmetry of the
rotular groove.

The patella is large, massive and of rather peculiar shape. It is high, quite broad

An Ps S—SVOll, IDK, BR

312 THE OSTEOLOGY OF ELOTHERIUM.

and thick in the middle portion, but with the distal part quite thin and narrow, and
tapering to a blunt point; the proximal portion is also narrow and rises above the
articular surface as a compressed, but thick and rugose process. The femoral surface is
convex transversely, and only very obscurely divided into external and internal facets by a
broad and low median ridge. This patella bears very little resemblance to the very thick
knee-cap of Ancodus and still less to that of Sws. In the latter the patella is a short, rather
narrow, but very thick bone, the posterior surface of which is of a regularly oval outline.
Hippopotamus also has a patella which bears but little resemblance to that of Hlotherium ;
it is short, but very broad and extremely thick, and sends off a long, horizontal process
from the internal border.

The tibia is a massive bone, considerably shorter than the femur, but relatively
heavier. The proximal end is very broad and thick; the condyles are of the usual
saddle-shaped form and have a rather small antero-posterior extension; the inner
condyle is somewhat more extended in this direction, while the outer one is wider trans-
versely, and projects over the external side of the shaft. The fibular facet is small and
is confined to the postero-external angle of the outer condyle. The tibial spine is low
and bifid. The enemial process is exceedingly heavy and prominent, and runs far down
upon the shaft, extending for nearly half the length of the bone; its proximal portion
displays a depression for the long patella, and the sulcus for the tendon of the extensor
longus digitorum is deeply incised. The shaft of the tibia is heavy throughout, not
diminishing much in diameter distally; it has a decided lateral and a slight anterior
curvature. The distal end is quite broad, but not very thick, and has an unusually
quadrate outline. The astragalar surface is divided by a low intercondylar ridge into
two facets, of which the external one is much the larger and the inner one more deeply
impressed. The intercondylar ridge, which pursues a very straight course across the distal
end, is remarkable for its bifid termination at the anterior margin. A considerable sulcus
is placed upon the intercondylar ridge, invading the articular surface on each side.
On the external side of the distal end of the tibia is a broad, rugose depression for the
fibula, but with only a very small external facet for the latter ; an additional fibular facet
forms a narrow band upon the distal surface, the tibia extending somewhat over this por-
tion of the fibula. The malleolar process is short and compressed, and has no great antero-
posterior extension.

The tibia of Anthracotherium (Kowalevsky, 73, Taf. X, Fig. 29) is much like that
of Elotherium, but is relatively shorter and heavier. Sus also has a similar tibia, differing
only in minor details. The tibia of Hippopotamus is of the same general type, but is
extremely short and massive.

The fibula is complete and is not codssified with the tibia at any point, but is, never-

THE OSTEOLOGY OF ELOTHERIUM. oe

theless, very much reduced. The proximal end is laterally compressed and very narrow,
but retains considerable antero-posterior extent, and bears a narrow, obliquely placed and
slightly convex facet for the tibia. The shaft tapers and becomes exceedingly thin and
delicate, though of very irregular shape; distally the shaft thickens much in the fore~
and-aft diameter, but remains very narrow. The distal end forms a large external mal-
leolus, but continues to be very narrow. The malleolus projects inward beneath the
tibia and has a narrow facet which presents proximally and articulates with the facet,
already mentioned, on the distal face of the tibia. The astragalar facet is quite large,
extending for almost the whole thickness of the malleolus and curving downward in
front ; the caleaneal facet, which occupies the entire distal end of the fibula, is narrow, but
has a very considerable antero-posterior extension. On the outer side of the malleolus are
two deeply incised sulci for the peroneal tendons. In Sus the fibula is very much stouter
and less reduced than in Hlotheriwm, while the distal end is less enlarged and does not
extend beneath the tibia. The fibula of Hippopotamus is relatively very slender, but it
differs from that of the White River genus in having a smaller proximal and yery much
larger distal end.

Measurements.

IPall Wits} SIGH Nscosccs000cbo9o ce codsc0codsosacocosdoDEDSOGODHINdOSdasaonOAodoaSobODSdHSoaGoCoNoaSUANESUNESEaEDBBN0NE000 0.495

Relvismantero-Vateniormbrend bivecesesecesesaseseonceteneesec nee eee ee ereeeee tee tece ee een CeCe meron 395
Relvismbpreadtbyabrace tab ulllumlecserrsecs snes esertnastcseccaeseh ees e hese ceeeoee steer eee tee eeeee eee eeereeeeee 191
WMA, TEEN socescoonoso0dgocdondo0ndegonsHO05N0 noaoDOGAoHSbOOAODSEHSoSOAOSEHAaERSCGaGNSAgsCEDADvsOaS woodUoSoseCDRCS C 280
UNTER, GAGE ESI H WAKEIUN -occososacs0b nqnong 900K sEsaDNOOBO CbodDODHDoEDocabASCOODISEReADUESq5O .cdodenosedbeoenSodacsS 197
Ischium, length 215
(OVD TERNIOW MOTTA, US IYGLN. ooes05cee eacda0ce Boocdo200 sono ccosaconcHGonSQooRHEasHRUDOOD saaKSODS eooSUCEUNELBLOCE .094
SHON ASIS, IGT socooces noape2a5030060c—s3000 abanSaaN DooDacoSonONCASSocoNEACEBABAEOOX-BoaRdo9DESODEGLOCdODESONE .190
IRemmMITAR, Wey EED dco pacosdqo0 nonn soos ooNNdODEoOdNdONDND antOcUDoOOSaEHOODN OR SoGNSaOOKEsa0NC donSodenDEnSDEDBeEDEodOI—BEC 405
Femur, breadth proximal end 115
Werrawne, Jonenelin Chisel Gravel, caspaccooasecqoQ0ede 2oDSoDSodDooSEHoDoAaCDoSODED BoROSDNCaDboAdH paoOSHOOBOCOnOoMNBSHONCS .100
BemurmohickmessdistalW en de..ccsces-scmnciemsereeransacsscnseelsnensranaheeaasiscseesseeresceseeeretarcaseocerchs 103
Femur, breadth of trochlea...

Pate llawyerhicalydiame beleescneaseassesessesstenseciascecseees=etaseere tech aeeee resp eseeee Cae eC a ores we usences .107
Patella mbranSverse) Glam CleLeeeseseseseseccerse sence /rassoreaceneertenaeesese sees steer eceseceerceececeser sees .056
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Tibia, breadth proximal end... -092
MibiassbnreadcMudistallwemd ceecsew ste cesceesl oasce (isa cseldestncnseGhcrmeereleresetecesseree ser eteseeeiercareseer .063
ibrar hicknessspLroxdmallenCeecncseaneeccsonetactnashacsecretenceeteeteeteseresstenereste taser enereca essere es .088
Tibia, thickness distal end........:...seecseeeeeeeee FEE EdH bonneboods se0G0 Gectabeoc SecbaroNcocEuEe Ro saeoLoosecen 054
TOL, Weta pococcoescegchc0n9se60000050008 600000009500000500020000 1onosaSsodNc oe sabSooHNS “innaanodogaasecHCoNgNEeaCes .305
Fibula, breadth proximal end.............cc:sseeeseeeeeees pooscoocdoonDcdcD sonabDDeDOGEDADOONDSGOESOSNOSODAENNS 012
En pula bread thedistaluendesccrmectscscessensecrencinccceceteesmrertecteeseet eee sse nec eneesceeeseee see ceerenes -016
Fibula, thickness proximal end 023

Fibula, thickness distal end...........-..-...12-+++- “opedooen occSoOSobScéncbcebastbecHeoSodtooasHeccoegocHoEe -040

314 THE OSTEOLOGY OF ELOTHERIUM.
1, tens, 12g.

The tarsus has undergone little specialization, although the hind foot, like the fore
foot, is didactyl.

The astragalus is elongate, though broad and massive as well. The proximal
trochlea is deeply but very broadly grooved and its two parts are unequal, the external
condyle rising much more, both proximally and dorsally, than the internal, but not pro-
duced so far distally. While the outer condyle is widely separated from the cuboidal
facet, the inner one is continued so far distally as to become confluent with the nayicular
surface. A very large and deep pit occupies a great part of the dorsal surface between
the proximal and distal trochlez. The distal trochlea is broad and is unequally diyided
into facets for the cuboid and navicular, the latter being much the wider and of a different
shape. The surface for the cuboid is strongly convex in the dorso-plantar direction, but
nearly plane transyersely, while the navicular facet is hour-glass shaped, and on the fibular
side of the median line has a distinct, though wide and shallow groove for a corresponding
ridge on the proximal side of the nayicular. The junction of the two facets forms a sharp
but not prominent edge.

The facets for the caleaneum somewhat resemble those which we find in Ancodus,
but they have not attained to such a degree of specialization as in the American species of
that genus. The proximal external facet is divided by a sulcus into two parts, both of
which are concaye and present distally, as well as laterally. The proximal portion is set
on a conspicuous prominence of the fibular side of the astragalus, and is clearly visible
when the bone is seen from the dorsal side, while the distal portion is also prominent, but
is concealed when looked at from the same point of yiew. ‘The sustentacular facet is very
large and is strongly conyex in the proximo-distal direction, but almost plane trans-
versely ; its external border projects as a shelf beyond the body of the astragalus, and
thus helps to enclose the large and deep sulcus which is found upon the external side of
the bone. The distal external facet for the calcaneum is very small. The fibular facet
is well extended in the proximo-distal diameter, but is narrow in the dorso-plantar
direction.

In Kowalevsky’s specimen (’76, Taf. X XVII, Fig. 34) the astragalus, so far as it is
preserved, resembles that of the American species, but the external part of the proximal
trochlea is too much damaged to show the characteristic external calcaneal facet. In
Anthracotherium (Kowaleysky, °75, Taf. XI, Fig. 59, de Blainyille, Ostéographie,
Anthraco., Pl. IL) the astragalus is proportionately much broader and lower than in
Elotherium, the ridge on the distal trochlea, formed by the junction of the two facets, is
more prominent and pursues a more oblique course. The sustentacular facet is narrower

and shorter and the proximal calcaneal facet projects less. The astragalus of Sus is quite

THE OSTEOLOGY OF ELOTHERIUM. 315

like that of Hlotherium, especially in the proportions of the distal trochlea. In Hippopo-
tamus the astragalus is remarkable for its extreme shortness, for the asymmetry of its
proximal trochlea, the outer condyle much exceeding the inner in size, and for the almost
equal division of its distal trochlea between the nayicular and cuboid facets.

The calcaneum has a long tuber, which is deeply channeled on the external side and
for most of its length is compressed and rather slender, but swells at the free end into a
massive, club-shaped expansion, which has a broad, shallow tendinal. sulcus on the
plantar face. From the free end the dorgo-plantar diameter of the caleaneum increases
gradually to the fibular facet, where it reaches its maximum, and from which it contracts
rapidly toward the distal end. The sustentaculum is yery prominent and bears a wide,
slightly concave facet for the astragalus. The distal astragalar facet is much more
extended in the dorso-plantar direction than is the corresponding surface on the astragalus
and indicates an unusual amount of movement between the two bones. The cuboidal
facet is narrow transversely, but much extended antero-posteriorly ; it is divided, though
very obscurely, into dorsal and plantar parts, of which the former is the larger and has
something of a saddle-like shape, while the latter is smaller and concave.

Kowaleysky does not describe the caleaneum of 2. magnum and his description and
figures of Anthracotherium do not furnish data for comparison. The caleaneum of Sus
resembles that of Hlotherium, but is broader and has a tuber of more uniform thickness,
not channeled on the outer side. The articular surface for the cuboid is yery distinctly
divided into two facets, the junction of which forms a sharp ridge. In Hippopotamus the
calcaneum has an exceedingly long and massive tuber, which is greatly swollen at the
free end.

The navicular is a large bone, not very broad, but of considerable dorso-plantar
diameter. The surface for the astragalus is hour-glass shaped, with two coneayities sepa-
rated by a broad, convex ridge, which on the dorsal side is marked by an eleyation of the
proximal margin. The concavity on the tibial side is the larger of the two and its plantar
border rises much higher than that of the external concavity. There are three facets for
the cuboid on the fibular side of the bone, one plantar and two dorsal; the former is very
strongly convex, projecting well outward, and is high yertically, but narrow antero-
posteriorly. The two dorsal facets are both small and plane, and are placed at the
proximal and distal margins of the navicular. The plantar hook is very much reduced,
forming hardly more than a roughened ridge. The distal end is occupied principally by
the large facet for the ectocuneiform, which extends across the whole dorsal side and
much of the tibial side also. Partially separated from this is a minute surface for the
mesocuneiform. The facet for the entocuneiform is much larger than the latter ; it stands

isolated at the postero-internal angle of the distal end and is somewhat saddle-shaped,

316 THE OSTEOLOGY OF ELOTHERIUM.

concaye antero-posteriorly and conyex transversely. In one species of Hlotheriwm, not
yet identified, a somewhat different proportion of these cuneiform facets is found ; the
mesocuneiform facet is larger and that for the entocuneiform smaller and in shape and in
position more as in the recent pigs.

Kowaleysky’s figures (76, Taf. XX VII, Figs. 34, 37) do not display any character-
istic differences in the structure of the navicular between the American and the European
species of Hlotherium. In Anthracotherium (Kowaleysky, ’73, Taf. XI, Figs. 48, 59) the
nayvicular has a long, massive and rugose hook, given off from the plantar side; the facet
for the ectocuneiform is relatively smaller and that for the mesocuneiform much larger
than in Llotherium, and the two surfaces are distinctly separated. Much the same
description will apply to Sus. In Hippopotamus the nayicular is very low and broad, and
its distal facets are well distinguished.

The entocuneiform is in shape not unlike the rudimentary, nodular metapodials ; it is
high, narrow and compressed, thickest proximally and tapering distally to a blunt point.
The navicular facet is relatively large, and is saddle-shaped, with curves the converse of
those which occur on the corresponding surface of the nayicular. Distally, there is a
facet on the fibular side for the plantar projection from the head of the third metatarsal.

This element has not yet been found in connection with Anthracotherium, or with
the European species of Elotherium. In Sus it is of quite a different form and decidedly
smaller, while in Hippopotamus it is broader, heayier and shorter than in the fossil form.

The mesocuneiform is firmly ankylosed with the ectocuneiform, but its shape is,
nevertheless, clearly distinguishable ; it does not extend quite so far distally as the latter and
is very small, especially transversely, and narrows toward the distal end. Its facet for the
second metatarsal is obscurely displayed and it has no contact with the third. In /. magnum
(Kowaleysky, Taf. XX VII, Figs. 35, 37) the two cuneiforms are even more completely
fused than in the American species. In Anthracotherium the mesocuneiform is separate
and has a large surface for articulation with the second metatarsal, as is also the case in
Hippopotamus. In Sus this element is likewise distinct, but higher and narrower, and
articulates with the second metatarsal more extensively than with the third.

The ectocuneiform is a large bone, of irregularly quadrate shape; its proximal
surface bears a large, plane facet for the navicular, and the distal end is occupied by a
still larger surface for the third metatarsal ; the latter is abruptly contracted toward the
plantar side. On the tibial side and distal to the mesocuneiform is a minute lateral facet
for the second metatarsal. The contact with the cuboid is restricted to two facets near
the proximal end, one dorsal and the other plantar, of which the latter is the smaller, but
the more prominent. In Z. magnum this bone is yery much as in the American species,

but the distal facet is of a different shape, not contracting so much toward the plantar

THE OSTEOLOGY OF ELOTHERIUM. Bl7/

side (Kowalevsky, Taf. XXVIJ, Figs, 35). In Anthracotherium (Kowaleysky, 73,
Taf. XI, Figs. 48, 59) the ectocuneiform is lower and has a more extended connection
with the second metatarsal. The ectocuneiform of Hippopotamus is low, but very broad,
in keeping with the great size of the third digit. In Sus this element is not so wide as in
Lilotherium, and differs from that of all the genera mentioned in having no contact with
the second metatarsal, from which it is cut off by the articulation of the mesocuneiform
with the third.

The cuboid is massive and large in all its dimensions, high, broad and thick. The
proximal surface is about equally divided between the facet for the caleaneum and that
for the astragalus, though the latter is slightly the wider. This facet, which is simply
concave antero-posteriorly, is widest near the dorsal border, and in the middle of its
course is deeply emarginated from the tibial side. The calcaneal facet is imperfectly
divided into two parts, of which the dorsal portion is much the larger, particularly in
width, while the plantar portion curyes inward so as to lie, in part, behind the astragalar
surface. The cuboid is firmly interlocked with the navicular by means of the deeply
concave facet on the tibial side near the plantar margin, which receives the projection
from the nayicular already described. Dorsally the contact between these bones is
limited to two small facets, one of which is proximal, and the other is distal on the navi-
cular, median on the cuboid, where it helps to form the projection between the navicular
and the ectocuneiform ; this prominence is, however, very short. The facets for the
ectocuneiform are also dorsal and plantar, and are just distal to those for the nayicular.
The distal end of the cuboid is taken up by the large facet for the fourth metatarsal, that
for the rudimentary fifth being very small and lateral in position, The plantar hook is
not long, but is very broad and massive, and bears on its tibial side a facet for the pos-
terior projection from the head of the fourth metatarsal,

In Elotherium magnum (Kowalevsky, 76, Taf. XX VII, Figs. 34-36) the cuboid is
not so high in proportion to its breadth as in the American species, and the tendinal
sulcus on the fibular side is deeper. The cuboid of Anthracotherium is broader and lower
and has, of course, a larger and more distal facet for the fifth metatarsal. In Sus similar
proportions recur, and the division of the calcaneal surface into two parts is complete. In
Mippopotamus the cuboid is very low and broad, and the astragalar facet is much wider
than the calcaneal.

The metatarsus, like the metacarpus, consists of two functional (iii and iy) and two
rudimentary members (ii and vy).

Metatarsal [Tis a small nodule, which is much compressed laterally and tapers to a
point at the distal end; the articulations are proximally with the mesocuneiform and
laterally with the ectocuneiform and mt. iil.

318 THE OSTEOLOGY OF ELOTHERIUM.

Metatarsal ITT is considerably longer than the corresponding metacarpal and of a
different shape, being much narrower transversely and thicker in the dorso-plantar dia-
meter. The head is of moderate width, but the long and massive projection from the
plantar side gives it great thickness. On the tibial side of the head is a depression in
which lies the nodular mt. ii. The plantar projection bears a rounded, plane facet on
each side; that on the tibial side is for the entocuneiform, and that on the fibular side is
for mt. iv; a second facet for mt. ivy is formed by a shallow depression near the dorsal
border. The shaft of mt. iii is long, straight and slender; it is flattened on the plantar
and fibular sides, rounded on the others. Toward the distal end the shaft gradually
expands both in width and thickness; a very prominent and rough tubercle is developed
on the fibular border of the dorsal face, just above the trochlea. The latter is rather
low and narrow and has a prominent carina, which is confined altogether to the plantar
face.

Metatarsal TV is a counterpart of mt. iii, with which it forms a symmetrical pair,
though the plantar projection is even larger and heavier than that of the latter and articu-
lates with the posterior hook of the cuboid. The connection with mt. iii is by means of
two facets, the dorsal one a low, rounded prominence which fits into the depression on mt.
iii already described, and the plantar one on the tibial side of the posterior projection.
The two metatarsals are held very firmly together, externally by the hook of the cuboid
and internally by the entocuneiform. A small depression on the fibular side of the head
lodges the rudimentary mt. y. The shaft and distal trochlea are like those of mt. 111.

Metatarsal V is even more reduced than mt. ii. It has a thickened club-shaped head,
which bears a facet for the cuboid and another for mt. iv, the two meeting at a very open
angle. What remains of the shaft is slender and styliform. The mode of digital
reduction in the pes, as in the manus, is entirely “inadaptive,” the rudimentary mt. 11
still clinging to the mesocuneiform and preyenting mt. iii from reaching that tarsal, which
is much diminished in size, while the ectocuneiform follows the enlargement of mt. 11.

Kowaleysky found no metatarsals associated with L. magnum. In Anthracotherium
(Kowalevsky, ’73, Taf. XI, Figs. 45, 55, 59) the lateral metatarsals are still large, fune-
tional and provided with phalanges ; the median pair are relatively shorter and heavier
than those of Elotherium, but in other respects resemble them closely. Hippopotamus
has very short and massive metatarsals, which do not exceed the metacarpals in length
and which retain the primitive mode of articulation with the tarsals. The metatarsals of
Sus differ from those of Hlotherium in much the same way as do the metacarpals of the
two genera. The laterals are still functional, though much reduced, and the medians are
short and very heavy, with the carinee completely encircling the distal trochlee ; mt. 11
has acquired an articulation with the mesocuneiform, cutting off mt. 11 from the ecto-

cuneiform,

THE OSTEOLOGY OF ELOTHERIUM. 319

The phalanges of the pes differ from those of the manus principally in their greater
slenderness. The first phalanx is a little longer than that of the fore-foot, and decidedly
more slender ; the proximal trochlea is less deeply concave and the groove for the carina
narrower and deeper. The second phalanx is of nearly the same length as in the fore-
foot, but is much narrower and somewhat less asymmetrical in form. As Kowaleysky
points out, the proportions of this phalanx are very exceptional among ungulates. The
ungual is smaller in every dimension than that of the manus and, in particular, is nar-
rower. Apparently, Anthracotherium (Kowaleysky, ’73, Taf. XI, Figs. 52, 53) displays
the same difference between the phalanges of the pes and those of the manus as does
Elotherium. In Sus and Hippopotamus the phalanges of the two extremities differ very
little.

Measurements.

ANSIESAIITIS, HOLY >>> 200000 000000000s9n9e0 doodNDsDASIDSZNGDDNs9ADbDOTORKCSoB0conq55Ka spo9ncONeONNdEDOAGEO500H0q550000d 0.083
Astragalus, width proximal trochiea....... -045
Weratewllere, IGT 01h, o902¢0000000000000009600080020000006000950000500N6854 aaR9HNNGe NSbocODbEAC sesadouGGeG0000N00050000 024
INFAVAIEMI ETE, WIELD Dodo cocdococoeasccacaqsn990D0EE00C: 480600000 SanSdaTaNSeeHOaSod josAToHDoSuAcSeABaoROgHSAdSOdenq000000 029
INFANBKCTUEN Dn CUKe) 610 2\SShanae deat ase dood eadsaeArianccadc ao Saanadtocdadedo-. Gb SE qneEaascsdosacodo oases amocBecasecsocee .044
IBMT UMAGA, XS FOOT socessaagoGsass cagdeooooEDBoAHO0cooDoRHadecoode sododsqdaoSAsegNosOUBADEGDNDGAsREaRan00G50 .037
Entocuneiform, width... -014
INIESOCTINGTIORTIN, INET EA RcaccocoscossecondeoDnonaso5ceasqabanDHSaGdoCONs DAncaDNSEDOD SEO soNSEOSaoKondaOGoENAEHOAOGOOE .016
JRO OUINE WIGAN, EKA F-sooao03o0050030000/d9N900097000009000000000500e000006000000000000000 >eoeconacqonadecoo500G6~ -022
PM CLOCTIN ETFO WIG bh x oc ce veateseenich swe suscecceeecale sccchnasebe ceatisues nadie sues seine soem eb casemate es aewemeee a 025
(CaOONGL, INES oi ecoscccoponnosccouesoscopandandEopodoSBecanEdeaoDHSENACOD Fe eee aioe er ee eee Ee .045
Cuboid, width... napooonasnoobancoodeo0 . .038
C@uboid thickness. 5 -cceeepeatnernst ssc cde ne'sd cia nu ancti see bale nea et ales Chioe mension etieneinciney octet eee oie cuiscwacecse 047
Metatarsallpint elem otleencectreesta-pems steerer ceattel a ssacrcerterecttstecte steer et serreerethe -eeeeeeseesse tees 181
Metatarsal iit) width proximal Cnd-..---...--.-0.c-++.cneeeaeceneseccnee \uescassccercassssesecsccessierseonees -029
Metatarsa laine wa dibhydistallWem derccmeccesrcasssedersehocasceenenacectoteceiecectceeitccsaseettadsccescsesiccecs .032
Metatarsal iii, thickness proximal end.... -O41
INIGIDTERESEM Tity, GIVEN dorccooncnece sscocodonncessonbenn vos093290300n00000095052000NETes0DEs vagEnDHceBNI00000 00000000 181
Metatarsal iv, width proximal end.....-..........+-sssscccceccsnesecsessccancsesccccceasssssanssccensescanees 033
NICHI ESAL Tiny, walsh Ghistell GrN(Cl..9000G009006000000000000000600000 600005900 050959n0N000 s9D0900H99B0600059005C00008 .031
Metatarsallivs thickness proximal Cndl-s-rcccr ere seecorersinnnstereasecsheasrcaeeesnassceressceseeeerc senses .046
Proximal phalanx, length 060
Proximal phalanx, width proximal Cnde...--..1....sccersaasereseliserecesesensee esreaccneceeesecaeecceses .032
Broximalysphalanxs width distally end scceccescesscecars-snerencecnseresseeereseeeiieceataseesscsesscseeseossec ss 027
Sieeoinel RIES, MEINE D ceescocd9d506000000000000800000900000005030 xzsoasnoscoODEnn||RDOvEOSORceASSdE2qG000Is FONEO .042
Second phalanx, width proximal end -030
Second phalanx, width’ distal! end)... .-..-..-2...eceensormsseeceeosenenseasmsersoserssosrcrsarsecsessecemcses| « 024
[URSA TAME, ETE Necascoscas oandocoss6s990oGaDAD20q00990G0 cOccDSeHJanSaConoNnqEdONSos0D0s86Ns0R05600080000008 -032
Uneualyphalanx. widbhsyproxamallenls.esss.-ssene en: -ceraasceaesscssteceesceeeeceeteasiescesteceseeeeceesel 022

A. P. S.—VOL. XIx. 2 0.

ey)
i)
S

THE OSTEOLOGY OF ELOTHERIUM.

X. Restoration or Erornertum (Plate XVII).

The skeleton of this genus has a remarkable and even grotesque appearance. As in
so many of the White River genera, the skull is disproportionately large, and the
immense, dependant projections from the jugals, together with the knob-like protuberances
on the mandible, produce a highly characteristic effect. The long, straight face, the
prominent and completely enclosed orbits, the short cranium, the high sagittal crest, and
the enormously expanded zygomatic arches give a certain suggestion of likeness to the
skull of Hippopotamus. The neck is-short, nearly straight and very massive, with
prominently developed processes for muscular attachment. The trunk is short, but
heavy ; the anterior thoracic spines are yery high and heavy, while those of the posterior
region are short and quite slender. In consequence of the sudden shortening of the
thoracic spines, a conspicuous hump is formed at the shoulders. The thorax is of
moderate capacity and the loins are short. The tail appears to be of no great length,
though the individual vertebrae are greatly elongated. The limbs are long and rather
slender, and the fore and hind legs are of nearly equal height ; the humerus and femur
are almost the same in length, as are also the radius and tibia, while the pes is somewhat
longer than the manus. The scapula is very large, especially in the vertical dimension,
which considerably exceeds the length of the humerus, and has a short but promment
acromion; the pelvis, on the other hand, is rather small, the ilium having a long and
slender peduncle, and only a moderate anterior expansion. The elongate limbs and
slender, didactyl feet are in curious contrast to the huge head and short, massive trunk,
and form a combination which would hardly haye been expected.

Prof. Marsh has published, with a very brief explanatory text, a restoration of
Elotherium (94, Pl. 1X) which differs in several details from the skeleton here figured.
It is difficult to tell from the data furnished exactly how much of this restoration is con-
jectural, or to determine how far the discrepancies to be mentioned are the result of the
association of parts of many different individuals in a single figure, and how far they are
due to actual specific characters. On comparing the two figures, one is struck by the
following differences: (1) In Marsh’s restoration the skull is somewhat smaller in pro-
portion to the length of the limbs. (2) The neck is more slender and the spines of the
cervical yertebree, notably those of the sixth and seventh, are much less developed. (3)
The trunk is decidedly longer and twenty thoraco-lumbar vertebre are figured. No
reason is assigned for this departure from the well-nigh universal formula of the artio-
dactyls, which is nineteen, and we are therefore ignorant of the evidence by which it is
supported. (4) The spines of the thoracic vertebree are much more slender and decrease
‘more gradually in length posteriorly, so that there is no such decided hump at the

THE OSTEOLOGY OF ELOTHERIUM. 321

withers. These spines are figured as having curious expansions at the tips, which are
either absent or much less distinctly shown in the skeleton described in the present paper.
(5) The lumbar region is longer and has neural spines which are lower and incline more
strongly forward. (6) The conjectural restoration of the presternum is entirely different
from the specimen herewith figured. (7) The scapula is relatively shorter and broader,
and has a less prominent acromion. (8) The ilium has a shorter neck, expanding more
gradually into the anterior plate and with the acetabular border of an entirely different
shape. The ischium is much more slender, is more everted and depressed at the posterior
end, and has a much less massive and prominent tuberosity.

Materials are yet lacking to detcrmine how wide is the range of variation in the
skeleton of the different species of Elotherium. So far as I have been able to observe,
there are no important differences between the species, save those of size and proportions,
the larger forms haying more massive as well as longer bones. In particular, the great

John Day species have exceedingly heavy limb and foot bones.

XI. Tue Retationsures oF ELorHEerium.

There has been a very general agreement, among those who have made a study of
this genus, regarding the systematic position of Hlotheriwm. The acute, compressed pre-
molars have, however, led some observers to see affinities with the Carnivora and de Blain-
ville went so far as to include the genus in his carniyorous family Subursi. Almost
every other writer has referred these animals to the suillines. Leidy says of it: ‘‘ Hlothe-
rium is a remarkable extinet genus of suilline pachyderms. .... Its allies among
extinct genera are Cheropotamus, Palwocherus, Anthracotherium, and among recent
animals the Hog, Peccary and Hippopotamus” (769, p. 174). Kowalevsky expresses the
same idea in a more definite and specific way: “Schon bei dem ersten Anblick der
Bezahnung bleibt kein Zweifel tiber die Familie zu der diese Form gehdrt, niimlich den
Suiden ; sie bildet aber darin wegen des auffallenden Baues der didactylen Extremitiiten
eine sehr eigenthtimliche Gattung. Pl6tzlich konnte eine derartige Form sich nicht
bilden, das Entelodon hatte gewiss Vorahnen, deren Knochenbau einen allmiiligen
Uebergang von der tetradactylen zu der didactylen Form vermittelten, bis heute aber
sind uns solche noch ginzlich unbekannt” (76, p. 450). Zittel refers the genus to the
Achenodontine, a subfamily of the Suid (94, p. 335). Marsh erects a separate family
for the genus, and says of it: “‘The Hlotheride were evidently true suillines, but formed
a collateral branch that became extinct in the Miocene. ‘They doubtless branched off in
early Eocene time from the main line which still survives in the existing swine of the old
and new worlds” (’94, p. 408). Schlosser has expressed a somewhat different opinion

322 THE OSTEOLOGY OF ELOTHERIUM.

and has referred the genus to the bunodont division of the family Anthracothervide, which
family he derives*from an Eocene stock common to the Anthracotheriide, the Anoplothe
rude, the Hippopotamide and the Suide (87, p. 80).

The complete account of the dental and skeletal structure of Hlotherium is now
before us and yet it is hardly less difficult than before to determine its phylogenetic
relationships and systematic position. The genus is so far specialized that it implies a
long ancestry, not a member of which is, as yet, certainly known, although there are
certain Eocene genera which throw some light upon the problem. In the absence of this
ancestral series, we are without any sure criterion by which to distinguish parallelisms
from characters of actual affinity, since only by tracing, step by step, all the gradations
of a differentiating phylum, can we safely determine the true position of its members.
However, some facts seem to bear a clear and definite significance. In the first place, it
is plain that Marsh is right in forming a separate family for this genus, as it belongs to a
line which diverged very early from the main stem, whatever that was. In the second
place, the relationship of this family to the Swidw must be a very remote one. When we
compare the skeleton of Hlotheriwm with that of the swine and peccaries, point by point,
the only notable resemblance between the two groups is found to consist in the bunodont
character of the molar teeth, and this resemblance, standing by itself, cannot be regarded
as at all decisive. The selenodont molar has been independently acquired by several
distinct lines, and so far as the artiodactyls are concerned, the bunodont pattern is almost
certainly the primitive one. That two widely separated families should each have
retained a common primitive character is too frequent a phenomenon to excite surprise.
In all other structures, skull, vertebral column, limbs and_ feet, no particularly close cor-
respondences between the Hlotheriide and the Suidw can be detected, though that a
common early Eocene progenitor should have given rise to both families is altogether
likely.

Between Llotherium and Hippopotamus, on the other hand, are many points of
resemblance. The likeness in the dentition is here quite as great or even greater than
between either of these genera and the Suide. In the skull there is much to suggest
relationship, though combined with many striking differences, which may perhaps be
referable to different habits of life, such as the enormous massiveness of the premaxillary
and symphyseal region in the modern genus, the peculiar development of the canines and
incisors and the elevated tubular orbits. In the skeleton the two genera are widely
separated ; Hlotherium is a long-limbed, long-footed, didactyl creature, with small thorax
and slender ribs, evidently of terrestrial habits. Hippopotamus, on the contrary, 1s a
short-limbed, short-footed, tetradactyl and isodactyl form, with immense thorax and
broad, almost slab-like ribs, which is chiefly aquatic in its habits. Whether the resem-

THE OSTEOLOGY OF ELOTHERIUM. 323

blances in skull and dentition indicate any relationship between the two families can be
determined only when their history has been worked out. In any event, it is not prob-
able that the relationship can prove to be closer than that both lines were derived from a
common stock which separated from the other Artiodactyla at a very early date.

As has already been observed, no direct ancestors of Hlotheriwm haye yet been
recovered, but there are certain Eocene forms which seem to be related to these unknown
ancestors in such a way as to suggest the character of the latter. The Achawnodon
(Elotherium) uintense of Osborn (95, p. 102) is such a form and differs from the
A. robustum of the Bridger in the “ great elongation of the face and the shortening of the
cranium, both of which characters relate it to Elotherium” (/. ¢., p. 103). This species
is more specialized in several respects than the White River Elotheres, and like its fore-
runners of the Bridger, A. robustum and A. insolens, it has but three premolars in each
jaw, and hence is not at all likely to be ancestral to the later genus. In the Wasatch
Achenodon is represented by A. (Parahyus) vagum Marsh, which likewise has but three
premolars, and, so far as it is known, differs from the Bridger species only in its smaller
size. There is some reason to think, as Osborn has pointed out, that even A. wintense had
four functional digits.

While it is very unlikely that Achwnodon can haye been the direct ancestor of Hlothe-
rium, there are, nevertheless, so many suggestive resemblances between the two genera, and
the types of their dentition are so nearly identical, that we can feel little doubt as to their
real phylogenetic relationship. In this case, Achenodon will represent a somewhat modi-
fied side-branch of the stem which culminated in Elotherium. <A species of Achenodon,
or of some closely allied genus, with unreduced dentition and unshortened face, may well
prove to be the desired ancestral form. If so, the line had already become distinct in the
Wasatch and the group thus has no subsequent connection with any existing artiodactyl
family, unless possibly with the Hippopotamide. Elotherium would then represent the
termination of an ancient and very peculiar line, which attained a remarkable degree of
specialization in many parts of its structure and which extended its range over the whole
Northern Hemisphere. At the same time, the cerebral development of the genus was
very backward and this was doubtless one, at least, of the factors which led to its extinc-
tion. After the John Day, the line disappeared, leaving no successors.

LITERATURE.

48. Aymard, A. Coll. id. Mém. Soc. Agric. Sci. et Bell. Lett. du Puy., Vol. XII, 1848.
79. Cope, EH. D. Observations on the Faun of the Miocene Tertiaries of Oregon. Bull. U.S. Geol. and Geogr. Survey
of the Territories, Vol. V, No. 1. -

74. Kowalevsky, W. Monographie der Gattung Anthracotherium. Paleontographica, Bd. XXII.

324 THE OSTEOLOGY OF ELOTHERIUM.

. Kowalevsky, W. Osteologie des Genus Entelodon Aym. Jbid., Bd. XXII, p 415.
3. Leidy, J. The Ancient Fauna of Nebraska. Smithsonian Contributions to Knowledge, 1853.
. Leidy, J. The Extinct Mammalian Fauna of Dakota and Nebraska, Phila., 1869.

Marsh, O. C. Notice of New Tertiary Mammals. Pt. If Amer. Journ. Sci., 3d Ser., Vol. V.

. Marsh, 0. C. Description of Miocene Mammalia. Ibid , Vol. 46.

Marsh, O. OC. Restoration of Elotherium. Jbid., Vol. 47.
Osborn, H. F. Fossil Mammals of the Uinta Basin. Bull. Amer Mus. Nat. History, Vol. VII.
Pomel, A. Sur un nouveau Pachyderme du Bassin de la Gironde (Elotherium magnum). Bull Soc. Géol. de
France, Tom. IV, 1846-7.
Pomel, A. Sur un nouveau genre de Pachydermes Fossiles (Elotherium). Bibl. Univ. de Geneve. Tom. V, 1847.
Schlosser, M. Beitrige zur Kenntniss der Stammesgeschichte der Hufthiere. Morph. Jahrb., Bl. XU.
Scott, W. B. On the Osteology of Elotherium. Comte-Rend. d. Séances du 3me Congres Internat. de Zoologie,
Leyden, 1896.

. Zittel, K. A. Handbuch der Paleontologie, Bd. 1V, Munchen and Leipzig.

EXPLANATION OF THE PLATES.

Plate X VII.

Skeleton of Hlotherium ingens Leidy, from the Titanotherium beds of South Dakota, about ;'; natural size. Only

the eighth thoracic vertebra and the distal ends of certain ribs are conjectural. The tail may well have been considerably

longer, as only the vertebre associated with the skeleton have been drawn.

ig. 10. Hlotheriwm ingens. Right ulna and radius.

Plate X VIII.

1. Hlotherium mortoni. Basal view of skull, } nat. size. Ty, tympanic bone ; ¢, canal opening above and behind
the posterior nares.

. Hlotherium mortoni. Occiput from behind, ; nat. size.

. Hlotheriwm ingens. Atlas, ventral side.

. Hlotherium ingens Axis, left side.

. Elotherium ingens. Fifth thoracic vertebra, from the front.

. Hlotherium ingens. Last lumbar vertebra, from behind. ¢s, episphenial process.

. Hlotheriwn ingens. Anterior caudal vertebra, from above.

Elotherium ingens. (?) Fifth caudal vertebra, left side.

OMIA AK WW

. Hlotherium ingens. Posterior caudal.

ig. 11. Hlotheriwm ingens. Right manus. ii, second metacarpal (conjectural) ; v, fifth’ metacarpal.
ig. 12. Hlotherium ingens. Right pes. ii, v, second and fifth metatarsals.

(Figs. 3-12 are approximately + nat. size and are of bones belonging to the skeleton figured in Plate X VIT.)

ARTICLE VIII.
NOTES ON THH CANIDAD OF THE WHITH RIVER OLIGOCENE.
BY W. B. SCOTT.

(INVESTIGATION MADE UNDER A GRANT FROM THE ELIZABETH THOMPSON FUND OF THE A. A, A. S.)
(Plates XIX and XX.)

Read before the American Philosophical Society, February 4, 1898.

The problems concerning the origin and mutual relationships of the various families
into which the Carnivora Fissipedia are divided haye not yet been satisfactorily solved,
principally because of the rarity of well-preserved fossils representing the earlier and
more primitive members of the families. Especially obscure are the questions dealing with
the derivation and systematic position of the Helid@, a family which by many authorities
is regarded as occupying an entirely isolated position, not directly connected with any
of the other groups. Hardly less puzzling, however, are many of the facts of canine
phylogeny, such as the relations between the two great series of the wolves and the foxes,
and the connection between the many divergent genera of successive geological horizons.
No satisfactory answer to these questions can be given until many complete phylogenetic
series of the Carnivora shall have been discovered, for so long as the numerous wide gaps
which now separate the known members of the various series remain unbridged, those
series must continue to be largely conjectural. At any time, new discoveries may call for
an entire readjustment of our views regarding the lines of descent of the different
families.

Recently, there has come into my hands some uncommonly well-preserved material for
the phylogenetic history of the Canide and is the occasion of the present paper. This
material was obtained for the museum of Princeton University by Messrs. Gidley and Wells,
who in the summer of 1896 made a collecting trip through the Bad Lands of Nebraska and
South Dakota. They had the good fortune to discover certain unworked localities where
the exposures of the White River Oligocene proved to be richly fossiliferous and, in par-
ticular, yielded-many unusually complete specimens of primitive dogs. A study of this
material has brought to ight some very remarkable and unexpected facts, which, to the
writer at least, seem to require a revision of some current views upon the phylogeny of
the carnivorous families, and to throw some light upon the obscure and difficult problems
relating to the origin of the cats. The most valuable of these specimens are referable to

326 NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE.

the genus Daphenus Leidy, which has long been known, though but very imperfectly, and
several partially preserved skeletons permit an almost complete account of its osteology to
be given.

DAPHAENUS Leidy.

Proc. Acad. Nat. Sci. Phil., 1853, p. 393. Amphicyon Leidy (non Pomel), ibid.
1854, p. 157; Het. Mamm. Fauna Dak. and Nebr., 1869, pp. 32, 359; Cope, Ter-
tary Vertebrata, pp. 894, 896. Canis Cope, Ann. Rep. U. S. Geolog. Surv. Terrs.,
1873, p. 505.

This genus represents nearly the most primitive type of dogs which has so far been
determined from the Tertiary deposits of North America. It was originally described
and named by Leidy, who afterward mistakenly referred it to the European genus
Amplhicyon, a reference which was also adopted by Cope. Though more than forty
years have thus elapsed since the first discovery of these animals, singularly little has
been known about them, for the material obtained has been very scanty and very badly
preserved. Fragments of jaws, a few very imperfect skulls and fewer limb-bones have
hitherto been the only specimens found, in spite of long and careful search, and beyond
the fact that Daphenus was apparently a primitive member of the canine phylum, little
could be predicated of it.

The new material gathered by Messrs. Gidley and Wells fortunately removes this
difficulty and gives us information regarding nearly all parts of the skeleton of these
curious animals. These skeletal characters are of a very surprising nature and their
interpretation is by no means easy. Especially remarkable are the many points of
resemblance which we find between the structure of Dapheenus and the corresponding
parts of such primitive Machairodonts as Dinictis. Aside from the dentition and the
shape of the mandible, these resemblances in structure between the primitive dogs and
the early sabre-tooth cats are ubiquitous, and recur in the structure of the skull, of the
vertebree, of the limbs and of the feet. To bring out the full force of these remarkable
characteristics, it will be necessary to enter into a detailed and somewhat tediously minute
description of the osteology of Daphanus, so that the means of comparison may be com-
pletely laid before the reader.

lL. Tee Dentirron.

The dental formula of the genus is I 3, C4, P 4, M 3, the same as that of Amphi-
cyon, a resemblance which caused the erroneous identification of the two genera already
referred to.

A. Upper Jaw (Pl. XIX, Fig. 2).—The incisors are closely crowded together and
form a nearly straight transverse row; they are smaller and occupy less space both

NOTES ON THE CANIDA OF THE WHITE RIVER OLIGOCENE. 327

transversely and antero-posteriorly than in most recent species of Canis. As in that
genus, the external incisor is much the largest tooth of the series, and forms with the
upper and lower canines a formidable lacerating apparatus. The diastema between the
incisors and the canine is somewhat greater than in Canis, and the premaxillary is quite
deeply constricted at that point, forming a groove for the reception of the lower canine.

The canine is of the usual compressed, oval section, but the compression is less
decided than in Canis, the longitudinal diameter not so greatly exceeding the transverse.
The fang of the canine is long and stout, producing a marked swelling upon the outer
face of the maxillary ; the crown is of only moderate length, but is both actually and
proportionately heavier than in the coyote (C. latrans).

The premolars are notably small and simple; they increase in size regularly from
the first to the fourth, the sectorial being, of course, much larger than any of the others.
The first premolar is implanted by a single fang, and has a small crown of compressed
conical shape, with much less conspicuous internal cingulum than in the recent species of
the Canide. 'The second premolar is decidedly smaller than in most of the modern dogs,
and is separated by longer interspaces from both the preceding and the succeeding tooth ;
it has a low, pointed, simple and much compressed crown, without the small posterior
tubercles which are found in nearly all the recent species of the family. The third pre-
molar is much longer and especially has a higher crown than p 2, but has a similar shape,
without posterior basal tubercles, and, like p 2, is inserted by two fangs. The sectorial (p +
is very primitive in character, as compared with that of the typical recent species of
Canis. Certain modern members of the family, such as Ofocyon and Canis corsac, for
example, have, it is true, even smaller and simpler sectorials than Daphenus, but as in
these forms this is doubtless due to a secondary simplification, they need not be drawn
into comparison. The primitive character of the sectorial in the White River genus is
shown in the thick, pyramidal shape of the antero-external cusp (protocone) which is less
compressed and trenchant than in the modern species, in the smaller size of the postero-
external cutting ridge (¢r7tocone) and in the unreduced internal cusp (dewterocone) which
is very much larger and more prominent than in Canis, and is carried upon a larger
fang. The position of this inner cusp with reference to the protocone is the same as in
the recent genus. As a whole, the sectorial is small and gives to the dentition a decidedly
microdont character.

The premolar series of the two sides diverge quite rapidly posteriorly, each tooth,
except p 1, being oblique in position, with reference to the long axis of the skull, thus
giving the bony palate its greatest width at the hinder edge of the sectorials. The
obliquity of the teeth and their divergence posteriorly are eyen more strongly marked
than in most recent dogs.

A, B §& VO, SIOX, Y P,

328 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

The upper molars are large and well developed, though the different species vary in
this respect, D. vetus haying larger tubercular molars than D. hartshornianus. The first
molar is, in general, like that of Canis, but differs in certain details. Thus, the two
external cusps are more conical in shape, more nearly equal in size, and are not placed
so near to the outer edge of the crown, resembling in this respect the upper molars of
certain creodonts, such as Sinopa; the large inner crescentic cusp is much as in Canis,
though hardly so prominent, especially in D. hartshornianus ; in D. vetus it is larger.
The second molar is much like the first in shape and construction, but smaller and some-
what simplified, the conules being minute or altogether absent. The third molar is very
small and has a low, transversely oval crown, in which separate elements are not distin-
guishable. This tooth is rarely preserved and none of the specimens at my disposal
possess it, though the alveolus for it is almost always present; it is well figured by Leidy
(Cosy Je Ih, lanes, 5)

B. Lower Jaw (Pl. XIX, Figs. 5, 6,7). In none of the available specimens are
the lower incisors sufficiently well preserved to be worth description.

The canine is very much the same as in the recent members of the family. The
premolars are somewhat more complex than those of the upper jaw. The first is very
small and simple, while p. 5, 3 and j, increase progressively in size and in the develop-
ment of the posterior basal cusps. In the more ancient and primitive species ? D. dodgei,
from the Titanotherium beds, the premolars are lower, thicker transversely and less
acutely pointed, and have larger posterior basal cusps than in the later species from
higher horizons. In all the species these teeth are more widely separated than in the
modern genera.

The molars are yery characteristic of the genus, but well-marked specific differences
may be observed. In ? D. dodgei the anterior triangle of the lower sectorial is of only
moderate height and the heel is but slightly concaye, the outer and inner ridges (hypo-
and entoconids) being very little raised. In D. hartshornianus the protoconid is high,
narrow and pointed, and the talon is more concaye than in the first-named species, and
has more prominent internal and external cusps. In D. vetus the inner cusp of the
talon (entoconid) is reduced and, as Cope has already pointed out (84, p. 898), there isa
tendency toward the formation of a talon with a single trenchant ridge, a tendency which
is fully carried out in the genera Temnocyon and Hypotemnodon of the succeeding John
Day horizon. In all the species of Daphenus the inferior sectorial is much more primi-
tive than in the typical modern Canida, as is clearly shown by the higher and more
conical protoconid, the lower and smaller paraconid and much less reduced metaconid.
In fact, both the superior and inferior sectorials of Daphaenus haye a close resemblance to
those of the creodont family Miacida, from which this genus could hardly be separated

upon the ground of the dentition only.

NOTES ON THE CANID#Z OF THE WHITE RIVER OLIGOCENE. 329

The tubercular molars are not preserved in the specimens of ? D. dodgei ; in D. vetus
they are proportionately larger than in D. hartshornianus. M, is relatively large,
especially in the antero-posterior diameter; it resembles the corresponding tooth of
Canis, except for the presence of the small paraconid, thus giving to the tooth all the
elements of a true sectorial, as is also the case in the creodont Miacide, though in the
White River genus all the cusps are lower and more tubercular. Mg is quite small,
though both proportionately and actually larger than in species of Canis of similar
stature, and is inserted by a single fang ; the crown is of oval shape and has an irregularly
ridged surface, without distinct cusps.

As a whole, the dentition of Daphenus is that of a primitive member of the Canide
and resembles the dentition of the recent members of the family in general plan and

structure.
Measurements.
No. 11421. No. 11424. No. 10538. | No. 11423. No. 11425. No. 11422.
| — =I _— =
Upper dental series, length C to M 2...........64- GOOB60000 | 0.069 0.076 |
«< incisors, transverse Width «.......---.seseseseceee-ee; .014 | .015 |
CS GTI, TETRA cenooccobosoapsoonoscoubds ponaaoongoonboonen 010 .0115 |
BARRE BAC width Bese ssn eat cso oi sks. sk ieeecetiees | .008 008 |
Cees Dpielongthy as ete meres <a etiats ete ease A OOS: .005 .006
Os IPB Ce .008 .0085 .0095
CFD GY GG .009 2.010 .009*
oG DA, 9c 2.014 0145 £015 £015
SREP A Mori dbase ces tcrccievstsocs ees Ree bees | 0085 | .009 .0105 .0105
“M1, length... cool! ON O11 011 .012
Sool Mowatlilieiecer et ee cert tes eee bea ere eee OLS maee lemeOls 015 .016
“MQ, length......... bate har tcePa eee ar eee | .0065 | .007 007 .007
UT IUD AGU hiaasee iocboocae eee oRCoE EERO eee ERCS ESE | .010 O11 011 O11
Lower dental series, length C to M 3....-0+.....:0eere0ee 078 -090* -090
«premolar series, length | .036 | (OBES OHO | KOS
“molar series, length .... .026 | .0245 .031* -030 |
Soman canines lenathiceereatt recess ae ice rece | Oil .O11* 012 | .010
fe GG! "ATT he es ea | 0085 .009* | .008 | .007
0G PS. SThesave hss saaade ueccosbanetan a osaaseeeeeeeroonc ean | .0045* .005* | .004 | .003
© PH a .0085 .008 .009* | .010 | .006
@ jpg ce .0095 .010 010 | .o11 | .008
Gi PDL 6 Om | (oe .012* 012 011
Come Vi [gas che ce tenn na SUP ae ea cy OL eer Ots .014* LOI || O14
GES ANC Sa AGRA ic ecto ti aN ese | .007 .007 | .009 | .0v08
BMS MeO Tenethts tot en ere, | 0085 | .008 0095 | .0095* |
CORE O8 widthiias .ccsvsek ss Buvstes Seat Bs ee: | ,006 0055 |
“M3, length Be O0S2 O04 .006* 004%
IM Sew bliss ss tact cesteeeen cron reacts soreness | .002* | .003* |

*Alveolus.

Oo
icy)
=)

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

II, THe Sxunn (Pl. XIX, Figs. 1-7).

The skull of Daphenus is exceedingly primitive in character and plainly shows
many traces of the creodont ancestry of the genus. Unfortunately, well-preserved skulls
are exceedingly rare and none of the species is represented by an altogether complete
specimen. However, seyeral more or less imperfect specimens haye been recovered,
which together give us information concerning nearly all parts of the skull.

As in the creodonts generally, the cranial region, reckoning from the anterior edge
of the orbits backward, is exceedingly elongate, while the face in front of the orbits is
very short, slender and tapering. The elongation of the cranium is not due to an enlarge-
ment of the cerebral fossa, which on the contrary is short, narrow and of relatively small
capacity. The postorbital constriction, which marks the anterior boundary of the cerebral
fossa, is notably deep and is remoyed much farther behind the orbits than in Canis. On
the other hand, the cerebellar fossa is long, and the postglenoid processes occupy a more
anterior position than in the existing species. In consequence of the elongate cranial
region, the zygomatic arches are very long, as in the more primitive types of creodonts.
The upper contour of the skull is nearly straight, the descent at the forehead being yery
slight and gradual, which gives to the skull an alopecoid rather than a thooid aspect.
This resemblance is, however, entirely superficial, for the frontal sinuses are large and
well developed, as in the thooid series of the modern Canide. The sagittal crest is low,
but varies in the different species, being decidedly thicker and more prominent in the
larger and heayier D. vetus than in the smaller and lighter D. hartshornianus.

Turning now to the more detailed study of the elements which make up the skull,
we shall find a number of striking and significant differences from the existing repre-
sentatives of the family, though the general aspect of the whole is distinctively canine.

The dasioccipital is broad and quite elongate and has a much more decided median
keel than Canis. All the occipital bones are firmly ankylosed in the specimens at my
disposal ; hence, in the absence of sutures, it will be necessary to deseribe the compound
bone as a whole, without much reference to the elements of which it is made up. The
occiput is of quite a different shape from that found in the existing members of the
family, being broader, lower, and with a wide, gently arched dorsal border or crest (see
Pl. XTX, Fig. 3); in Canis this crest is pointed and somewhat like a Gothic arch in
shape. The occipital crest is thin, but much more prominent than in Canis, which is
due to the larger and deeper depressions of the cranial walls behind the occipital lobes
of the cerebral hemispheres, the shape of which is plainly visible externally. The
foramen magnum has much the same low and broad outline as in Canis. The

condyles are low, but well extended transversely, and on the ventral side they are. sepa-

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. 351

rated by a wider notch than in Canis. The depression, or fossa, external to the condyle
is very much deeper and more conspicuous than in the modern genus, in consequence of
which the condyles project more prominently backward from the occiput than in the
modern dogs. ‘The paroccipital processes are short, but quite stout and bluntly pointed ;
they project much more strongly backward and less downward than in the liying forms,
and are less compressed laterally. Another difference from the modern genus consists in
the fact that, while in the latter the paroccipital process has quite an extensive sutural
contact with the tympanic bulla, in Daphenus there is no such contact, the minute bulla
being widely separated from the process. The direction taken by the paroccipital process
in its course is thus evidently not determined by the size of the bulla, for in the John
Day genera, Temnocyon, Hypotemnodon and Cynodesmus, in which the tympanic is greatly
inflated, the shape and direction of the paroccipital are the same as in Daphenus, with
its insignificant bulla. A considerable portion of the mastoid is exposed on the surface
of the skull, but it is rather lateral than posterior in position, a difference from Canis, in
which the mastoid is hardly visible when the skull is viewed from the side. The mastoid
process is slightly larger than in the existing genus and is channeled on the inner side
by a groove leading to the stylo-mastoid foramen.

The limits of the dasisphenoid are not clearly shown in any of the specimens, but
this element appears to have much the same broad and flattened form as in the recent
dogs. The presphenoid is long and narrow and, as in the existing species, is almost
concealed from yiew by the close approximation of the palatines and pterygoids along the
median line. The ali- and orbito-sphenoids are not well displayed in any of the speci-
mens, but so far as they are preserved, they differ little from those seen in the more
modern members of the family.

The auditory bulla of Daphenus is very remarkable and differs from that of any
other known carnivore. Its principal peculiarities were obseryed and noted by Leidy, but
the material at his command was insufficient to enable him to describe these peculiarities
with confidence. The tympanic is exceedingly small, and is but slightly inflated into an
inconspicuous bulla, the anterior third of which is quite flat and narrows forward to a
point. There is no tubular auditory meatus, the external opening into the bulla being a
mere hole, but the anterior lip of this opening is drawn out into a short process, some-
what as in existing dogs. Behind the bulla is a large reniform vacuity or fossa, of which
Leidy remarks: “ At first, it appeared to me as if this fossa had been enclosed with an
auditory bulla and what I have described as the latter was a peculiarly modified auditory
process” (769, p. 33). Several specimens representing both the White River and John
Day species of Daphenus show that the fossa is normal and was either not enclosed in
bone, or, what seems less probable, that the bony capsule was so loosely attached that it

)

B82 NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE.

invariably became separated from the skull on fossilization. At the bottom of the fossa
(7. e., When the skull is turned with its ventral surface upward) is seen the exposed
periotic, or petrosal, which is only partially overlapped and concealed by the tympanic.
Such an arrangement is far more primitive than that which is found in any other known
member of the canine series, and is not easy to interpret. A clue to its meaning may,
however, be found in the mode of development of the bulla in the recent Canidw. Here,
as is well-known, the structure consists of an anterior membranous and posterior carti-
laginous portion, which eventually ossify and coalesce into a single bulla. Reasoning
from this analogy, we may infer that in Daphwnus the bulla was also composed of two
portions, but that only the anterior chamber was ossified, the posterior one remaining
cartilaginous. Communication between the two chambers was provided for by the space
which separates the hinder edge of the anterior chamber from the petrosal. If this
interpretation be correct, it supplies an interesting confirmation of the results derived
from the ontogenetic study of the recent genera. At all eyents, it seems much more
probable that we have to do here with a primitive rather than a degenerate structure.
The parietals are large and roof in most of the cerebral fossa; they are much less
convex and strongly arched than in Canis, in correspondence with the smaller size of the
cerebral hemispheres, and posteriorly the depressions behind the hemispheres are much
larger and deeper. As already remarked, the sagittal crest varies in the different species,
and is much thicker and more prominent in D. vetus than in D. hartshornianus. The
frontals are more or less damaged in all the specimens and in none of those at my disposal
is it possible to determine the posterior limits of these bones, though from the position of
the postorbital constriction. we may confidently infer that they formed a smaller proportion
of the cranial roof than in the modern members of the family. The supraciliary ridges are
feebly developed, especially in D. hartshornianus, and the postorbital processes are like-
wise much less prominent than in most of the recent dogs; from this process a ridge de-
scends downward and backward to the optic foramen, which, though not prominent, is yet
more so than in Canis. The frontal sinuses are large and yet in spite of them the forehead
is nearly flat, both longitudinally and transyersely, with a very shallow depression along
the median line. The nasal processes of the frontals are long, narrow and pointed,
and are separated by only a short interval from the ascending rami of the premaxillaries.
The squamosal is of moderate size and differs only in subordinate details from that
of Canis. One such difference is the presence of a broad shelf-like projection, the pos-
terior extension of the root of the zygomatic process, which overhangs the auditory
meatus and is doubtless to be correlated with the lesser breadth and conyexity of the
brain. The glenoid cavity is like that of the recent species, but has a much more
distinct internal boundary, due to an elevation of the squamosal at that point. ‘The

NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE. 399

zygomatic process is stout and well-developed, especially in D. vetws, which has heavier
arches than a large wolf, while in D. hartshornianus the zygoma is lighter and more
slender, much as in the coyote. The jugal is strongly curved upward, as well as out-
ward, and is shaped quite as in Canis, forming nearly the whole anterior and inferior
boundary of the orbit ; the postorbital process is very feebly indicated, being even less
prominent than in the modern genus, so that the orbit is more widely open behind. The
lachrymal is rather larger than in Canis, forming more of the anterior orbital border, and
has a quite well-developed spine.

The nasals have a general resemblance to those of Canis, but, in correspondence with
the shortness of the whole facial region, they are considerably shorter, and somewhat
broader and more conyex transversely ; their posterior ends are more simply rounded and
have a less irregular suture with the frontals, while the anterior, free ends are much less
deeply notched. 4 :

The maxillary is somewhat peculiar in shape, corresponding to the remarkably
constricted, narrow muzzle. The facial portion of the bone is relatively higher than in
existing representatives of the family, especially in front, its anterior border rising in a
steeper and bolder curve. Just in advance of the orbits the maxillaries expand quite
suddenly in the transyerse direction, much more abruptly than in Canis. The infra-
orbital foramen occupies nearly the same position, with reference to the teeth, as in the
latter genus, being above the front edge of the sectorial, but it is very much nearer to
the orbit, which occupies a more anterior position. The palatine processes of the maxil-
laries follow the shape of the muzzle, and are long, narrow for most of their length, but
broadening much behind; anteriorly they are emarginated in an unusual degree to
receive the long premaxillary spines.

The premaxillaries, especially their alveolar portion, are somewhat narrower than in
Canis, and behind the external incisor the alveolar border is constricted on each side,
forming well-marked grooves for the reception of the lower canines. The exposed part
of the ascending ramus is much narrower than in the modern genus, forming a mere
strip on the side of the narial opening. At the same time, this ascending ramus is
relatively longer than in existing dogs and extends almost to the nasal process of the
frontal. The anterior narial opening is somewhat larger proportionately than in the
recent members of the family, especially in the vertical direction, and its borders are less
inclined ; the floor, formed by the dorsal surface of the horizontal rami of the premaxille,
is more simply and deeply concave, and the horizontal rami themselves are less massive.
The palatine processes of the premaxillaries are distinctly smaller than in Canis, while
the spines are relatively longer and more slender. The incisive foramina are large and
from them quite deep grooves are continued forward to the alveolar border, while in the
modern genus these grooves are yery shallow and feebly marked.

334 NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE.

The palatines are shaped yery much as in Canis, As a whole, the bony palate
differs from that of the latter genus in the greater and more abrupt expansion of its
posterior half, beginning at p 2; it is also somewhat more concave transversely and has
a more prominent ridge along the median line. The palatine foramina are likewise
somewhat different from those of recent dogs; one conspicuous opening on each side
occupies the same position as in the latter, opposite the middle of the sectorial, but instead
of a single opening opposite m 1, is a group of two or three minute foramina.

The Cramal Foramina. Unfortunately, none of the specimens are sufficiently
well preserved to permit a complete account of the cranial foramina, though the more
important facts concerning these structures may be determined. Leidy states that in
D. vetus “the anterior condyloid, Eustachian and oval foramina present very nearly the
same condition as in the Wolf” (’69, p. 33). The specimen upon which Leidy’s descrip-
tion was founded, belonging to the Academy of Natural Sciences of Philadelphia, has been
mislaid and is not at present available for comparison, but the description cited above
does not altogether apply to the cranium of D. hartshornianus, of which an account has
been given in the foregoing pages. In this specimen the condylar foramen is widely
removed from the condyle, much more so than in Canis, and is placed near the edge of
the reniform fossa which lies behind the tympanic bulla. The existence of this fossa
removes the necessity for a distinct foramen lacerum posterius, which is indicated only
by a notch in the hinder margin of the fossa; similarly, the stylomastoid foramen is an
open groove, only partially enclosed by bone. The postglenoid foramen is large and
conspicuous and is not concealed by the anterior lip of the auditory meatus as is the case
in the John Day Cynodesmus. The foramen lacerum medium appears to occupy a
somewhat more internal position than in Canis, though this is not altogether certain,
because of the unfavorable condition of the fossil just at this point. The Eustachian
canal is more concealed under the long anterior process given off from the tympanic
bulla than in the existing genus, and the foramen ovale is separated from the entrance to
the canal by a much more prominent bony ridge, so that the foramen presents forward
instead of downward.

By a curious coincidence all the crania of Daphenus in the Princeton museum are
damaged in such a way that none of them displays the alisphenoid canal, the foramen
rotundum or the foramen lacerum anterius, though there is no reason to doubt that all of
these foramina were present and corresponded in position to those of Canis. The optic
foramen is overhung by a ridge, already described, which is much more prominent than
in the latter, and the lachrymal foramen is decidedly larger and more conspicuous. The
parietal is perforated by a yenous foramen which opens in the depression behind the

cerebral hemispheres ; this foramen, the postpariectal, is not found in the modern genus.

90;

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. OOo”

The mandible differs considerably in the yarious species, though the comparison
between them can as yet be but partially made, for the only specimen known to me in
which the angle and coronoid process are preseryed, is that figured by Leidy (/. ¢., Pl. I,
Fig. 2), which belongs to D. vetus. In ? D. dodgei (Pl. XIX, Figs. 6, 7.) the horizontal
portion of the mandible is thick, heavy and relatively short; the inferior border is very
far from straight, rising beneath the masseteric fossa almost to the level of the molars and
descending forward from this point in a bold, sweeping curve, quite as in the modern
Canis aureus ; the masseteric fossa is very deep and its ventral border forms a prominent
ridge, distinct from the lower border of the jaw; the symphysis is short and the chin
abruptly rounded and steeply inclined.

In D. vetus the horizontal ramus is of an entirely different shape (see Pl. XTX,
Fig. 5) being longer, more compressed and slender and with a decidedly straighter
ventral border; the symphysis is longer and the chin more gently rounded, rising more
gradually from the inferior margin of the ramus. The masseteric fossa is quite deeply
impressed, though less so than in ? D. dodge, and is very large, extending far up upon
the ascending ramus. The angle is a stout hook, which is less elevated above the general
level of the horizontal ramus than in modern wolves or foxes. The condyle also has a
low position, below the level of the molars, while in recent species the condyle is raised
above the molars, and in some species very much so. The ascending ramus has great
antero-posterior extent, by which the condyle is remoyed far back of the last molar.
This is a primitive feature which recurs in most creodonts and is evidently correlated
with the characteristic elongation of the cranium and zygomatic arches. The coronoid
process is high and wide, and has a bluntly rounded end; it inclines much more strongly
backward than in Canis and has a much more concave posterior border. The condyle
resembles that of the recent dogs, but is set upon a more distinct neck, is more extended
transversely, and is less cylindrical in shape, tapering more toward the outer end.

In D. hartshornianus the mandible, so far as it is preseryed in the various speci-
mens, resembles that of D. vetus, save that the horizontal ramus is somewhat shallower
and more slender.

The Brain. Very little can be said concerning the brain, since no complete cast of
the cranial cavity is available for study. The general shape and development of the
brain are, however, indicated in the specimen of D. hartshornianus already described
(Pl. XIX, Fig. 1). Its proportions are very different from those found in existing
members of the family, a difference which may be briefly stated as largely consisting in
the much greater relative size of the cerebral hemispheres and smaller size of the olfac-
tory lobes in the modern species. In Daphenus the brain is narrow and tapers
rapidly toward the anterior end; the cerebellum and medulla oblongata are long, the

A. P. S.—VOL. XIX. 2Q.

DY)

306 NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE.

hemispheres narrow and short, and the olfactory lobes very large. The partially exposed
cast of the cerebral fossa shows that the cerebral conyolutions are fewer, simpler and
straighter than in any known species of Canis, and are even more primitive than those of
Cynodesmus (see Scott, 94, Pl. I, Fig. 2). The only sulcus visible in the specimen is
apparently the suprasylyian, which is short and pursues a nearly straight course, but
curving downward slightly at both ends. From the external character of the skull it is
clear that the hemispheres overlap the cerebellum but little.

Measurements.

| No. 11421. No, 11424, No. 10538. No, 11423. No. 11425. No. 11422.

SSP RTIUL, Teta AoococoonnacanonsooasDanos0n59600R9cnooDKoHAEaoCOND0be 20.151
Cranium, length fr. occ. condyles to preorbital border) 108
Face, length in front of Orbits ..-----....-ssseseeeeneresseeee 065 ?.050 073
Zygomatie arch, length 080
Talkie, Mei ERH] A, s0sq000ds00s 9 onoopoDAdSIHODDGaNONDSACOSEGsNd0000000 076 092
GE WAKGlidn th 7D SoosocopenccocccoddadaqnocdonoocDecqas2000000 .044* 047%) 052
Mandible, length from chin to masseteric fossa.........| .084 | -093 -096 2.079
a -020 -018 -023 -025 025
se 0175 0:5 017 020 019
$ 010 .009 | -010 012 012

* Approximate.

I. Tur Verresrat Conumn.

The vertebral column is remarkable in many ways. All the regions of the column
are well represented by several specimens of D. vetws and D. hartshornianus, but no com-
plete backbone belonging to a single individual has as yet been recovered.

Cervical Vertebre. The collection contains only a single imperfect specimen of the
atlas and this belongs to D. vetus. Imperfect as it is, this atlas displays some important
differences from that of Canis and most of these differences are approximations to the
feline and yiyerrine types of structure. In Daphenus the atlas is elongate in the
antero-posterior direction, the anterior cotyles are small and only moderately concaye,
and are somewhat more widely separated on the ventral side than in Canis. When
viewed from aboye, the cotyles are seen not to project so far in front of the neural arch
as in the cats, but farther than in the dogs. The posterior cotyles for the axis are small,
nearly plane, and but slightly oblique in position, with reference to the fore-and-aft
median line of the vertebra. These cotyles are more distinctly separated from the

articular surface for the odontoid process of the axis than in the modern dogs, in which

NOTES ON THE CANID2 OF THE WHITE RIVER OLIGOCENE, ow”

all three facets are confluent. The neural arch is low and broad, considerably elongated
from before backward, and without ridges of any kind, save an inconspicuous tubercle,
which represents the neural spine. Near its anterior border the arch is perforated by
the usual foramina for the first pair of spinal nerves. The inferior arch is very slender,
forming a more curved bar and has a much less antero-posterior extension than in Canis.

Wortman (794, p. 137) has pointed out that the foramina of the atlas display certain
characteristic features in the various carnivorous families. ‘In all of the Felidae which
I have had the opportunity of studying, the [vertebrarterial] canal pierces the transverse
process at its extreme posterior edge, where it is thickened and joins the body of the
bone. The superior edge of this posterior border slightly overhangs the inferior edge.
.... This character appears to be very constant in the Melide and so far as we know
the structure of the atlas in the more generalized Nimravide [Machairodonts], it is true
of them also. In the Canide, upon the other hand, the foramen for the vertebral artery
is situated well in advance of the posterior border of the process, and instead of having
a fore-and-aft direction, as in the cats, pierces the process almost vertically from above.
In the Viverride and Hyenide the position of the foramen is very much as in the cats.
There is, however, an important difference between these two families and the felines
where the artery enters the suboccipital foramen in the anterior part of the atlas. The
difference consists in the formation of a bony bridge in this situation, which gives to the
suboccipital foramen a double opening in the hyzenas and civets, whereas it is single in
the cats.”

In Daphenus, it is interesting to observe, the foramina of the atlas are in all respects
like those characteristic of the cats and thus depart in a very marked way from the
arrangement found in the recent Canide. The transverse processes are broken away, so
that their shape is not determinable, but enough remains to show that the atlanteo-diapo-
physial notch is not converted into a foramen, thus agreeing with the canines and felines
and differing from most of the hyzenas and civets.

The azis is likewise feline rather than canine in its general character and appear-
ance. The centrum is elongate, narrow and depressed, with a thin and inconspicuous
hypapophysial keel, running along the ventral surface, and has a slightly concave posterior
face. The articular facets for the atlas are convex and rise higher upon the sides of the
neural canal than in Canis, and on the ventral side they project below the level of the
centrum, so that they are separated by a broad notch, which is not present in the modern
dogs, and is not well marked in the cats. The odontoid process is a long, slender, bluntly
pointed peg, with a heavy, rounded ridge upon its dorsal surface, which is continued
back along the floor of the neural canal. The transverse processes are quite long and
relatively very stout; they are shorter and heavier than in Canis, and keep more nearly

338 NOTES ON THE CANID#Z OF THE WHITE RIVER OLIGOCENE.

parallel with the centrum, not diverging so much posteriorly. As in the felines, the ver-
tebrarterial canal is longer than in the modern dogs, and its posterior opening is not vis-
ible when the vertebra is seen from the side; the anterior opening is larger and is placed
farther forward than in the recent Canide. The neural canal is proportionately larger
than in the latter, both vertically and transversely, nor does it contract so much toward
the hinder end. The neural spine forms the great, hatchet-shaped plate usual among the
Carnivora, and in its details of structure it is feline rather than canine. In the latter
group, the spine is not continued back of the postzygapophyses into a distinct process, but
its hinder borders curve gently into them. In Daphenus, as in nearly all the cats and
viverrines, the spine is drawn out into a blunt and thickened process behind the zyga-
pophyses, from which it is separated by a deep notch. The zygapophyses are rather
small and do not project so prominently from the sides of the neural arch as they do in
Canis.

The other cervical vertebrae are more slender and lightly constructed than in the
existing Canide of corresponding stature. The centra are long, narrow, depressed and
very feebly keeled in the ventral median line; in most of the species this keel does not
terminate in a posterior hypapophysial tubercle, such as is found in the existing dogs.
In the largest species, however, D. felinus, the keels are more prominent, especially on the
third and fourth vertebree, and there is some indication of the tubercle. The centra are
slightly opisthoccelous and the faces are somewhat oblique in position. In very few of the
specimens are the transverse processes sufficiently well preserved to require description,
and in such cases as they are present (as, for example, on the fifth and seventh cervicals
of one individual of D. hartshornianus) they display no noteworthy differences from the
corresponding processes of Canis. The vertebrarterial canal is, however, somewhat longer
than in the latter.

The neural arches are very different from those seen in the modern representatives
of the family. In them the dorsal surface of the neural arch is very broad and on each
side projects outward as an overhanging ledge, which connects the prezygapophysis with
the postzygapophysis of the same side ; ridges and rugosities for muscular attachment are
well marked and in the large species often yery prominent; the zygapophyses, and
especially the posterior pair, project but little in front of and behind the arches, and those
of each pair are separated by notches of only moderate depth. In consequence of this
arrangement, there are but small interspaces visible between the successive arches, when
the vertebree are in position. In Daphenus, on the other hand, the dorsal surface of the
neural arch is relatively narrow, somewhat convex transversely and usually smooth, with-
out ridges or tubercles ; the overhanging ledge which gives such an appearance of breadth

to the arch in Canis is little developed ; the zygapophyses project far in advance of and

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. 339

behind the arch, and between each transverse pair is a deep notch which greatly reduces
the antero-posterior length of the bony arch in the median line. When the yvertebree are
placed in position, the openings between the successive arches, on the dorsal side, are
very large and are longer antero-posteriorly than broad transversely. In these peculiari-
ties of the cervical vertebree of Daphenus we find no approximation to the structure of
the cats or the viverrines. .

The neural spines are also quite differently developed from those of the recent dogs.
The third cervical has no spine, merely a very faintly marked keel, the overhanging
spine of the axis leaving no room for the development of one on the third vertebra.
The fourth cervical has a very low spine, and on each successive vertebra the spine
becomes higher and more pointed; that of the seventh is very high and slender, very
much more prominent than in Canis, being almost as high, though not nearly so stout,
as the spine of the first thoracic vertebra in the modern genus. The length of the spines
in the neck constitutes another similarity to the structure of the felines.

Thoracic Vertebre—The number of trunk vertebre characteristic of Daphenus
cannot as yet be definitely determined for any of the species, for no specimen has been
found with complete backbone. In one specimen of PD. vetus are preserved twelve
thoracic and five lumbar vertebre and the type of D. felinus contains six lumbars. It is
altogether probable that the extinct genus agreed with the existing dogs in having
thirteen thoracics and seven lumbars. The first thoracic has a broad, very much
depressed centrum, with anterior face convex and posterior face deeply concave. The
prezygapophyses project forward very strongly and, as in the cervicals, the notch between
them is very deeply incised, invading the base of the spine, a very different arrangement
from that seen in Canis; these processes are relatively larger and more concave in
D. vetus than in D. hartshornianus. The postzygapophyses are much smaller, but
project prominently from the hinder end of the neural arch, extending both laterally
and posteriorly ; the articular faces are somewhat convex transversely and have an
oblique position, presenting outward rather more than downward. The neural spine is
high and compressed, shaped yery much as in Canis, but somewhat more slender. The
transverse processes are very long, prominent and heavy, especially in the large species,
D. felinus ; at the distal end of the process is a large and deeply concave facet for the
tubercle of the first rib.

The second thoracic very much resembles the first, but has a smaller, narrower,
lighter, and much less depressed centrum ; the prezygapophyses are smaller, less concave
and less widely separated, while the postzygapophyses are larger and present downward,
instead of obliquely outward, as they do on the first. The transverse processes are much

smaller in every dimension than those of the first thoracic, and spring from the neural

3x40) NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

arch at a higher level, though they are still very prominent and carry large, concave
facets for the second pair of ribs. The neural spine is somewhat heavier than on the
preceding vertebra, and was probably higher, as well, but in none of the specimens is the
spine preserved for its entire length.

The other vertebree in the anterior part of the thoracic region have rather small
centra, and in general character are very much like those of Canis. The (?) sixth
vertebra has a curiously shaped spine, which exaggerates the condition seen in the modern
genus; its proximal portion is inclined very strongly backward, while the distal portion
is curved so as to project upward; the other thoracics, as far back as the (?) tenth, have
similar spines. One yery marked difference from the recent Canide consists in the deep
notch which, in Daphenus, separates the two prezygapophyses. The anticlinal vertebra
is probably, as in the existing dogs, the tenth, and at this point the thoracic vertebra
undergo an abrupt change of character, assuming more the appearance of lumbars. In
Canis the spine of the tenth thoracic is exceedingly small and much lower than those of
the ninth and eleventh, but in Daphenus, on the other hand, the spine is much better
developed, both in length and thickness; the postzygapophyses are small, somewhat
convex and placed high up upon the neural arch, presenting outward. The (?) eleventh
thoracic is not preserved in any of the specimens. The (?) twelfth and thirteenth are
much like lumbars, except for the smaller and lower spines, thickened at the distal
end, and for the entire absence of transverse processes, which in Canis are present, though
very short, even on the thirteenth; the anapophyses are remarkably long and stout,
being much heavier and more prominent than in the recent dogs, and high, massive
metapophyses rise above the prezygapophyses.

The lumbar vertebrae (P|. XIX, Fig. 8) were probably seven in number, though not
more than six haye been found in connection with any one specimen. These vertebre
are remarkable for their relatively great size and massiveness, and for the length of all
their processes, being in these respects feline, rather than canine in character and appear-
ance. Assuming that seven is the full number, the missing one will then be the third,
and the following description is made upon that assumption. The centra increase in
length posteriorly, reaching a maximum in the fifth and sixth, but the seventh is no
longer than the first, though much broader and heavier. Compared with those of Canis,
these centra are longer, stouter, less depressed and more rounded. The transverse pro-
cesses are longer and heavier than in Canis and less so than in the large species of Melis.
The neural spines are likewise intermediate in character between those of the recent dogs
and of the larger felines; they are much higher, more extended antero-posteriorly, more
thickened at the distal end and more steeply inclined forward, than in the former. In

D. felinus especially, the great height of these spines is very striking and the resemblance

NOTES ON THE CANID OF THE WHITE RIVER OLIGOCENE. 341

of the lumbar vertebrie to those of the contemporary Machairodont Dinictis is very
great. Another similarity in the structure of the lumbar vertebrae between Daphenus
and the felines consists in the great height and heaviness of the metapophyses, which are
much better developed than in the recent Canide ; on the last lumbar these processes
become very much reduced and are, in fact, almost rudimentary. The anapophyses are
smaller than on the thoracic vertebree and diminish in size on each successive vertebra
posteriorly ; only on the first and second are they yery large and prominent. In the
existing representatives of the Cunide these processes are rudimentary, except on the
first lumbar, where they are small. This constitutes another point of resemblance
between Daphenus and the cats, and emphasizes the statement already made, that the
posterior thoracic and lumbar vertebree of this Oligocene dog, for as such it must be
regarded, are decidedly more feline than canine in appearance, using those terms only
with reference to their modern application.

The sacrum (Pl. XX, Fig. 14) consists of three vertebrae, and, in correspondence
with the great development of the tail, it resembles that of the larger cats in many
respects. Only the first sacral vertebra has any contact with the ilium and bears massive
pleurapophyses. The centra are much larger and heavier than in the modern dogs and
the postzygapophyses much more prominent. The resemblance between the sacrum of
Daphenus and that of the large cats is not very close, and the following differences may
be noted: (1) the neural spines are much lower and weaker; (2) the neural canal is
smaller ; (5) the transverse processes of the second, and especially of the third vertebra,
are decidedly shorter, so that the posterior portion of the sacrum appears much narrower.
From the sacrum of the recent dogs that of Daphenus differs particularly in its greater
proportionate length and massiveness.

Caudal Vertebre (Pl. XIX, Figs. 9, 10).—In none of the specimens of the collection
is the tail completely preserved, the largest number of vertebrae found being thirteen of
one individual and eleven of another, but enough remains to satisfactorily demonstrate
its character. The tail is remarkably long and stout and is, in fact, almost as well
developed as in the leopard or tiger, and, consequently, is much longer and thicker than
in any of the existing Canidae.

The first caudal vertebra is quite like that of the lion, but is relatively lighter and
more slender in all its parts, and has a short but distinct neural spine; the zyga-
pophyses are very prominent, and even the metapophyses are distinctly shown ; the
transverse processes are very long, but are not so broad proportionately as in the lion,
and are quite strongly recurved. Posteriorly the caudal vertebrae become successively
more and more slender and elongate, while all of the processes are gradually reduced in
size. The middle region of the tail is made up of extraordinarily elongate yertebree,

342 NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE.

which are very much like the corresponding caudals of the long-tailed cats, but are
decidedly longer and more slender proportionately. Near the tip of the tail the vertebree
become very small.

The vids are represented only by fragments, which, so far as they are preserved, do
not differ materially from those of the modern Canidw. From the character of the pos-
terior thoracic vertebree, it may be inferred that the eleventh, twelfth and thirteenth pairs
of ribs did not possess tubercles.

Of the sternum very little is preserved. One segment of the mesosternum is asso-
ciated with the type specimen of D. felinus ; it has much the same shape as in modern
dogs, but is somewhat thicker transversely and shallower vertically, in proportion to its
length. Another segment accompanies a specimen of D. vetus (No. 11424) and is much
wider and more depressed than in any of the existing fissipedes, except certain hyenas.
As the association of this weathered fragment with the skeleton of Daphenus may be

accidental, no great stress can be laid upon it.

Measurements.

No. 11421, No. 11423. No. 11425.
INTER, IGTYGTD cooccoosencecsanqnoncnsecococnees Hopsconbo.esce0o900E80 sasDa noc UD .OsENSEEqNEDIDSBOHOoOSEOSESORRaAES 0.031
Axis, length (exel. of odontoid) 041
oi «« of odontoid process 013 | -014
Commnwa Obbpmibenl OG ace sercectcetsctenesenechteeste-trciscseciers iacesesiansesecsise sane spencoooanancas sosasccce .028 .031
Third cervical vertebra, length ........s.cesccesssensecccnmseeeesecueeaeecnnsucaeserecencessevecmec==eess 030 | .031
a 2 - Width Of anterlOr face.--.........--c+sccenecosensccerancenseecnceeesecerene= .014 | -016
Fourth “ ee length.... 030 .030
Fifth ‘ ff G8 ocpnandocab2000 soncnssoncooouecHacdacHcansasoncoaDtosncooaN0sTNIoC epon000 030 |
Sixth <“ ee GO 3900000 8edeer0 So COnSOHORE BEE EeEDJDbeDIaH donaasbes00000 LchebcusdoDerdDEEDeIC 024 028 | 031
a ag s WAG Ob AMbELLOL LACE) sacscncersescaseeelecwasirr ceeaunteneesnesiatiecs reer 012 014 016
Seventh ‘“ cc(e Wilenath wees eese eee ese .022 024 | —.026
First thoracic vertebra, length.........-.0c2sc++ cseccnsercecsssscesesensecracsesseee sanssr. seeecceeeens -017 .020 | 021
Thirteenth thoracic vertebra, length.....--.....:00.20...0ceneceencnsseeenecaeseeesceasecneeeserccserces -021 024 | 024
as ae cs WHGKHD GAO LEIMKOIE IY Goode Godcangonocdoos acaosobacaESbAUoCSEMSdODScOO0e -017 021 | -021
First lumbar vertebra, length.......-.-..2:-ssececsseececnsececnteccenseeecsseeseeecceeasessaenesereenerses -028 | 028
Ks w Width anterior £aCe.........c020ceseceece sere raceee GeacangodooansoDoqaDSDbeONNN .020 | -020
Sixth “ CD elon erhhites vevsc censure succssace- cc esesehe Soon coesscaten aac ouce eaten cer cea 037 | .037
y “ cs width anterior face 021 -021
Last lumbar, length......:--.ccccsseecnssesccenceeereeeeccaneceessseceuersceverseesaceaeseensesesnuesecaeees 030 -028
ee €€ "Width ANGELION £ACC.....00ceeesecsesevesseseseccnccscenacsecanccasenscnccncrseanssasernenae .022 -025
Sacrum, length .....-...ssecescecnecnnccesseeeccnscenseceeeeceseesscesectesencccneceaseeatensessecesecenecees | 058
«« width across pleurapophyses .051 051
First caudal vertebra, length ..........cccccsessescececcccssesetessnsnecccececceceseseeeevenscesssvananeess 020
ic ‘« width across transverse processes .060
Median caudal, length ..ccsccoessesssesrcessceenesersecnsenncsecsserseeseese ososccssanssrtuenesnascessessne -040

NOTES ON THE CANID@ OF THE WHITE RIVER OLIGOCENE. 345

IV. Tue Fore Lines.

Of the scapula no part has yet been recovered.

The humerus (Pl. XX, Fig. 15) differs in several important respects from that of
the recent Canide. Unfortunately, in all of the specimens the proximal end of the bone
is broken away, so that nothing can be determined with regard to the head, tuberosities,
or bicipital groove. The shaft is rather short and stout, and is arched strongly forward,
though less so than in Canis; the deltoid ridge descends low upon the shaft and is very
prominent, much more so than in the existing canines or felines, though it does not attain
the exaggerated development seen in the early Machairodonts, such as Dinictis and
Hoplophoneus. The distal end of the humerus is remarkably cat-like in appearance, and
does not suggest any relationship with the modern Canide. The supinator ridge is very
prominent and extends far up upon the shaft, while in Canis this ridge is almost obso-
lete. The internal epicondyle is very much larger, more rugose and more prominent
than in the modern genus, quite as much so, indeed, as in the cats, and there is a large
entepicondylar foramen, bridged over by a stout, straight bar of bone. The anconeal
fossa is lower, broader, shallower, and altogether more cat-like than in Canis, and does
not perforate the shaft to form a supratrochlear foramen. The humeral trochlea is
extremely low, its vertical diameter being conspicuously less than in Canis and less even
than in Felis, resembling in this respect the humerus of the sabre-tooth Hoplophoneus.
The shape of the trochlea is of feline appearance, haying a simply convex surface for the
capitellum of the radius, and no such distinctly marked intercondylar ridge or convexity
as is found in the recent Canidw. The internal border of the trochlea is prolonged
downward into a large flange.

The radius (Pl. XX, Fig. 16) is also singularly eat-like in structure and in all its
parts is much more feline than canine. The proximal end bears an oval and somewhat
concave capitellum, for articulation with the humerus; its transverse diameter only
slightly exceeds the antero-posterior dimension. The anterior notch of the humeral
surface is somewhat more deeply incised than in Felis, but not more so than in Hop-
lophoneus, which has an entirely similar capitellum. The articular facet for the ulna
surrounds more than half the circumference of the head of the radius, which is in
remarkable contrast to the small size of this facet in Canis. The shape and mode of
articulation of the bones which enter into the formation of the elbow-joint show that
Daphenus possessed unimpaired powers of pronation and supination of the manus. In
the existing members of the Canidae, on the contrary, this power is lost, the head of the
radius being so much expanded transversely, as to occupy nearly the whole width of the
humeral trochlea, and interlocking with it in such a way as to allow only the movements
of flexion and extension.

A. P. S.—VOL. XIX. 2 R.

344 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE,

The shaft of the radius in Daphenus is slender and has a similar shape to that
which we find in-the cats, although it is not so much expanded distally; it is thus very
different from the broad, antero-posteriorly compressed and almost uniform radial shaft
of the modern dogs. The distal portion of the radius is likewise very feline in appear-
ance, but is rather lighter and narrower in proportion to the length of the bone ; it is
convex anteriorly and quite deeply concave posteriorly, with well-marked sulci for the
extensor tendons upon the dorsal face. The distal facet for the ulna is small and of sub-
circular shape and forms quite a projection upon the ulnar side; upon the inner side of
the distal end is a tubercle, which is even more rugose and prominent than in Felis, and
more distinctly set off from the carpal surface. This carpal facet has a shape like that
seen in the cats, and is more concaye transversely and narrower in the dorso-palmar
diameter than in the existing forms of Canide, and its internal border is more prolonged
distally into a downward projecting flange.

Had this radius been found isolated, one would hardly have hesitated to refer it to
one of the Machairodont genera, so completely does it differ from the radius of the modern
dogs. Fortunately, there is no room for scepticism regarding the reference of this bone
to Daphenus, for several of the specimens, representing different species, have radii of
the same type. In this connection, it may be of interest to note that the Eocene creodont
genus, Miacis, which has a remarkably canine type of dentition, has a very cat-like form
of radius.

The wna is hardly less characteristically feline than the radius. In marked con-
trast to the creodonts, which have a very long olecranon, that of Daphenus is rather
short ; its antero-posterior diameter is proportionately less than in Felis, or even than in
Canis, and its postero-superior angle is thickened and rugose, though somewhat less so
than in either of the modern genera mentioned, which gives its proximal border a
straighter contour than in them. The tendinal sulcus is wider and deeper than in the
recent dogs, less so than in the cats. The sigmoid notch is deeply incised, but describes
a parabolic curve rather than a semicircle; the proximal humeral facet is relatively much
wider than in Canis, and is continuous with the broad distal internal facet, which is like-
wise broader than in the existing dogs and is shaped much as in the cats, while the
external distal facet is nearly or quite obsolete. The radial facet is large, quite deeply
concave, and continuous or single, while in Canis it is much smaller and is divided by a
sulcus into two portions.

The shaft of the ulna is stout and, in the proximal portion, laterally compressed,
tapering toward the distal end, where it becomes trihedral in section. In shape this
shaft is very much like that of the cats and differs entirely from the ulnar shaft of the
recent Camda, which has become very much more slender, reduced and styliform, a

NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE. 345

change which is obviously correlated with the increased size of the radius. The distal
end of the ulna in Daphenus is narrow and carries a continuous convex articular surface,
which is not divided into separate facets for the pisiform and pyramidal. The distal
radial facet is raised upon a prominent’ projection, another point of resemblance to the
cats and of difference from the existing representatives of the Canide.

Measurements.
No. 11424. | No. 11425.
Humerus, width of distal end.............c.cccccscescccceseneccceesesenccncnsccnccecnsecsscessesenseecnesensensesseseonece 0.050
oS uo “« trochlea....... -033
Radius, ant.-post. diameter of head -016
““ transverse a ee 021
CS Jonemialiln ost GIST GiVEl -osccocoacacasasdesbsoceRenabocacsgacoqsEsoonoSn0e9C90090000 SosoDaSooUSBSbIEO5ORGeG0N00 Bag000a08 -022
oY €¢ “© Carpal FACE -..gsceseessensearseenees Beemer c cia seeinalseneteciseiienieitonceeicisciecienstemsteieietseswia sie .014
Ulna, Seen di stale Da seeet ot acc ener occesel ck sees canscaccatecssscesaireancec seca nonecwstinns ausavanctcecaciersesasiet .013
y G6 96 erp TENCE tracccogasoococadacdoo 6d0000506C0 0600000930007 .bo5ene 4959p oCH9ebscaR>uccaDpecNdesebadGnD000 .008

V. THE Manvs.

Of the carpus the only element preserved is a single scapho-lunar of J). vetus, inter-
esting as showing that the coalescence of these elements had already taken place. This
bone differs in a marked way from that of both recent canines and felines, but resembles
the seapho-lunar of the White River sabre-tooth, Hoplophoneus. It is broad transversely
and thick in the dorso-palmar diameter, but very low proximo-distally, even more so
than in Canis; the tubercle at the postero-internal angle of the bone is well marked, but
smaller than in the felines or modern dogs. The radial facet is simply convex in both
directions, not having the postero-internal saddle-shaped extension which occurs in the
recent dogs. This radial facet is reflected far oyer upon the dorsal and internal surfaces
of the bone, converting the inner side into a thin edge, formed by the junction of the
radial and trapezial facets.

On the distal end of the scapho-lunar are three plainly distinguished facets, for the
unciform, magnum and trapezoid respectively. The very deeply excavated unciform
surface reduces the ulnar side of the scapho-lunar to an edge, not yery much thicker
than the radial border, and hence there is no well-defined facet for the pyramidal, such
as occurs in Canis. The shape and proportions of the unciform and magnum surfaces
are very much as in the latter genus, but that for the trapezoid is not demarcated from
that for the trapezium, though there can be little doubt that the latter element articulated
with the scaphoid, as it certainly does both in Cynodictis and in Canis. The general

346 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

shape of the seapho-lunar, recalling that which we find among the mustelines, strongly
suggests that Daphanus had a plantigrade or, at least, al semiplantigrade gait.

The metacarpus (Pl. XX, Fig. 17) consists of five members, which bear little resem-
blance to those of the recent Canide. Schlosser (88, p. 24) has pointed out the essential
characteristics of the metacarpus among the modern forms, and it will be well to
quote his description, in order to make clear how widely Daphenus departs from the
arrangement which has been attained by the later representatives of the family.

“Die Metapodien haben sich auffallend gestreckt und sind zugleich kantig
geworden. Sie zeigen nahezu quadratischen Querschnitt, in Folge ihres gegenseitigen
... Die distalen Gelenk-
fiichen haben das Aussehen yon sehr kurzen Walzen und sind beiderseits scharf

Druckes ; sie liegen einander niimlich ungemein dicht an.

abgestutzt. Es lisst sich eine freilich sehr entfernte Aehnlichkeit mit dem Fusse von
Hufthieren, namentlich yom Schweine—nicht verkennen. .... Die Anordnung der
Carpalien ist scheinbar primitiver als bei den tibrigen Raubthieren, wenigstens als
dieselben unter einander und mit den Metacarpalien nur reihenweise artikuliren, statt
wechselseitig in einander zu greifen. Auch hat nur das Scapholunare eine etwas
betriichtlichere Grisse erreicht, Magnum sowie Trapezoid und Trapezium bleiben sehr
kurz und enden sowohl oben als auch unten simmtlich in einer Ebene. Demzufolge
liegen auch die proximalen Facetten der Metacarpalien so ziemlich in einer einzigen
Ebene.”

This description of the structure of the manus in the recent Canid@ does not at all
apply to Daphenus. In this genus the metacarpals are remarkably short and quite
slender; they are not very closely approximated, but diverge somewhat toward the distal
end, and hence they have not acquired the quadrate shape which Schlosser mentions as
so characteristic of the modern dogs. The general appearance and character of the meta-
earpals, and their mode of articulation with each other and with the carpals are very
much as in the wolverine (G'wlo).

The first metacarpal, even of the large D. felinus, is actually not much longer than
that of the coyote (@. latrans), but is much longer in proportion to the other metacarpals,
as well as much stouter and in every way better developed. The proximal end is
thickened both transversely and antero-posteriorly, and bears a large facet for the trape-
zium, which must have been a relatively large bone; this facet is convex in the dorso-
palmar direction and is very slightly concave transversely, while in Canis it is deeply
concave in this direction. In OD. vetus the articular surface for the trapezium is more
oblique and inclined toward the radial side than in D. felinus. There is no other well-
defined facet for any carpal but the trapezium, nor for me. ii. The shaft is

short, slender, of oval or subcireular section, and arched toward the dorsal side.

NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE. BAT

The distal end is large and has a well-developed trochlea, which is much more strongly
convex than in Canis and of a different shape, the modern genus haying here a trochlea
which is more like that of a phalanx than of a typical metacarpal. In Daphwnus, but
not in Canis, there is a well-defined palmar carina, and the lateral processes for ligamen-
tous attachment are more prominent than in the recent type.

The second metacarpal is much longer and stouter than the first, though very short
with reference to the size of the animal and to the length of the other segments of the
fore limb. The proximal end is not much expanded transversely, but has a great dorso-
palmar extension, the head projecting much farther behind the plane of the shaft than in
Canis. The facet for the trapezoid is less concave transversely than in the modern genus
and is of more uniform width, narrowing less toward the palmar side ; the ulnar border
rises more above the head of me. iii and has a more extensive contact with the magnum.
Though larger than in the recent Canide, this contact with the magnum is much smaller
than in existing felines, and is of about the same proportions as in the early sabre-tooth,
Hoplophoneus. The combined facets for the magnum and for me. iii form a broad,
curved band upon the ulnar side of the head, which is made slightly concave to receive
the adjoining metacarpal. No distinctly marked facet for the trapezium is visible upon the
radial side. The shaft is short, weak, of transversely oval section, and is arched toward
the dorsal side. The distal end is expanded, and made broad by the large, rugose pro-
cesses for the attachment of the lateral metacarpo-phalangeal ligaments, processes which
are much better developed than in Canis. The distal trochlea is of a quite different shape
from that seen in the modern genus, being narrower, higher and of more nearly spherical
outline, and is demarcated from the shaft by a deep depression, such as does not occur in
the existing members of the Canide. The palmar carina is prominent and thins to a
narrow edge.

The third metacarpal is incomplete in the only manus found in the collection
(D. felinus, No. 11425, Pl. XX, Fig. 17) as it lacks the distal end. The portion pre-
served is, however, as long as the whole of me. ii and the complete bone was evidently
considerably longer. The shape of the proximal end is much as in Canis, except for the
relatively greater dorso-palmar diameter. The magnum facet is narrow, but deep, some-
what concave transversely and strongly convex antero-posteriorly, but less so than in
existing dogs. The facet on the radial side for me. ii is larger, more oblique and more
prominent, and is more extensively overlapped by me. ii than in the latter, and the
surface for me. iv, while not so deeply concave, is larger. When the third and fourth
metacarpals are placed together in their natural positions, it is seen that the former rises
higher proximally than the latter and has a contact with the radial side of the unciform,

which, though narrow, is larger thanin Canis. The shaft is somewhat more slender than

348 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

that of me. ii and is of a more quadrate section, the dorsal and lateral surfaces forming
distinct angles.

The fourth metacarpal has a narrow, but deep head, which projects prominently
behind the plane of the shaft; the facet for the unciform is slightly concave in the
transyerse and strongly convex in the dorso-palmar direction. Compared with the cor-
responding bone of Canis, the following differences in the shape of the facets for the
adjoining metacarpals may be observed. The surface for me. ili is, as in the recent
animals, divided into dorsal and palmar portions, but they are not completely separated ;
the dorsal moiety is much larger, but not nearly so prominent, and the palmar portion is
much smaller. The facet for me. y is of about the same shape in both genera. ‘The
shaft is slender and nearly straight, but slightly arched toward the dorsal side; though
relatively short, it considerably exceeds me. ii in length. The prominence of the lateral
ligamentous processes gives great proportionate breadth to the distal end. The trochlea
is like that of me. ii, except for its greater size and presents the same differences from the
modern type.

The fifth metacarpal has been lost from the specimen.

The phalanges are yery remarkable, but can be most conveniently described in con-
nection with the pes, with which the most complete specimens are associated.

Measurements.
No. 11424. No, 11425.
Scapho-lunar, breadth 0.015
24 «depth (Gorso-palmar)......ccccceccecesececeeseseerennscssneeeecceeseccesccccscccnseesecasccesessenssseseces 011
Metacarpal i, length......cccccccceceecceceeecececnecearteeeeeceeecnseeccesescaseesenessssseassseacesessteaeeneseeeercrsreccercs .023 .026
i breadth of proximal end........... Fecer CBRE cee aceccp sbonasenancspadsnaconqgceooe00g .007 .009
oo GE GINS HAIL G16 | po6conpc00snocob canon ooASEN gosDODODOSDSODOOsHCODSCODHODOsbODAbdSSGONoOSoCoNSSq9000056" .006
os €€ Gistal trochlea........sscsssessseecessceesencccesccssuceccaaeseceecuscsessesenssenserssccsscsens .0045
Metacarpal ii, length.........sccccccnceceeccnneeeeeecceneccesccescneanscecssseeccececesersrssesecesseesannnenss poac00t000000005 0395
es “ breadth of proximal end .009
UG ef 6G install @iaVél-coosoonneeanndcqn 000005000 pnodenodOD DO sDODODONdHEUOSDUSOUOSUBNBAOGSAOSDDASDOQcOSEN000 -012
a af ie GG TRGYEL HIKE jronconcccenscn nonoDononoDDqGOsODSNDSECENBDOS0R00CC Suneeaevesecseceseccesseseces .009
Metacarpal iii, breadth of proximal end........--eeceteeesssesesseceeeeeeeeeeneeeseeceranecasenaaeereeeeeeseeenecererces -0105
Metacarpal iv, length -050
se Dreadth of proximal end.........cccssseeecccccseseececsrsneeececeeececeseserencnsecsescassaseeessecenans .0095
ss ae GB GIS EAT (1116! coococooopacdbeug2o0dD50aHeccKO6 oc¢nod danoacansobapAAabocoacconopdaneocq—ICo0adaGn 030 .012
o ns a BO OCHeatenrateseeantecteeeesnetsester-teerane Spqnodqoaadqassonnq50bcanSSNOnDDSEeNeS .010

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. 349
VI. Tue Hinp Line.

The pelvis is represented by several specimens belonging to D. vetus, D. hartshornianus
and D. felinus, all of them incomplete, but so supplementing one another, that the shape
of the os innominatum may be determined, with the exception of the anterior border of
the ilium, which is unfortunately missing from all the individuals.

So far as it is preserved, the pelvis is rather feline than canine in character, both in
its general outlines and in its details of structure. The neck or peduncle of the ilium is
wider and shorter than in Canis, narrower than in Fe/is; the anterior plate expands to
its full width somewhat more abruptly than in the latter, but enough of the broken
fossils remains to show that the iliac plate has the narrow form which is found in the
cats and does not expand so much at the free end as in the modern dogs. The gluteal
surface is not simply concave, as it is in the two recent genera mentioned, but is divided
into two unequal fossee by a prominent longitudinal ridge, such as occurs, though not so
prominently developed, in certain viverrines. This feature is repeated in another White
River dog, Cynodictis, and is almost duplicated in the contemporary sabre-tooth, Dinictis,
another of the many correspondences between Daphenus and the early Machairodonts.
The sacral surface is placed much less in advance of the acetabulum than in Canis, and
oceupies about the same relative position as in the cats. The ischial border of the ilium
is, for most of its length, nearly straight and parallel to the acetabular border, but
descends more abruptly than in either the recent dogs or cats, and follows a course more
like that seen in Viverra. As in Canis, the acetabular border is more distinctly defined
than in the true felines, and ends near the acetabulum in a long, roughened prominence,
the anterior inferior spine. The pubic border is very short, and hence the iliac surface
is not well defined. The acetabulum is of moderate size and has somewhat more elevated
borders than in the cats.

The ischium, which in the existing Canide is much shorter than the ilium, is very
elongate, and is proportionately even longer than in the felines. The anterior portion of
this element is straight, rather slender, and of obscurely trihedral section; behind the
acetabulum the dorsal border is arched upward into a convexity, the spine of the ischium,
terminated abruptly behind by the ischiadic notch, which is as conspicuous as in the cats,
while in Canis it is very faintly marked. The posterior part of the ischium is expanded
into a broad and massive plate, which is very rugose upon the external surface. This
posterior portion is not so strongly everted and depressed as in the modern dogs, and
there is no such stout and prominent tuberosity, which, again, constitutes a resemblance
to the cats.

The pubis is L-shaped and its anterior, descending limb is unusually long, broad
and thin, much more so than in the felines or modern dogs. The obturator foramen is

350 NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE.

very large, forming an oval, with its long axis directed antero-posteriorly, in shape and
size agreeing much more closely with the condition found in the cats than with that of
the recent dogs.

The femur (Pl. XX, Fig. 18) is stout, and long in proportion to the length of the
fore-limb bones, but not very long as compared with the size of the animal. While not
differing in any very marked fashion from the thigh-bone of Canis, it yet has some
resemblances to that of the felines. The small, hemispherical head is set upon a longer
neck than in recent dogs and has a smaller, deeper and more circular pit for the round
ligament, than in the latter. As in Canis, the head projects more obliquely upward and
less directly inward than in Felis. The great trochanter is large and has a very rugose
surface, but it has no such antero-posterior extension, does not rise so high and is not so
pointed as in the existing forms of Canide. In consequence of this shape of the great
trochanter, the digital fossa is smaller and much shallower than in the cats or recent
dogs. From the great trochanter a sharp and prominent ridge, the linea aspera externa,
descends along the external border of the shaft. Whether a third trochanter was present
cannot yet be definitely determined, because in the only two femora preserved in the
collection, the outer edge of the shaft is broken away at the point where the third
trochanter would be, if present. In all probability, however, Daphenus did possess this
trochanter, at least, in rudimentary form, as may be inferred from the analogy of the
sabre-tooth Dzinictis, and still more from the little contemporary dog, Cynodictis, which
in many respects approximates the structure of the modern Canide more closely than
does Daphenus. The lesser or second trochanter is larger, more prominent, and of more
decidedly conical shape than in the recent species of either Canis or Felis.

The shaft of the femur is long, slender and nearly straight, though slightly arched
toward the dorsal or anterior side; it differs from that of the modern dogs in its lesser
curyature, and in broadening and thickening more gradually toward the distal end, and
from that of the true cats in being more slender and of more nearly cylindrical
shape. The rotular trochlea is rather narrower transversely than in the true cats,
or eyen than in Dinictis, but is characterized by the same shallowness, and resembles
that of the latter genus in its shortness vertically and lack of prominence. Trans- |
versely, the groove is but slightly concave, and it has much less prominent borders
than in the existing species of Canis ; these borders are slightly asymmetrical, the external
one rising a little higher and being a trifle more prominent than the internal. A decided
difference from both Canis and Felis consists in the fact that the trochlea hardly projects
at all in front of the plane of the shaft, the anterior face of the latter gradually swelling
to the level of the groove. In both of the recent genera mentioned, and especially in the
canines, the trochlea projects prominently in adyance of the shaft.

Jt
—_

NOTES ON THE CANID#® OF THE WHITE RIVER OLIGOCENE. De

The femoral condyles are feline rather than canine in shape; they are small and of
nearly equal size, though the outer one is slightly the larger of the two, and project
much less strongly behind the plane of the shaft than in Canis. They are also less
widely separated and less expanded transversely than in the latter genus. As in so
many features of the limb bones, the whole distal end of the femur is more like that of
Dimetis than it is like the corresponding part of the modern dogs or cats. In Dinictis,
however, the rotular groove is shorter proximo-distally and broader, and the condyles are
even less prominent.

The patella is very different from that of the recent Canide, in which group this
bone is small, narrow and thick, but has more resemblance to that of Dinictis. It is
quite broad, but very thin in the antero-posterior dimension; the anterior face is more
roughened than in the Machairodont genus and the proximal end is more pointed, not so
abruptly truncated. The facet for the rotular trochlea of the femur is, in correspondence
with the shallowness of that groove, but slightly convex transversely and slightly concave
proximo-distally.

The tdia (Pl. XX, Figs. 19, 20) is relatively short and slender, and bears consider-
able resemblance to that of Dinictis, more than to that of Canis. The proximal facets
for the femoral condyles are small and but little concave ; the outer facet is somewhat
larger than the inner, and projects farther beyond the line of the shaft, both posteriorly
and laterally. On the distal side of the overhanging shelf thus formed is a facet for the
head of the fibula, which is much larger than in the recent dogs and more rounded in shape
than in Dinictis. The spine of the tibia is very low and is more distinctly bifid than in
the Machairodont genus, though much less so than in Canis. As in the former, the
cnemial crest is not very strongly developed ; it is far less prominent than in the existing
Canide and does not descend so far upon the shaft as in them.

The tibial shaft is slender and nearly straight, not displaying the lateral and antero-
posterior curvatures seen in Canis ; proximally the shaft is of trihedral section, becoming
approximately cylindrical below and transversely oval at the distal end. The latter is
shaped much as in Dinictis and is conspicuously different from that of Canis; the
astragalar facets are less deeply incised, and the intercondylar ridge is less elevated than
in the latter, but the facets are deeper and the ridge higher than in the Machairodont, in
correlation with the deeper grooving of the astragalus. The large transverse sulcus,
which in the recent dogs invades these astragalar facets, is not shown in Daphenus.
The internal malleolus is very large and resembles that of Dinictis, save that its posterior
border is more inclined and the process is thus distally somewhat narrower. The sulcus
for the posterior tibial tendon is very distinctly marked, more so than in Canis. The

ANS 1, SE —AVOlby SADE, MHS

302 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

distal fibular facet is quite large, beimg much as in Dimctis and consequently much
larger than in ‘the recent Canide.

The fibula (Pl. XX, Figs. 19, 20), which is greatly reduced in the modern dogs, is
in Daphenus much stouter and has heayier ends, both proximal and distal. In Canis
these ends have the appearance of being reduced and simplified from the condition seen
in the White River genus. In the latter the proximal end of the fibula is relatively very
large, especially in the fore-and-aft dimension, in which it considerably exceeds that of
Dinictis, though the excess is principally due to a large tuberosity which projects from
the hinder border, and which is present, though much less prominent, in the Machairo-
dont. The facet for the head of the tibia is longer antero-posteriorly and narrower
transversely than in the latter, forming a long, narrow, irregular oval. The shaft of the
fibula is slender, though very much thicker both actually and proportionately than in
Canis, and has about the same proportions as in Dinictis ; it is laterally compressed, the

‘principal diameter being the antero-posterior one, and of oval section, though its size and
shape vary from point to point in an irregular fashion.

The distal end of the fibula resembles that of Dinictis, though it is somewhat smaller,
in proportion to the length of the bone. ‘The enlargement is both antero-posterior and
transverse and gives rise to a very stout outer malleolus, at the postero-external angle of
which is a deep sulcus for the peroneal tendons. The distal tibial facet is rather larger
than that of Dinictis, while the surface for the astragalus is somewhat smaller, the two

together making a high narrow band.

Measurements.

No. 11421. | No. 11424. | No. 11423.

Femur, length (fr. head) .........:scecccccecnseecececccces sesseeesecseneeenreoes adi toe acy ea 0.195
«« ‘Dreadth of proximal end........ ....0cceceeececcceeecececeececcecerencnerneeeeenereneeseeeseenee: - 044
es GO" _GbETHIL r0\6 ld scosocnbsoomnpnossodcodadedodonsos ods OD0ocd osa0DodDede: NoaDobaosOI0oDNGoD00 .038
“fs €€ POtUAT ZTOOVE.--------eeeeceeeeereneeeeeeeeccnessnenceseneccsetcserecessraeeseneeeees .014

Bis DAN Terie thiva set emma Sk es ai Boncas i tcc antalat cae ang ae ean ee ema 149°
¢ “‘preadth of proximal end .........-.-cscccsneeeccecensercnscecensercnesererscsensserensstecensesee .031 .036
cs OF AIP EYL Bia ly cacona nodoeacosnonadcosqecmedodaogss0N060579-0 nos oDsoOSaONBODaROnUBeNOSNOSS .021 .021 -025
Fibula, ant.-post. diameter prox. CN .....eeeceeeeeseeeeeeneeeesccereeeeseseescssaeceseetaareessecees 019
ef as ee Alii CO Sggecoeeesonencengoodbdbascodabononcocdscuencosasddcesocca00q000 .0145 .017

VII. Tue Pes (Pl. XX, Figs. 21, 21a, 22).

The pes, which displays structures of the highest interest, is much better represented
in the collection than the manus and may be more adequately described. As a pre-

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. d00

liminary, it will be useful to cite Schlosser’s account of the salient characteristics of the
hind foot among the recent Canide.

“Die Anordnung der Tarsalien und Metatarsalien weicht natiirlich weniger ab yon
jener der tibrigen Carnivoren als jene der Carpalien und Metacarpalien, doch finden wir
auch hier immerhin einige nicht unwesentliche Modificationen. Es hat sich das Navi-
culare ziemlich betrichtlich yverschmiilert, so dass es nicht mehr die Aussenseite der
unteren Astragalus-Partie umhtillen kann. Das Metatarsale II, das sonst nur von zwei
Punkten mit dem Mt. ITI in Beriihrung kommt, legt sich hier seiner ganzen Breite nach
an das Oberende desselben. In Folge der Verkiirzung des Tarsus ist auch der aufstei-
gende Fortsatz des Mt. V sehr kurz geworden. Die Phalangen haben gleich den Meta-
podien nahezu quadratischen Querschnitt, die Krallen sind sehr spitz, aber wenig gebogen,
haben jedoch ziemlich bedeutende Liinge. Die Hunde sind die ausgesprochensten Zehen-
gadnger unter allen Carnivoren ” (’88, p. 22).

In Daphenus the astragalus is decidedly different both from the astragalus of
Dinictis and from that of Canis, but approximates more the latter. The trochlea is low
and but moderately grooved, decidedly more than in Dinictis, but less than in the modern
dogs, and the articular surface does not descend so far upon the neck as in the latter.
The trochlea is asymmetrical, the outer condyle considerably exceeding the inner in size.
The neck of the astragalus is much longer than in Hoplophoneus, Dinictis, or even than
in Camis, and is directed more strongly toward the tibial side of the foot; the head is
depressed, but yery convex. The external calcaneal facet is hardly so large or so
oblique in position as in Dinictis, but it is more like the facet seen in that genus than
like the facet of Canis. The sustentacular facet is shorter and wider than in the
latter, and the sulcus separating it from the external facet is very much shallower. In
Dinictis the sustentacular facet has a posterior concave prolongation, such as is not found
in Daphenus, nor does the latter possess the distal accessory facet for the caleaneum
which is so distinctly shown in Canis. The navicular facet is depressed, but very convex,
and there is a small facet for the cuboid.

The caleaneum is more like that of Dinictis than that of the recent dogs ; though the
tuber calcis is longer, thinner and more compressed than in either of those groups, and
its dorso-plantar diameter is more uniform, increasing less toward the distal end; its free
end is less thickened and more deeply grooved by the suleus for the Achilles tendon.
Along the outer edge of the dorsal border is a quite deep and conspicuous groove, which
occurs also in Dinictis, but not in Canis. ‘The external astragalar facet is very like that
of the Machairodont, being more angulated and more oblique in position than in the
modern dogs, presenting inward as much as dorsally. The sustentaculum also resembles

that of Dinictis in being less oblique,much more prominent and in having its facet much

54 NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE.

(Js)

more widely separated from the external astragalar facet than in Canis. In the latter
genus occurs a third astragalar facet, which is distal to the sustentaculum, and which is
found in neither Dinictis nor Daphenus. The distal end of the calcaneum is occupied
by the large cuboidal facet, which is more regularly oval in outline and much more deeply
concave than in the existing forms of Canide. In these forms we find a facet for the
nayicular, which adjoins and forms a right angle with the accessory astragalar surface
already mentioned, but is not present in either of the White River genera. On the
external side of the caleaneum, near the distal end, is a prominent projection for liga-
mentous attachment. This process is not present in Canis, but it recurs in Dinictis, less
markedly in Hoplophoneus, and is found in many of the recent viverrines, mustelines
and raccoons.

The cuboid is not peculiar in any noteworthy way; it is longer proximo-distally
than in Dinictis and is proportionately narrower and thinner (7. ¢., in the dorso-plantar
diameter). The long, thick and rugose ridge which on the fibular side of the bone over-
hangs the suleus for the peroneal tendons is more prominent, especially on the plantar
face, than in the Machairodont, but lacks the great, rugose plantar protuberance, which
occurs in the recent Canide. The facet for the caleaneum is more convex than in
Dinictis, very much more so than in Canis, in which this surface is almost plane. On
the tibial face of the cuboid are three facets, a narrow proximal one for the navicular,
and a median and minute distal facet for the ectocuneiform. The facet for the head of
the fourth metatarsal is very much more coneaye than in the modern dogs, while that
for mt. v is smaller than in the recent forms, and lateral rather than distal in position.

The navicular, as compared with that of Canis, is short proximo-distally, but broad
transversely, not having undergone the reduction in width which Schlosser mentions as
characteristic of the recent members of the family. The astragalar facet is not more
concave than in the latter, and there is no such stout tubercle on the plantar side of the
bone as occurs in them. Two very small facets articulate with the cuboid, one near the
dorsal and the other near the plantar border of the fibular side. The distal facets for the
three cuneiforms have nearly the same shape and proportionate size as in Canis, but they
are more in the same transverse line, the surface for the entocuneiform being less dis-
placed toward the plantar side.

The entocuneiform is of similar shape, but relatively better developed than in Canis,
as would naturally be expected from the presence of a complete hallux in Daphenus.
The bone is long proximo-distally, thick antero-posteriorly, and narrow, though broader
than in Canis, and its proximal and distal facets, for the navicular and first metatarsal
respectively, are relatively larger and more coneaye. The only other facet is an obscurely

marked one on the tibial side for the mesocuneiform.

NOTES ON THE CANID& OF THE WHITE RIVER OLIGOCENE. = BOO

The mesocuneiform is a very small, wedge-shaped bone, broadest dorsally and thin-
ning to an edge on the plantar side. The nayicular facet is concave and yery different
from the curious oblique surface which we find in Dinictis, As is well-nigh universal
among the Carnivora, the proximo-distal diameter of this bone is much less than that of
either of the two adjoining cuneiforms, an arrangement which allows the head of the
fourth metatarsal to rise above the level of the first and third.

The ectocuneiform is, as usual, much the largest of the three, though it is not so
large proportionately as in Dinictis. The shape of this element is very much as we find
it in Canis, but with certain minor differences. Thus, the proximal end is less extended
in the dorso-plantar diameter, and the navicular facet is more concave; the plantar
tubercle has a more constricted neck and enlarged, rugose head; the facets on the tibial
side for the mesocuneiform and second metatarsal, and on the fibular side the inferior
facet for the cuboid are more distinctly developed, while the distal facet for mt. iii is more
concave and has a shorter plantar prolongation.

As a whole, the character of the tarsus is rather more machairodont, or viverrine,
than canine. A conspicuous difference from the tarsus of the modern Canidw, is to be
seen in the fact, that the articulations which in the latter are nearly plane (e. g., the
cubo-calcaneal) in Daphenus retain their more primitive concayo-conyexity.

The metatarsus consists of five members, which are longer and relatively more
slender than the metacarpals, though an exact comparison between the two cannot yet be
made, because the collection contains no specimens in which both metacarpals and meta-
tarsals are represented by anything more than fragments.

The first metatarsal is considerably longer and stouter than the corresponding meta-
carpal. In this case we can determine the true proportions, for of the species to which
the finely preserved hind foot (Pl. XX, Fig. 21) belongs, D. hartshornianus, we also
possess a pollex, though associated with a different specimen. The almost exactly similar
skulls of the two individuals show that the animals were of approximately equal size.
The head of mt. iis enlarged in both the transverse and dorso-plantar diameters, and bears
a roughened tubercle upon the plantar side. The proximal facet, for the entocuneiform,
is large, and strongly convex antero-posteriorly, nearly plane transversely; no other
facets are visible on the proximal end. The shaft is slender and arched toward the dorsal
side; in section it is transversely oval, expanding somewhat at. the distal end, where the
breadth is increased by the prominent tubercles for the lateral ligaments. The distal
trochlea is small, but well developed, and of irregularly spheroidal shape, with plantar
carina. The first metatarsal of Dinictis is like that of Daphenus, and certain viverrines,
such as Cynogale, also have a hallux of much the same proportions, but in all the
recent Canidae, with the exception of certain domesticated breeds, mt. 1 is reduced to a

nodule.

306 . NOTES ON THE CANIDHZ OF THE WHITE RIVER OLIGOCENE.

The second metatarsal is much longer and stouter than the first, but it is much
shorter and weaker than mt. ii in Canis, and rather resembles that of the viverrine genus
Cynogale, though it does not have the peculiar shape of the proximal end which charac-
terizes that genus. In Dinictis mt. ii is somewhat heayier than in Daphenus, but is other-
wise similar. In the latter the proximal end of mt. ii rises considerably above the level
of mt. i and iii, owing to the shortness, proximo-distally, of the mesocuneiform, and is
firmly wedged in between the ento- and ectocuneiforms, an arrangement common to all fami-
lies of the fissipedes and already general among the creodonts. On the fibular side is 2
wedge-shaped projection which is received into a corresponding depression on mt. iii,
thus making a very firm and close connection between the two bones. Above this pro-
jection are two facets for the tibial side of the ectocuneiform, one near the dorsal border
and the other on the plantar projection. The shaft is straighter than in Canis, but is
slightly arched dorsally, the distal end not curving toward the tibial side, as it does in .
the modern genus. In section the shaft is transversely oval, while in the recent dogs it
has become trihedral for most of its length, owing to its close approximation to the shaft
of mt. iii. The distal trochlea resembles that of Dinictis and differs from that of Canis
in its more spheroidal and less cylindrical shape, and in its demarcation’ from the
shaft by a deep depression ; the lateral ligamentous processes are likewise more symmetri-
cally developed.

The third metatarsal is much longer and stouter than the second, the difference
between the two being greater than in Dinictis or the viverrines, or even than in Canis.
The proximal end bears a facet for the ectocuneiform, of the usual shape, but the plantar
prolongation of this facet is shorter and broader than in the last-named genus, and it
resembles that of Dzinictis in being oblique to the long axis of the bone, inclining
decidedly toward the tibial side of the foot. The tibial side of this facet is deeply incised
to receive the wedge-shaped prominence of mt. ii, an incision which does not appear in
the recent dogs, but occurs, though somewhat less conspicuously, in Dinictis. On the
fibular side are two facets for mt. iv; one near the dorsal border, which is a deep
spherical pit, and the other a small, plane surface placed upon the plantar prolongation
of the head. The shaft, when viewed from the front, appears quite straight, but when
looked at from the side is seen to have a slight curvature toward the dorsal side. The
distal end displays the same differences from Canis as do the other metatarsals.

The fourth metatarsal forms a symmetrical pair with the third, very much as it does
in the recent dogs and cats, though in Daphenus they are relatively shorter and weaker.
In Canis these two metatarsals are closely pressed together for most of their length, and
their shafts have thus acquired a more or less trihedral section, with the approximate

surfaces flattened, while the distal ends curve away from each other, somewhat as in

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. 307

Poebrotherium. In Daphenus it is only the proximal portions of the two shafts which
are thus closely pressed together ; for the greater part of their length they are not in
contact, and thus preserve the primitive oyal section. As their divergence is due to the
relative positions of the tarsal bones, there is no necessity for the lateral curyature of the
distal ends. The two metatarsals are very closely interlocked and in much the same
fashion as in Canis. On the head of mt. iv are two facets for mt. ili, of which the dorsal
one is a stout hemispherical prominence, which is received into the pit on the head of
mt. ii, already described. The plantar facet is actually upon the plantar rather than on
the tibial face of the bone ; the prolongation from the head of mt. iii extends around and
embraces this facet, and by means of the double articulation a very firm interlocking of
the two bones is effected. On the fibular side of mt. iv is a large and deep depression
which receives the projection from mt. y. The facet for the head of the latter is large,
slightly concave, and continues without interruption from the dorsal to the plantar
border, while in Canis there are two distinct and quite widely separated facets. The
shaft resembles that of mt. iii, but is somewhat more slender. In both of these meta-
tarsals the distal carina is placed symmetrically with reference to the trochlea, but is less
compressed and prominent than in Canis.

The fifth metatarsal is not completely preserved in any of the specimens, the only
representative of it being the proximal end, belonging to a large individual of D. vetus
(No. 11423). As the specimen is incomplete, nothing can be determined respecting its
length, but probably this was equivalent to that of mt. ii, the two forming a symmetrical
pair, much as in Dinictis, though mt. y, so far as it is preserved, seems to be somewhat
the stouter of the two. On the fibular side of the head is a very prominent projection,
ending in a roughened thickening, and directed obliquely outward and upward, the
“ascending process ” (aufsteigender Fortsatz) of which Schlosser speaks in the passage
already quoted. In the recent dogs this process is very much reduced, while in Dinictis
it is of quite a different shape. In the Machairodont the process is a long and promi-
nent ridge, extending along the whole dorso-plantar thickness of the head, and projects
much more proximally than externally, while in Daphewnus it is a blunt hook which
projects more outward than upward. ‘The Machairodont Hoplophoneus has the process
developed in very much the same way as in Daphenus.

The facet for the cuboid differs from that of Canis in being quite concave transversely
and in presenting as much toward the tibial side as it does proximally, while in the
modern genus the facet is small, plane, subcircular in outline and altogether proximal in
position. On the tibial side is a rounded protuberance which fits into the pit on the head
of mt. iv; this protuberance is more prominent than in Canis and decidedly more so than
in Dinictis. What little of the shaft is preserved is transversely oval in section, with a

308 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

sharp ridge running down the fibular side, and is thus quite different from the trihedral
section, with flattened tibial side, which is found in Canis, and is much more like the
corresponding metatarsal of Dinictis.

The parallel arrangement of the metatarsals which we observe in the modern
Canide is in Daphenus replaced by a radiating arrangement, the bones diverging
toward the distal end. This distal divergence is, however, less decided in the pes than in
the manus.

The phalanges display a very curious and surprising combination of characters.
They are long, both actually and proportionately ; compared with the tibia as a standard,
they have about the same length as in the recent species of Canis, but they are decidedly
longer than in that genus when compared with the length of the metatarsals.

A proximal phalanx of one of the median digits is long and depressed, but quite
strongly arched upward or dorsally. The metatarsal facet has quite a different shape
from that seen in Canis, the transverse diameter being relatively greater and the dorso-
plantar less. The facet is also somewhat more oblique to the long axis of the phalanx,
presenting rather more dorsally and less entirely proximally ; the notch for the meta-
tarsal carina is less deeply incised. Similar differences are observable in the body of the
bone; its breadth being proportionately greater and its thickness less. The distal
trochlea, which in Canis describes a semicircle from the dorsal to the plantar surface, is
in Daphenus much more restricted, projecting less prominently from the plantar side and
not reflected so far upon the-dorsal face. On the other hand, this trochlea is more deeply
cleft in the median line than in the modern genus and the tubercles for the attachment
of the phalangeal ligaments are larger.

Tn all the differences from the modern Canidew which have been mentioned, we may
observe resemblances to the corresponding phalanx of Dinictis, in which the bone is
somewhat shorter and broader than that of Daphenus, and has rather more prominent
ligamentous tubercles, but is otherwise very like it.

The proximal phalanges of the lateral digits differ from those of the median pair
only in being shorter, more slender and less symmetrical, and in having a lateral curva-
ture which becomes yery pronounced in the hallux.

The second phalanx is of about the same length, with reference to the first, as in
Canis, but is broader, more depressed, and more asymmetrical than in that genus. The
proximal facet, for the first phalanx, is more distinctly divided into two depressions by a
more prominent median ridge, and the beak-like process of the median dorsal border is
much more pronounced. The distal trochlea is reflected farther upon the dorsal side and
projects more”from that side, but extends less upon the plantar face ; it is thus more con-
vex in the dorso-plantar direction, but much less concave transversely than in Canis.

259

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE.. oo

The asymmetry of this phalanx is quite marked: its tibial side is straight, while the
fibular border is quite coneaye, and the dorsal surface is hollowed, or cut away, near the
distal end, allowing a retraction of the claws, to a limited extent, as may be readily seen
when the second and third phalanges are put together. This asymmetry of the second
phalanx is much less conspicuous than in Dinictis, not to mention the modern felines,
but it is, nevertheless, unmistakable and is certainly one of the most surprising features
in the whole structure of Daphenus.

That an animal with the skull and dentition of a primitive dog should prove to pos-
sess even imperfectly retractile claws is not what our previous knowledge of the early
carnivores would have led us to expect. So unlooked for was this character, that at first
I was strongly inclined to believe that the association of the hind foot shown in Pl. XX,
Fig. 21, with the skull of D. hartshornianus was an accidental one, and that the pes
must belong to some genus of felines or Machairodonts as yet unknown. Fortunately, how-
ever, the collection contains a number of other individuals with more or less well-pre-
served hind feet, and the agreement among them all is complete. Curiously enough, the
characteristic second phalanges are preserved only in connection with the specimen
figured, but other specimens haye parts of the tarsus, metatarsus, proximal and ungual
phalanges, and a comparison of them shows that the reference of this particular hind
foot is not open to question. The fact that the pes and the skull were found enclosed
in the same block of matrix corroborates this inference, though, of course, such a fact is
not of itself entirely conclusive.

The ungual phalanx is hardly less peculiar than the second, being short, very much
compressed laterally, and bluntly pointed ; it is very little decurved and has a plainly
marked groove on the plantar face near the distal end. The narrowness, compression
and straightness of this claw are in very decided contrast to the heavy and strongly
decurved ungual phalanges of the modern Canidae, though among the latter there is con-
siderable variation in these respects. The articular surface for the second phalanx is
much more strongly concave than in Canis, permitting a greater freedom of motion in
this joint, as was necessary in order to provide for the retraction of the claw. The sub-
ungual process is not so large as in the modern genus and does not project so promi-
nently upon the plantar face of the bone, but it is produced much farther proximally,
extending beneath the distal end of the second phalanx, when the two are in their nat-
ural position. The long hood which envelopes the base of the claw is of about the same
size and shape as in Canis, though the space between this hood and the body of the
ungual phalanx is narrower. The ungual phalanx of Dinictis is shorter, more compressed,
but deeper in’ the dorso-plantar diameter than in Daphenus, and has a decidedly larger
subungual process, in correlation with the more complete retractility of the claws. The

A. P. S.—VOL. XIX. 27.

360 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

few specimens of these phalanges which I have seen are without the bony hood around
the base of the claw, having much the appearance of the unguals in the viverrine genus
Cynogale. It is possible that the apparent absence of the hood may be due to the break-
ing away of that delicate structure, but this does not seem very likely.

Measurements.

No. 10546. | No, 11421. No. 11424. No. 11423. No. 11425.
is
C@alcanenmemlen otiltscanae eaeeare) leben series eee tentel see seiaeeeeleee rena eter | 0.045 0.044 0.051 0.055
of dorso-plantar Ciameter.....-..-..cceeeeeeeeeeseeeeee eee eee 016 | -015 -020 020
“ Tenet Grit aber ee eee ie. eee tee coche ie. | 028 036 040
gs extreme distal bread th........-.0..sssseceescecereeeenneneee Ole | 017 -022 022
Astragalus, length ........... os gdaQoasdounocamnaqassoanoTH0OO usanSCo nEReeDOoG -027 -031 031
m PLOXimal Dread th. .....-......2.0-c-veceeeees eacceseerecneeere .018 .021 -022
= width of head..... nace 014 -016 -019
CITTSORGL, THEI] 118, .oxoscn9c0060 426500000 AnasaqnocAobsoanARoEtaasboDNSsDsedns0CO 015 .016
G0 SGVAKE I a cosace osedosasnnosqnoo¢bono pescSayaacan 59ND Den GaboSssaxED|TOOOIe | O11 -012
Navictlar, Width..........-:.cccesccsscneeecenccnereennecuseesenewnccsrecenees | O17 O19
Ectocuneiform, width ..... 010 O10
Metnbarsal to longthe.dvss.cte.ctee st teeta there echroes ccna] Weneost
s Dreadth prox. CNd..........cccscccsecneceneeensereerecscnees -009 010 |
i oe dist, °° 007
Metatarsal ii, length. .-...-2...0...0.c0ssccnssesecaecceecessccnercoenecrenre .044
a breadth prox. end ........c.sccecceeeceeeeceererereeecseneee .006 007 |
“ ia) Gddsbaoek hed arate ec are meee | .009 |
Metatarsal iii, length | -054
= Dreadth prox. CNd......-...ssccessceseceeeeeceseneceecerens -009 O11
" eFIn Hi cEMEIND naa us atte TE eae 0105 |
Metatarsal iv, length ....--...scceesssccneecccsceceneesceccectensssenseeseane | -056
ss breadth prox. end... 006
ie G6 GbE, © joononsmocoocadanoccbcesaceHbcanacaqccNeC .010
Metatarsal vy, breadth prox. Cn) .......-..ceeeeeereeeereeccneeneeesseneers -O11

The species of Daphenus hitherto recognized are three in number, two of them, D.
vetus Leidy and D. hartshornianus Cope, from the White River stage, and the third, D.
cuspigerus Cope, from the John Day. Two additional species are described in the sequel,
one of which, however, can be referred only provisionally to the genus, until more com-
plete material has been obtained, though the species in question is evidently very closely
allied to Daphenus, if not actually referable to it.

Dapuenvs vetus Leidy.

Dephanus vetus Leidy, Proc. Acad. Nat. Sci. Phila, 1853, p. 393. Amphicyon vetus
Leidy, ibid., 1854, p. 157; 1857, p. 90.. Hxtinet Mamm. Fauna of Dakota and
Nebraska, pp. 32, 369. Cope, Tertiary- Vertebrata, p. 896.

This species has a skull about equal to that of the coyote (Canis latrans) in size,

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. 361

but the vertebrae are much larger and the tail is longer and stouter. The tubercular
molars of both jaws are relatively larger than in the other species. The inferior sectorial
has a low anterior blade, and the internal cusp of its talon is reduced in size. The hori-
zontal ramus of the mandible is long and slender and has a nearly straight inferior bor-

der. White River.

DAPHENUS HARTSHORNIANUS Cope.

Daphenus vetus Leidy, Amphicyon vetus Leidy, in part, loc. cit. Canis hartshor-
nianus Cope, Synopsis New Vert. from Colorado, 1873, p. 9. Ann. Rept. U.S.
Geolog. Surv. Terrs., 1873, p. 505. Amphicyon hartshornianus Cope, Tertiary
Vertebrata, p. 896.

This species is somewhat smaller, and the tubercular molars of both jaws are propor-
tionately smaller than in the preceding species; the anterior triangle of the lower secto-
rial is high and acute, and its talon is basin-shaped, with the internal cusp as large as
the external. The horizontal ramus of the mandible is straight and slender. Both this
species and the preceding one have been found in the middle division (Oreodon beds) of
the White River formation, but not as yet, to my knowledge, in the lower (Titanothe-
riam beds) or the uppermost division (Protoceras beds).

DaAPHENUS CUSPIGERUS Cope.

Canis cuspigerus Cope, Proc. Amer. Phil. Soc., 1878, p. 70. Amphicyon entoptychi
Cope, wid., 1879, p. 872. Amphicyon cuspigerus Cope, Bull. U. S. Geolog. Surv.
Terrs., Vol. vi, p. 178; Tertiary Vertebrata, p. 898.

D. cuspigerus is much the smallest known species of the genus. The sagittal crest
is very short and inconspicuous ; the cranium is fuller and more rounded, the postorbital
constriction is shallower and more anterior in position than in the White River species,
and the mandibular ramus is nearly straight and very slender. The inferior sectorial is

very robust and has a low anterior triangle and basin-shaped heel. John Day stage.

DAPHENUS FELINUS, sp. nov.

The inferior dental series of this species slightly exceeds in length that of D. vetus
and the sectorial is larger. The lower tubercular molars are inserted in the border of
the ascending ramus of the mandible, and, judging from the alveoli, were reduced in size.
The horizontal ramus is not much longer, but much heayier than in D. vetus, and has a

more sinuous ventral border, which rises more beneath the masseteric fossa. The limb

362 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

bones and vertebrae are somewhat larger and heavier than those of D. vetus, and the neu-
ral spines of the lumbar vertebree are very high and incline strongly forward. In size
D. felinus is the largest and most massive species of the genus. The type specimen
consists of a fragmentary skeleton (No. 11425) with which are associated both mandibu-
lar rami, and which was found by Mr. Gidley in the Oreodon beds of Hat Creek Basin,
_Neb., in 1896.

? DapHznus Doneet, sp. noy.

As already intimated, the reference of this species to Daphenus cannot yet be defin-
itely made, but the material so far obtained, consisting of lower jaws, affords no sufficient
ground for separating it from that genus. The inferior dental series is relatively short ;
the premolars are much smaller, especially in the antero-posterior dimension, than those
of the later species from the Oreodon beds, but, at the same time, they are proportion-
ately thick and heavy. The lower sectorial has a low, massive anterior triangle and a
basin-shaped talon, with the inner cusp much smaller than the outer. The horizontal
ramus of the mandible is short, but relatively much stouter than in any of the other
species, and has a more sinuous ventral border, which rises steeply toward the angle.

This species is dedicated to my friend, Mr. Cleveland H. Dodge, of New York,
whose liberality has made possible much of the work undertaken by the Princeton
Museum and to whose kindness I am under the greatest obligations.

The type specimen (No. 11422) was found by Mr. Gidley in the Titanotherium
beds of the Hat Creek Basin.

Before proceeding to an examination of the next genus of White River Canide,
Cynodictis, it will be necessary to introduce a brief description of a species which has
been found in the Uinta stage of the upper Eocene (or lower Oligocene) and which ap-
parently represents the forerunner of Daphenus, though more perfect specimens will be
required before its position in the canine phylum can be definitely determined,

MIACIS Cope.

This form differs from Daphenus in the construction of the upper tubercular
molars. M1 has an exceedingly broad external cingulum, forming at the antero-exter-
nal angle a very large projection ; the internal unpaired cusp found in Daphenus and
in all subsequent genera of the Canid@ is absent in both m+ and m2. The upper secto- -
rial is of yery primitive and undeveloped character in the shortness of the posterior cut-
ting ridge and the great transverse breadth of the crown,

NOTES ON THE CANIDE® OF THE WHITE RIVER OLIGOCENE. 363
Mracis vrinrensis Osborn.
Bull. Am. Mus. Nat. Hist. N. Y., Vol. vu, p. 77.

Size rather less than that of D. hartshornianus; upper sectorial relatively small and
tubercular molars large ; premolars short and thick.

Measurements.
MM.
Men cths pe scOMMeniMClusivetc-sseccons cecccsetanascneeressuccacssceessescecasccsssceescscccostcvecuitecservoseans 37

P * length
P+ length
P= widthes:-<---ce Sa0dD 5 ROESDOE COSC ANS ISHSOS OS DUCEE SAEED” -o/aucH UNE BoC Gac0 Bocas Bode Su SuSoCeaae eee aceaee aaEeReEeere 11
M + length...
Me Dev dthv eens faeces ae cea behest RELL. Se oatbee ee gd i tad Feil ceded lal ty
aU EAS ay Tea fl Nogebeaecoaaho I eaco- bo eas ageo NSO RO Repo TeSUer ao Jon ec ERE EE ne Ere SRS EERE
INCE TIGh it soccccs bo aencacbSdanc sad See qeande Yondos ed aacosaconcc EISEN GELS a cicee ecSCeHCuG Ie ne SUR OEE ene eee Hed)

Fie. A.—First upper molar of the left side :
1, of ? Miacis wintensis. 2, of Daphxnus hartshornianus. 3, of Canis latrans. x, cusp usually regarded as the

protocone.

If Ihacis be rightly regarded as having a place in the canine phylum, then the
structure of its upper tubercular molars is of great interest and will require a revis-
ion of the current views concerning the homologies of the cusps in the upper molars of
the dogs. In Cunis, according to the usual interpretation, m 1 is composed of two external
cusps, the para- and metacones, and at the apex of the triangle of which the para- and
metacones form the base, an unpaired internal cusp, the protocone, with the proto- and
metaconules on the anterior and posterior sides of the triangle respectively. Internal
and somewhat posterior to the protocone is a large crescentic cusp, which is commonly
regarded as an enlargement of the cingulum, although in unworn teeth a faint cingulum
may be traced all around this crescentic cusp and is continuous with the prominent cin-
eulum which bounds the anterior wall of the crown. If this interpretation of the cusps
be correct, and further, if Jfacis is ancestral to the Canide, them min the Uinta
genus is without a protocone and has only the para- and metacones, minute conules and
the large inner crescentic cusp. Itseems much more rational to conclude that the lat-
ter is really the protocone and that the cusp which has been so named in Canis is an

additional element subsequently developed. In Daphenus this inner crescentic cusp and

364 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

the conules are relatively smaller than in the modern representatives of the family, which
goes to confirm the conclusion that the name protocone should be given to the innermost
cusp and that in Canis the middle part of the crown has undergone a special increase in
complexity.

CYNODICTIS Gervais.
Amphicyon Leidy, Marsh, in part. Canis Cope, in part. G'alecynus Cope, non Owen.

It is with much hesitation that I employ the name of this Kuropean genus for North
American species, for there are certain constant differences which Schlosser (’88,)
appears to consider as being of generic value. An actual comparison, however, of the
American forms with specimens of Cynodictis lacustris, Gervais’ type species, and from
the typical locality, Débruges, has failed to reveal any important differences between the
two, and, therefore, for the present at least, I retain the name of the European genus for
the American species, which are very closely allied, if not positively referable to it.

The structure of these small carnivores, especially of the John Day species, is much
better known than that of Daphenus, though our knowledge of the White River species
has hitherto remained yery incomplete, and even of the better known John Day forms —
only Cope’s brief descriptions have as yet been published. Despite the fact that Cyno-
dictis is one of the commoner White River fossils, well-preserved specimens are com-
paratively rare and of these the greater part consist only of skulls. The bones of the
skeleton are so small and so fragile that it is exceedingly difficult to obtain more than
fragments of them. By dint of great care and attention paid to these small formis,
Messrs. Hatcher and Gidley have succeeded in gathering some very fine specimens for
the Princeton Museum, and others I owe to the kindness of Mr. John Eyerman.
Together, these various individuals represent nearly all parts of the skeleton and enable
us to reconstruct the animal and to compare it with the better preserved and more
abundant species of the succeeding John Day formation.

I. The Dentition.

The dental formula of Cynodictis is: I 3, C4, P 4, M 2, differing from that of
Daphenus only in the absence of the third upper molar.

A. Upper Jaw.—The incisors are very ‘small, simple and antero-posteriorly com-
pressed, giving them chisel-shaped crowns; they increase in size from the first to the
third, but the latter does not greatly exceed the others; not nearly so much, for exam-
ple, as in Canis or Daphenus, and hardly more than in the viverrines. A very short
ciastema separates the lateral incisor from the canine.

The canine has a stout, gibbous fang, which produces a marked convexity upon the

side of the maxillary ; its crown is quite elongate and somewhat recurved and much com-

NOTES ON THE CANIDEH OF THE WHITE RIVER OLIGOCENE. 365

pressed laterally. The tooth is relatively smaller than in the recent dogs and thinner
transversely, and has therefore quite different proportions from those seen in Daphenus.

The premolars increase in size posteriorly ; in the unworn condition they have high,
compressed, thin and very acute crowns, but in old individuals, without showmg much
appearance of wear, these teeth have low crowns, elongated in the fore-and-aft direction.
The first premolar is very small and simple; it is inserted by a single fang and follows
immediately behind the canine, without a diastema, which is a difference from Daphenus.
The second premolar is much larger than p+; it is implanted by two fangs and has a
perfectly simple crown, without posterior basal tubercle, though the cingulum is thick-
ened at that point. The third premolar is still larger, especially in the vertical height
of the crown, and is distinguished by the presence of a posterior tubercle in addition to
the thickening of the cingulum already found in p 2. The fourth premolar is a very
effectively constructed, though small, sectorial blade, being much more compressed and
trenchant than in Daphenus. The anterior cusp of the shearing blade (protocone) is
relatively higher and thinner and has a sharper point and edge than in the latter genus,
and the posterior cutting ridge (tritocone) is better developed and more efficient. On
the other hand, the internal cusp (deuterocone) is very much smaller (hardly larger
proportionately than in Canis) and occupies a more posterior position. In the Euro-
pean species of Cynodictis the deuterocone is not so much reduced and is placed as far
forward as in Daphenus.

The first molar is large, particularly in the transverse dimension, and is of subquad-
rate outline. The outer cusps are high and quite acutely pointed, and the central cusp
(usually called the protocone) is lower and of crescentic shape, and the internal cusp is
a broad, crescentic shelf, which occupies about the same position as in Canis. The
ecnules are very small, but of nearly equal size, a difference from the modern genus, in
which the metaconule is large, while the protoconule is rudimentary or absent, and eyen
in Daphenus the posterior conule is much the larger of the two. The cingulum is very
prominently developed upon the outer side of the tooth and forms a large projection at
the antero-external angle, as in Daphenus, though not in Canis, a reminiscence of creo-
dont ancestry.

In the John Day species, C. geismarianus and C. lemur and still more in C. lati-
dens, the first upper molar has a much more distinctly quadrate crown, due to the enlarge-
ment of the metaconule, which has become as large as the central cusp, and to the more
symmetrical development of the internal cusp (? protocone). In the typical European
species, C. lacustris, on the contrary, the crown of this tooth retains a more trigonodont
character.

The second molar is very small, being relatively much more reduced than in Daphe-

366 NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE.

nus. It is composed of the same elements as m4, but has a different shape, owing to
the greater proportionate length, antero-posteriorly, of the inner portion of the crown.
In appearance this tooth is a miniature copy of that of Canis.

B. Lower Jaw.—The incisors are very small and closely crowded together, so that
the fang of i 5 is pushed back out of line with the other two.

The canine, which is even more compressed laterally than the upper one, is long and
recurved ; it is separated from p ; by a very short diastema.

The first premolar is a very small, simple cone, inserted by a single fang. The sec-
ond is much larger and is supported by two roots; it has an anterior basal cusp, which
is formed by the cingulum and is subject to considerable variation, being much larger in
some individuals than in others. The third premolar has a high, compressed and sharp-
pointed crown and bears three accessory cusps, anterior and posterior basal cusps formed
by the cingulum, and a third developed upon the posterior edge of the protoconid, very
much as in Canis. The fourth premolar is slightly larger than p 5 and has more dis-
tinetly developed accessory cusps, but on both p 3 and p q these cusps are subject to much
variation and in some specimens they are feebly marked or eyen absent.

The European C. intermedius has very similar premolars to those of C. gregarius,
and in both species the anterior basal cusps (which are not present in Daphenus) give a
somewhat viverrine character to the dentition.

The first molar has a quite elevated anterior triangle, with a high, pointed proto-
conid and a well-developed paraconid, both of which are more compressed and trenchant
than in Daphenus. The metaconid is smaller than in the latter and is placed lower
down and more posteriorly, so that it is visible from the outer side, much as in the mod-
ern dogs. The heel is basin-shaped and is composed of a large, crescentic external cusp
and a smaller internal cusp. In the European species may be observed certain differ-
ences in the structure of the lower sectorial from the White River form, though these
differences are not great. In the Old World species the anterior triangle is higher and
the protoconid less compressed, while the metaconid is larger and occupies a more ele-
vated and anterior position; in other words, the anterior triangle resembles that of
Daphenus. Another difference from the American forms consists in the presence of a
second internal cusp in the heel of the sectorial, which may be observed in most of the
individuals figured by Schlosser and Filhol. Howeyer, in a specimen of C. /acustris from
Débruges, which the Princeton Museum owes to the courtesy of Prof. Gaudry, this sec-
ond cusp is not visible. In perfectly unworn teeth of Daphenus hartshornianus a feeble
indication of this second cusp may be seen.

The second molar is tubercular and of a narrow and elongate oval shape ; in consti-
tution it entirely resembles that of Canis; the paraconid has disappeared, while in

NOTES ON THE CANIDH® OF THE WHITE RIVER OLIGOCENE. 367

Daphenus it is still distinctly visible, though very small. The proto- and metaconids
are of equal size and placed on nearly the same transverse line; these cusps are higher,
more sharply pointed and more slender than in the recent Canidw. The talon, which is
somewhat lower than the anterior half of the tooth, retains a distinctly basin-like form.
In the European species we find a more primitive character of m 5 in the retention of the
paraconid. The third molar is yery small ; it has an oval, roughened crown and is car-
ried upon a single fang. As Cope has pointed out, this tooth is usually missing in the
fossils, and occasionally a specimen is found which has not even an alveolus for it.

The dentition of Cynodictis gregarius is, on the whole, a little more modernized and
advanced than that of the European representatives of the genus. This adyance is shown
in the reduction of the inner cusp of the upper sectorial ; in the somewhat more quad-
rate outline of m 1; in the less elevated shearing blade and more posterior position of
the metaconid on the lower sectorial, and, finally, in the more complete reduction of the
paraconid of m 3. In the John Day species, especially in C. geismarianus and C.
latidens, the departure from the European type is even more marked.

Measurements.

No. 10193. | No. 10513. | No. 10939. | No. 11012. | No. 11382. | No. 11432.

Upper dentition, length I1 to M 2... 0.044 0.044 | 0.0485 0.0435
Upper canine, ant.-post. diameter............sseeeeeeeeees 005 005 005 -0045

is «« transverse Mae roasts Se ste fesesevstee dente: 0035 .003 003

Dee premolar series, lena this--c-esensebeese)le-ecerereseo== 025 | 923 025

“¢ molar series, length 010, | ~— 010 O1L LOOM OLO -010

aeeSPe lel erie tilivesscstyss-snsee cacatsseoveseeavepauesnersoscee: .0035 003 | .003 .003

C8 IPB SFO capsdiaansoodsoscodcnsadsoncobsnsocHnsnoesnonnoce -005 .0045 2.004 | .0045 003

SpTep svt) use Bae ales NES Toe Dery, 2 0055 005 | 0055

CIP AL GB Aeaeesacancctcopercosesccbanonue sedabsdooonecee -010 | -009 -009 .0095 | 0085 -009

COL SETBIL STORE LN ec pO Eocene 006 | 2.004 .0055 | .005

Sri Mp (og lerieet tio sstecet xeon sect cet tce,tace Se coco: 0065 | 007 .006 006 = —-.006 .006

“« M1, breadth..... ae ccd| 00) | .008 008

OE WB Ueya1N2550c 3:0ce odcccaauaneeouensocs! OsnaqoedaKeecos .0035 | .003 -004 004 -003 -003

CC WEB Toren chD cocossosconcosocecosoabs sonopososacesabecadHe -006 -006 0055 | .004 -005
Lower premolar series, length.....--- 02.00.00. +:11eeeeeeeeees | | 1021S eae OL9

‘“« molar series, length... 017 ~+| ~ .016 017 -015

4 SP evlenathvans tn. tac Sana ee enn | 003 | 003 |

TBO NRCP ae oBS sec cooper RSE ee aE S005 gm en 005i ia 004: ~ |

oC SI2Gy ete ei taie seteeisetetelentetseiseteetciseecicecissiemettcstetest -0055 -005 | .006 -005

OC aad colt: OC Sectodins SaQsos0Oc Uo oTOSHaSdCScosEC CaS BASOSONeCS -0065 -007 -006 | 0065 | .006

PaREAMIV aT pactne: Kates Say eee Ni 010 | .0095 0095 | .010 | .009

BO NED OG nance seocascacdoosbongaconoagsagsDasbAGECeGa5 005 -005 5005-00 | 0045 |

«<M 2, breadth -003 | 003 .0035 |

A. P. S.—VOL. XIX. 2 U.

368 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.
JUL, “Abisns}ysituney (IEG SSID Ieee Ly by)

The skull of Cynodictis is decidedly primitive and in general appearance resembles
that of such viverrine genera as Paradoxurus, rather than that of the modern Canidae.
Among the latter the alopecoid series have skulls more resembling the type of Cynodictis
than do the thooids, though the Brazilian bush-dog (Jcticyon) is, on the whole, most like
the fossil in the proportions of its skull.

In Cynodictis, as in Daphenus, the facial or preorbital region of the skull is very
short and the cranial portion very long. The occiput is low and the upper contour of
the skull rises steeply from the inion to about the middle of the parietals, whence it
descends in an almost straight line to the anterior nares, the only departure from straight-
ness being a hardly noticeable concavity or “dishing” of the nasals about midway in
their length. In Vulpes the profile is quite similar, but the posterior rise from the occi-
put is much shorter and less steep, and the dishing of the nasals is more conspicuous.
The sagittal crest is low and weak, and in the John Day C lemur, the smallest species
of the genus, the crest is replaced by a lyrate sagittal area. The cranium, though slen-
der, elongate and contracting anteriorly, is relatively fuller and more capacious than in
Daphenus, and the postorbital constriction, though much deeper, is as near the orbit as
in the modern foxes, and is, therefore, much farther forward than in Daphenus. ‘The
John Day specimens, which Cope has referred to C. gregarius (85, Pl. LX VIII, Fig. 6),
have an eyen fuller cranium and shallower postorbital constriction, which should, per-
haps, be a reason for separating these animals specifically from the White River forms.
The muzzle in Cynodictis is very slender, but tapers gradually and is not so abruptly
constricted at the line of the infraorbital foramina as in Daphanus. In the European
representatives of the genus the skull is much like that of the American species, but is
somewhat more primitive and like that of Daphenus. Thus, the muzzle is more abruptly
constricted, and the postorbital constriction is deeper and occupies a more posterior posi-
tion.

A more detailed examination of the skull brings out the following facts :

The occiput is low, very broad at the base and narrowing toward the summit less
than in the large wolves, but more than in Vulpes or Urocyon ; a well-marked median
conyexity is produced by the vermis of the cerebellum. The crest of the inion is low
and weak, much less prominent than in Daphenus. 'The foramen magnum differs some-
what in shape in the different individuals, being in some low and broad, and in others
of subcireular outline, a difference which may, in part, be due to a slight crushing. The
dorsal margin of the foramen projects much more prominently than in the recent Canide.

The basioccipital is long, broad and of nearly uniform width throughout; it is

NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE. 369

slightly concave transversely, but has a low median conyexity, with very feebly devel-
oped keel, the convexity being much less prominent than in Daphenus.

The exoceipitals are low and wide and so conyex in the median line that this por-
tion projects much behind the sides. The condyles are low and depressed and are
separated on the ventral side by a narrower, deeper and more V-shaped notch than
in the modern wolves or foxes. The paroccipital processes are very small and_ project
almost directly backward, as if to avoid the auditory bulla, with which they are not
in contact at any point.

The supraoceipital isa large bone, both high and broad; dorsally it is reflected
over upon the cranial roof, and in this region is thickened and diploétic.

The mastoid is exposed quite extensively upon the occipital surface, somewhat more
so than in the modern representatives of the family, and as the distance between the
paroccipital process and the postty panic process of the squamosal is greater than in the
latter, the mastoid occupies a rather more lateral position. The mastoid process is very
small, almost obsolete.

The sphenoid bones cannot be described, as none of the specimens allow the limits
of these elements to be gletermined.

The tympanic differs in very important ways from that of Daphenus. In the first
place it is inflated into a very much larger auditory bulla, filling out the entire fossa
and leaving no part of the periotic exposed ; and in the second place, the posterior cham-
ber of the bulla is ossified and fused with the anterior chamber. The line of junction
between the two elements which compose the bulla is very plainly marked by a groove
upon the external surface, and shows the posterior chamber to be considerably the smaller
of the two. I have not been able to detect any, even partial, septum between the two
chambers, but such a septum as that of Canis may well haye been present. The bulla
is relatively as elongate as that of Canis, but is much narrower and more compressed,
and therefore has a less inflated appearance. ‘The external auditory meatus isa very large,
oval aperture, without any tubular prolongation, the borders being flat, except the ante-
rior one, which forms a more prominent lip than in Canis and partially conceals the
postglenoid foramen. The auditory bulla of Cynodictis is thus thoroughly cynoid in
development and displays no resemblance to the characteristic viverrine type.

The parietals are proportionately very large bones and make up the greater part of
the sides and roof of the cranium. Throughout their length they unite to form a very
low and weak sagittal crest, which becomes moderately prominent only at the concavity
of the cranium formed between the occipital crest and the hinder wall of the cerebral
fossa. Owing to the larger size and backward extension of the cerebral hemispheres, as

well as to the lowness of the occipital crest, this concavity is shorter and much shallower

370 NOTES ON THE CANID#Z OF THE WHITE RIVER OLIGOCENE.

than in Daphenus. In some specimens, even aged ones, the anterior half of the parietals
carries a very narrow sagittal area, rather than a crest, but only in the little C /emur
from the John Day does this area assume the lyrate form. ‘This fact is of importance in
determining the primitive or secondary nature of the sagittal crest, concerning which
there has been some dispute.

The frontals form relatively as much of the cranial roof as in Canis and haye, when
viewed from above, an hour-glass shape, which is due to the deep postorbital constriction,
though the depth of this depression varies considerably in different individuals. The
postorbital processes are very small and owe their prominence entirely to the constric-
tion. The forehead is slightly convex, both transversely and longitudinally, though in
some specimens it has a narrow and shallow depression along the median line, such as is
found, though much more distinctly, in modern species of both Canis and Vulpes. The
forehead is bounded by the obscurely marked supraciliary ridges converging posteriorly
to the sagittal crest, which is entirely upon the parietals, none of it being formed by the
frontals. Anteriorly the frontals are emarginated to receive the narrow nasals, and send
forward slender nasal processes, which are separated by short interspaces from the
ascending rami of the premaxillaries. A noteworthy difference from Daphenus consists
in the absence of frontal sinuses, in which respect Cynodictis agrees with the alopecoid
series of the modern Canidae, as Daphenus does with the thooid series. The significance
of this fact will be discussed in a subsequent chapter.

The squwamosal has a relatively small extension upon the side of the cranium, and
this portion of it has a different shape from that seen in the modern dogs, the pari-
etal suture descending very steeply forward from the occipital crest, while in the modern
genera this suture pursues a nearly horizontal course. From the base of the zygo-
matic process to the posttympanic process of the squamosal runs a projecting shelf,
which overhangs the auditory meatus and is much wider than in Canis or Vulpes,
though not so broad as in Cynodesmus, Hypotemnodon or Daphenus. The posttym-
panic process is not larger than in Canis, but is made more conspicuous by the absence
of any tubular meatus auditorius. The zygomatic process is relatively somewhat heavier
than in Vulpes, and in shape and proportions much like that of the wolves, though not
so strongly arched upward; anteriorly it extends to the postorbital process of the
jugal. The glenoid cavity is broad and the postglenoid process is proportionately heavier,
more extended transversely and its distal end is more curved forward than in Canis.
There is no preglenoid ridge. :

The jugal also resembles that of Canis, though it displays some differences. Thus,
it is not quite so long as in the modern genus and does not extend so near to the glenoid
cavity ; it has a less decided upward curvature, and the postorbital angle (it can hardly

be called a process) is even less conspicuous ; the masseteric surface is broader, more lat-

NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE. 37/1

eral and less inferior in position, and is bounded above by a distinet crest; the antero-
inferior, or maxillary, process is shorter, and the ascending, or frontal, process is narrower,
but extends farther upward along the margin of the orbit. As a whole, the zygomatic
arch is of nearly the same proportionate length as in Canis latrans, but has a straighter
fore-and-aft course, being much less strongly arched upward, though curving outward
quite as decidedly from the side of the skull. This comparative shortness of the arch,
in association with the very elongate cranium, is due to the anterior position of the zygo-
matic process of the squamosal, which is placed much farther in advance of the occi-
pital condyle than in the recent members of the family.

The dachrymal forms but a very smail portion of the anterior rim of the orbit and
carries a rudimentary spine. Within the orbit the bone is relatively more extended and
occupies a more eleyated position than in the modern dogs, while the ascending or fron-
tal process is much shorter ; the lachrymal foramen is large and is farther removed from
the frontal suture.

The nasals are short, narrow and slender, splint-like bones, which are convex trans-
versely and very slightly concave antero-posteriorly ; their general shape is much the
same as in Vulpes, except for the much less distinct fore-and-aft concayity and their lesser
elongation.

The premaxillaries are small ; the alveolar portion is weak, in correspondence with
the smallness of the incisors, and is not produced anteriorly in the spout-like form which
characterizes Daphenus ; the groove for the reception of the inferior canine is much less
deeply incised than in the latter. The ascending ramus is long and slender, but forms a
wider strip upon the side of the muzzle than in the last-named genus. The anterior narial
opening is small, oval in shape and more oblique in position than in either Canis or Vul-
pes. The palatine processes of the premaxillaries are short and very narrow, and the
incisive foramina are small. This portion of the palate has an entirely different appearance
from that found in Daphenus ; the premaxillaries are not nearly so much extended in
front of the canines, the incisive foramina are shorter and have no such grooyes extending
forward from them ; the spines are very slender and much shorter, reaching only to the
canines and not to the line of p 4, as they do in the larger genus. In most of these
respects Daphenus is nearer to Canis and Vulpes than is Cynodictis.

The mazillaries are relatively very short, much shorter than in the existing genera,
a statement which especially apples to the facial or preorbital portion. At the same
time the vertical height is proportionately great. Except for the swelling produced by
the root of the canine, the facial surface of the maxillary is simply conyex, there being
no distinctly marked fovea maxillaris. Owing to the shortness and height of the facial
portion, its superior and anterior margin, formed by the sutures with the frontal, nasal

and premaxillary, is more strongly curved and descends much more steeply in front than

372 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE,

in Canis. As in Daphenus, the infraorbital foramen is placed very near to the orbit,
while in the modern genera it is much in adyance of the orbit. The arrangement seen in
Cynodictis is due chiefly to the anterior position of the orbit and in much less degree to
the backward shifting of the foramen itself. The palatine processes of the maxillaries are
short and narrow, corresponding to the shortness and slenderness of the muzzle, and they
resemble those of Daphenus in being slightly concave transversely, with a faintly marked
median ridge along the line of suture.

The palatines have nearly the same shape and proportions as in Canis latrans (though
they are relatively somewhat narrower) and extend forward to the anterior edge of p +;
the palatine notch is more deeply incised than in either Canis or Vulpes and is nearly as
deep as in Urocyon. Only a single posterior palatine foramen is visible on each side.
As a whole, the bony palate resembles that of Canis more than that of Daphenus in its
much less abrupt narrowing at the level of the sectorials. The posterior nares have
about the same shape and position as in Vulpes and have a similar median spine-like
process on the anterior border.

The pterygoids terminate in longer, more distinct and more thickened hamular pro-
cesses than in the recent genera, some of which, like Urocyon, have no vestige of such
processes. From the descending process of the alisphenoid is given off a prominent
lateral spine, which, in Canis and Vulpes, is represented only by a low ridge.

The mandible has a slender and compressed horizontal ramus, which tapers rapidly
toward the anterior end; it forms a long symphysis with its fellow of the oppo-
site side and curves very gently upward at the chin. The ventral border describes a
somewhat sinuous course, curving downward beneath the sectorial, from which point it
rises yery gradually and regularly to the symphysis, while beneath the masseteric fossa
it is coneaye. There is no trace whatever of the lobation which is found in so many
of the existing Canide, both alopecoids and thooids. The ascending ramus, which forms
an obtuse angle with the horizontal, has a proportionately smaller antero-posterior width
than in Daphenus, though a greater one than in the modern genera; the coronoid
process, in particular, is much narrower than in the former, and the sigmoid notch is
wider than in the living forms. The masseteric fossa is very deeply impressed, but it
has no such definitely marked upper boundary and it does not extend forward so far
beneath the molars as in Canis, features of resemblance to the alopecoids. The angle
is formed by a short, slender and blunt, hook-like process. The condyle, which is not
in any way peculiar, is elevated much more above the level of the molar teeth than in
Daphenus.

The cranial foramina are yery minute and hence are often difficult to detect, save
in exceptionally well-preserved specimens, a very slight degree of crushing being often

sufficient to obliterate them. In general, they may be described as characteristically

NOTES ON THE CANIDE® OF THE WHITE RIVER OLIGOCENE. Bile:

eynoid. The condylar foramen is an opening, hardly larger than a pin-hole, which per-
forates the ridge running mesially from the paroccipital process ; its position is just as in
Canis. ‘The foramen lacerum posterius is rather smaller than in existing representatives
of the family, which is due to the greater proportionate elongation of the auditory bulla,
and for the same reason the stylomastoid foramen is less conspicuously displayed. An
important difference from Canis and Vulpes consists in the presence of a well-defined
external opening of the carotid canal, which grooyes the inner side of the auditory bulla
somewhat behind the middle of its course ; it is much better shown in some specimens
than in others. In the modern Canidae, “the carotid canal is complete and of tolerable
dimensions ; but its external opening is not visible on the surface of the bulla, being
deep in the foramen lacerum posticum” (Flower, ’69, p. 24). The other carnivorous
fantilies, however, have the carotid canal with visible opening, but varying in position in
the different groups.

The foramen lacerum medium and the Eustachian foramen are very much as in
Canis, but the glenoid foramen is somewhat concealed by the prolonged anterior lip of
the auditory meatus. The foramen ovale is a narrow slit which may be readily over-
looked, and is closed by even a slight distortion of the skull. An alisphenoid canal is
present, and the other openings, the optic, anterior lacerated and round foramina, are as
in the recent cynoids. The whole structure of the cranial basis and its foramina are thus
canine in character, with only a single difference, the distinctness of the carotid canal.
There is nothing to suggest relationship with the viverrines.

Measurements.

| No, 10493. No. 10513, | No. 10939. | No. 11012. | No. 11381. } No. 11382. No. 11432.
Skull, length (fr. oce. condyles).-.--..-..-..seseesseeceecereeres 0.092 | 0.092 | 0.086 | 0.089
Cranium, length (occ. condyles to preorbital border)....- -062 | ?.062 064 -064 -059 -063
Pace, preorbital lenoth........ ......02....2..-ce.-sccesccons nee: .032 030 | 030 028
Occiput, breadth across mastoid processes ..-...----+-++-++++- | 033 | .034 .034 .038 0385 .032 .033
Brain case, greatest breadth...-.....-:--csseceeeceeeeeeceereeeeees | .081 | .032 | .032 | .035 | .033 | .033 033
Skull, width across ZYGOMAS «....-..0.-ceeseeeeceecseeceeeneeeeeee .052 | | 055
Zygomatic arch, length...-...-....-...2.-0sce-secoeeeeneennsennee- | .042 | .043 -043 -043 | 042 | .014
TDE@S, SUNT GP 0 2 cocecshee ce cacseeeeece ecco secouseenecboooecesenece | .026 .026 .026 .030 | .025
ne He 8S GNTTINE = co scesteggnce co sca snonacosaeeBataoeadecoseo3e 016 OL | 018 | .015
Mandible, length (fr. condyle) ne 063 | 060
EME des itive ric ate cba ct zea coe cones eases O09 ye eOten Pe Olie | OTT = | 010
se ASCE) S5-aancsconocadoosaeeaoscscosecebacctonsnocces | 010 | 008 007
x thickness at mM y--.---seeeee-eeceeceeeeeeeeree esse ness -0045 | -0055 .0055 | .005 | .005
SS height of coronoid process (from ventral |
ond en cee lee ee caaks Hhtne.. 027 | .029 | 2.027 | 029

height of condyle fr. angle........-..-sseesseeeeee 5 .014 | | -013

O74 NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE.

TWO “Weas) eestor (Ie 2s, Itinee, 114),

The brain of Cynodictis has already been described by Bruce (’83, p. 41), but as I
wish to consider it from a different standpoint, some account of it will be necessary. In
this genus the brain is relatively smaller than in any of the recent Canidw. The olfac-
tory lobes are large and are left exposed by the hemispheres, with which they are con-
nected by short and thick olfactory tracts. The cerebral hemispheres are pear-shaped,
broad behind, but tapering rapidly forward, where they decrease in vertical as much as
in transverse diameter. The frontal lobe is short, narrow and of small vertical depth,
while the parietal lobe much surpasses it in every dimension ; a transyerse depression
marks the boundary between the two. The temporo-sphenoidal lobe is also quite well
deyeloped and adds materially to the dorso-ventral diameter of the brain in this region.
Posteriorly the hemispheres slightly overlap the lateral lobes of the cerebellum (which
appears not to be the case in Daphwenus), but leave the yermis entirely uncoyered. The
shape of the cerebrum is thus alopecoid rather than thooid in character. In the former
series the hemispheres are wide behind and taper anteriorly, with slight incurvations at
the sylvian and presylvian fissures, while in the thooids the cerebrum is narrower behind
and at the presylvian fissure the sides are abruptly incurved almost at a right angle ;
the frontal lobes are much larger relatively than in the foxes (see Huxley, ’80, pp. 245—
247). The hemispheres of Cynodictis agree well in shape with those of the alopecoids,
and when compared with the brain of the later and more adyanced genus Cynodesmus
from the John Day, the greater width of their posterior region is distinctly to be seen.
The whole character of the skull makes it evident that Cynodesmus is a thooid, while
both brain and skull structure approximate Cynodictis more to the alopecoids.

The hemispheres are very simply conyoluted and the sulci are few, simple and short,
though it should not be forgotten that the brain-cast very probably fails to reproduce all
of the fissures. In the recent Canide the conyolutions are numerous and complex, and
the sulci pursue a remarkably curved course, giving to the conyolutions, when seen from
the side, the appearance of a succession of U-shaped, concentric coils, grouped around
the sylvian fissure as a centre. In Cynodictis, on the other hand, the visible sulci are
few, shallow, short and nearly straight. On the dorsal surface of the hemisphere only
two fissures are to be observed, the lateral and the suprasylvian, the former of which is
short and almost straight, dying away before it reaches the hinder part of the parietal
lobe. If the coronal sulcus is present at all, it is in the same fore-and-aft line as the
lateral, and has not the outward sweep around the crucial fissure which is so characteris-
tic of Canis. No trace of the crucial fissure is preserved in the brain-cast, and if it was
present in the brain, it must haye been short, as is indicated by the straight course of the

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. 375

lateral sulcus. The suprasylvian sulcus is likewise very short and but little curved, and
is not divisible into anterior and posterior portions. The sylvian fissure itself is but
feebly marked upon the cast, but the rhinal sulcus, on the contrary, is very distinctly
shown and extends for nearly the whole length of the hemisphere. Making all due
allowance for the fact that a cast of the brain-case can but imperfectly reproduce the
features of the brain itself, yet it is clear that the cerebrum of Cynodictis was convolu-
ted in a much simpler way than in any of the existing Canidae, and that it retains char-
acteristics which among the modern dogs are embryonic and transitory.

The cerebellum is rather large and is less overlapped by the hemispheres than is the
case among the recent members of the family. The vermis is narrow, but prominent,
and is quite clearly divisible into three lobes, corresponding apparently to the lobus cen-
tralis, lobus monticuli and declivus of Canis. The vermis is less regularly curved in the
antero-posterior direction than in the modern genus, the posterior surface forming nearly
a right angle with the dorsal. The lateral lobes of the cerebellum have quite a different
appearance from those of the recent Canide. Thus, the lobus quadrangularis. is less
extended transversely and narrows less toward the external side, while the lobus lunatus
inferior is very imperfectly developed, and the lobi semilunares appear not to be repre-
sented at all, or, if present, they must be exceedingly small. This latter point is difficult
to decide definitely, because a small fragment of the skull, which cannot be removed with-
out danger to the specimen, covers the place where the semilunar would be if present.
A small additional lobe, nét represented in Canis, lies upon the dorsal surface of the
lobus quadratus and near to the vermis. Complex as it looks, the cerebellum of Cyno-

dictis is simpler than in the recent dogs.

Measurements.

e No. 10513.
Brain, length fr. cerebellum to olfactory lobe (incl.)...........sceseceeeseeeeseecnseesdecseececsensesenensees 0.045
Olfactory lobe, fore and aft diameter -005
s SSeS VELLICAl (AIAMELCTSa-cechese< sat rccee se ce sccee com sk ccceec acme cae eaten coee ane eer conecee bent anna -O11
Cerebrum, length in median line.......... ets we ate e clorsesjonts «le ne Sentecense Wetea spec Seca es Se oR ENN se cnienre see teweasaces -030
oo height at temporo-sphenoidal lObE........-cceceeeeesccccceeseceeseseeececcscceseeecesceuneeesesees .025
i width ‘‘ “ a Uo i e0 -030
Cerebellum, length in median line...-....-.-.------:-0--2-02e-s-+-eenveneesesrsceeecescccccccnscacsnanenscesseess 013
ce PUL E Ms msec oceanaet ac cee cabs Saw Nou sempea a casa ceetoe tu emat ee eaeeee cites cela oes aut oeecauckmeduiawehe .024
86 vertical height 018
.012

Medulla oblongata, width.....
IV. THe VERTEBRAL CoLuMN.
The backbone is not preserved entire in any of the specimens, but by the aid of

the more complete individuals from the John Day, the numbers of the various categories

of vertebree may be inferred.
A. P. 8—VOL. XIX. 2 V.

376 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

The atlas (Pl. XIX, Fig. 13) is somewhat more canine in character than that of
Daphenus, haying a short and broad body and moderately developed transverse pro-
cesses. The anterior cotyles are shallower and more depressed than in Canis ; the neu-
ral arch is well extended in the antero-posterior direction and is quite smooth, without
ridges or tubercles of any kind; it is very strongly convex, giving to the neural canal
an almost circular shape. The inferior arch is very slender and has but a rudimentary
hypapophysial tubercle. The posterior cotyles for the axis are somewhat more concave
than in Canis and present more obliquely toward the median line. The transverse pro-
cesses are rather small and are much less extended antero-posteriorly than in Canis, not
reaching so far behind the surfaces for the axis, nor so far forward upon the neural arch ;
in consequence of this, the atlanteo-diapophysial notch is less deeply incised. The pos-
terior opening of the vertebrarterial canal presents backward, as it does in Daphanus,
but has shifted a little more toward the dorsal side of the transverse process, thus show-
ing a tendency to assume the position which is characteristic of the recent Canide.

The axis is not especially canine in appearance, but rather resembles that of Viverra.
The centrum is long, narrow and very much depressed anteriorly, becoming somewhat
deeper vertically toward the hinder end, which has a transversely oval and nearly flat
face for the third vertebra; the ventral keel is relatively better developed than in
Daphenus. The articular surfaces for the atlas are low and wide, but project much less
outside of the pedicels of the neural arch than they do in Canis, and are more conyex
than in that genus. The odontoid process is slender and elongate, more so than in
Viverra, and the articular surface on its ventral side is not, as in Canis, continuous with
the lateral facets for the atlas, but is separated from them by a feebly marked ridge.
The transverse processes, which are very thin and compressed, are of no great length;
they are perforated by the vertebrarterial canal, which is relatively longer than in the
recent dogs. The pedicels of the neural arch are short from before backward, but are
quite high, and the neural canal is proportionately much larger in both dimensions than
in the existing dogs. The neural spine, at least in the White River species, resembles
that of Daphenus much less than it does that of Canis. It is long, not very high, and
in front extends far in advance of the pedicels, but posteriorly it does not project
behind the zygapophyses, as it does so conspicuously in Daphenus; as in the modern
genus, the dorsal border of the spine is continued into the hinder margins of the neural
arch. The zygapophyses are rather small and do not extend out so prominently from
the sides of the neural arch as in Canis.

The axis of the John Day species, C. geismarianus, as figured by Cope (’85, Pl.
LX Xa, Fig. 12), differs from that of C, gregarwus in haying a much higher neural spine,

which is continued posteriorly into a pointed projection, similar to but shorter than that
seen in Daphenus.

NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE. 2)

The third cervical vertebra is markedly different from that of Daphenus and quite
like the corresponding vertebra of Canis. The centrum is moderately elongate (though
shorter with reference to the axis than in most of the modern dogs), quite depressed and
slightly opisthoccelous, and has a stout, prominent ventral keel, which is better developed
than in Daphenus, or eyen than in Canis, and ends behind in a tubercle. The ante-
rior face is broad, depressed, quite conyex and yery oblique in position with reference
to the fore-and-aft axis of the centrum, while the posterior face is more nearly circular
in outline. The transyerse process is, in general character, quite like that of Canis,
but has a relatively smaller extension from before backward, and is less obviously
divided into anterior and posterior projections, the ventral margin of the process being
nearly straight. The vertebrarterial canal is proportionately much longer than in Canis,
being nearly as long as the entire centrum. The neural canal is relatively larger and
especially wider than in the modern genus, while the neural arch is long and broad and
but slightly conyex on the dorsal surface. One noteworthy difference from Canis con-
sists in the fact that the arch does not project oyer the sides, or pedicels, as an oyerhang-
ing shelf, or does so but slightly. The neural spine is represented only by an incon-
spicuous ridge.

The zygapophyses are small and extend but little in front of and beliind the neural
arch, which constitutes a very marked difference from Daphenus. In the latter, it will
be remembered, the neural arches are deeply emarginated between each transverse pair
of zygapophyses, so that when the vertebre are placed in their natural position, large
vacuities occur between the successive neural arches. In Cynodictis, as in Canis, these
interspaces are very narrow and in certain parts of the neck they are hardly at all visible.

The fourth vertebra is somewhat shorter than the third, but is otherwise yery much
like it and also like the corresponding vertebra of Canis. The transverse process is some-
what larger and heavier than on the preceding vertebra, and the greater antero-posterior
extension of its outer portion makes the vertebrarterial canal relatively longer than in
Canis ; the inferior lamella is very thin and light. The neural spine is short and slen-
der, but is relatively better developed than in most of the modern representatives of the
family.

On the fifth cervical the neural spine is higher but more slender than on the fourth.

The sixth is not preserved in connection with any of the specimens.

The seventh cervical is almost a miniature copy of the same vertebra in Canis ; the
neural spine is relatively higher, more slender and more pointed than in most species of
the existing genus, and the transyerse processes are proportionately longer and thinner,
but otherwise the resemblance is very close and detailed.

The number of thoracic vertebre cannot, as yet, be definitely stated, because in

378 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

none of the specimens is the series preserved entire. Probably, however, these vertebree
numbered thirteen, as is commonly the case among the recent representatives of the
family. The specimen of C. geismarianus: figured by Cope (’85, Pl. LX Xa) has the
posterior ten thoracics in place, and there must have been at least three additional ones.
The anterior vertebre of this region have very small, contracted centra, but long and
prominent transverse processes and neural spines which are relatively higher and more
slender than in Canis, and are also inclined more strongly backward than in the latter.
Posteriorly the centra become longer, broader and more depressed, and are quite distinctly
keeled in the median ventral line. In addition to this median keel are two shorter and less
prominent lateral ridges, which, however, terminate behind in distinct tubercles and thus
give avery characteristic appearance to these vertebrae. The transverse processes become
more and more shortened and the neural spines lower, less strongly inclined, but more
compressed and broadened at the base (antero-posteriorly). The antepenultimate thoracic
(presumably the eleventh) is the anticlinal vertebra, of which the neural spine is low,
broad, compressed and erect. The penultimate (? twelfth) and last (? thirteenth) thora-
cies are very much like lumbars in appearance and structure, but have no transverse
processes, while in Canis these processes, though small, are quite distinct on the twelfth
and thirteenth thoracies. Large, heavy and prominent anapophyses and metapophyses
are present on the last two thoracics.

Of lumbar vertebre this genus probably possessed seven, that many being preserved
in position and in connection both with the thoracies and with the sacrum in Cope’s speci-
men of C. geismarianus. In the White River material at my command not more than
five lumbars have been found in association with any one individual, but the series is
obviously incomplete, and there is no reason to suppose that C. gregarius differed in this
respect from the John Day species. The lumbar region is proportionately long and stout
and the individual vertebrae are quite massively constructed (7. e. for so small an animal),
indicating a powerful musculature in this region. The centra increase in length up to
that of the penultimate vertebra, while the first and the last are the shortest of the
series. These centra are broad and depressed, and bear distinct median ventral keels,
while the lateral ridges and tubercles are present on the first two vertebrae, but not on
the last three. The faces are kidney-shaped, slightly convex in front and concave
behind, and are placed obliquely with reference to the long axis of the centra. This
obliquity is to provide for the curvature of the loins, which rise to the pelvis, the rump
standing considerably higher than the shoulders. The transverse processes, which are
quite short on the anterior lumbars, increase steadily in length up to the sixth, where
they become yery long; they are slender, depressed, pointed and curyed forward. The
neural spines are low, compressed and thin, broad at the base, narrow and pointed at

NOTES ON THE CANID#H OF THE WHITE RIVER OLIGOCENE. 379

the tip, and are inclined forward rather more decidedly than in Canis. Anapophiyses
are quite prominent on the anterior lumbars, but diminish posteriorly, becoming rudi-
mentary on the fifth, while the metapophyses are conspicuous in all. The zygapo-
physes are but moderately concave and convex respectively. The general aspect of
the lumbar region is not canine in character, but rather resembles that of the civets
and mustelines.

The sacrum is quite short and consists of three vertebrae, only the first of which has
a contact with the ium. The first sacral has a broad and much depressed centrum and
large, expanded pleurapophyses, which give considerable width to the vertebra. The
neural spine is a mere feebly marked ridge, while the spines of the second and third are
higher and separate. The transverse processes of all the sacrals are fused into a continu-
ous lateral ridge, but that of the third vertebra extends outward much farther than the
others and ends in a point, an arrangement which gives to this sacrum an appearance
quite different from that of Canis. The prezygapophyses of the first vertebra are large
and conspicuous, but all the other zygapophyses of the sacrum are small. The neural
foramina are remarkably small. The centrum of the last vertebra is almost as large as
that of the first and the widely extended transverse processes make the sacrum nearly as
broad behind as it is in front.

The caudal vertebre are not preserved entire in any of the specimens, nor, indeed,
ean all of them be recovered from all the individuals combined, so that the number of
tail vertebree is, as yet, conjectural. However, enough remains to show the character of
the tail and of the various elements which compose it. The tail was evidently very well
developed, being relatively longer and stouter than in any of the recent Canida, and
much like that of some of the long-tailed yiverrines, such as Herpestes. ‘The anterior
caudal vertebrae have short, but heavy centra and very long, broad and depressed trans-
verse processes, which extend out nearly at right angles with the line of the centrum.
The breadth of the first caudal across the transverse processes about equals that of the last
sacral. The zygapophyses of the anterior caudals are large and prominent. ‘The ante-
rior caudals are succeeded by a number of vertebrae with very elongate centra, which
resemble in miniature the corresponding vertebree of Daphenus, having distinct remnants
of the various processes. Toward the tip of the tail the vertebrae become very slender
and of a cylindrical shape, the centra being slightly contracted in the middle and
expanded at the ends.

The ribs, so far as they are preserved in the yarious specimens, are remarkable
chiefly for their length and slenderness and for their subcylindrical shape. Tubercles
appear to be absent from the twelfth and thirteenth pair. The sternum is of the usual
carnivorous character, without being especially like that either of the dogs or of the

380 NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE.

civets. The manubrium is long, more so than in Canis, as well as narrower and more
compressed. The first pair of ribs is attached to a pair of wing-like processes, which
are unusually far from the second pair. In front of these processes the bone is com-
pressed and yery narrow. For much of its length the manubrium possesses a ventral
keel. The segments of the mesosternum, so far as they are preserved in the various
specimens, are more elongate, more slender and depressed and more contracted in the
middle than in the recent Canide.
Measurements.

|
No. 10493. No. 11012. No. 11381. | No. 11382. | No, 11432.
|

| zh |

WATE acer eEH tesa ccc crete ae Be Pence 0.016 | |

2 TURE GEA De cosccsnanse no ssooseoctnans ante t ene ssanstenecesnnc suosonscetonses -034
Axis, length (excl. of odontoid) ..........2.-......0secseseeeeeeeeeneeeees 019 | .020

“ ** of odontoid process...- 005 |

my £0 — “Width pOst. £ACE <..ceccce--cas--nsenerecnssenscucdscosesanss- -010 -011 -009 |
Sixth <€ Vength........------seeeneeceeneeceeeecee ees tensseeeescnarsccens® -015 201 St -014
Seventh ‘ re 013 | -013 -012
Sacrum, length | -024 -| -026
First sacral, width across pleurap.-.-----+-ss-sssseeeeseeeeeeeeseeeseees |}. .024 | .024
Third ‘ s CO TREE TIP -cocanacgconceaentaneconseacesccccon | 021 |
First caudal, length ......-.......2--..-.s----0-se-enc--neneennersrreerecnnnn= 007 | .008 ?.010

ce St Width ACrOSS ELANSV. PI----------2---202-seeeeecoerereeene= 021 | — .026 |
Médian ‘caudals lengthoae ot 6 het catia ot iS eee | | |

2 ct TEGRHD DnB, FRYOD coonceccoscencaccasasoancneceonescosecnces -005

V. Tue Fore Lime.

The scapula is quite remarkable and is in character rather yiyerrine or raccoon-like
than canine. The shoulder blade is rather low and broad and is divided by the spine
into pre- and postscapular fossee of nearly equal breadth, while in the modern dogs the
scapula is high, narrow and of subquadrate shape, and has the spine so placed as to
make the postscapular fossa much the larger of the two. The glenoid cavity is moder-
ately concave, and is elongate antero-posteriorly, but narrow transversely. The coracoid

NOTES ON THE CANID#Z OF THE WHITE RIVER OLIGOCENE. 381

process is unusually large, forming an incurved hook, which, however, does not appear
prominently when the scapula is viewed from the external side ; in the recent Canida the
coracoid is reduced to much smaller proportions. A resemblance to the shoulder-blade
of Canis is to be found in the broad neck of the scapula and in the absence of any well-
defined coraco-scapular notch. The coracoid border is slightly concave at the neck, but
then curves forward and upward, giving great width to the prescapular fossa; the gle-
noid border is, as usual, straight and is steeply inclined, so that the postscapular fossa,
which is very narrow distally, becomes very broad proximally. The spine is high and
ends in a very long and prominent acromion, which descends below the level of the gle-
noid cavity, which suggests that in this genus the clavicles were much better developed
than in the existing dogs. A very large metacromial process is also present. The meta-
cromion may be observed in most of the existing families of Carnivora, but it is seldom
so large and so prominent as in Cynodictis ; perhaps, the nearest approach to it among
modern genera is in Arctictis.

The humerus is much more suggestive of viverrine than of canine affinities. As
compared with the bones of the forearm, or even with the femur, the humerus is elon-
gate, but it is short in proportion to the length of the back or loins. The head is
strongly convex and projects farther behind the plane of the shaft than in the modern
dogs ; the external tuberosity is a heavy, but low ridge, which barely conceals the head
when the bone is viewed from the front; a large, irregularly circular area near the
hinder end of this ridge plainly indicates the insertion of the infraspinatus muscle.
The external tuberosity is both lower and shorter than in the modern dogs, but the inter-
nal one is rather more preminent, and the bicipital groove is more widely open, more
internal in position and more of it is visible from the anterior side. The shaft is rather
long, and, when seen from the side, exhibits a sigmoid curvature, which is somewhat
better marked than in Canis. For most of its length, the shaft is laterally compressed
and has but a very short cylindrical portion before expanding laterally at the distal end.
Most of the ridges and prominences for muscular attachment are well developed, more
so than would be expected in so small an animal. The deltoid ridge is much more
prominent than in the recent dogs, and is more like that of the cats and viverrines ; the
supinator ridge is likewise very much more prominent than in Canis, in correlation
with the power of rotation of the radius, which Cynodictis appears to have retained in
almost undiminished degree. On the other hand, the rough ridge, which runs down
from the head upon the outer side of the shaft (spina humeri) and serves for the attach-
ment of the teres minor, anconzus externus and brachialis internus muscles, is much
fainter than in Canis and the linea tuberculi minoris is very feebly marked. The supra-
trochlear fossa is very shallow and the anconeal fossa is much smaller and shallower
than in the modern representatives of the family, there being no perforation of the shaft

AR.
382 NOTES ON THE CANID#Z OF THE WHITE RIVER OLIGOCENE.

at this point. The internal epicondyle is much more prominent and more massive than
in Canis, and a conspicuous epicondylar foramen is present, in the form of a long, nar-
row slit. The external epicondyle, on the contrary, is rather smaller than in the recent
genus.

The humeral ¢rochlea has a much smaller proximo-distal diameter than in the exist-
ing Canide, in which respect it preserves a primitive character and resembles the troch-
lea of such viverrine genera as Cynogale and Viverra. ‘The radial surface is small and
simply convex, while the ulnar facet is much larger than in the recent dogs; the inner
flange of the ulnar facet is also more produced distally and forms a sharper edge than in
the latter.

The radius is not at all suggestive of canine affinities, but rather resembles the cor-
responding bone of the cats and viverrines. The capitellum is small and of subdiscoi-
dal shape ; while it is somewhat more extended transversely than in Felis, it is much less
so than in Canis ; its articular surface is moderately concave and is slightly notched on
the anterior border. The proximal facet for the ulna is a simple, convex band, separated
from the humeral surface by a distinct angle and entirely resembling that of Daphenus.
The character of the articulation at the elbow-joint and the large development of the
supinator ridge on the humerus would seem to imply that in Cynodictis a considerable
degree of freedom in the rotation of the manus had been preserved, though probably less
than in the cats and in many viverrines. The bicipital tubercle is prominent, but occu-
pies a more posterior position than in either the cats or the recent dogs, and is not visible
when the radius is looked at from the front.

The shaft of the radius is relatively short, slender and rounded, very different from
the broad, oval and antero-posteriorly compressed shaft seen in Canis; it has a slight
double curvature, arching anteriorly and externally, and is of almost uniform thickness
throughout its length, except at the distal end, where it broadens considerably. A very
striking difference from Canis consists in the very great size and prominence of the sty-
loid process, which forms a relatively enormous tuberosity ; it is even much larger pro-
portionately than in the cats or civets and is as large as in Me/hvora, though of a differ-
ent shape. In Daphenus, as we have already learned, the styloid process is very promi-
nent and of a generally feline appearance, but it is proportionately smaller than in Cyno-
dictis. The radius figured by Schlosser (’89, Taf. VII, Fig. 8) and by him attributed to
one of the European species of the latter genus has a styloid process in the form of an
enormous, recurved hook, much longer and much more slender than in the American
species and of an entirely different appearance. The distal tendinal sulci are not very
well marked, though that for the abductor and extensor muscles of the pollex is a deep
eroove. The distal facet for the ulna is smaller and less deeply impressed than in Canis.
The carpal facet is small and slightly concave, narrowing toward the internal side; it

NOTES ON THE CANIDH® OF THE WHITE RIVER OLIGOCENE. 383

does not extend over upon the styloid process, from which it is separated by a broad and
deep notch.

The w/na is, in its way, as peculiar as the radius. The olecranon is quite typically
fissipede in character and differs from that of the creodonts in its comparative shortness
and breadth ; though proportionately somewhat longer than in Canis, it is hardly so long
as in Daphenus, and the sulcus for the tendons of the anconeal muscles is more distinct
than in the former. The sigmoid notch is hardly so deep as in Canis, and, in particular,
the internal facet for the humerus projects less in front of the plane of the shaft, and the
external process is very feebly developed. The radial facet is narrower and less deeply
concaye than in the modern Canide, but has a somewhat greater vertical diameter.

The shaft of the ulna is decidedly less reduced than in the recent representatives of
the family, and for most of its length is little or not at all more slender than that of the
radius. In its proximal portion the shaft is much more compressed laterally and thicker
antero-posteriorly than in Canis, in which genus this portion of the shaft is trihedral.
The middle and distal portions are of triangular section, none of it haying the subcy-
lindrical shape which characterizes the distal one-third of the shaft in the recent genus.
The distal end has quite a different shape from that seen in Daphenus, a difference which
is due to the much greater prominence of the radial facet in the latter. In Cynodictis
this facet is almost sessile and projects but little more than it does in Canis. The car-
pal facet is very small and quite simply convex.

Measurements.

| No. 10493. | No. 11012. | No. 11381. | No. 11382, | No. 11432.
| | | |
Scapula, length seeceeee tone | 0.054
sf greatest Width...-...---ce..-esece-nneenoeeeeneenceeecennereren ners 2.049 |
ag TRAHAN.) (a0 <senbpe cnneroshdspecetato baduoconcasccesAceacecsencras | 013
ue ant. post.-diameter of glenoid cavity ....-.-++.+-.2---..+-0 .012 | .0095 .0095
Ge transverse sf Be ue | -008 007 | .007
TETMITIVERES METH... cocesascononnccduncoecbadgnonocooasnooossHosodcossoCersceKs 075) | .070 070
fed. antdpost.. dani. proxsend! ais. /isatiived Saaeeees 012 | 015 1OLSINn |e Ols 015
i transy. SS fe OO aso cule secs ennneaeseosernins anne | 014 | .016 | 013 0125
& preadihnko te dustallendeesseeseeceeceseetesesesseseeeceseeeseeeee | 016 .020 | 015
£ epee Cetrochll eaiescerecrccncnacetes sane cesasse-teeee ener 012 0145 | O11
TREGUTIS, TWerepal 1 205-000 nanan oceongoosa5no ada q5HGeDonecacHbcaoemacesNecoHESHEaS | .057 061 | |
«« ant post. diam. prox. end 005 .006 007. | .005 005
“<— transy. ce ee 1007 |) 007 | 2009 007 007
Pm are cidital endemic ek eee [Perot sors lose | 009
“ eee eA call MACE biases eeeeessa ese eae sae ears 0055 | .606 007 0055
Ulna, length | ee ore |
fs CORE Ofer OleECrAn OMe ectomtes cs ecees ton aacceMocere cocci soc-leace aes | | .007 -010 .0095 -009
Co {Tp RMERS Wt CECE AGI cosonaccocemencenssaconaoasesagsocoureAsnooc | .010 .008 .008

A, 2 §& SVG, MIO Zr

384 NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE.

VI. Tse Manus (Pl. XX, Fig. 23).

By a fortunate discovery of Mr. Hatcher’s, 1am enabled to give an account of an
almost complete carpus belonging to Cynodictis, which has hitherto been entirely
unknown.

A scapho-lunar is present, formed by the coalescence of the scaphoid, lunar and
central, which distinguishes Cynodictis from the creodonts. This bone resembles that of
Canis in general character, but displays quite a number of differences in points of detail,
and these differences are, at the same time, approximations to the structure found in
Daphenus. The seapho-lunar has a very small vertical (proximo-distal) diameter,
especially on the radial side, where it thins away to a mere edge, the facets for the radius
and the trapezium almost meeting. As compared with the corresponding carpal of Canis,
this bone has a somewhat greater transverse and smaller dorgo-palmar diameter. The
radial facet is simply convex both transversely and antero-posteriorly, and has not the
saddle-shaped extension at the interno-palmar angle which is found in the recent dogs.
This facet descends quite low upon the dorsal side of the bone, as is also the case in
the modern plantigrade and semiplantigrade carnivores. The hook-lke process which
arises from the postero-internal angle of the scapho-lunar is much shorter and less mas-
sive in every dimension than that of Canis. Another difference from the modern genus
consists in the absence of any distinct articular surface for the pyramidal, the facet for
the radius and that for the unciform almost coming into contact along the ulnar side of
the bone.

On the distal side of the scapho-lunar are four facets, for all the carpal elements of
the distal row. That for the unciform is relatively smaller than in Canis, and is con-
fined to a narrow strip near the ulnar border; the magnum facet is much the same as in
the modern genus, but is somewhat more oblique in position. The surface for the tra-
pezoid is fairly large and keeps more nearly parallel with that for the magnum than in
the recent dogs, while the trapezium facet is small and of almost circular shape.

The pyramidal is a very different-looking bone from that of the modern dogs,
being broad, depressed and scale-like in shape; its vertical (or proximo-distal) diameter is
very small and relatively much less than in Canis, and there is no such process from
the ulnar side of the bone as in the latter, in which the pyramidal articulates with the
head of the fifth metacarpal by a much more extensive facet than in Cynodictis.
The recent viverrines have the pyramidal shaped very much as in the White River
genus. The proximal surface is divided into two narrow and somewhat concave facets
for the ulna and pisiform respectively, of which the latter is slightly the larger. On

the distal side is a single large and concave facet for the unciform, and posterior to this

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. 585

a very narrow surface which appears to be destined for articulation with the head of
the fifth metacarpal.

The pisiform differs very decidedly in shape from that of Canis. This carpal is
small and light; its proximal (7. ¢., articular) end is greatly depressed, but much extended
transversely (in the existing genus the principal diameter of the proximal end is the
vertical one) and the facets for the pyramidal and ulna are correspondingly broad-
ened transversely and narrowed vertically. The pyramidal facet is the larger of the
two and is quite deeply concave, while that for the ulna is small and nearly plane; the
two facets together form an acute angle and are separated only by an inconspicuous ridge.
The distal end of the pisiform is moderately expanded, but in the yertical dimension, so
that the proximal and distal expansions are almost at right angles with each other.
Between the two expansions the body of the bone is much contracted and very slender,
which is in marked contrast to the shape seen in Canis.

A so-called “radial sesamoid” appears to have been present; at least, there occurs
in the same block of matrix through which the carpals of one individual were scattered,
a small, irregularly wedge-shaped bone, to which I can give no other interpretation.
Assuming that this reference is correct, we find in the relative size and shape of this bone
another resemblance to such yiyerrine genera as Herpestes, Cynogale and Paradoxurus,
ete. The radial sesamoid also occurs in Canis, at least in certain species, but is very
minute.

The trapezium is very small and differently shaped from that of Canis; its princi-
pal dimension is the dorso-palmar, while the transverse diameter is the least. The sur-
face for the scaphoid, which in Canis is a very oblique, convex facet, is in Cynodictis
entirely proximal in position and nearly plane, and there is no such large concave facet
for the trapezoid on the ulnar side as in the modern genus ; the distal facet for the head
of the first metacarpal is less distinctively saddle-shaped than in the latter. In view of
the well-developed pollex, the small size of the trapezium is somewhat surprising.

The trapezoid is shaped very much as in the existing dogs, but with certain minor
differences, especially noticeable in the very small vertical diameter and in the thinning
of the bone to an edge on the ulnar side. The proximal end bears a simply convex facet
for the seapho-lunar, while the distal facet, for the second metacarpal, is very slightly
saddle-shaped ; on the palmar side the trapezoid contracts to a point.

The magnum is small and that portion of it which is visible from the dorsal side,
when all the carpal elements are in their natural positions, is minute, especially in its
proximo-distal dimension. In shape the magnum does not differ materially from that of
the recent dogs, but the proximal surface is narrower and rises more abruptly to the

“head,” and on the palmar side the bone broadens out in a fashion not repeated in Canis.

386 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

The unciform facet is large and plane and does not rise so high upon the head as in the
modern genus. On the radial side we find no distinct facet for the trapezoid, which, as
already mentioned, thins to a mere edge toward the magnum, but there is a well-defined
facet for the projection fromthe head of the second metacarpal, which is proportionately
larger than in Canis. On the distal end of the magnum is a narrow facet for the third
metacarpal, a facet which is less concave in the dorso-palmar direction than in the case
of the last-named genus.

The wneiform is viverrine rather than canine in character, being much narrower
in proportion to its vertical height than in the recent dogs. The facet for the seapho-
lunar, which in Canis has an almost entirely proximal position, is in Cynodictis much
more nearly lateral. The pyramidal facet is also decidedly more steeply inclined than
in the existing genus, the two articular surfaces meeting at a very acute angle and mak-
ing the proximal end of the unciform narrow and wedge-shaped. On _the radial side is
a large facet for the magnum and a small one, confluent with it, for the extension from
the head of the third metacarpal. The distal facets, for the fourth and fifth metacarpals
respectively, are narrower than in Canis, contracting especially toward the palmar side.

The metacarpals, five in number, are remarkably short, slender and weak and.haye
but little resemblance to those of the recent dogs.

The first metacarpal is very small, but is, nevertheless, proportionately much less
reduced than in Canis, taking the length of me iii in each genus as a standard of
comparison. The head is thicker and relatively heavier than in Canis and on the radial
side, internal to the trapezium facet, is a tubercle for the attachment of the lateral liga-
ment. The facet itself is much less deeply concave transversely than in Canis, but
more convex in the dorso-palmar direction. The shaft is short, slender, arched toward
the dorsal side, antero-posteriorly compressed and of oval section, tapering considerably
toward the distal end. The distal trochlea is very small, but formed entirely like those
of the other metacarpals ; it is strongly convex, almost hemispherical and bears a dis-
tinct carina upon the palmar face, just as in Daphenus. In Canis, on the other hand,
this structure is of an entirely different character, forming an asymmetrical hemicy-
linder, with a broad shallow groove placed somewhat internal to the median line, and
thus resembles the trochlea of a phalanx rather than that of the other metacarpals.

The second metacarpal is represented in the collection only by a single imperfect
Specimen, consisting of the proximal end. This shows a much stouter shaft than me i,
being of about the same diameter as the corresponding portion of me iy, and more slen-
der than that of me iii. The head is narrow and bears a saddle-shaped facet for the
trapezoid, but sends out a projection which rises more above the head of me iii than in

Canis and articulates with the magnum by a larger facet than in that genus.

NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE. 387

The third metacarpal, though short and slender,-is somewhat the longest and heay-
iest of the series. The proximal articular surface for the magnum is shaped very much
as in Canis, but is slightly broader in proportion and rather more concave transversely ;
on the radial side of the head is a large facet for me ii, which has a more oblique
position than in the modern genus. On the ulnar side is a small projection which
abuts against the unciform and is relatively larger than in Canis. The shaft, and
indeed the whole metacarpal, has a viverrine rather than a canine appearance ; it has not
acquired the prismatic, quadrate shape which is so characteristic of the modern dogs,
but is of oval section and is of almost uniform width throughout, but broadens slightly
at the distal end. The distal trochlea, though much lower in the vertical diameter, is
yet of decidedly more canine character than is that of Daphenus, being broad and hemi-
cylindrical in shape instead of subspherical. The pit above the trochlea, which is absent
in Daphenus, is distinctly marked and the lateral processes for ligamentous attachment
are much less prominent. All of these conditions are approximations to the conditions
seen in Canis.

The fourth metacarpal is not completely preserved in any of the specimens, but. it
appears to have been of about the same length as me iii and to have formed with it a
symmetrical pair, although the two metacarpals are not so closely appressed as in Canis,
but diverge slightly toward the distal end. The head has a simply convex facet for the
unciform and is somewhat narrower proportionately than in the existing members of the
Canidae, owing to the overlapping of the head by me ii, in order to reach the unciform.
So far as it is preserved, the shaft is rather more slender than that of me i and of a
more cylindrical, less compressed shape.

The fifth metacarpal is remarkably short, much more so in proportion to the length
of me ii than is that of Canis. The head is less broadened and thickened than in the
latter genus, and carries a simple, convex facet for the unciform. In the modern genus
there is likewise a large facet for the pyramidal, which extends down over the unciform
and comes into contact with me y. In Cynodictis there appears to be a facet of a simi-
lar kind, but if so, it is very small and obscurely marked and may be regarded as in only
an incipient stage of development. The shaft is slender proximally and broadens dis-
tally, the reverse of the proportions which obtain in Canis, and the distal trochlea is
small and is of somewhat more spherical, less cylindrical, shape than in the existing
members of the family.

The phalanges. It is unfortunate that in all of the specimens in the collection the
phalanges are in such a fragmentary state that only an incomplete account of them can
be given, and some important questions must be left unanswered for the present. The

proximal phalanx of one of the median digits is short, slender and straight, and is rela-

388 NOTES ON THE CANID& OF THE WHITE RIVER OLIGOCENE.

tively broader but more depressed than in Canis. As in Daphenus, the proximal articu-
lar surface is somewhat more deeply concave and presents more obliquely toward the dor-
sal side than in the recent genus. The distal trochlea likewise resembles that of Daphe-
nus in having a deeper median groove and in being more confined to the palmar aspect
of the bone than in Canis, which has the distal trochlea reflected well over upon the dor-
sal side of the phalanx.

Of the second phalanx only the proximal half is preserved in any of the specimens,
and I have so far failed to find eyen a fragment of the distal end. So far as can be
judged from the material at hand, Cynodictis would appear to have differed from Daphe-
nus in the very important respect that the claws were not at all or only very imperfectly
retractile. In Daphenus the asymmetry of the second phalanx is clearly displayed even
in its proximal portion, while in Cynodictis the proximal end is quite symmetrical and
does not possess any depression or excavation upon the ulnar side. However, a certain
resemblance to Daphenus and difference from Canis may be observed in the greater con-
cavity and more marked separation of the two pits into which the proximal facet is
divided, as well as in the greater prominence of the beak-like process which rises from
the dorsal margin and fits into the median distal groove of the first phalanx. In the
absence of the distal end of the second phalanx, it cannot be positively stated that
Cynodictis had lost (or had never possessed) all trace of the retractility of the claws, but
it does not seem unlikely that such was the case.

Measurements.

|
| No. 10493. | No. 11012,

Carpus, height in median lime ......--..eeceeeeeseeeeeteneeecce cece eecec ee eereeneecesaaeeeseeceseasccesaeeceusesecen easene | 0.006
GO” TODEYEtHN cocdoccdocsta0annancdanaosconoencmasos uahsea qs0Gbse05 shHeGHouno0 Sano GDADaEEbOsOUencdoDDOoNrO capcadavEG9NbaGoe | | O11
Wigan 1, WEWGTYI cocccocospnseconsononaadaoonmncaooonaneanbadoceossosseCescocaysded09D05000030 Bee ea aconene Co eatrosct ad | | .012
oe “ width of proximal end 0035 | 004
Bap, GL Ae. CPMar Sia steinendy. Cees eon hte 2 hide... s OBE SCN ai Ahan, cd ee mene nmne ne | 2003

Metucarpal ii, width of proximal end..........-. . cesseececeeeceseeeceeececeeeerceceeeceneesceeeceseesccceeeseeecesee: | .0035

Metacarpall ii, Vength--.-.-------<nc-cesarereeee-eemseecccneecce=r set ea-raceee ene 2 Wiad ay EOE Re cea dees Be me 50220 .0215

os ‘* width of proximal end... -004 | .0035

fe G hs CEG bei Rall TS 116 scaRebnoddodo secre oonoccscrascaarcropbacactiosaradconadsvoasacadéc: sndonnqodbod nosdo soc | .005 | 0045

Metacarpal iv, width of proximal end...........:-..cssc..ceeseecceecccsncececactersseerecseceneertceneesrcaeoserevecss | .004 | -0035
Mietacanpallayamlen piney rseses eres =tee rer ee -1sliesty aeeeileel ee eae eller eet leech teeters | -017 | 016
S G6 Sychiln OF joRORMe)! GiMGl coosooconbocovoDooDoboDbongDeDoDosEHONDSOO HOSEN HeaCaDooO DAErieodaoNcaceAsasocORAAOAE 004 | 004

eT WARE atcMUistall cenidee es Coke ae eck eens AEE ASE UTS by tee nat | 004 | 0045

The ungual phalanx differs in several not unimportant details both from that of

Daphenus and that of Canis, and is, on the whole, intermediate in character between

NOTES ON THE CANID#® OF THE WHITE RIVER OLIGOCENE. 389

the phalanges of the two genera. As compared with the ungual of Daphenus, it has a
somewhat less concave proximal trochlea, a smaller subungual process, and a much less
extensive bony hood reflected over the base of the claw. Indeed, this hood is rudi-
mentary and can hardly be said to exist at all. The phalanx is also slightly thicker and
has more conyex faces. Comparing this ungual with that of Canis, we find it to be
decidedly sharper, narrower and more compressed and to haye a more deeply concave

trochlea. In the modern genus the bony hood is almost as well developed as in Daphenus.

? VII. Tur Hinp Live.

The pelvis approximates more nearly to the modern canine type than does that of
Daphenus, though still retaining a number of primitive characters. A conspicuous
difference from the recent members of the family consists in the elongation of the post-
acetabular portion of the pelvis, which in Canis is short, and in the consequent change
of shape of the obturator foramina. The ilium is fairly elongate and in shape is rather
more yiverrine than canine ; the peduncle is short and laterally compressed, but of con-
siderable dorso-ventral breadth. The anterior expansion of the ilium is less extensive
than in Canis, in which genus the ilium widens gradually to the free end, or erista, while
in Cynodictis it attains nearly its full width immediately in front of the peduncle, and
from this point forward the dorsal and ventral (or ischial and acetabular) borders pursue
an almost parallel course. The widening is almost confined to the ischial border, being
very feebly marked on the acetabular border, and owing to this the shape of the ilium is
much as in the modern Herpestes. The gluteal surface does not display the wide and
simple concavity which is seen in Cunis, but, as in Daphenus and Dinictis, there is a
narrow dorsal depression and beneath this a convex ridge, but this ridge is not so
prominent as in the other White River genera which haye been mentioned. The iliac
surface is short and narrow, and the sacral surface is small and placed far back, so that
the ium projects well in front of the sacrum. When viewed from above, the two ilia
are seen to curve outward less, and to diverge less anteriorly than in the modern dogs.
The acetabular border ends in a well-marked tubercle and the ilio-pectineal process is
also quite prominent. |

The ischium is relatively long and its anterior portion is slender, but posteriorly it
expands into a broad plate. This posterior portion is much less decidedly everted and
depressed and oceupies a more vertical position than in Canis, and the ischial tuberosity,
just as in Daphenus, is much more feebly developed than in the existing Canide. On
the other hand, the spine of the ischium and the ischiadic notch are much more distinetly
shown and are placed farther behind the acetabulum than in the latter, though not so far
back as in Herpestes, The obturator foramen is narrower and more elongate than in

390 NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE.

Canis, and its anterior border is notched by the obturator sulcus. The acetabulum is
small, deep and nearly circular.

The anterior or descending ramus of the pubis is long and slender and encloses with
its fellow a broad anterior pelvic opening. The horizontal ramus is proportionately
longer and stouter and the symphysis is longer than in the recent dogs, almost as long as
in the cats. The horizontal ramus is less flattened and depressed than in the former,
forming a prominent ridge along the ventral side of the symphysis.

The os penis may be conveniently described in connection with the pelvis. - In none
of the White River specimens that have fallen under my observation is this bone pre-
served, but in the beautiful specimen of C. geismarianus figured by Cope (85, PI.
LXX) it is present and in nearly its natural position, though Cope has omitted any
mention of it in his description. Flower (69) has pointed out the characteristics of this
bone in the three sections into which he divides the fissipede carnivores. The Arctoidea
“all have a large penis with a yery considerable bone, which is usually more or less
curved, somewhat compressed, not grooved, dilated posteriorly and often bifurcated or
rather bilobed in front” (p. 14). The cats and viverrines “all have a comparatively
small penis, with a more or less conical termination, and of which the bone is small,
irregular in shape, or not unfrequently altogether wanting” (p. 22). To this statement
Cryptoprocta forms an exception, haying a bone relatively long, “slender, compressed,
slightly curved, not grooved or divided anteriorly, rounded and slightly dilated at each.
end, but thickest posteriorly ” (p. 23). In the hyzenas the bone is wanting. The dogs
resemble the raccoons, weasels, ete., in having a large os penis, “though the os is of a
different form, being straight, wide, depressed and grooved” (p. 26). In Cynodictis this
bone is entirely different from that of the modern Canide ; it is long, slender, compressed
laterally and strongly curved and is slightly grooved upon the sides, but not on the dorsal
border ; the anterior end is so broken that the presence or absence of a bilobation cannot
be determined. The resemblance in the character of the os penis between Cynodictis,
on the one hand, and Cryptoprocta and the mustelines, on the other, is an important fact,
the significance of which will be discussed later.

The bones of the hind limb proper considerably exceed in length those of the fore
limb, more so than in Canis, though the difference is rather between the proportions of
the radius and tibia than between those of the humerus and femur. :

The femur is slender and quite elongate and in essentials differs but little from that
of Canis. The head is small, of hemispherical shape, and is set upon a somewhat longer
and more distinct neck than in the modern genus, projecting more directly inward and
less upward ; the pit for the round ligament is deeply impressed but very small. The
great trochanter is lower than in Canis and is separated from the head by a narrower,

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. 391

shallower notch, while the digital fossa is relatively much smaller. The second tro-
chanter oceupies nearly the same position as in the modern genus, though somewhat more
posterior, so that it is almost or entirely concealed when the femur is viewed from the
front; it is of about the same prominence as in the existing dogs, but rather more slender
and pointed. The intertrochanteric ridge, which connects the greater and the second
trochanters, is rather better developed than in Canis, especially in the larger and longer-
limbed individuals. What may fairly be regarded as a remnant of the third trochanter
is present in the form of a low, short, thickened and rugose ridge, which is placed a short
distance below the great trochanter. The third trochanter is all but universal among
the Creodonta, and in rudimentary form it persists in many of the earlier and more
primitive carnivores, such as Dinictis, but it is somewhat surprising to find it retained in
so advanced a genus as Cynodictis. It is true that in certain muscular and powerful
domestic breeds of dogs the third trochanter recurs, though it is not distinctly shown in
the existing wild species of Canidae.

The shaft of the femur is long, slender, arched strongly forward and slightly toward
the internal or medial side. As would naturally be expected in so small an animal, the
ridges for muscular attachment are not so prominent as in the modern species. On the
anterior face no ridge for the vastus externus muscle is distinguishable and on the poste-
rior face the linea aspera is neither so long nor so prominent as in Canis. The distal end
of the femur has quite a different appearance from that seen in the existing members of
the family ; a difference which is principally due to the smaller size and less prominent
projection of the condyles and rotular trochlea. The trochlea resembles that of the
viverrines in being shallow and in having the two borders of nearly equal height and
length, and also in the absence of any distinctly marked suprapatellar fossa. On the
other hand, this trochlea is relatively narrower and extends farther up the shaft than in
the civets. The condyles are small, of nearly equal size and prominence, and are sepa-
rated by an intercondylar space which is relatively narrower than in Canis ; small sesa-
moid bones were evidently, as in the existing species, attached to the proximal faces of
the condyles.

The patella is viverrine, or more accurately herpestine, rather than canine in char-
acter. It is a short, rather wide, thin and scale-like bone, of subquadrate more than
ovate shape. The articular surface for the femur, in correlation with the shallowness of
the rotular groove, is but slightly concave proximo-distally, and even less convex trans-
versely.

The tibia, as in Canis, is of about the same length as the femur. Compared with
the radius, the tibia seems to be very long, but that this is due rather to the shortness of
the radius than to the elongation of the tibia, appears from a comparison with the verte-

A, 2) S— VOL, EX. 2X,

392 NOTES ON THE CANIDEH OF THE WHITE RIVER OLIGOCENE.

bral column, whence it becomes evident that all the limb bones of Cynodictis are propor-
tionately shorter than those of Canis, and that the bones of the forearm are especially
short. The tibia of Cynodictis differs from that of the modern canines in several par-
ticulars. The proximal condyles are of nearly equal size, but the external one projects
much farther behind the plane of the shaft than in Canis, and on the distal face of the
overhanging shelf thus formed is a facet for the head of the fibula, which is much larger
and more distinct than in the recent genus. The tibial spine is bifid and very low, but
the two parts are closely approximated, the condyles being less widely separated than in
Canis. The cnemial crest, though stout and prominent, is much less so than in the mod-
ern forms, and the sulcus for the extensor longus digitorum is much less deeply incised.
In its proximal portion the shaft is stout and trihedral, but for most of its length it is
slender and subcylindrical, expanding moderately at the distal end; it has a double cur-
yature, arching forward and outward. ‘The various ridges which serve for the attach-
ment of muscles are much the same as in Canis and are, consequently, better developed
than those of the femur. The distal articular surfaces of the tibia are intermediate in
character between those of Daphenus and those of Canis. The grooves for the astraga-
lar condyles are deeper and the intercondylar ridge higher than in the former, less so
than in the latter, and the suleus which in Canis invades the articular surface has not
yet been developed. The internal malleolus is somewhat smaller than in Daphenus,
but, as in that genus, it forms a heavy, prominent ridge, which extends across the whole
dorso-plantar diameter of the bone, while in Canis the process has not half this exten-
sion. The groove for the tendon of the long flexor muscle is very distinctly marked and
has more elevated borders than in the modern dogs. The distal fibular facet is some-
what larger than that of Canis and differs from it in haying its principal diameter trans-
verse instead of longitudinal. The resemblance in the structure of the distal end of the
tibia between Cynodictis and Daphenus, on the one hand, and the primitive sabre-
tooth Dinictis, on the other, is very marked and very suggestive, though Cynodictis has
already begun to change in the direction of the modern Canidw. Among living forms
the tibia of Herpestes offers a close analogy to that of the White River genera which
have been mentioned.

The fibula is relatively much less reduced than in the existing Canidae, and both the
shaft and the terminations are larger. The proximal end of the fibula is much larger
and heavier proportionately than in Canis, and though smaller than in Dinictis, it has
a very similar shape; its principal diameter is the antero-posterior one, while trans-
versely it is narrow and compressed ; the thickening of the anterior and posterior border
is present, as in Dinictis, but much less conspicuous. The facet for the head of the
tibia is large, subcircular in shape and proximo-lateral in position. The shaft, though

NOTES ON THE CANIDAH OF THE WHITE RIVER OLIGOCENE. 393

slender and delicate, is relatively very much less so than in Canis, in which genus the
fibula has undergone a more extensive reduction than in Cynodictis. Another difference
from the recent forms is to be found in the fact that the fibula is not so closely applied to
the tibia, the two bones coming into contact only at their proximal and distal extremities.
The distal end is expanded and thickened to form a stout external malleolus, which is
somewhat smaller than in Daphenus or Dinictis, but of much the same shape, and has
on its outer side a deep suleus for the peroneus tertius tendon. The distal tibial facet is
a narrow band, with its long diameter directed antero-posteriorly ; obscurely separated
from it is the larger, subcireular facet for the astragalus.

Measurements.

No. 10493. No. 11012. No. 11381. No. 11382. No. 11482.
elivista eng thisacenessw sees senses seeeceih fae cactcvveceee cot tseseecoskenscseoees | 20.064
name Dueadthyatracetabulumsecs::-coce:.nccocsseresseckcseneceeececsceace eeeeOsG 037
Ilium, length fr. acetabulum..............2...00. sesceesceeesenenseseeeeee ?.033 037
eee OLeACthvotspedunCleracercersereceriaasenet sticneesecsessecesesen ees -011 -010 009
of See as v amlbenplatencesaa-cseccrossecccesssncseoserereaves cnemen ; -013
Ischium, length fr. acetabulum............0...sseesescsseneeeeceeeeeeeeees | 027 26
Acetabulum, fore-and-aft diameter ..........2.c2ceecccecesesees avceseee | .008 -O11
IDET, NOMEPH Ncceceseoc cencaanc6 -coossqueqcacouaoDoDNcHopcooooSenpSEDDo gHONIEGCS 095 | .085 -086
Smee breadthtoMproxmend ees testce tee ceee ence eee ee | | 017 020. | 015 .016
se «« _ ** distal end... | .016 017 -014 014
PATS, Toray hase cess oa er i ge | 089 .099
Cembreadthvofproxaendeessmerse tie eee eters cece | | ls 018 | .014 | .014
‘« thickness of prox. end | 013 016 -012 012
«« breadth of distal end........ iat 009 .O1L 012 | 009 -009
Fibula, thickness of prox. end -007 |
os ns Sex dastaliendissccesc.ssecesescosseccteeseceesstectec crs 0065 009 |

VIII. Tux Pres (Pl. XX, Fig. 24).

The general appearance of the hind foot recalls that of the viverrines. The astra-
galus is quite like that of Daphenus, but with some differences which tend in the direc-
tion of the modern Canidae, this bone in Cynodictis standing intermediate in structure
between the two extremes, though somewhat nearer to Daphenus. The proximal or
tibial trochlea is but little more deeply grooved than in the latter genus, and is therefore
much shallower than in Canis, but its borders have the same clean-cut angularity as in
the modern forms, instead of curving gradually into the facets for the tibial and fibular
malleoli. Im Canis the tibial trochlea is extended over upon the dorsal side of the neck,
but this is not the case in either of the White River canines. The neck of the astraga-

394 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

lus is relatively longer than in Canis or eyen than in Daphenus, resembling that of such
viyerrine genera as Paradoxurus, but is not directed so strongly toward the tibial side of
the foot as in Daphenus. The head with its convex navicular facet is shaped much as
in Canis, except that it is more depressed in the dorgo-plantar dimension. In Daphe-
nus there is a distinct facet for the cuboid, which meets the navicular facet nearly at
right angles; in Cynodictis this cuboidal facet is very much smaller and sometimes it is
altogether wanting, while in Canis the astragalus and cuboid are not in contact. As in
Daphenus, the external caleaneal facet is more oblique in position and more simply con-
rave than in Canis, but the sustentacular facet is different from that of both the genera
mentioned ; it agrees with that of Daphaenus in being shorter and wider than in the
modern forms, but while in the former this facet is separate from that for the nayicu-
lar, in Cynodictis, as in Canis, it is confluent with it, but at a different point ; i. €., more
toward the tibial side. The interarticular sulcus is somewhat deeper than in Daphenus,
but shallower than in Canis. In the latter we find a third calcaneal facet which forms a
narrow band upon the fibulo-plantar side of the head and is connected at one end with
the sustentacular facet. This accessory caleaneal facet does not occur in either of the
White River genera.

The calcaneum, like the astragalus, is more viverrine than canine in general appear-
ance and quite closely resembles that of Paradoxurus, but the resemblance to Daphenus
is even more marked. The tuber is slender, compressed and proportionately much
shorter than in Canis; in the latter the tuber makes up more than two-thirds of the
total length of the caleaneum, while in Cynodictis it is about two-fifths of this length.
The free end of the tuber is moderately thickened and club-shaped and is deeply grooved
by the sulcus for the plantaris tendon. As in Daphenus, the dorsal and plantar borders
of the tuber are nearly parallel and its dorso-plantar diameter is thus almost uniform
throughout, not increasing toward the distal end as it does in Canis. Near the distal end
of the caleaneum and on the fibular side is a very prominent process for the attachment
of the lateral ligaments. This process is not present in the recent Canidae, but is very
conspicuous in the primitive carnivores, such as Dinictis ahd Daphenus, and it recurs
among modern plantigrade and semiplantigrade forms, such as Procyon, G'ulo, Para-
doxurus, ete. Usually, however, it is smaller and less prominent in the fossil than in the
recent genera. The facets for the astragalus are somewhat different from those of both
Daphenus and Canis. In the latter the external astragalar facet is in two parts, one of
which presents distally and the other dorsally, the two meeting at an angle which does
not much exceed 90° ; in the former the whole facet forms one continuously curved con-
vexity, not divided by an angulation. In Cynodictis the two parts are distinguishable as
in Canis, but they meet at a much more open angle. The sustentaculum is of moderate

NOTES ON THE CANIDA OF THE WHITE RIVER OLIGOCENE. B95

prominence and, as in Daphcenus, it carries a subcircular facet for the astragalus ; in the
modern genus this surface is narrower and more elongate. The sustentaculum also agrees
with that of Daphenus in not being so obliquely placed, with reference to the long axis
of the caleaneum, as in the existing members of the family. On the plantar side,
between the sustentaculum and the body of the bone, is a groove, the sulcus flexoris hal-
lucis, which is better marked in Canis than in either of the White River genera. This
is curious, in view of the fact that the latter possess. a well-developed and functional
hallux, while in the former this digit is reduced to the merest rudiment. In Canis we
find a third facet for the astragalus, a small plane surface distal to the sustentaculum,
from which it is separated by a narrow sulcus; continuous with this accessory facet, but
at right angles to it, is a small facet for the navicular. Neither of these articular surfaces
is to be found in Cynodictis. The facet for the cuboid, which in the recent dogs is almost
plane and semicircular in shape, is quite deeply concave and of nearly circular outline.

The cuboid is relatively high and narrow, differing from that of Canis principally
in the smallness of its transverse and dorso-plantar diameters. The proximal surface is
occupied by a large facet for the calcaneum, which, as in Daphenus, is much more con-
vex than in the existing dogs. The hook-like projection from the plantar side, which in
Daphenus is very large and prominent and in Canis is even more massive, in the present
genus is quite inconspicuous and is continuous with the projection from the fibular side
which overhangs the deep tendinal suleus. The astragalar facet is small and is confined
to the dorsal side of the cuboid, being much less extensive than in Daphenus. ‘The facet
for the nayicular is not so prominent as in Canis or eyen as in Daphenus, and is con-
tinuous with that for the ectocuneiform. The distal end of the cuboid resembles that of
Daphenus in haying quite a concave facet for the head of the fourth metatarsal, while
that for the fifth is lateral in position. In Canis, on the other hand, the surface for mt.
ly is almost plane and that for mt. vy occupies an entirely distal position ; the plantar
portion of the facet for mt. iv is much narrower than in the two White River genera,
and has thus quite a different shape and appearance.

The navicular. is almost a miniature copy of that of Daphenus and presents the
same differences from that of Canis. Seen from the proximal end, it is of more regularly
oval shape and is less contracted on the plantar side than in the modern genus. The
position of the navicular in the tarsus is likewise different. In Canis this bone has been
somewhat rotated, so that its principal diameter is the dorgo-plantar one, and on the
plantar border it has been brought into contact with the calcaneum, for which it has
acquired a special facet. It is of interest to observe that a similar but more extensive
rotation of the tarsal elements has been carried out in the horses, as Riitimeyer has
shown. In the White River genera, on the other hand, the principal diameter of the

96 NOTES ON THE CANIDEH OF THE WHITE RIVER OLIGOCENE.

©o

navicular is transverse, and owing to the elongation of the neck of the astragalus, it is
carried so far distally that it can have no contact with the caleaneum, the astragalus
articulating with the cuboid. The astragalar surface is concave, but somewhat less so
than in Canis, and the facet for the cuboid is small and confined to the dorsal moiety of
the fibular side. The distal end displays the usual facets for the three cuneiforms, which
do not require any particular description.

The entocuneiform has much the same shape as in Canis, elongate in the proximo-
distal diameter, but very narrow and much compressed. The nayicular facet is rela-
tively smaller than in the modern genus and there is no such distinct facet for the meso-
cuneiform. The distal surface, for the head of the first metatarsal, is no wider but much
more deeply concave than in Canis.

The mesocuneiform is a minute bone and, as in the fissipede Carnivora generally, its
vertical or proximo-distal diameter is much less than that of the adjoining ento- and
ectocuneiforms, forming a depression or recess in the distal row of the tarsus, into which
the head of the second metatarsal is tightly wedged. The only articular surfaces visible
on the mesocuneiform are the proximal and distal, for the nayicular and the second meta-
tarsal respectively.

The ectocuneiform is much the largest of the three. Compared with that of Canis,
it is narrower in proportion to its height and is also less extended in the dorso-plantar
dimension, but the projecting process from the plantar surface is even more prominent,
and is more thickened and club-shaped at the free end. On the tibial side is a minute
facet (not double as in Canis) for the side of mt. ii. The facet for the cuboid is much
smaller than in the modern dogs and is confined to the dorsal border, while at the infero-
external angle of the bone is a minute facet for the head of mt. iv, which is not repre-
sented in Canis. The distal end of the ectocuneiform is taken up by a facet for mt. 111,
which is less concave and has a shorter plantar prolongation than in the modern genus.

The metatarsus consists of five well-developed members. Unfortunately, there is
not a single complete metatarsal preserved in connection with any of the specimens, but
enough remains to show that these bones were much longer and stouter than the meta-
carpals, and that the disproportion in size and length between ‘the fore and hind feet
was much greater than in the recent dogs and quite as great as in many viyerrines, such
as Herpestes and Paradoxurus or as in Daphenus.

The first metatarsal is sufficiently weli preserved to indicate that the hallux was
well developed and functional, though somewhat more reduced than in Daphenus, or in
such recent viverrines as Cynogale or Paradoxurus. The head bears a narrow, convex
facet for the entocuneiform and upon its tibial side is a large, rugose prominence for the

attachment of the lateral ligament. The shaft is very slender and is arched slightly

NOTES ON THE CANIDH® OF THE WHITE RIVER OLIGOCENE. 397

toward the fibular side of the foot, making the tibial border somewhat concave. ‘The
length of the bone, as already intimated, is not determinable, but the portion preserved
in one specimen is nearly as long as the entire fifth metacarpal of the same individual.

The second metatarsal is much stouter than the first and more slender than the
third. The head is very narrow, being slightly excavated on the tibial side. Owing to
the shortness of the mesocuneiform, the head of mt. 11 rises above the level of mt. i
and iii and is firmly held between the ento- and ectocuneiforms, though there are no such
distinct lateral facets for these tarsals as we find in Canis ; a stout prominence occupies
the plantar side of the head. The shaft is slender and of oval section, not having
acquired the trihedral shape characteristic of the recent dogs.

The third metatarsal is the stoutest of the series; the head is broad dorsally but
very narrow on the plantar side, where there is a large, projecting process, more promi-
nent than in Canis. The facet for the ectocuneiform is convex (in the recent dogs it is
slightly concave) and oblique in position, inclining downward toward the tibial side.
Deep sulci invade the head on both sides; on the tibial side the sulcus is narrow, but
that on the fibular side is broad. A deep pit on the fibular side of the head receives a
corresponding prominence from mt. iv, and an additional facet for the same metatarsal is
found on the plantar projection, so that the two median metatarsals are very firmly inter-
locked. The shaft, for most of its length, is of transversely oval section, very different
from the squared, prismatic shape seen in Canis, though an approximation to this shape
occurs in the proximal portion of the shaft, where mt. iii and iv are closely appressed.
The distal end is broadened and antero-posteriorly compressed ; the trochlea resembles
that of the corresponding metacarpal, save that it is larger and relatively somewhat
lower.

The fourth metatarsal is of nearly the same thickness as mt. ili, though a trifle
more slender. The head is narrow and the facet for the cuboid is slightly convex in
both directions ; the plantar extension is neither so broad nor so prominent as in Canis.
On the tibial side is a rounded protuberance, which is received into the depression
already mentioned, in the head of mt. iii, while on the fibular side is an exeayation for a
prominence on mt. v, and proximal to this excavation is a narrow but well-defined facet
for the same metatarsal. Very little of the shaft is preserved, and this proximal por-
tion has much the same tetrahedral shape as in the recent dogs. Doubtless, however,
the distal part of the shaft assumes a transversely oval section, as does that of mt. iu,
though the digits of the pes evidently diverge less distally than do those of the manus.

The fifth metatarsal is entirely missing from all of the specimens, so that the inter-
esting question regarding the reduction of the external ascending process cannot be

answered,

398 NOTES ON THE CANID#® OF THE WHITE RIVER OLIGOCENE.

The phalanges of the pes do not differ from those of the fore foot, except in their

considerably greater size.

Measurements.

| No. 10493. | No. 11012. | No. 11381.
is |
Tarsus, height (excl. caleaneum) | .021
CH KeRMeTIT, HEMEL N 2550060000 00009 000000030 0005a0000900000BnD25a0090000CDDOS. DascORDOHDDODEOSanoNODDAoDAoG0N00 | 0195 .020
ig IGiTELH OF HWIORT -ocsss0000y 92000000 qos 2009 nND00000000090 209d .0EEN.8DGs0D09GEKONRADD9baB00=00 012 012
uC GlomsO=jy Farms, GDR 50ccsncco5e epo006.9990020000G0000090000 359509 {cb ona503HE3s6q05NRDIEINNEO00]| .007 .008
Agim me}, WET E3§)N 5000009000 900000590202900200090000090 903003020 DN9DaD9D0EI IDI nIODUBTOIIcNUBASZDgTOBHIDODONE 013 .013 014
“ STATO RAE EOCTIN Gay mess eee ares aes a ta ae ane tae aan ee Le TOC eeeo0s 0055 | 006
od IgTRRR HN OY WEE] Rococoessscocans coosougDsassHondEDsEHUDS eooCURLODJasbaDIDEGEDDOsBDSONUALOUOBEDESNO .006 .006 | .006
ag SyiLOHUaWCoye 10Y 27 Gnaceno recone kaseacaccaes meaasepecine cee ccosntncedtenceicchy sasnacceee cere 007 | .007 | .008
INAAGTIE, TREE ococoscon00 cons ocadscosasnooesce00Us0c0Ie209=.esasro0009 SonEEEDoSecanCoSosas0NRLON.ELADDODHS .003
es PVT OL tibaicisesceriets wrsio eis cteteisiotvie slats e a eiicletesicia's sisiectovs laters ate stese mia a mets siatete sats cieinle sla oa wiselnsneisecivtalceaiomeces .006 |
Hetocuneiforme Neieht.csssinccuesessndeccesessnccccessoecsanoneteesesevennasdaneccnecassancensnectnsresackt | .0045
of width dist. end ... | .0045
INGEN! Thy VaAGlt TOROS, Gil joooosc0n.5ce00s00n0oD.ocs0000bccnoSEEDIONG0D snoDoDOADOD SEH 0ONNDONoELODONS .0045
a CR 003 | .003
a6 TiS sae ahs Ces | 005 | .005
“< iii, width dist. end... 005
os iv, width prox. end -0035 |

IX. ReEsrorarion.

The general appearance of the Cynodictis skeleton has little about it to suggest
canine affinities, but has some resemblance to the civets and especially to the herpestine
section of that family. This resemblance is not merely a general one of outline and pro-
portions, but may also be traced in many of the details of structure. The small head,
with its elongate and narrow cranium and short, tapering muzzle, is of strikingly viver-
rine character. So is also the neck, which is relatively long and stout, the vertebrae hay-
ing heavy centra and well-developed processes. The resemblance to the civets continues
into the thoracic region, where the vertebrae are small, especially in the anterior portion,
and have short, slender neural spines. The thorax itself, with its slender and moderately
curyed ribs, is narrow and compressed, as in the Carnivora generally, while the prominent
and compressed manubrium has a somewhat viverrine appearance. The lumbar region is
long and is strongly curved upward; the vertebre are much elongated, with stout
depressed centra, very long, slender and anteriorly directed neural spines, which are not
like those of modern dogs or civets and most resemble the spines of Lynx. The trans-
verse processes are likewise peculiar in their length and slenderness. The tail is unlike

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. 599

that of the modern dogs, being much longer, stouter and in every way better developed ;
it was not, perhaps, quite so long proportionately as in Herpestes, but nearly so. This,
however, is a primitive feature, which is common to the greater part of the earlier carni-
vores and ungulates, and is even more conspicuous in Daphenus than in Cynodictis,
while the White River Machairodonts, Dinictis and Hoplophoneus, have very long and
massive tails.

The limbs, though not so long proportionately as in the recent dogs, are much more
so than in the John Day species, C. geismarianus, the hind legs being especially elon-
gate. The scapula is not at all canine in character, being relatively very large and
having the broad blade and irregularly curved coracoid border of the viverrines ; the
great length of the acromion and the unusual size of the metacromion are peculiar.
The humerus is short but quite heavy, and with its low trochlea, prominent deltoid and
supinator ridges, and large epicondyle and epicondylar foramen, has an exceedingly
viverrine appearance. The ulna and radius are relatively short and slender, and the
discoidal head of the latter shows that the power of rotating the manus had been but
little diminished; the great styloid process of the radius is very characteristic. The
‘arpus 1s low and the metacarpals are exceedingly short and weak, resembling in their
proportions those of Paradoxurus. The phalanges are elongate and the claws sharp
and compressed.

The pelvis has a viverrine appearance in its shape and in the elongation of its
posterior portion, while the os penis resembles that of the mustelines in size and curya-
ture. The femur is long and the tibia is somewhat longer than the femur, bearing much
the same relation to that bone as in Canis, while the fibula is much stouter than in the mod-
ern genus. The pes is far larger in all its dimensions than the manus, the difference in
size between the two being much greater than in Canis. It is often exceedingly difficult
to determine from the bones alone whether a given animal was plantigrade or digiti-
grade in gait, but from the resemblance of the limb and foot bones of Cynodictis to those
of the civets, it seems very probable that the former had a similar semiplantigrade gait.

The John Day species, C. geismarianus, is considerably larger than the White River
forms, but resembled the latter in proportions. Cope says of it: “ Although the skull
and pelvis of this species have about the size of those of the fisher, the vertebrae and
humerus are more slender and the anterior foot is decidedly smaller. It is probable that
the Galecynus [7. e., Cynodictis] geismarianus resembled a large Herpestes in general pro-
portions rather than a Canis. It stood lower on the legs than a fox and had as slender
a body as the most ‘ vermiform ’ of the weasels, the elongation being most marked in the
region posterior to the thorax. The tail was evidently as long as in the Ichneumons.
Its carnivorous propensities were as well developed as in any of the species mentioned,

A, B: & VO, SOX, DW:

400 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

although, like all other Canidw of the Lower Miocene period, the carnassial teeth are

relatively smaller than in the recent types ” (’89, p. 929).

The White River species of this genus are probably two in number.
CYNODICTIS GREGARIUS Cope.

Syn. Amphicyon gracilis Leidy (non Pomel), Proc. Acad. Nat. Sci. Phila., 1856, p. 90 ;
1857, p. 90; Ext. Mamm. Fauna Dak. and Nebr., p. 36. —Amphicyon angustidens
Marsh, Amer. Journ. Sci. and Arts, 3d Ser., Vol. Il, p. 124. Canis gregarius Cope,
Ann. Rept. U.S. Geolog. Surv. Terrs., 1873, p. 506. Gralecynus gregarius Cope,
Tertiary Vertebrata, p. 916.

This is the species which has been described so minutely in the foregoing pages. It
is one of the commonest White River animals and is very much more frequently met
with than any of the contemporary carnivores. Despite this abundance of individuals,
well-preserved specimens are rare and eyen these consist mostly of skulls only. As will
be seen from the tables of measurements, the different specimens vary little in size or in
the proportions of the various parts of the skeleton. One apparent exception to this
statement may be found in the case of No. 11381, which is remarkable for the length of

its hind limb, but this probably belongs to the following species :

CYNODICTIS LIPPINCOTTIANUS Cope.

Canis lippincottianus Cope, Synopsis of Vertebrata Collected in Colorado ; Miscell. Publ.
U.S. Geolog. Surv. Terrs., 1873, p. 9; Ann. Rept. U.S. Geolog. Surv. Terrs., 1873,
p- 006. Galecynus lippincottianus Cope, Tert. Vert., p. 919.

The status of this species is still a matter of some uncertainty ; Cope, who estab-
lished it upon mandibular rami, describes it as haying “dimensions half as large again as
in C. gregarius,” and adds: “ Unfortunately there is not enough material in my hands
to render it clear whether the specimens represent a distinct species or a large variety of
the C. gregarius” (85, p. 920).

Among the specimens described in the foregoing pages is one (No. 11381) in which
the limb bones decidedly exceed in length and thickness those of the other individuals,
while the cranium is but little larger. Probably this specimen should be referred to C.
lippincottianus, but in the absence of teeth the reference can be only provisional.

In the John Day formation Cynodictis is represented by more numerous and more
varied species than in the White River beds; from the former horizon Cope has deter-

mined C. gregarius, C. lemur, C. latidens and C. geismarianus.

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. 401

Still another species should be mentioned in this connection. In the American
Museum of Natural History, New York, are the remains of a small eynoid animal from
the Uinta beds, which may belong to Cynodictis, or if not, should be referred to some
closely allied genus. It is important to observe that in the Uinta stage (uppermost
Kocene or lowest Oligocene) we find that the two canine series, represented in White
River times by Daphenus and Cynodictis, had already been established.

Tur PHYLOGENY OF THE CANIDA.

It seems probable that the fossil genera of this family already known are sufficient
to indicate to us the main outlines of its phylogenetic history. The problem of recon-
structing the series is, however, obscured by two circumstances ; first, the variety and
multiplicity of nearly allied genera, the mutual relationships of which are very complex
and difficult to disentangle ; and in the second place, by the fact that only rarely do we
obtain satisfactory material of any of the genera. Most of the forms are known only
from the skull and teeth, and the skeleton has, so far, been found in but few of the
species. Cynodictis, Daphenus, Temnocyon and dlurodon are now known from more
or less complete skeletons, but we shall need to learn far more than we know at present
concerning the structure of the other genera before we can reach a solution of the many
problems of canine phylogeny.

Before taking up the discussion of these phylogenetic problems, it will be conveni-
ent to establish the order of geological succession in which the various genera make their
appearance. We haye seen that in the Uinta there appear to be two distinctly sepa-
rated canine series, one of which is represented by ? Iacis and the other by a genus
which is very closely allied to, if not identical with Cynodictis. The former series would
seem to be continued into the White River by Daphenus and the latter, of course, by
Cynodictis. The latter genus may well prove to be of Old World origin, for in the
European Oligocene it attains such a variety and fullness of development as it never
reached in America, although, on the other hand, the American creodont genus MJiacis,
from which Cynodictis probably took its origin, has not yet been found in Europe. In
the John Day stage the canine phylum underwent an extraordinary expansion. Daphe-
nus persisted, but is represented only by a single small species, D. cuspigerus, while the
series branched out into several distinct and more or less specialized genera, such as
Temnocyon, Hypotemnodon, Cynodesmus, Enhydrocyon, and perhaps even the little known
Hycnocyon. No new genera of the Cynodictis series haye yet been detected, but that
genus itself became differentiated into many more species than occur in the White River,
and some of these may, on better knowledge, prove to be generically distinct. On the

other hand, Oligobunis probably represents, as Schlosser has suggested, an immigrant

402 NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE.

from the Old World, belonging to the series which leads from the Oligocene Cephalogale
to the Phocene Simocyon. The dogs of the Loup Fork, with the exception of the aber-
rant d/urodon, are very imperfectly known and the remains of them which have been
found are not, according to present knowledge, generically separable from Canis, though it
hardly seems probable that the modern genus had actually been differentiated so early as
the upper Miocene, and we may regard it as extremely likely that these supposed repre-
sentatives of Canis will eventually prove to belong to more primitive genera. None of
the forms which haye hitherto been found in the Loup Fork beds can be referred to the
Cynodictis line.

The mutual relationships between the two canine series, which are already so well
distinguished in the Uinta, are quite obscure and puzzling, although there is nothing to
forbid the ‘assumption that both series converge to a common ancestor in the Bridger, per-
haps the genus JMacis. The Cynodictis series, when we first meet with it, is decidedly
more adyanced than the other phylum, as is shown in the deyelopment of the skull, the
reduction of the dentition, the character of the limbs and feet and the digitigrade gait.
Continuing through the White River age and, so far as North America is concerned, at-
taining its maximum of deyelopment in the abundance and yariety of its species in the
John Day, the line apparently disappears and can be traced no farther. Whether the
series actually died out at the end of the John Day, or whether it continued farther and
possesses representatives even at the present time, are questions which cannot yet be defi-
nitively answered. Schlosser (88, p. 247) has suggested that some of the species of Cyno-
dictis may, perhaps, be of phylogenetic significance in the canine stem, but if so, they
can hardly be placed in the thooid series, which apparently has no place for them. M.
Boule (89, p. 321), in an article upon the Pliocene Canis megamastoides Pomel, comes to
the conclusion that the modern Canidw are diphyletic, and have arisen by a process of
convergence, the thooids and the bears being divergent groups derived from Amphicyon,
while the alopecoids and viverrines are descended from Cynodictis. In discussing the
affinities of the Pliocene form Boule says :

“Ta description précédente nous montre que le fossile de Perrier se rattache de plus
pres aux Renards qu’ aux autres représentants actuels de la famille des Canidés. Par
son crane, le Canis megamastoides ressemble beaucoup le Renard de nos pays. Par la
forme de sa mandibule, il se place au contraire pres des Renards américains (Canis
cancrivorus, C. azarae, C. cinereoargentatus) et prés de ? Otocyon megalotis de V Afrique
australe. Ces espoces, notamment la derniére, sont regardées par tous les auteurs com-
me des formes primitives.

“Tout en ratifiant ce premier rapprochement, la dentition presente des caractéres
particuliers que nous retrouyons en grande partie dans les Cynodictis et Cephalogale du

Miocéne (p. 527).

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. 403

“Les belles récherches de M. Filhol nous ont révélé la richesse en especes de ces
genres si curieux, placés aux confins de plusieurs familles de Carnassiers. Les Cynodic-
tis et les Cephalogale avaient la formule dentaire des Chiens actuels, mais leurs dents
presentaient un aspect particulier qui a yalu a ces animaux fossiles le nom de Chiens
viverriens. Or en ¢tudiant les pieces originales de la collection du Muséum et les livres
de M. Filhol sur les Phosporites du Quercy, j’ai été frappé de retrouyer, comme parsemés
dans diverses especes de Cynodictis beaucoup des characteres présentés par le Canis mega-
mastoides ” (p. 328).

“Tl semble done que les Renards actuels représentent une branche emanée du buis-
son touffer des Cynodictis, duquel se serait également detachée la branche des Viverridés.
Je suppose que lorsqu’ on connaitra suffisament les membres des diverses espéces de Cyno-
dictis, on trouvera des formes de passage allant d’un cété aux membres des Viverridés et
Wun autre c6té aux membres des Renards.

“Si ces considerations sont exactes, les Chiens ont une origine differente des Renards.
Les Amphicyons représentent les anecétres communs des Ours et des Chiens, comme les
Cynodictis représentent les ancétres communs des Ciyettes et des Renards ” (p. 329).

M. Boule’s argument as to the derivation of the foxes from Cynodictis is not a very
convincing one and is open to several obyious objections. In the first place, M. Boule does
not define the sense in which he uses the term fox ; it is evidently. not the same as Hux-
ley’s alopecoid, for C. canerivorus and C. azare are called foxes, while Huxley regarded
them as typical though primitive thooids. M. Boule does not say whether C. megamas-
toides possessed a frontal sinus, but from the statement that “le frontal est saillant, 4 sur-
face arrondie” (pp. 324, 525), one would infer the presence of a sinus, and if so, CL mega-
mastoides is not an alopecoid, but a thooid. The presence or absence of frontal sinuses
and the shape of the cerebral fossa are the only diagnostic characters which Huxley could
find definitely distinguishing the two canine series from each other. In the second place,
the resemblances in tooth structure between Cynodictis and Canis megamastoides, upon
which M. Boule places such emphasis, are in themselves of no great value, because the
resemblance of the latter species to Cephalogale is eyen greater, and Cephalogale, as
Schlosser has shown, probably belongs in a totally different line, which has no existing
representatives. In any event, the gap between the Pliocene and Oligocene forms is
still so wide that no determination of the taxonomic value of their resemblances and
differences can yet be made.

Again, it is highly improbable that the yiverrines can be descended from Cynodictis,
for the latter, though having certain marked resemblances to the civets, is in all essen-
tials of structure distinctly a member of the Canid@, and is no more ancient than cer-

tain unmistakable viverrines. Indeed, the genus Viverra itself is reported from the

404 NOTES ON THE CANIDA OF THE WHITE RIVER OLIGOCENE.

upper Eocene of Europe, occurring in the same horizons as those in which Cynodictis
first appears. For similar reasons, it is very difficult to believe that Amphicyon can be
the ancestor of the thooids, for that genus has already begun to become differentiated in
the direction of the bears and is contemporary with or even younger than certain Ameri-
can genera, such as Temnocyon and Cynodesmus, which are undeniable thooids.

M. Boule’s hypothesis involves some rather startling consequences ; if true, we shall
be forced to conclude that the two series of modern Canidw have been separated ever
since the close of Eocene times and that they had no common ancestor nearer than the
middle Eocene or Bridger stage. This conclusion would imply such an extreme and
remarkable degree of parallelism or convergence as has hardly been believed possible,
an exact parallelism in all parts of the dentition, skeleton and soft parts, terminating in
almost complete identity of structure. Indeed, many systematists regard most of the
modern foxes and wolves as belonging to the single genus Canis, and Huxley speaks of
the differences between them as being so slight, that a generic separation can be justi-
fied only on the grounds of convenience. Is it conceivable that two series of mam-
mals which were already separated in the Eocene should have converged into what is
practically a single genus ?

Unlikely as it may appear, I am inclined to believe M. Boule’s hypothesis concern-
ing the relationship of Cynodictis to the alopecoids is not to be summarily dismissed, but
that it may eventually prove to be well founded. It is certainly a suggestive fact that
Cynodictis, like the foxes, is deyoid of any frontal sinus, while all of the other Ameri-
can genera, from Daphenus onward, have well-marked sinuses, as in the wolyes. Fur-
thermore, whatever conclusion we may reach with regard to the single or dual origin of
the Canidae, there is much reason to believe that such extreme cases of parallelism and
convergence have occurred among mammalian phyla and that they may be more fre-
quent than is commonly supposed. One very striking example is that of the true cats
(Feline) and the sabre-tooth series (Jachairodontine) originally pointed out by Cope
and elaborated in much detail by Adams (96).

Unfortunately, complete demonstration is lacking in this very extraordinary case of
parallel development, because the early stages in the phylogeny of the true cats have not
yet been recovered, but the successive genera of the Machairodonts are fairly well known,
and they form a connected series. None of these machairodont genera, not even the ear-
liest and most primitive of them, can be regarded as ancestral to the true eats, for with-
out exception they all display the characteristic and unmistakable features which place
them in the sabre-tooth series. The more primitive genera, such as Dinictis, possess a
dentition which is but slightly modified in the direction of the cats, and cranial foramina
resembling those of the early dogs in the presence of an alisphenoid canal, the separa-

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. 405

tion of the condylar foramen from the foramen lacerum posterius, ete.; the femur has a
third trochanter and the humerus an extremely prominent deltoid ridge; the feet are
plantigrade and pentadactyl and, like those of many of the viverrines, they are supplied
with partially retractile and very incompletely hooded claws. In all probability these
structural characters also occurred in the ancestral Feline, but what distinguishes even
the earliest Machairodonts is the elongation and compression of the upper canines, the
reduction in size of the inferior ones and the development of bony flanges from the ven-
tral border of the mandible for the protection of the superior tusks. From such begin-
nings the sabre-tooth series may be traced, with various divagations and side branches, to
the Pleistocene Smilodon, which in all parts of its structure is extraordinarily like Felis,
the only important differences consisting in the dentition (which is of similar type) and
in the modifications of the skull, which are necessarily correlated with the enormous
enlargement of the upper canine tusks. |

Seeing, therefore, that the machairodont series is well-nigh complete and that none
of its known members is at all likely to prove ancestral to the true eats, there can be
little reasonable doubt that the remarkably close resemblance which we observe between
Felis and Smilodon is not directly due to their relationship, but has been independently
acquired in the two series and is the outcome of a parallel course of development, con-
tinued from the Oligocene to the Pleistocene. If this be true, there can be no @ priori
ground for denying that the same phenomena may have been repeated in the dogs and
that Boule’s suggestion concerning the derivation of the alopecoids from Cynodictis may
possibly prove to be correct. In this case, however, the final identity of the two series is
even more striking than in the cats and Machairodonts ; to verify the suggestion, it will
be necessary to recover the missing links of the alopecoid phylogeny and to show that it
has followed a course parallel to but independent of that of the thooids.

Another alternative possibility is that the foxes became separated from the principal
canine phylum at a comparatively late date, and that, consequently, Cynodictis and its
allies represent but an abortive side-branch from the main stem. That the separation is
of considerable antiquity is shown by the parallel arrangement of the two series to which
Huxley has called attention. In both wolves and foxes we find species with microdont
and macrodont dentition, with sagittal crests and lyrate sagittal areas, with lobate and
non-lobate mandibles. So far, at least, we are almost certainly dealing with indepen-
dently acquired characters. From the standpoint of present actual knowledge it is more
probable that the separation did not take place before the end of the Miocene than that
it had already been accomplished in the Eocene, though this conclusion involves the
admission that Cynodictis had anticipated the foxes in quite a remarkable way. While
very far from denying the possibility of such convergence as is implied in Boule’s

406 NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE.

hypothesis, I think it should not be assumed in a given case except upon the clearest
evidence. Whichever of these alternatives be true, it is, in any event, probable that the
alopecoids are not of American origin.

Still a third possible solution of the problem concerning the mutual relationships of
the wolves and foxes is that Cynodictis, or some similar form, is the common ancestor of
both lines, and that the supposed early thooids, such as Daphenus and Cynodesmus, are
devoid of permanent phylogenetic significance. This is decidedly the least probable of
the three alternatives, for the thooids of the American Oligocene and Miocene seem to
form a truly connected series, in which Cynodictis has no place. Further, this view
involves the assumption that the supposed thooids have independently run a course par-
allel to that of the true thooids and thus encounters the very difficulty which it was
intended to avoid. The conclusion which we reach is, therefore, that the thooids are
probably of American origin and that the alopecoids are a branch which the wolf stem
gave off after certain of its representatives had established themselves in the Old World.

The thooid genealogy itself is by no means free from difficulties. In a former paper
(94), I suggested that the line begins in Daphanus of the White River, and is con-
tinued by the John Day Cynodesmus, but now that we haye learned the remarkable char-
acters of the skeleton, especially of the limbs and feet, of the former genus, this view no
longer appears so simple and natural, and its acceptance carries with it some far-reaching
and unexpected consequences. In particular, it might be objected to this view that the
peculiar differentiation of the feet in Daphenus would exclude that form from any place
in the direct canine phylum, for it seems @ prior? unlikely that the dogs should first have
acquired the power of retracting the claws and should then have subsequently lost it.
Indeed, many morphologists are inclined to deny altogether the possibility of this method
of evolution. In the present state of knowledge, however, such a denial is at least prema-
ture, and there is a considerable body of evidence which goes to show that it does not
properly apply in the case of the canine phylum.

In the first place, the John Day genus Zemnocyon, the osteology of which has been
very fully described by Eyerman (’96), appears to be a direct descendant of Daphnuse,
with which it agrees in the essentials of structure, though, at the same time, it displays
many marked changes and advances. One of the most striking of these changes in the
later form is in the great elongation of the limbs and the assumption of a digitigrade
gait, both limbs and feet quite closely approximating those of the modern Canide. Yet
even in Temnocyon a reminiscence, as it were, of the partially retractile claws of Daphe-
nus may be observed in a certain asymmetry of the second phalanges of both manus and
pes, which are slightly excayated on the ulnar and fibular sides respectively. While
Daphenus was a short-limbed, plantigrade or semi-plantigrade form, which, in all

NOTES ON THE CANIDA OF THE WHITE RIVER OLIGOCENE. 407

probability, was not cursorial in habits, Zemnocyon, on the other hand, was undoubt-
edly cursorial and probably essentially resembled the modern wolves in appearance and
habits. In this change to a digitigrade gait and cursorial habit, it seems juite reasonable
to suppose that the mode of using the claws should have been changed likewise, the feet
being used almost exclusively for purposes of locomotion and the claws losing their
importance as weapons and grasping organs. Under these circumstances the power of
retraction would become superfluous and tend to disappear, although, as we have seen,
Temnocyon retains recognizable traces of the structure which permits retraction of the
claws. It is true that Zemnocyon itself isnot in the direct line which leads up to the
modern Canidae, for the heel of the lower sectorial and the whole of m z have become
trenchant through the loss of the internal cusps, a curious specialization ; but, on the
other hand, there is no reason to suppose that it differed in any other important respect
from its contemporary Cynodesmus, which appears to be a member of the direct phylum.

In the second place, a similar loss of the power of retracting the claws has almost
certainly occurred among the Fe/ide. The hunting leopard or cheetah (Cynelurus) has
acquired something of the proportions and appearance of the wolves, haying very elon-
gate limbs and feet and a running gait which is described as quite different from that of
the ordinary cats. Comparing the phalanges of Cynelurus with those of Helis, some
marked differences are at once apparent; in the lateral digits the second phalanx is
quite symmetrical and is not excavated on the ulnar (or fibular) side; the excavation
is distinctly shown only in the third digit and is much less marked in the fourth. The
bony hood of the ungual phalanx is much reduced, leaving more than half the length of
the phalanx exposed, and the subungual process is much smaller than in Fe/is. The tar-
sus, in fact the skeleton of the entire pes, has a canine aspect, and the retractility
of the claws is very partial and imperfect. Now, there can be little doubt that Cyne-
lurus is not the remnant of a very ancient group, given off from the feline stem at a
time when the power of retracting the claws had been but partially attained, but that it
was derived from ancestors which differed little from Felis. If such a transformation
could take place among the cats, there would seem to be no good reason for denying that
it might also occur in the dogs.

Unfortunately, the phylogenetic history of the dogs is not made clearer and more
intelligible by reason of the new material of Daphenus, which has been described in
the foregoing pages, and which raises more problems than it solves. I am inclined to
believe, however, that Daphenus should still be given a place in the canine phylum,
for the differentiation of its limbs and feet is hardly of that radical kind which would
prevent a subsequent change in the trend of development, and its many resemblances
to the early Machairodonts are, at least in part, survivals of primitive conditions, sey-

A. P. S.—VOL. XIX. 2 Z.

408 NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE.

eral of which, like the shape of the radius, reeur in Cynodictis. Tending to the same
conclusion is the fact that what little is known of the structure of the ecreodont Miacis
is of similar composite canine-feline character and it is to that creodont family to which
most of the lines of fissipede Carnivora appear to lead back. It may be hoped that the
problem will receive its definite solution when we shall have recovered the as yet miss-
Ing or very imperfectly known dogs from the Uinta, uppermost White River and lowest
John Day formations, and are thus enabled to trace the successive changes step by step.
Assuming, then, as probable that Daphenus should have a place in the direct canine
phylum, the larger question at once arises: What was the relation between the early
members of the Caunide and Felide, and of both of these groups to the other fissipede
families? It seems to be a comparatively rare phenomenon among the mammals that
parallelism or convergence of development should be manifested in all parts of the struc-
ture of two independent lines, though that this may happen is shown by the case of the
Machairodonts and felines, to which reference has already been made. Usually, however,
parallelism is displayed in a few structures only, such as the dentition, or the feet, or the
vertebrae, and the more widely separated any two phyla are at their point of origin, the
less likely are they to develop along similar lines. It will be sufficiently clear from the
foregoing descriptions that the resemblances between Daphenus and the more primitive
Machairodonts, such as Dinictis, are not only exceedingly close, but that they recur in
all parts of the skeleton. The skull, the vertebral column, the limbs and the feet are
all so much alike in the two series that, in the absence of teeth, it is often very difficult
to decide to which of the two a given specimen should be referred. Such close and gen-
eral resemblance is prima facie evidence of relationship, even though it should have been
independently acquired, because parallelism is much more frequent between nearly allied
than between distantly related groups. In the present instance, however, there is no rea-
son to infer that the resemblances were separately attained ; on the contrary, the evidence
now available seems to favor the conclusion that the dogs and cats are derivatives of the
same Eocene stock. It cannot be pretended that this conclusion is, as yet, a well-estab-
lished one, nor can it be so established until we recover the missing links of the canine
and feline genealogies. Daphenus may eventually prove to be merely an abortive side-
branch without phylogenetic significance, though this seems unlikely in view of its rela-
tionship to the John Day dogs. On the other hand, when we have learned more of the
Uinta dogs, it may appear that all the many resemblances of Daphenus to the Machai-
rodonts haye been separately attained ; but existing evidence does not favor this sug-
gestion either. It seems exceedingly likely that the dogs and cats are more closely
related than has hitherto been believed and that they were derived from a common mid-

dle or late Eocene progenitor.

NOTES ON THE CANID® OF THE WHITE RIVER OLIGOCENE. AQ9

On the assumption that the dogs and cats are thus quite closely connected, what can
be said concerning the relations of the other fissipede families with these groups and with
one another? Of the derivation of the Procyonide nothing is yet known; the family
may be traced back into the Loup Fork without finding essential changes, but beyond
that period we lose track of it altogether. The position of the bears and hyenas is rea-
sonably clear, the latter being late derivatives of the viverrines and the former of the
dogs, neither family making its appearance until long after the other fissipede groups
had become clearly differentiated. The Viverride have a great many characters in com-
mon with both the early dogs and the early Machairodonts ; almost all the structural
features which are found in both Daphenus and Dinictis recur also in the viverrines,
and the latter again have many points of similarity to Cynodictis, as has often been
remarked. That the viverrine features of Cynodictis are more numerous and apparent
than those of Daphenus is largely due to the small size of the former, which agrees
much better with the stature usual in the recent viverrines. The viverrines thus seem
to be derivatives of the same Eocene stock as that which gave rise to both the dogs and
the cats, though, perhaps, they are more nearly allied to the latter than to the former,
and apparently they have departed less from that primeyal fissipede stem than has either
of the other families. Aside from the peculiar character of the auditory bulla and the
reduced number of the molar teeth, such a genus as Viverra would seem to differ but
little from the hypothetical Eocene ancestor of all the fissipede families. The Justelide
represent a quite specialized branch of the fissipedes, but between its earlier and more
primitive members and the corresponding representatives of the viverrines are so many
structural resemblances that Schlosser does not hesitate to derive them from a common
stem. An interesting and significant example of this community of characters among
the early representatives of the different fissipede families is given by the os penis of
Cynodictis, which resembles that of the mustelines much more closely than that of the
modern dogs. This probably indicates that all of the earlier fissipedes had this bone
shaped very much as in the existing mustelines, which have thus retained the primitive
form, while in the other families it has become much modified in shape and size. This
would explain the apparent anomaly of the very large os penis of Cryptoprocta which is
so different from that of the other viverrines. According to this way of looking at the
subject, there was a middle Eocene group of flesh-eaters, perhaps the creodont family
Miacide, which rapidly diverged into four principal branches, the cats, dogs, viverrines
and mustelines, all of which families were established in the late Eocene or early Oligo-
cene, and to these should perhaps be added a fifth family, the Procyonide, though of this
we know nothing definite. The Fissipedia are thus probably a monophyletic rather
than a polyphyletic group, which was derived from a single creodont family.

410 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

It is exceedingly difficult to unravel all this complicated mesh-work of similarities
and definitely to distinguish those characters which are due to genetic relationship from
those which are merely phenomena of parallelism or convergence. But the important
fact remains that in the late Eocene and early Oligocene all of the families of fissipede
Carnivora which had then come into existence were very much alike and in all parts of
their structure resembled one another much more closely than do their modern repre-
sentatives. They are obyiously converging back to a common term, and the only ques-
tion is what that common term was and whether we are to look for it in the middle or
the lower Eocene. It must be reiterated, however, that natural and probable as this con-
clusion appears to be, it is only tentative and cannot be demonstrated until the successive
phylogenetic stages of each family are much better known than they are at present.

SUMMARY.

1. Daphenus, so named in 1853 by Leidy and afterwards referred to Amphicyon, is
very different from the latter and an entirely distinct genus.

2. The dental formula is: I 3, C41, P 4, M3; the premolars are small and sim-
ple and are set well apart in the jaws; the sectorials are small and primitive, especially
in ? D. Dodgei, and the molars relatively large, most so in D. vetus. ‘The dentition is
more like that of the creodont family Iacide than of the typical modern dogs.

3. The skull is of a very primitive character, with short face, very elongate cranium
and high sagittal crest; the cranial cavity is of small capacity and the postorbital con-
striction is placed far back of the eyes. Large frontal sinuses are present.

4. The occiput is low and broad, with very prominent crest; the paroccipital pro-
cesses are short and blunt and are widely separated from the tympanic bull.

5. The auditory bulla is minute and does not fill up the fossa, exposing the periotic ;
it probably represents only the anterior chamber, the posterior chamber was either not
ossified or was very loosely attached, so that it is lost in all the known specimens.

6. The cranial foramina differ very little from those of Canis.

7. The mandible has a short horizontal ramus, varying in its proportions in the
different species ; the ascending ramus is low and very broad.

8. The brain is remarkable for the small size and simple convolutions of the cerebral
hemispheres and the large size of the cerebellum and olfactory lobes.

9. The foramina of the atlas differ from those of: the recent dogs and resemble those
of the cats.

10. The axis is also of feline character, especially in the shape of the neural spine.

11. The other cervical vertebrxe have more prominent zygapophyses, narrower neu-

ral arches and higher neural spines than in Canis,

NOTES ON THE CANIDE OF THE WHITE RIVER OLIGOCENE. 411

12. The thoracic vertebre probably numbered thirteen ; they resemble those of the
modern dogs, except for their longer neural spines, and for the much more prominent
anapophyses on the last three vertebre. i

13. The lumbars, probably seven in number, are remarkably large and massive and
all their processes are very long; the appearance of these vertebrae is feline rather than
canine.

14. The sacrum is composed of three vertebre and resembles that of the larger cats
in its size and weight.

15. The tail is very long and stout, resembling in its proportions and in the deyel-
opment of the individual vertebree that of the leopard.

16. The humerus is in most respects like that of the Machairodonts, Dinictis and
Hoplophoneus, haying very prominent deltoid and supinator ridges, very low trochlea,
large epicondyles and an entepicondylar foramen.

17. The radius is very feline in character, as is seen in the discoidal head, the slen-
der curyed shaft and expanded distal end.

18. The ulna is much less reduced than in the modern dogs, and its shape, espe-
cially that of the distal end, is much more feline than canine.

19. The only carpal element preserved is the scapho-lunar which is very like that
of the Machairodont Hoplophoneus.

20. There are five metacarpals which are not at all like those of modern dogs, the
pollex being far longer and all of the metacarpals having short, slender, rounded shafts,
spheroidal distal trochlez, and a divergent instead of a parallel arrangement. The con-
tact of me. ii with the magnum and of me. iv with the unciform is much less than in
the true felines and about as in the Machairodonts.

21. The pelvis is machairodont rather than canine, the ilium béing relatively short
and narrow, the ischium long, with inconspicuous tuberosity, and the obturator foramen
large ; the pubic symphysis is elongate.

22. The femur is not very long in proportion to the size of the animal ; its troch-
lea is very low and shallow ; a third trochanter appears to have been present.

23. The patella is like that of Dinictis, being broad, thin and almond-shaped.

24. The tibia is short and slender and bears considerable resemblance to that of
Dinictis ; its distal end bears a very large internal malleolus and feebly grooved astra-
galar trochlea.

25. The fibula is much stouter than in Canis and has more thickened ends.

26. The tarsus is, on the whole, of machairodont or viverrine character, but with
not a few canine features.

27. The metatarsus has five members, a well-deyeloped hallux being present ; the

412 NOTES ON THE CANIDZ OF THE WHITE RIVER OLIGOCENE.

character of these is intermediate between those of the dogs and those of the Machairo-
donts.

28. The phalanges are long and depressed; the second one is excavated on the
fibular side, showing that the claws were partially retractile, though much less completely
so than in the cats; the unguals are straight, compressed and bluntly pointed, and with
bony hoods much as in Canis.

29. The known species of Daphenus are: D. vetus Leidy, D. hartshornianus Cope,
D. felinus, sp. noy., ? D. Dodger sp. noy., all from the White River beds, and D. cuspi-
gerus Cope, from the John Day.

30. The cynoid from the Uinta beds, Jfacis wintensis, is regarded as the forerunner
of Daphenus.

31. The small American cynoids of the White River and John Day, and, perhaps,
of the Uinta, should be referred to the European genus, Cynodictis.

32. The dental formula of Cynodictis is: I 3,C 4, P 4, M2; the premolars are
small, the sectorials microdont and quite viverrine in appearance, but more trenchant
than those of Daphenus, and the tubercuiar molars are small.

33. The skull has a very viverrine look; the face is short, the cranium long, though
shorter and fuller than in Daphenus, and the postorbital constriction is near the orbit ;
the sagittal crest is low and weak, and in the small C. lemur is replaced by a lyrate area.

34, There are no frontal sinuses.

30. The occiput is low and broad, the crest inconspicuous and the paroccipital pro-
cesses are small and not in contact with the bulle.

36. The auditory bulla is very large and the posterior chamber fully ossified.

37. The cranial foramina are like those of Canis, save for the visible carotid canal.

38. The mandible has a short, slender horizontal ramus and the ascending ramus is
much narrower than in Daphenus.

39. While the cerebral hemispheres are larger and better convoluted than those of
Daphenus, they are smaller and have fewer, straighter sulci than in the modern Canide ;
the olfactory lobes are large and the cerebellum complex.

40. The atlas has short transverse processes and its foramina are feline in character.

41. The axis is much like that of Viverra.

2. The other cervicals are of canine type.

43. The thoracic vertebre are small and have high, slender spines ; on the last two
are prominent anapophyses.

44, The lumbar region is long, heavy and arched upward ; it is composed of seven
yertebree, which have very long transverse processes and low, slender spines. Anapo-

physes are large anteriorly, but disappear on the sixth,

NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE. 413

45. The tail was very much as in such viverrines as [erpestes.

46. The sternum is of a generalized fissipede character, without special resemblance
to either dogs or viverrines.

47. The scapula has little resemblance to that of Canis, being low and broad, with
spine placed nearly in the middle of the blade; the metacromion is very large and the
acromion exceedingly long and prominent, from which it may be inferred that the clayi-
cles were less reduced than in the modern dogs; the coracoid is very large.

48. The humerus is much more viverrine than canine in appearance, having, like
Daphenus, very prominent deltoid and supinator ridges, a low trochlea and entepicon-
dylar foramen, but no supratrochlear perforation.

49. The radius is like that of Daphenus, except for the immense styloid process.

50. The ulna is much stouter than in the recent dogs and differs from that of
Daphenus in having the distal radial facet sessile.

51. The carpus contains a scapho-lunar which is quite like that of Canis ; the pyra-
midal is viverrine and the pisiform quite peculiar in shape ; a radial sesamoid appears to
have been present ; the trapezoid and magnum are canine, while the unciform is viverrine.

52. The metacarpus has five elements, which are very short and slender like those
of the civets.

53. The pelvis is, in general, canine, but primitive in the elongation of the post-
acetabular portion.

54. The os penis is very large and shaped like that of Cryptoprocta aud the muste-
lines.

55. The femur is elongate and differs little from that of the recent dogs, except
in the presence of a small third trochanter and in the narrow, shallow rotular trochlea.

56. The patella is wide, thin and scale-like, herpestine in shape.

57. The tibia is of nearly the same length as the femur, and its distal end is like
that of Daphenus and Dinictis, but more deeply grooved.

58. The fibula is relatively stout.

59. The general appearance of the pes is viverrine and has many resemblances
to that of Daphenus and some to that of Canis.

60. A well-developed hallux is present and the metatarsals exceed the metacarpals
in length much more than they do in Canis.

61. The phalanges differ materially from those of Daphenus in that the claws
are little or not at all retractile ; the unguals have but rudimentary hoods.

62. The skeleton of C. geismarianus was very herpestine in proportions, while that
of C. gregarius was more like that of a very small fox in which the hind leg much
exceeded the fore leg in length.

414 NOTES ON THE CANIDH OF THE WHITE RIVER OLIGOCENE.

63. The known American species of the genus are: C. gregarius Cope and C.
lippincottianus Cope (the latter doubtful) from the White River, and C. gregarius Cope,
C. geismarianus Cope, C. latidens Cope and C. lemur Cope, from the John Day.

64. The dogs are represented in the Uinta by two lines, ? Cynodictis and Miacis, the
former continued through the White River and John Day and the latter apparently
passing into Daphenus of the White River, and through this into Temnocyon, Hypo-
temnodon, Cynodesmus and Enhydrocyon of the John Day, Oligobunis of this formation
being probably an immigrant from the Old World.

65. M. Boule’s hypothesis that the alopecoids are derived from Cynodictis and the
thooids from Amphicyon implies an improbable degree of convergent development, but
it is not to be rejected as impossible. According to present evidence the alopecoids
arose relatively late from the thooid stem.

66. The thooid line appears to be Miacis—Daphenus— Cynodesmus—Canis, the re-
tractile claws of Daphenus having been changed when the digitigrade gait and cursorial
habit were assumed.

67. The very many resemblances between Daphenus, Cynodictis and Dinictis were
probably not independently acquired, but point to a common Eocene ancestor.

68. The early members of the canines, felines, mustelines and yiverrines all have a
great many more structural features in common than do their existing representatives
and would seem to converge to a single Eocene type, which may prove to be the
creodont family Miacide. The hyznas and bears belong to a later cycle of develop-

ment and were derived, the former from the viverrines and the latter from the dogs.

LITERATURE.

ApAms, G.I. 796. The Extinct Felide of North America. Amer. Jour. Sci., 4th Ser., Vol. I.

Boutz, M. ’89. Le Canis Megamastoides du Pliocene moyen de Perrier. Bull. Soc. Géologique de France, Tome
XVII.

Bruce, A. T. ’83. Observations upon the Brain Casts of Tertiary Mammals. Ball. Princeton Geol. Museum, No. 3.

Corr, E. D. 84. Tertiary Vertebrata, Washington, 1884.

EYERMAN, J. ’96. TheGenus Temnocyon, etc. American Geologist, Vol. XVII.

Frower, W. H. ’69. On the Value of the Characters of the Base of the Cranium in the Clas-ification of the Order
Carnivora, etc. Proc. Zod]. Soe., London, 1869.

Huxtety, T. H. ’80. On the Cranial and Dental Characters of the Canide. P. Z. S., 1880.

Leipy, J. °69. The Extinct Mammalian Fauna of Dakota and Nebraska, Philadelphia, 1869.

ScHLossER, M. ’88. Die Affen, Lemuren, etc., d. europ. Tertiairs. II. Th. Beitrige zur Paleontologie Oesterreich-
Ungarns, Bd. VII. 2

Scuiosser, M. 789. Ditto, III. Theil, cdéd., Bd. VIII.

Scorr, W. B. ’94. Mammalia of the Deep River Beds. Trans. Amer. Phil. Soe., Vol. XVII.

WortMAn, J. L. 794. Osteology of Patriofelis, ete. Bull. Amer. Mus. Nat. His., Vol. V.

NOTES ON THE CANID# OF THE WHITE RIVER OLIGOCENE. 415

EXPLANATION OF THE PLATES.

Plate XIX.

Fig. 1. Daphenus hartshornianus Cope. Side view of skull.

Fig. 2. oe ne RY Palate and teeth of a second specimen.

Fig. 3. “ ss Ge Occiput ; same specimen as Fig. 1.

Fig. 4. a ie Ss Basis cranii of same individual : ty., tympanic ; f., fossa behind bulla; c. f.,
condylar foramen.

Fig. 5. Daphenus hartshornianus Cope. Right lower jaw.

Fig. 6. Daphenus Dodget, sp. nov. Lower teeth, crown view.

Fig. 7. G a sc «Side view of right lower jaw.

Fig. 8. Daphenus vetus Leidy. Lumbar vertebra, from the side.

Fig. 9, fs “ a Anterior caudal vertebra from above ; same individual.

Fig. 10. a a is Posterior caudal vertebra from the side ; same individual.

Fig. 11. Cynodictis gregarius Cope. Side view of skull (lower canine broken away).

Fig. 12. p of ay Brain cast from the right side: olf., olfactory lobe; r#., rhinal sulcus; f.,
frontal bone, showing the absence of sinus.
Fig. 13. Cynodictis gregarius Cope. Atlas from above.

(All figures natural size.)

Plate XX.

Fig. 14. Daphenus vetus Leidy. Sacrum from above ; same specimen as Figs. 8, 9, 10.

Fig. 15. Daphenus felinus, sp. nov. Lower end of humerus, front view.

Fig. 16. a ge sc «« ~~ Proximal end of radius ; same individual.

Fig. 17. ef “ cc «* Metacarpals i-iv of left manus ; same specimen.

Fig. 18. Daphenus vetus Leidy. Right femur, front view; same specimen as Fig. 14.

Fig. 19. Daphenus hartshornianus Cope. Lower half of right tibia and fibula.

Fig. 20. a as «Distal ends of same.

Fig. 21. ss co cs Right pes ; same individual.

Fig. 21a. se ss Gr iii digit, from tibial side ; same individual.

Fig. 22. Daphenus vetus Leidy. Left caleaneum and astragalus ; same specimen as Fig. 14.
Fig. 23. Cynodictis gregarius Cope. Left manus, front view.

Fig. 24. ss ee Left pes, front view. (Specimens seen since this plate was drawn show that the

metatarsals should have been made considerably longer.)
(All figures natural size.)

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ARTICLE IX.

CONTRIBUTIONS TO A REVISION OF THE NORTH AMERICAN BEAVERS,
OTTERS AND FISHERS.
(Plates XXI-XXV.)

BY SAMUEL N. RHOADS.

Read before the American Philosophical Society, May 6, 1898.

An unusually fine series of the skins and skulls, with reliable data and measure-
ments, of the beavers, otters and fishers of the United States and Canada having lately
come into the custody of the writer, it is thought advisable to publish the results of a
study of the various nominal forms of these mammals and briefly discuss the nomencla-
ture involved. Owing to a lack of specimens from some regions whose faunal condi-
tions are known to produce in many other mammals well-recognized geographic varia-
tions, this paper must be considered rather as a contribution to the subject, and in no
sense a complete synopsis. The area covered by this study comprises solely that part of
North America north of Mexico, no attempt being made to discuss the relationships of
the tropical species.

To Mr. Outram Bangs the author acknowledges his gratitude for a most valuable
loan of skins and skulls of nearly every species and race recorded in these pages. To
the kindness of Mr. F. W. True, of the National Museum, is due the loan of a series of
skulls of the Alaskan otter.

The North Carolina Department of Agriculture has courteously loaned two skins
and four skulls of beavers recently killed in Stokes county of that State through the
kind offices of Mr. H. H. Brimley, the Curator of the State Museum.

Aid has likewise been generously given by Dr. J. A. Allen, Dr. C. Hart Merriam,
Dr. T. 8. Palmer, Mr. Gerrit S. Miller, Jr., Dr. M. W. Raub and Mr. C. 8. Brimley.

THE BEAVERS OF NORTH AMERICA.

Contrary to evidence which must eventually be accepted by all zodlogists, the Ameri-
can beaver, Castor canadensis Kuhl, is still considered by many eminent authorities as

418 CONTRIBUTIONS TO A REVISION OF THE

specifically the same as the Castor fiber Linnzeus of Europe. In 1897, Dr. E. A. Mearns
described* a subspecies of the typical Canadian animal, naming it Castor canadensis
frondator and assigning its habitat to the “southern interior area of North America,
ranging north from Mexico to Wyoming and Montana.” This appears to be the first
attempt in literature to formally subdivide the American beaver, a species whose con-
stancy of characters over the vast and varied habitat which it frequents had hitherto been
unquestioned. There can be no doubt as to the tenability of Dr. Mearns’ “ Broad-tailed
Beaver ” as distinguished from the Hudson bay animal, whose habitat Kuhl designated
as “ad fretum Hudsoni” in his original description of canadensis.

It is probable that the beavers inhabiting the Carolinas, Georgia, Alabama, Missis-
sippi and Tennessee are equally entitled to subspecific rank. So rare has the beaver
become in these States, however, it would probably be impossible to verify such a predic-
tion with specimens now in our museums.*f

From what we know of the relationships of the representatives of our eastern species
inhabiting the Pacifie slope, we are led to expect that the beaver of that region would
also prove separable from canadensis. A very complete series of skulls, with three adult
and three young skins from the Cascades of Washington and Oregon, shows this to be
the case.

Fortunately the synonymy of the American beayer is not inyolved and requires no
elucidation in this connection, as is shown by reference to Dr. J. A. Allen’s Monograph
of the North American Rodentia. A synopsis of the American forms is herewith pre-
sented.

CaNADIAN Beaver. Castor canadensis Kuhl.
Plate XXI; Fig. 3. Plate XXII; Fig. 3.

Castor canadensis Kuhl, Beitr. Zool., 1820, p. 64.

2“ Castor americanus F. Cuvier, Hist. des Mam. du Mus., 1825” (fide Brandt in Kennt.
Sdugt. Russl., 1855, p. 64).

Castor fiber americanus Richardson, Faun. Bor. Amer., 1, 1829, p. 105.

Castor fiber var. canadensis J. A. Allen, Monog. N. Amer. Rod., 1877, p. 444.

Type Locality. Hudson bay (“ad fretum Hudsoni” Kuhl).

Geographic Distribution.—Northeastern North America, from the northern limit of
trees south to the United States and west to the Cascade mountains ; intergrading east
of the Mississippi river into subspecies carolinensis, south-centrally into subspecies fron-
dator and westwardly into subspecies pacificus.

* Proc, Nat. Mus., Vol. XX (adv. sheet, March 5, 1897).
+ As will be seen later, such specimens have since come to hand and are described as Castor canadensis carolinensis.

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 419

Color.*—Winter pelage, above, including sides, dark bay or blackish brown, tip-
ped with chestnut or russet, becoming pure chestnut on top and sides of head and on
chin, jaws and sides of neck. Rump and thighs purer chestnut. Ears black. Hair of
feet, legs and under parts seal brown.

Anatomical Characters.—Size, smallest of the American forms. Scaly portion of
tail more than twice as long as wide; hind foot with claw about 175 mm. Skull wide
for its length ; maximum size of skull 136 by 99 mm. ina New Brunswick example, No.
31, collection of E. A. and O. Bangs. Rostrum and nasals relatively short and wide,
the nasal bones averaging more than half as wide as long and extending but little
behind the premaxillaries. Upper molar dentition wide and heavy, the crowns oblique,
triangular and very wide anteriorly.

Measurements.—Of a large, typical, adult male specimen from Quebec, No. 3825,
collection of E. A. and O. Bangs (measurements made by collector from newly killed
specimen). Total length, 1130 mm.; tail vertebre, 410 mm.; scaly portion of tail
(dry meas. from skin), 263 by 122 mm.; hind foot, 176 mm.; length of skull,
152 mm.; breadth of skull, 93 mm.; length of nasal bones, 46 mm.; breadth of nasals,
21.4 mm.t+

Remarks.—The above diagnosis is taken mainly from the Quebec specimen, because
of the authentic measurements and superior condition of the skin and pelt. The aver-
age beaver from the Hudson bay regions, however, is somewhat lighter colored than this
specimen, which, in its darkness and richness of shade, rivals the best examples of paci-
jicus. In size, and ratio of length to width, the skull of the Quebec specimen is typical,
but the nasals are too narrow to serve as a standard for canadensis, whose nasals average
wider than pacificus and narrower than frondator. In general terms, canadensis differs
from frondator in smaller size, narrower tail, much darker coloration and narrower nasals.
It differs from carolinensis in smaller size, narrower, longer nasals and somewhat darker
coloration. From pacificus it differs in smaller size, lighter coloration, wider nasals and
broader skull. Subspecies pacificus differs from frondator in larger size, greatly nar-
rowed and lengthened tail-paddle, rostrum and nasals, and in its dark coloration. In
color frondator is decisively and uniformly lighter than eastern canadensis and carolinen-
sis and western pacificus, but darkened canadensis (not melanistic) are nearly as dark as
pacificus. In size, pacificus is much the longest of the three, with very long hind foot
and tail. Its skeleton is slenderer and weaker in every part as compared with the massive
frame of canadensis and frondator of same age. Cuarolinensis is nearly of the color of

* Ridgway’s Womenclature of Colors is the standard used throughout this paper.

+ The narrow nasals of this specimen are an exception, the average of several east Canadian specimens showing the

ratio of length to breadth as less than two to one.

420 CONTRIBUTIONS TO A REVISION OF THE

lighter hued canadensis, but agrees with all the other characters of frondator, to which
it seems most nearly allied in cranial and caudal characters.

Specimens EHxamined.—New Brunswick, 1 skull; Quebec, 1 skin with skull ;
Canada (?), 3 skulls, 1 skeleton, 2 mounted skins; Ft. Simpson, N. W. T., 1 mounted
skin; Idaho, 1 skin with skull.

CaRoLiniAN Beaver. Castor canadensis carolinensis, subsp. nov.
Plate XXIII; Figs. 1 and 2.

Type Locality—Dan river, near Danbury, Stokes county, North Carolina. Type
No. 2.607, old ad. 3, in the collection of the North Carolina State Museum, Raleigh,
N.C. Collected by a trapper in flesh for the Museum, April, 1897.

Geographic Distribution.—Carolinian fauna, south into the Austroriparian.

Color.—Of type and topotype: Overhair of upper head, neck, back and sides,
bright hazel. Underfur of same parts, seal brown. Hinder back and rump lightening
from hazel to cinnamon rufous and then to tawny olive near base of tail. Vent and
under base of tail, dark, rich burnt umber. Ears pale blackish. Sides of head below
eyes light hair brown, shaded with pale cinnamon rufous. Feet bistre. Below, from
throat to vent, dark broccoli brown with wood-brown tips to overhair.

Anatomical Characters.—Size large, larger than canadensis, with relatively much
broader tail, as in frondator. :

Skull large and broad, with very short, broad nasals. In the type the base of
nasals does not reach back to the line connecting the anterior walls of the orbits. Ros-
trum very short and broad. Audital bulle remarkably contracted laterally, with a
strongly developed osseous column on the outer wall and the transverse diameter less
than the longitudinal. Incisors weak, narrowed; molars large, with triangular crowns.
Pelage short and harsh as compared with canadensis.

Measurements.—Ot the type, from carcass: Total length, 1130 mm.; scaly portion of
tail, 279 by-158 mm.; hind foot, 184 mm.; ear, from crown, 21 mm.; length of skull,
148 mm.; breadth of skull, 107 mm.; length of nasals, 43.5 mm.; breadth of nasals, 29
mm. Of the topotype (ad. 3’): Total length, 1080 mm. ; scaly portion of tail, 260 by 146
mm.; hind foot, 174 mm.; ear from crown, 23 mm.

Remarks.—The two skins and four skulls upon which the above diagnosis of caroli-
nensis is based were secured, just before the completion of this paper, from the authorities of
the State Museum of North Carolina. They are intended to form a group exhibit in the
State Museum, and have been carefully measured by the curator, Mr. H. H. Brimley,

while yet in the flesh, The old male which forms the type had lost one of its fore feet,

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 421

apparently in a trap, some years previous to its final capture, but its evident health and
great size show that it had suffered little inconvenience from the loss of the member.

The strong cranial and caudal affinities which this beaver shows to frondator as dis-
tinguished from canadensis indicate that it is more closely related to the western form.
In color, howeyer, it shows a nearer approach to canadensis, as, in fact, do many other
animals of similar distribution and racial differences. The Mississippi and Louisiana
beayers are undoubtedly, from what I can hear from the furriers, the darkest and thin-
nest pelted of our American beavers, but their separability from what I have named
carolinensis is not probable. They may be considered as belonging to carolinensis rather
than to frondator.

Specimens Hxamined.—Stokes county, North Carolina, 4.

Sonoran Beaver. Castor canadensis frondator Mearns.

Plate X XI; Fig. 2. Plate XXII; Fig. -2.
Castor canadensis frondator Mearns, Proc. U. S. Nat. Mus., XX, adv. sheet, Mar. 5, 1897.

Type Locality.—San Pedro river, Sonora, Mexico, near monument No. 98, of the
Mexican boundary line.

Geographic Distribution.—Southern interior of North America from Mexico to
Wyoming and Montana, intergrading northwardly into canadensis, southeastwardly into
the trans-Mississippian carolinensis and westwardly into pacificus.

Color.—Much paler than canadensis or carolinensis. “ Above russet, changing to
chocolate on the caudal peduncle above and to burnt sienna on the feet ; toes reddish
chocolate. Below grayish cinnamon, brightening to ferruginous on the under side of
caudal peduncle. Sides wood brown enlivened by the tawny-olive color of the over-
hair.”* A specimen from Red Lodge, Montana (No. 32, collection of E. A. and O.
Bangs), taken in November, is wood brown above and below, the longer overhair of
upper pelage washed with pale rusty.

Anatomical Characters.—Size large, exceeding average of Hudson bay beaver, with
a longer foot and broad tail. Scaly portion of tail less than twice as long as wide, hind
foot with claw about 185 mm. Skull massive, large, with short rostrum and very wide,
short, tumid nasal bones, the average skull probably exceeding canadensis in size, cer-
tainly exceeding it in relative width to length and in the relative breadth of the nasals.
Upper molar dentition as in canadensis.

Measurements.—Of the type: Total length, 1070 mm.; tail vertebrae from anus, 360
mm.; scaly portion of tail, 290 by 125 mm.; hind foot, 185 mm.; length of skull, 133

* Quoted from Dr. Mearns’ original description (/. c.) of type.

422 CONTRIBUTIONS TO A REVISION OF THE

mm.; breadth of skull, 99 mm. Maximum length of old males, measured by Dr.
Mearns, 1130 mm.; of the tail paddle, 285 by 155 mm.

Remarks. hye Mearns’ comparisons of frondator with canadensis were evidently
not made with the largest specimens of the latter, as I have examined some whose cra-
nial and body measurements are about equal to the maximum recorded by him for
frondator. Nevertheless, there is little doubt that the larger size of average frondator is
well established. Its long hind foot, broad tail and light coloration distinguish it
immediately from canadensis. Its approach to pacificus is solely along the line of great
size as indicated by the length of body and hind foot, but in cranial characters, as also
in color, it is farthest removed from that race. The close anatomical relation of frondator
to carolinensis has been mentioned.

Specimens Examined.—Montana, 1 skin with skull; Wyoming, 1 skull.

Paciric Beaver. Castor canadensis pacificus, subsp. noy.
Plate XX Mies Plate ®XOxcd ities

Type Locality.—Lake Kichelos, Kittitass county, Washington ; altitude about 8000
feet. Type, No. 1077, ad. 2, in the collection of S. N. Rhoads; collected in April,
1893, by Allan Rupert.

Geographic Distribution.—Pacifie slope, of America, from Alaska to California.

Color.—Above with very uniform, dark and glossy reddish chestnut overhair,
almost concealing along dorsum the seal-brown underfur. Top of head like back ; sides
of head, throat, rump, thighs and vent not decidedly lighter than back and belly as in
the other forms, these parts paling to walnut brown. Ovyerhair of sides and under parts,
between seal brown and broccoli brown ; under fur of belly drab gray at the roots ; hind
feet dark seal brown ; fore feet and limbs, dark wood brown. Ears black.

Anatomical Characters.—Size, largest of the canadensis group, but of more slender
build, the skeleton throughout being of much greater longitudinal and lesser lateral
dimensions than in the other forms. Tail and hind foot relatively long. Skull large,
relatively narrow, with long, narrow rostrum and nasals, the latter with outer margins
nearly parallel and reaching basally decidedly beyond the premaxillaries. Upper molar
dentition weak, the crowns of molar teeth rectangular.

Measurements.—Of the type from carcass: Total length, 1143 mm.; tail vertebre,
330 mm.; (from relaxed skin) scaly portion of tail, 295 mm. by 122 mm.; hind foot, 185
mm.; rene th of skull, 142 mm.; breadth of skull, 101 mm.; length of nasals, 53.6 mm.;
breadth of nasals, 24 mm.; average length and breadth of five skulls from Tacoma and
Lake Kichelos, Washington, 144 mm. by 99 mm.; average nasal length and breadth of
same, 04 mm. by 23 mm.

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 423

Remarks.

Reliable measurements of only one adult skin specimen (the type) of
pacificus were accessible. An adult mounted specimen from Josephine county, Oregon,
in the Wagner Institute, Philadelphia, confirms the color and measurements of the type
so far as the latter can be ascertained from the stuffed animal.

Pacificus, like its associates, Mustela americana caurina and M. canadensis pacifica
of the Pacific slope regions, is distinguishable by its rich and deep coloration from its
darkest trans-Cascadian representatives. No specimens have come to hand from Alaska,
but undoubtedly, from what we know of other species found there as well as from the
accounts of trappers and furriers, the Alaskan coast beaver represents the maximum of
size* and the greatest richness and depth of fur coloration seen in American beavers.

Specimens Examined.—Washington, Tacoma, 1 skeleton, 1 skull; Lake Kichelos,
1 adult skin with skull, 3 young skins with skulls, 1 skeleton, 12 separate skulls ; Ore-

gon, Josephine county, 2 mounted specimens ; British Columbia, (2) Sumas, 1 skull; +
Victoria, 1 skull.

THE OTTERS OF NORTH AMERICA.

As Mr. Oldfield Thomas has shown in his “ Preliminary Notes on the Species of
Otter,” published in 1889 in the Proceedings of the London Zoblogical Society, the charac-
ters and nomenclature of the North American species are in great need of study. Dr.
Elliot Coues has elucidated with sufficient clearness, in his Monograph of the Mustelide,
the habits and characters, and, to some extent, the synonymy of the typical Canadian
otter, Lutra hudsonica Lacépéde. Its relations, however, to other nominal species,
especially to the otters of the Pacific slope of America from California northward,
demand investigation.

As in the case of the American beaver, just treated, this paper has to do solely with
one central Canadian type and its subspecies found in America north of Mexican terri-
tory.

Avoiding a general preliminary discussion of the rather perplexing questions of
nomenclature and geographic variations and distribution, I will present these in order in
the more formal and detailed synopses which follow.

* Dr. Allen’s measurements of Alaskan skulls, page 447 of the Monograph of N. A. Rodentia, do not indicate

unusual size, but as we have no precise locality given they may not have come from the coast region, and, therefore, do not
. Tepresent pacificus.

{ Thisskull (No. 5545, 3, coll. of E. A. and O. Bangs) is the largest of which I find any record, measuring 154 by
108mm. The next in sizeis No. 2146, U.S. Nat. Mus., from Nebraska, recorded by Baird. Its size was 147 by 105.5
mm. Unlike all my pacificus specimens, No. 5545 has very wide convex nasals.

A. P. S.—VOL. XIX. 3B.

494 CONTRIBUTIONS TO A REVISION OF THE

Hupsonran Orrer. Lutra ‘hudsonica (“ Lacépéde,” Desmarest).

Plate XXIV; Figs. 1 and 2.

Mustela lutra Linn., canadensis Schreber, Stugt., III, Pl. CX XVI, B. (dated 1778 on
title-page, but, according to Sherborn, the text of Vol. III was published in 1777
and this plate in 1776).

Mustela (lutra) canadensis Kerr, Linn. An. Kingd., 1, 1792, p. 173 (see Thomas, Proc.
Zool. Soc. Lond., 1889, p. 197, and Allen, Bull. Amer. Mus. N. Hist., VII, 1895,
p- 188).

“ Mustela hudsonica Lacép.[éde],” Desmarest, Nouv. Dict. d’ Hist. Nat., XIII, 1803, p.
384 ; (Wow. Hd.) 1817, p. 219.

Lutra canadensis J. Sabine, App. Frankl. Jour., 1823, p. 653, and of nearly all subse-
quent authors (not L. canadensis F. Cuvier, Dict. Sci. Nat., 1823, p. 242; see O.
Athomasse/ ace spall):

Lutra hudsonica F. Cuvier, Suppl. Buff., 1, 1831, p. 194; Merriam, NV. Amer. Fauna,
No. 5, 1891, p. 82.

Lataxina mollis Gray, List Mamm. Brit. Mus., 1843, p. 70.

Lutra destructor Barnston, Canad. Nat. and Geolog., VIII, 1863, p. 147, F igs. 1 to 6.

Type Locality.—“ Ou la trouve au Canada sur les bords de la mer.”

Geographic Distribution—Northern North America from the Arctic ocean south-
ward into the United States and from the Atlantic ocean to the Cascade mountains ;
intergrading southeastwardly into subspecies /ataxina F. Cuvier and vaga Bangs, south-
centrally into subspecies soronw Rhoads, and westwardly into subspecies pacifica Rhoads.*

Color (taken from two specimens in the Bangs collection, No. 5638, yg. ad. 3,
Annapolis, Nova Scotia, November 23, 1896, and No. 4190, ad. 2, Upton, Me., Octo-
ber 25, 1895).—Above, dark seal brown from nose to tip of tail, darkest posteriorly,
below from breast to tail between broccoli and vandyke brown in the Nova Scotia speci-
men and between seal and yandyke brown in the Maine specimen. Head and neck
below a line running from nose to lower base of ear and base of foreleg light Isabella
color anteriorly darkening on lower neck to wood brown in the Nova Scotia animal. In
the Maine specimen the neck is Prout’s brown. Feet, legs and tail corresponding to
darker shades of upper and lower body. A summer specimen from New Brunswick
is dark, vandyke brown, but little paler below than on back, and darker than winter

specimens of /ataxina from Maryland.
Ee .
* The otters of Louisiana and Mississippi are stated by furriers to be very dark and light-pelted, resembling South

Florida and Gulf-coast skins. No specimens having been examined, they are referred to vaga.

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 425

Anatomical Characters.*—Size, medium (exceeded by vaga, sonora and pacifica).
Tail relatively short. Inferior webs of feet and interspace between posterior and ante-
rior callosities of manus, densely haired. Hind foot with claw about 125 mm. in old
adults ; but so variable as to have little diagnostic value. Total length rarely exceeding
1100 mm. Skull—size, medium (greatly exceeded by vaga and pacifica). Teeth large,
crowded longitudinally upon each other and obliquely overlapping. Postorbital neck
of frontals relatively short and wide, its superior ridge on a plane with nasals and occi-
pital crest. Mastoid width much less than zygomatic width. Postorbital processes short
and stout. Audital bulle large, tumid, rising abruptly from the sides of basioccipital.

Measurements.—See tables.

Remarks.—V ariations in the size of adult otters from apparently the same region
seem remarkable at first sight, but I find that these are not always to be attributed to sex
(for the female otter sometimes reaches near to the average size of the males), but to
environment. The otters of the Alleghany mountain streams are uniformly smaller
than those of the tide-water creeks and rivers of the Atlantic seaboard. This rule
applies from Labrador to Florida and is undoubtedly the result of the relative difficulty
of obtaining food and securing shelter from enemies in the two kinds of habitat. On
the other hand, this difference lies wholly within the limitations of individual variation
and in no sense affects the well-defined cranial and other characters which distinguish
the races and species hereafter defined. It has to do solely with size, not with propor-
tions. In a letter from Mr. C. 8. Brimley, of Raleigh, North Carolina, the same feature
is alluded to where he states: ‘‘ A trapper of our acquaintance says that otters from the
saltmarshes of eastern North Carolina average considerably larger than the otters of the
small streams of the central part of the State.”

There is rarely to be found a case in mammalian nomenclature more puzzling than
that of the first tenable name of the Hudsonian otter. Its synonymy inyolves that of
the mink and the fisher as well as the questions of priority of publication of Erxleben’s
and Schreber’s great works on the Mammalia, and the tenability of plate names. I have
consulted Drs. C. H. Merriam and T. 8. Palmer at length on these questions and have
accepted their ruling as to the first tenable name of the Hudsonian otter being Lutra
hudsonica Lacépéde and that of the northeastern mink to be Putorius vison Schreber.
In regard to the name of the fisher, however, I prefer to abide by Canon XLIII of the
Code of the American Ornithologists’ Union, which accepts, under certain conditions,
the names of species originally published on plates, which Drs. Merriam and Palmer
and Mr. Sherborn do not accept. Returning now to the abstract of synonymy as given
above for the Hudsonian otter, the case may be concisely stated thus: Mustela lutra

* The diagnostic value of the nose pad has no significance in this study of the relationships of a monotypic group,

426 CONTRIBUTIONS TO A REVISION OF THE

canadensis Schreber is a plate name published (jide Sherborn) in 1776, and is the ear-
liest applied to this otter. It would stand (A. O. U., Canon XLII) were it not unques-
tionably applied and intended by Schreber merely as a geographic name without refer-
ence to its specific relations to “ M/ustela lutra Linn.” For this reason alone it should be
discarded. Furthermore, the name Mustela canadensis was used by Schreber on a pre-
vious plate in the same volume (Pl. No. 126) in the specific sense for the fisher. This
plate was also (fide Sherborn) published in 1776, one year before the text, which was
published in 1777, and the bound volume of text and plates were dated 1778. In 1777,
Erxleben published a description of the fisher and named it Mustela pennanti, by which
name it has been since designated by authors generally. As this name is antedated by
the tenable plate-name Mustela canadensis of Schreber by one year, I adopt it as the
name of the fisher of Pennant from the northeastern United States. Erxleben pub-
lished in the same work a description of an animal which he named Mustela canadensis,
and which Baird and Coues have considered applicable to the mink, and the accept-
ance of the dates on the title-pages of Schreber’s (1778) and Erxleben’s (1777) works
would give priority to Erxleben’s name and displace Mustela vison of Schreber. But
Sherborn’s emendation of these dates makes IM. canadensis of Erxleben for the mink
untenable, it being preoccupied by Schreber’s plate-name JL. canadensis for the fisher,
as stated above. Besides this fact, Dr. Merriam considers that Erxleben’s description of
M. canadensis also applies to the fisher and the marten in such a way as to make it
untenable for any species.

Returning to the search for a first name for the otter, we find Kerr’s name, J. cana-
densis of 1792, to be unavailable because he placed it under the old genus Mustela. Next
in order appears to be the name hudsonica, which is accredited to Lacépede, in an article
on the Canadian otter in the first edition of the Nouvelle Dictionaire d Histoire Natur-
elles, which is signed “Desm.” Ihave not examined this reference personally, but am
indebted to Dr. J. A. Allen for a transcript of these facts from the only known copy of
the work in America which appears to be available, belonging to the library of the
American Museum of Natural History. In agreement with my previous rendering of
manuscript names, and on the supposition that Desmarest was the real author and pub-
lisher of this name and description of hudsonica, I cite it as Lutra hudsonica (“ Lacé-
pede,” Desmarest). I agree with Dr. Merriam that this name should stand for the otter
of eastern Canada. Frederick Cuvier seems to have been the first to place this animal
in the genus Lutra under the Lacépéde-Desmarest name hudsonica in 1831.

The Lataxina mollis of Gray and the Lutra destructor of Barnston are no doubt
synonyms of hudsonica.

Specimens Examined.—Labrador, Okak, 1 skull ; Grand river, 1 skull ; New

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 427

Brunswick, Restigouche river, 1 skin; Nova Scotia, Annapolis, 1 skin with skull;
Maine, Upton, 1 skin with skull; Bucksport, 1 skull; Massachusetts, Kingston, 1 skin
with skull; Westford, 1 skull; Canton, 1 skull; Missouri, 1 skull; British Columbia,
Vernon, 1 skull; Alaska, Tanana river, 1 skull.

CarouintaANn Orter. Lutra hudsonica lataxina (F. Cuvier).
Plate XXIV; Fig. 4.
Lutra lataxina F. Cuvier, Dict. des Sci. Nat., 1823, p. 242.

Type Locality.—South Carolina.

Geographic Distribution.—Carolinian faunal region, intergrading through the Tran-
sition region northward with hudsonica and southward through the Austrariparian into
vaga of southern Florida.

Color.—Much lighter than hudsonica. Above (from a specimen taken at Liberty
Hill, Conn., No. 4252, ad. 3, Nov. 19, 1895, collection of E. A. and O. Bangs*), dark
vandyke brown, tipped on upper head, neck and shoulders with wood brown, darkening
posteriorly. Upper feet and limbs dark bistre. Below, from lower breast to end of tail,
between Prout’s brown and broccoli brown. Head, neck and breast, including ears,
below a line connecting nose, upper eyelid, upper ear and upper base of fore leg, grayish
wood brown, lightest on head, darkening posteriorly to color (é. c.) of breast. The ayer-
age Carolinian winter specimens from Maryland southward are somewhat lighter and
some are Prout’s brown above, the wood brown of lower head and neck becoming a pale
grayish buff.

Anatomical Characters.—Size, smallest of the hudsonica subspecies. Inferior webs
of feet and interspace between callosities of manus, sparsely haired. Hind foot with
claw about 120 mm. Total length rarely exceeding 1100 mm. Skull relatively small,
with very large teeth, and weak postorbital processes. In other respects like the hud-
sonica type.

Measurements.—See tables.

Remarks.—The relations of this subspecies to northern Audsonica on the one hand
and to the southern vaga on the other are rather peculiar. It is without question a
nearer ally to hudsonica than vaga in the territory between Connecticut and South Caro-
lina, but, as Mr. Bangs has implied in his remarks on vaga, there is a tendency in the
Georgia (and we may infer in the South Carolina) otter to the large size and peculiar

* This specimen comes from the northern edge of the Carolinian region. No equally good skins from more southerao

localities being available, it is used as typical of the Carolinian race. It corresponds closely to two fine 1897-8 winter
pelts of Maryland otters, examined through the courtesy of Mr. 8. E. Shoyer, of Philadelphia.

428 CONTRIBUTIONS TO A REVISION OF THE

skull and color characters of the south Florida animal. There is so much evidence of
the intergradation of /ataxina both north and south that the specific separation of vaga
from it is not permissible. On the other hand it is impossible to ignore the decided
racial differences of the Carolinian otter from the Hudsonian type.

Cuvier’s original description of /atawina gives “Caroline du Sud” as the locality
where the type was taken ; it is, therefore, permissible to restrict this name to the Caro-
linian form as typified in the otters found in the Carolinian lowlands of the eastern
United States from south of the ‘“ Transition Zone” of Dr. C. Hart Merriam, as far
as middle South Carolina, Alabama and Mississippi, where it merges into vaga of the
Gulf or southern “ Austroriparian Realm ” of Dr. J. A. Allen.

I know of no restricted synonyms of lataxina. Dr. Coues quotes in his Hur-bear-
ing Animals a“ Latax lataxina Gray, Ann. Mag. N. H., 1, 1837, p. 119.” The work
referred to contains no such name. Cuvier’s description of Jataxina gives its color as
“dark blackish brown, a little paler beneath. Cheeks, temples, lips, chin and throat
pale brownish gray, and under side of tail grayish brown, the hair tips reddish.” He
compares the skull of datawina with his Lutra enudris, “ Loutre de Guiane ” of the pre-
ceding page and remarks on the “straight line, even concave or depressed,” joining the
nasals and occiput. This is significant, as one of the peculiarities separating vaga from
lataxina and hudsonica is the convexity of the frontal plane in the former.

Specimens Hxamined.—Connecticut, Liberty Hill, 1 skin with skull; Pennsylva-
nia, Clinton county, 2 mounted specimens; Monroe county, 3 skulls; New Jersey,
Tuckerton, 1 skull; Mickleton, 2 disarticulated skeletons ; Maryland, 2 fresh cased
winter furs; North Carolina, Raleigh, 2 skulls. ;

Fiorina Orrer. Lutra hudsonica vaga Bangs.

Plate XXV; Fig. 2.
Lutra hudsonica vaga Bangs, Proc. Bos. Soc. Nat. Hist., XX VIII, 1898, p. 224.

Type Locality.—Micco, Brevard county, Florida.

Geographic Distribution.—Florida, southeastern Georgia and the Gulf regions of
Alabama, Mississippi and Louisiana, intergrading (?) northwardly into dataxina.

Color.—Dark ; less black than hudsonica, darker and redder than /atazina. Breast
and belly nearly unicolor with back. Paler area of head and neck, scarcely reaching
breast. Above and below, dark, rich chestnut, scarcely paler on belly. Lower head and
anterior throat below line from nose to and behind ears, strongly tipped anteriorly with
tawny Isabella color darkening to raw umber on throat, the underfur darker than over-

fur, instead of lighter as in /atawina.

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 429

Anatomical Characters.—Size, large. Tail relatively long (fide Bangs). Inferior
webs of feet and interspace of palms nearly naked. Hind foot with claw reaching
maximum (No. 4998 Bangs Coll., yg. ad. #, Citronelle, Florida) of 1830 mm. Total
length (maximum of No. 4998, /. ¢., 1285 mm.) exceeding 1200 mm. Skull large,
teeth relatively small, not crowded longitudinally. Postorbital neck of frontals long and
narrow, suddenly constricted at base. Frontal plane strongly upraised above a line con-
necting occipital crest with base of nasals and above the level of postorbital processes.
Mastoid width nearly equaling the zygomatic width in very old specimens, in young adult
skulls the mastoid width is the greater. Wings of mastoid processes strongly developed
and flattened laterally. Audital bullee as in hudsonica and lataxina ; well developed,
tumid at basioccipital margins. Postorbital processes relatively weak and _ slender.
Underfur short, sparse.

Measurements.—See tables.

Remarks.—This subspecies just described by Mr. Bangs in his most valuable

paper on Florida and Georgia mammals is, as already noticed, quite different from
lataxina, its nearest geographic ally. In color it comes nearer hudsonica intermediates
from New England. In size and color and lack of hair on the webs and palms it shows
approach to the remote pacifica, but its peculiar long-waisted and broad-based skull dis-
tinguishes it from all other American forms except, perhaps, those of the northern Cen-
tral American and South American otters which I have examined. The yellowish
and reddish shades of south Florida vaga suggest affinity with what we find published of
the characters of the otters of the Caribbean coasts. In essential respects Mr. Bangs’
diagnosis of this animal is very good. He, however, used the skull of a young adult
male for cranial comparisons, and while it is true that the ratio of the mastoid to the
zygomatic width is much greater in vaga than hudsonica it is not as great as would
appear by Mr. Bangs’ figure. In crania of old adult vaga in my collection the mastoid
and zygomatic widths are about equal, the latter slightly wider. In hudsonica, however,
the excess of zygomatic width and slight development of the mastoid wings is marked.

Specimens Examined.—Florida, Tarpon Springs, 1 adult pelt, 3 young skins with
skulls and 2 extra skulls; Salt Run, St. John’s river, 1 skull.

Pactric Orrer. Lutra hudsonica pacifica, subsp. nov.
Plate XXIV; Fig. 3. Plate XXV; Figs. 1 and 3.

Lutra paranensis and aterrima Thomas, P. Z.S.,l.c., p. 199; Trouessart, Catal. Mamm.,
1897, pp. 286, 287 (not of Pallas, Zoogr. Ross. Asiat., 1811, p. 81).

LIutra californica Baird, Mamm. N. Amer., 1857, p. 187 (not of Gray, Mag. Nat. Hist.,
I, 1855, p. 580, which is L. felina ; see Thomas, /. ¢., p. 198).

430. CONTRIBUTIONS TO A REVISION OF THE

Type Locality.—Vake Kichelos, Kittitass county, Washington ; altitude about 8000
feet. Type No. 616, yg. ad. 3, in the collection of 8. N. Rhoads; collected in fall or
winter® of 1892—93, by Allan Rupert.

Geographic Distribution.—Pacitic slope of North America, from Alaska to Cali-
fornia.

Color.—Of type: Lighter than hudsonica, with a browner cast, approaching nearly
to lataaina. Average of coast specimens from Puget Sound northward, ruddy seal
brown, sometimes very dark in Alaskan coast specimens. Lower parts from breast to
end of tail much lighter (Mars-brown) than back. Ventral region conspicuously lighter.
Lower head, neck and breast very pale wood brown, almost dirty gray.

Anatomical Characters.—Size, very large.t Tail normal. Inferior webs of feet
and palmar interspaces nearly naked. Hind foot not recorded in type, the caleaneum
missing ; no measurements of other specimens available. Skull largest of the North
American otters (reaching a maximum of 119 mm. in occipito-nasal length and 83 mm.
in zygomatic expanse in an Alaskan coast example) ; teeth relatively weak, less crowded
longitudinally than in hudsonica. Interorbital width relatively very great, nearly 13
times postorbital constriction ; postorbital processes long and stout. Mastoid and zygo-
matic proportions as in hudsomca. Audital bulle remarkably flattened.

Measuwrements.—See tables.

Remarks.—The type specimen, though taken in the mountains and not fully mature,
is large and has a skull which would have, perhaps, eventually equaled the maximum
size recorded above for an Alaskan specimen of much greater age. A very old female
skull from the vicinity of Puget Sound confirms fully the diagnostic characters of
pacifica as given.

In treating of the otters of the Pacific slope of America we are confronted with
two nominal species to which they have been doubtfully referred by authors. In point
of time the first to be considered is the Viverra aterrima of Pallas,{ described from a
hunter’s skin, lacking skull and feet, taken in northeast Siberia, “ between the Uth
and Amur riyers.” Schrenck and Middendorff listed this animal in their works on
Siberian Zodlogy with the remark that they were unable to verify its existence or clear
up the mystery of its strange characters as given by Pallas. Mr. Thomas (P. Z S.,
l. c., p. 199) queries, on the basis of a mistaken suggestion of Dr. Coues, whether it may

* The season of capture was not recorded, but the pelt indicates that it was taken in full winter fur.

{Ihave no measurements of Alaskan otters, but judging by the great size of the skulls from there they. must
greatly exceed any known species of Lutra. On the basis of the skull they must attain a maximum length of over 1400
millimeters.

t Zoog. Rosso. Asiat., I. ¢.

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 431

not prove to be the same as the so-called Lutra paranensis Rengg. which he assumed
might occur throughout the whole Pacific coast regions of America. The close relation-
ship of our Pacific coast otters to hudsonica will effectually remove them from any com-
plication with paranensis, but as regards aterrima we must devote sufficient space to show
the impossibility of referring the Alaskan land otter to that animal, as Trouessart. has
lately done.*

A eareful study of Pallas’ original description, together with the fact that no later
author or explorer has been able to explain or rediscover the animal, convinces me that
it is either unidentifiable or will prove not to belong to the Lutrine but to the Musteline.
Pallas states it to be intermediate in size between the European otter and the European
mink. He states the length of the skin to be 19 inches, 3 lines, and of the tail 5 inches
with a brush of 12 inches! The color of the animal is said to be very black and shin-
ing, except the sides of the head between the eyes and ears, which change from black to
“subrufescent.” The absurdity of applying such a description to the animal which I
have named pacifica, or, indeed, to any member of the genus Lutra, is certainly evident.
So far as any animal now known to zodlogists is concerned, the Viverra aterrima of Pallas
should be consigned to oblivion.

Another name which has given trouble to those who had to deal with the Pacific
coast otter is the Lutra californica of Gray. Fortunately, Mr. Thomas has effectually
exposed the history and at the same time the inapplicability of that name to a North
American animal of the hudsonica type. He has shown in his paper in the Proceedings

of the Zoological Society (1. c., p. 198) that Gray’s type of californica did not come from
California, but most likely from Patagonia, in which case he makes it a synonym of
Lutra felina Molina.
Specimens Examined.—Washington, near Tacoma, 3 skulls ; Lake Kichelos, 1 skin
with skull, 1 skull; Oregon, 1 skull; British Columbia, Sumas, 1 skull; Alaska
(coast?), 3 skulls; Kodiak Island, 2 skulls; Mission, 1 skull; Queraquinat} Island, 1
skull.

Sonoran Orrer. Lutra hudsonica sonora, subsp. noy.

LIutra canadensis Mearns, Bull. Am. Mus. Nat. Hist., 111, 1891, pp. 253-256.

Type Locality.—Montezuma Well, Beaver creek, Yavapai county, Arizona. Type,
ad. 9, No. 3212 in the collection of the American Museum of Natural History. Col-
lected December 26, 1886, by Dr. Edgar A. Mearns.

* Catalogus Mammalium, 1. ¢.
+ It is conjectured that this skull came from the North Pacific. It has Capt. T. J. Turner’s name on it. I cannot

find an island of this name on the maps.

A. P. S.—VOL. XIX. 3.

452 CONTRIBUTIONS TO A REVISION OF THE

Geographic Distribution.—Arid southern interior of North America, from Mexico,
probably to Wyoming.

Color.—Of type, fide Mearns, J. c.: “ Above dark brown, without reddish tinge ; this
color changing gradually to a light grayish brown below, being palest (almost whitish)
upon the sides of the head below the level of the eyes and upon the under side of the
head and neck as far back as the fore limbs. . . . . The long hairs of the lighter por-
tions of the body are pointed with yellowish gray and upon the upper surface of the
head and neck the tips of the hairs are yellowish brown, giving a paler cast to that part
of the dorsum.”

Anatomical Characters.—Size, large, with a very long hind foot, the body length
measurements exceeding those of any other specimen of North American otter exam-
ined or recorded.* Webs of feet not densely haired beneath. Hind foot, 145 mm. Total
length reaching 1300 mm. Skull—size, large, nearly as great as in largest Alaskan
pacifica, but small for the great relative length of body, “less massive, broader, with
more evenly rounded zygomatic arches and with the brain case more convex or bulging
in its outlines.” ‘‘ Arizona skulls differ from all others in the slender, attenuated postor-
bital processes and in the greater height of the lower jaw from angle to condyle, or to
summit of coronoid process. From its geographically near neighbor, L. felina of Cen-
tral America, it presents many cranial and dental differences; in fact, skulls of the lat-
ter are so very distinct [in their inferior concavity, frontal depression, short muzzle,
narrow postorbital constriction and absence of the heel in front of the antero-internal
cusp of the last upper molar] from any known specimens from North America, north of
Mexico, as to be distinguishable from them at a glance.”

Measurements.—Of type: “Total length, 1300 mm.; head and body (measured from
tip of nose to anus), 815 mm.; tail measured from anus to end of vertebrae, 472 mm.

. ear, height above crown, 15 mm.” No skull measurements given.

Remarks.—I have accepted Dr. Mearns’ very full and satisfactory diagnosis of the
Arizona otter, given in the Bulletin of the American Museum of Natural History, as
conclusive eyidence of the existence of a recognizable race in arid interior America,
south of Montana. Its great size and light color together form a combination not found
in any other known or named otter.

It has been thought unnecessary to examine the type, as, owing to the author’s
remoyal from Philadelphia during the completion of this paper, such an examination
would have caused a greater risk to the type specimens than the facts warranted.

* The great size of the type, as compared with an adult male also recorded by Dr. Mearns from Arizona, indicates
that the sex of the type may have been wrongly determined. If correct, the size to be expected of a full-grown male

sonora would be extraordinary.

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 433
Newrounpianpd Orrer. Lutra degener Bangs.
Plate XXIV; Fig. 5.
Lutra degener Bangs, Proc. Biol. Soc. Wash., XII, 1898, p. 35.

Type Locality —Bay St. George, Newfoundland.

Geographic Distribution.—Confined to Newfoundland (?).

Color —Of type, ad. g, taken April 22, 1897: Above, black with seal brown reflec-
tions. ars, seal brown. Lower head and neck areas grayish wood brown, becoming
seal brown on breast; the remainder of lower parts nearly as dark as back. Tail uni-
color. Feet seal brown and densely haired on under side of webs and palmar interspaces.

Anatomical C haracters.—Size, much smaller than any of the hudsonica group.
Hind foot small, with ¢law averaging about 112 mm.* long in the two specimens exam-
ined. Total length about 1000mm. Tail relatively short. Skull very small, narrowed,
weak and fragile; the brain case wide anteriorly ; the frontal and interorbital widths
narrow and the postorbital processes weak and slender, strongly grooved on their supe-
rior face. Sagittal crest not developed even in old specimens. Interorbital constric-
tion about equal to postorbital constriction. Teeth weak, with normal cuspidation.
Audital bulle normal. ;

Measurements.—See tables.

Remarks.—The type specimens of degener, so generously loaned to me by Mr.
Bangs, when compared with the large series used in the preparation of this paper, con-
vince me that this depauperate insular form has no intercourse with the larger typical
hudsonica of Labrador and New Brunswick. A skull from Grand river, Labrador, shows
no approach to the degener type, and another from Okak, Labrador, agrees in the same
differences. A young adult skull and skin of hudsonica from Nova Scotia, and an adult
summer skin from New Brunswick, show that the maritime otter of the mainland some-
times attains a size nearly one-third larger than the largest known specimens of old,
-adult degener.

Specimens HKxamined.—Newfoundland, Bay St. George, 2 skins with skulls, 1 extra
skull.
THE FISHERS OF NORTH AMERICA.

Apology must be made for the inferior series of skins and skulls which form the
basis of the subjoined remarks on the Pekan. They serve, however, to elucidate some
* The collector’s measurement of the hind foot of type is given on label as ‘(126 mm.’’ This is certainly incorrect,

as the length determinable by feeling the caleaneum in the dry skin could not haye exceeded 115 mm, This accords with
the small size of the hind foot and the length of other specimens of degener.

A3B4 CONTRIBUTIONS TO A REVISION OF THE

questions sure to be soon brought up in the active advance of monographic work in
American mammalogy.

The synonymy of Pennant’s Fisher has already been discussed under Lutra hud-
sonica, and I have there given reasons for my adoption of the plate-name canadensis of
Schreber as having priority over the long-accepted name pennanti of Erxleben for
this animal.

Pennant’s Fisner. Justela canadensis Schreber.

Mustela canadensis Schreber, Saugt., HI, p. 492, Pl. CXXIV. Text published in
1777, plate in 1776 (fide Sherborn).

Mustela pennant Erxleben, Syst. An., 1777, p. 470.

Mustela melanorhyncha Boddaert, lene: An., 1784, p. 88.

Viverra piscator Shaw, Gen. Zodl., 1, 1800, p. 414.

Mustela nigra Turton, ed. Linn. Syst. Nat., 1, 1802, p. 60.

Mustela godmani Fischer, Syn. Mamm., 1829, p. 217.

Type Locality —< New York and Pennsylvania,” Pennant.

Geographic Distribution—Northern North America, east of the Cascade moun-
tains, from the northern limit of trees to Colorado and North Carolina in the mountains.
Intergrading on the Pacific slope into subspecies pacifica, and probably in the southern
Rocky mountain region into a paler race. Probably represented in the Hudsonian
faunal region by a subspecies.*

Color—From an adult, male, winter specimen He near Lancaster, Pa., March
11, 1896, and in the possession of Dr. M. W. Raub, of that city, who furnished
the description : ‘‘ Head and one-half of the length of body, gray and black mixed, gray
predominating ; throat darkest, with snout from tip to line of eyes dark brown. The
hinder half of body gradually darkens into a deep chocolate color until it reaches the
tail, which is almost black with a tip entirely black. Hind legs and tail, viewed at a
distance of six feet, look very dark, almost pure black. The fore legs are black but not
so deep. Tips of ears, darkest.”

Two specimens from the Bangs: collection, one from Moosehead lake, Maine, the
other from Idaho county, Idaho, seem to answer closely the above description. The
light upper and forward portions of body are a grizzled grayish brown, the long hairs
black tipped. The basal half of hairs of anterior back are hair brown. I can discover
no color characters to separate the Idaho specimen from the one from Maine, nor do the
skulls indicate any reliable differences. The Maine skin (of an animal two-thirds grown)

* Typical canadensis must be restricted to the Alleghenian form.

NORTH AMERICAN BEAVERS, OTTERS AND FISHERS. 435

has white patches on lower fore leg, breast and vent, and an immature specimen of paci-
jica has white spots on throat, arm-pits and vent. The four adult specimens examined
are not thus pied. Dr. Coues, in his Fur-bearing Animals, says that the fisher is an
exception to the marten, mink and weasel in not having these patches. They may dis-
appear with age in the fisher, but they do not in the other species.

Anatomicel Characters.—Size, smaller than subspecies pacifica. Skull small ; nasals
relatively short, less elongate at basal apex. Posterior upper molar relatively small, its
inner lobe not greatly developed longitudinally so as to only slightly exceed the breadth
of outer lobe ; neck of crown of same tooth but slightly constricted.

Measurements —Of Dr. Raub’s Pennsylvania specimen, old ad. 3, /. .c.: Total
length, from end of nose to end of tail hairs, 965 mm.; tail vertebre, 318 mm.; hind
foot, 115 mm.; ear from crown, 27 mm. A mounted specimen, No. 507, Academy
Natural Sciences, adult %, from “ Pennsylvania,” has a total length of 1000 mm., with
tail (minus brush), 390 mm., and hind foot, 112 mm., taken from the dry mount. The
Idaho specimen, No. 6964, young adult 2, coll. of E. A. and O. Bangs, is 978 mm. long,
with tail, 369 mm., and hind foot, 117 mm. Skull of No. 7437, yg. ad. 3, Greenville,
Me., total length, 117 mm.; zygomatic width, 63 mm.; mastoid width, 54 mm.; mesial
nasal length, 22 mm.

Remarks.—The characters of the Pennsylvania fishers above enumerated, so far as
they are based on reliable measurements and color diagnoses, may be considered as repre-
senting typical canadensis, based on Pennant’s original notice of the animal. Whether
a series of Alleghenian fishers will show the Hudsonian animal to be separable is an
interesting question probably to be decided in the affirmative. The Idaho and Maine
specimens examined, though not contrasted by me with Dr. Raub’s specimen, must be
very close to it. No skulls of Pennsylvania fishers have been examined, but the close
resemblance of the Idaho skull to those from Maine, as indeed to pacijica also, strongly
indicates that no cranial differences exist between the east American fishers of the north
and south. The “saturated” color characters of pacifica are alone sufficient to distin-
guish it from all fishers found east of the Cascades.

Specimens Examined—Pennsylvania, 1 mounted specimen (fide Dr. Raub, 1
mounted specimen) ; Maine, Mooseland lake, 1 skin with skull; Greenville, 2 skulls;
Lincoln, 1 skull; Idaho, Idaho county, 1 skin with skull. Other specimens from east-
ern North America, 1 mounted, 2 old ad. skulls.

Pactric FisHer. JJustela canadensis pacifica, subsp. nov.

Type Locality.—Lake Kichelos, Kittitass county, Washington ; altitude about 8000

436 CONTRIBUTIONS TO A REVISION OF THE

feet. Type, No. 1074, old ad. 9, in the collection of 8. N. Rhoads; collected in the fall
or winter of 1892-93, by Allan Rupert.*

Geographic Distribution.—Pacific slope of America, from Alaska to California.

Color.—Above, from between eyes to middle back, grizzled, grayish ochraceous
heayily lined with black, becoming hazel black on hind back and dark black on rump,
thighs and tail. Whole head, behind eyes cloye brown basally, strongly grizzled with
dirty white. Snout to eyes blackish seal brown. Chin, throat, breast and belly between
dark chestnut and hazel, obscured with black. Legs and feet black, the fore legs show-
ing the vandyke brown bases of hairs. Basal half of hairs of anterior back are Prout’s
brown as contrasted with the hair brown of canadensis.

Anatomical Characters.—Size, large, skull very large, with relatively long nasals.
Posterior upper molar large, with spreading inner lobe much wider longitudinally than
outer section of same tooth; the crown suddenly constricted at the middle.

Measurements.—Of type from relaxed skin: Total length, 1090 mm.; tail, 350
mm. without brush ; hind foot not determinable, as the bones are missing. Measure-
ments of a specimen two-thirds grown, No. 295, coll. 8. N. Rhoads, from near Tacoma,
Wash.: Total length (relaxed skin), 970 mm.; tail, 400 mm.; hind foot, 112 mm.;
ear from crown, 21 mm. Skull of type: Total length from hinder end of sagittal crest
to front end of premaxille, 125 mm.;. zygomatic expansion, 73 mm.; mastoid expansion,
54 mm.; interorbital constriction, 28.5 mm.; postorbital constriction, 20 mm.; mesial
length of nasals, 27 mm.

Remarks.—The dimensions of the type skull, when we consider it was from a
female, show that the fishers of the Cascade mountains attain a much greater size than
those of the Appalachian chain. Young adult skulls of the same age from western
Washington and Maine show the same distinctions. The younger specimen from Tacoma,
while approaching nearer to Idaho and Maine specimens in grayer color, is very much
darker than they, the difference in shade between the anterior and posterior dorsal areas
of the former being slight, while in the latter it is striking. The tawny suffusion so
deeply marked in the type of pacifica and which separates it at a glance from canadensis
is also noticeable in the Tacoma specimen.

Specimens Examined.—Washington, Lake Kichelos, 1 skin with skull, 2 skulls;
near Tacoma, 1 skin, 1 skull ; British Columbia, Sumas, 1 skull.

* Mr. Rupert, whose business is hunting and trapping, first sent me the fresh skull of a very old Q fisher, which
was entered in my catalogue as No. 621. I wrote him immediately that I would like to have the pelt belonging thereto,

and in a later shipment the skin, which forms the type of pacifica, was sent on without Jabel. As it is also from a female

and a very old animal, I consider the skin and skull as belonging to the same individual.

4357

FISHERS.

NORTH AMERICAN BEAVERS, OTTERS AND

Skull Measurements

of North American

Otters (in millimeters)

Collection.

E. A. and O. Bangs

do.
Acad. N. Sci. Phila.
Smithsonian Inst.
E. A. and O. Bangs
do.
Acad. N. Sci. Phila.
8. N. Rhoads
do.
do.
E. A. and O. Bangs
i, ~
do.
do.

WagnerInst., Phila

S. N. Rhoads
do.
do.
Smithsonian Inst.
do.
do.
E. A. and O. Bangs
do.
do.

Catalo,
Number.

Sex.

g. ad.

old ad.
old ad.
old ad.
old ad. of
old ad. S)
old ad.

yg. ad.

yg. ad.
old ad. oj
yg. ad. 9

yg. ad.
ad. 2

ad.

old ad.
yg. ad. S
old ad. 2
old ad.
old ad.
old ad.
yg.?ad.o
old ad. 2
yg. ad. 2

{
|

Locality. Species. é 4 33 8. Ee EE . z si | Remarks.
meee |S") Be SB] aa] ee
ers Se 5 eae hae Sg eS Sad EB es Paleeteos ut :
| Nova Scotia, Annapolis | L. hudsoniea (‘‘La- | 113.5 | 72 | 68 | 27.7] 23 | 35 | 15 | Large, coast form.
cép.,’’ Desm.)
| Labrador, Okak do. 74.5] 67 | 25 19 | 35 | 13.5) Coast form.
| Labrador, Grand River do. 105 | 72.5) 65 | 20.8) 20 | 29 | 10.5) Inland form.
Alaska, Tanana River do. 102 | 72 | 63.5) 24 | 18 | 82 | 12.5) Inland form.
Maine, Bucksport do. 109 | 73.5] 66 | 95.5| 21.5' 37 | 14 | Coast form.
Massachusetts, Canton do. 112 76 | 69 | 26 | 22 | 38 | 15. | Intermediate.
Pennsylvania, Monroe Co. L. h. lataxina(F.Cuv.)] 100 | 69.5) 65 | 22.8) 20 | 31 | 13 | Inland interm., prob. 2.
do. do. 104.5] 68 | 61 | 21.5| 19 | 28.6) 12 | Probably ©.
New Jersey, Tuckerton do. 104 70 | 63.5) 24.5| 23 | 83:5) 11 |
New Jersey, Mickleton do. 107 70 | 63 | 23 12
North Carolina, Raleigh do. 104 71 | 62 | 22) 22 | 38 | 18.5
do. do. 103 65.5) 61 | 21.5) 21 | 30.5 11
Florida, Mieco L. h, vaga Bangs 108 | 71 | 71.2) 24 | 18.6 35 | 16 | Type (fide Bangs).
‘Florida, Roseland do. [101] | 70.3) 67 | 21.8) 17.8) 30 (fide Bangs.)
Florida, St. John’s Riv., Vol- do. 105 72 | 67 | 24 | 22 | 34 | 18.2
usia Co,
Florida, Tarpon Springs | do. 116 | 79 | 76.5) 27 | 20.5] 39.5) 20
Washington, L. Kichelos L. h. pacifica Rhoads | 115.5) 72.5) 69 | 25 | 20 | 36.5) 12 | Type. :
Washington, near Tacoma do. 110.5 | 77 | 70 | 29 | 215) 43 | 16
Alaska (coast ?) do. 115.5 | 74.5) 70.4) 27.3] 24 | 41 | 16 | Col. by Dr. T. T. Minor.
do, do. 119 | 83 | 76 | 34 | 25 | 49 | 14 do.
do. do. 110 78 | 73 | 27 | 18 | 41.5) 15 do.
Newfoundland, Bay St. George | L. degener Bangs 101 | 66 | 60 | 22 | 19.5) 32.5) 11.5) Type.
do. : do. [98] | '70 | 63 | 22.8) 19 4) 33.6 Topotype (fide Bangs).
do. do. 93 64 | 56 | 19 | 18.8) 25.8) 10 | Topotype.

CONTRIBUTIONS TO A REVISION OF THE

438

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NORTH AMERICAN BEAVERS, OTTERS AND FISHERS.

~
co

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Or

EXPLANATION OF PLATES.

Plates XXTI and XXII.

(Seale slightly less than two-thirds natural size. )

Figs. land 1. Castor canadensis pacificus Rhoads. Topotype ; No. 1865, col. of S. N. Rhoads; old adult ot’, from

Lake Kichelos, Kittitass county, Wash. Superior and inferior, vertical aspects of same skull.

Figs. 2and 2. Castor canadensis frondator Mearns. No. 32, col. of E. A. and O. Bangs ; young adult 9, from Red

Lodge, Mont. Superior and inferior, vertical aspects of same skull.

Figs. 3and 3. Castor canadensis Kuhl. No. 31, col. of E. A. and O. Bangs ; old adult (probably 3’), from New Bruns-

wick. Superior and inferior, vertical aspects of same skull.

Plate X_XTIT.

(Seale four-fifths natural size. )

Figs. land 2. Castor canadensis carolinensis Rhoads. Type; No. Z. 609, col. of State Museum of N. Carolina; old

adult <j, from Dan river near Danbury, Stokes county, N. Carolina. Superior and inferior, vertical

aspects of same skull.

Plate XXIV.

(Seale six-sevenths natural size. )

-Lutra hudsonica (‘‘ Lacépede,’’? Desmarest). No. 4188, col. of E. A. and O. Bangs ; old adult <j, from Canton,

Mass. Superior, vertical aspect of skull.

Lutra hudsonica (‘‘ Lacépeéde,”? Desmarest). No. 1201, col. of E. A. and O. Bangs, old adult 3, from West-

ford, Mass. Inferior aspect of skull.

Lutra hudsonica pacifica Rhoads. No. 8686, col. of Smithsonian Institution ; old adult, from (the coast of ?)
Alaska. Inferior aspect of skull.

Lutra hudsonica lataxina (F. Cuvier). No. 3537, col. of E. A. and O. Bangs ; old adult <j\, from Raleigh, N.
Carolina. Superior, vertical aspect of skull.

Lutra degener Bangs. Type; No. 6965, col. of E. A. and O. Bangs ; adult <j', from Bay St. George, Newfound-
land. Superior, vertical aspect of skull.

Plate XX V.

(Seale slightly less than five-sixths natural size.)

Lutra hudsonica pacifica Rhoads. No. 8687, col. of Smithsonian Institution ; old adult (probably ©’), from
(the coast of ?) Alaska. Superior, vertical aspect of skull.

Lutra hudsonica vaga Bangs. No. 1580, col. of S. N. Rhoads; old adult <j’, from Tarpon Springs, Fla.
Superior, vertical aspect of skull.

LIutra hudsonica pacifica Rhoads. No. 303, col. of S. N. Rhoads ; old adult 9, from Tacoma, Wash. Superior,

vertical aspect of skull.

Trans. Am. Phil. Soe., N.S XIX

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Trans. Am. Phil. Soe., N. S. XIX

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Trans. Am. Phil. Soe., N. S. XIX HPL UE

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Ss lbc-cpri

TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE IV.

Hital moment} of momentu

ads
1

Fig.E.

DIAGRAM FOR THE CURVES OF A SYSTEM OF EQUAL STARS, UNDER THE INFLUENCE OF TIDAL FRICTION.

Lower Curve illustrates increase of Eccentricity as the Stars separate.

TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE V.

H, 1408

H. 20°02

Drawings of double nebule according to Sir John Herschel

TRANS. AM. PHILOS. SOC., N.S XIX. PLATE VI.

GLOSSOPHAGA SORICINA.

—

sei a 2 ras rig

TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE VII.

GLOSSOPHAGA TRUEI.

ss
wail

ae
E

ciese

TRANS. AM. PHILOS. SOC., N.S. XIX. PLATE VIII.

MONOPHYLLUS REDMANI.

TRANS. AM. PHILOS. SOC., N.S. XIX. PLATE IX.

BRACHYPHYLLA CAVERNARUM.

TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE X.

BRACHYPHYLLA CAVERNARUM.

TRANS. AM. PHILOS. SOG., N. S. XIX. PLATE Xl.

LEPTONYCTERIS NIVALIS.

7 = ze

= See ae od _—
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Per Sh. ee

TRANS. AM. PHILOS. SOC., N.S. XIX

PLATE XIil.

WERE a
a

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Shoe

ASSES

53 54
CHGERNYCTERIS MEXICANA

PLATE XIll.

TRANS. AM. PHILOS. SOC., N. S. XIX.

LONCHOGLOSSA CAUDIFERA.

TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE XIV.

Lilia
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ANURA WIEDII.

TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE XV.

Wed Hy

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PHYEEONYCPERIS SEZBCORNIT

TRANS. AM. PHILOS. SOC., N. S. XIX.

PLATE XVI.

ECTOPHYLLA ALBA-CEPHALOTES PERONI.

patric OncDhil Soa. WS aa Viol ttt Lie XVI.

a inth.v Werner & Winter, Frankfort°M

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TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE Xx\l.

RHOADS—-NORTH AMERICAN BEAVERS.

TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE XxXIil-

RHOADS-NORTH AMERICAN BEAVERS.

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TRANS, AM. PHILOS. SOC., N. S. XIX. PLATE XxXiill.

RHOADS—NORTH AMERICAN BEAVERS.

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Raat
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TRANS. AM. PHILOS. SOC., N. S. XIX. PLATE XXIV

RHOADS—-NORTH AMERICAN OTTERS.

TRANS. AM, PHILOS. SOC., N. S. XIX. PLATE XXvV.

RHOADS—-NORTH AMERICAN OTTERS.

NOTICE,

ee Ree eee

A few complete sets can be obtained of the Transactions, New Series, Vols.
JI—XIX. Price, ninety-five dollars.

Address, THE LIBRARIAN.

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